Thelentil Botanyproductionanduses 140718234710 Phpapp01 PDF

The Lentil
Botany, Production and Uses
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The Lentil
Botany, Production and Uses
Edited by
William Erskine,1 Fred J. Muehlbauer,2 Ashutosh Sarker3

and Balram Sharma4
1Centre
for Legumes in Mediterranean Agriculture (CLIMA), The University of

Western Australia, Australia
2USDA-ARS, 303 Johnson Hall, Washington State University, USA
3International Center for Agricultural Research in the Dry Areas (ICARDA),
Aleppo, Syria
4Division of Genetics, Indian Agricultural Research Institute, New Delhi, India
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Library of Congress Cataloging-in-Publication Data
The lentil : botany, production and uses / William Erskine . . . [et al.], editors.
p. cm.
Includes bibliographical references and index.
ISBN 978-1-84593-487-3 (alk. paper)
1. Lentils. 2. LentilsResearch. I. Erskine, William. II. Title.
SB351.L55L46 2009
635.658dc22
2008045710
ISBN-13:
978 1 84593 487 3
Typeset by AMA Dataset Ltd, Preston, UK.

Printed and bound in the UK by the MPG Books Group.
The paper used for the text pages in this book is FSC certied.
The FSC (Forest Stewardship Council) is an international network
to promote responsible management of the worlds forests.
Contents
Contributors
Foreword
vii
xi
1. Introduction
W. Erskine, F.J. Muehlbauer, A. Sarker and B. Sharma
2. Global Production, Supply and Demand

W. Erskine
3. Origin, Phylogeny, Domestication and Spread

J.I. Cubero, M. Prez de la Vega and R. Fratini
13
4. Plant Morphology, Anatomy and Growth Habit

M.C. Saxena
34
5. Agroecology and Crop Adaptation

M. Materne and K.H.M. Siddique
47
6. Genetic Resources: Collection, Characterization, Conservation

and Documentation
B.J. Furman, C. Coyne, B. Redden, S.K. Sharma and M. Vishnyakova
7. Genetics of Economic Traits
B. Sharma
64
76
8. Genetic Enhancement for Yield and Yield Stability

A. Sarker, A. Aydogan, S. Chandra, M. Kharrat and S. Sabaghpour
102
9. Breeding for Short Season Environments

M. Matiur Rahman, A. Sarker, S. Kumar, A. Ali, N.K. Yadav and M. Lutfor Rahman
121
10. Improvement in Developed Countries

F.J. Muehlbauer, M. Mihov, A. Vandenberg, A. Tullu and M. Materne
137
11.
155
Advances in Molecular Research

R. Ford, B. Mustafa, M. Baum and P.N. Rajesh
vi
12. Breeding and Management to Minimize the Effects of Drought and
Improve Water Use Efciency
R. Shrestha, K.H.M. Siddique, D.W. Turner and N.C. Turner
Contents
172
13. Soil Nutrient Management

S.S. Yadav, D.L. McNeil, M. Andrews, C. Chen, J. Brand, G. Singh,
B.G. Shivakumar and B. Gangaiah
194
14. Cropping Systems and Production Agronomy

M. Ali, K.K. Singh, S.C. Pramanik and M. Omar Ali
213
15. Biological Nitrogen Fixation and Soil Health Improvement

M.A. Quinn
229
16. Mechanization
J. Diekmann and Y. Al-Saleh
248
17. Diseases and their Management

W. Chen, A.K. Basandrai, D. Basandrai, S. Banniza, B. Bayaa, L. Buchwaldt,
J. Davidson, R. Larsen, D. Rubiales and P.W.J. Taylor
262
18. Insect Pests and their Management

R. Ujagir and O.M. Byrne
282
19. Virus Diseases and their Control

S.G. Kumari, R. Larsen, K.M. Makkouk and M. Bashir
306
20. Weed Management

J.P. Yenish, J. Brand, M. Pala and A. Haddad
326
21. Parasitic Weeds

D. Rubiales, M. Fernndez-Aparicio and A. Haddad
343
22. Seed Quality and Alternative Seed Delivery Systems

Z. Bishaw, M. Makkawi and A. Aziz Niane
350
23. Nutritional and Health-benecial Quality

M.A. Grusak
368
24. Postharvest Processing and Value Addition

Albert Vandenberg
391
25. Food Preparation and Use

R.S. Raghuvanshi and D.P. Singh
408
26. The Impact of Improvement Research: the Case of Bangladesh and Ethiopia
A.A. Aw-Hassan, S. Regassa, Q.M. Shaqul Islam and A. Sarker
425
Index
447
The colour plate section can be found following p.224
Contributors
Ali, Asghar, Pulse Programme, National Agricultural Research Centre, Park Road, Islamabad,
Pakistan, asgharapk@yahoo.com
Ali, Masood, Indian Institute of Pulses Research, Kanpur 208024, Uttar Pradesh, India, masoodali53@
rediffmail.com
Ali, Mohamed Omar, Pulses Research Centre, Bangladesh Agriculture Research Institute, Ishurdi,
Pabna, Bangladesh, aliomarprc@yahoo.com
Al-Saleh, Yahya, University of Aleppo, Agricultural Faculty, Rural Engineering Department, Aleppo,
Syria, s-yahya@scs-net.org
Andrews, Mitchell, School of Sciences, University of Sunderland, Sunderland SR1 3SD, UK, mitchell.andrews@sunderland.ac.uk
Aw-Hassan, Aden A., International Center for Agricultural Research in the Dry Areas (ICARDA),
PO Box 5466, Aleppo, Syria, a.aw-hassan@cgiar.org
Aydogan, A., Central Research Institute for Field Crops (CRIFC), Ankara, Turkey, akadir602000@
yahoo.com
Banniza, Sabine, Crop Development Centre, University of Saskatchewan, 51 Campus Drive,
Saskatoon, SK S7N 5A8, Canada, sabine.banniza@usask.ca
Basandrai, Ashwani K., CSK Himachal Pradesh Agricultural University, Hill Agricultural Research
and Extension Centre, Dhaulakuan, Dist. Sirmour-173001, India, bunchy@rediffmail.com
Basandrai, Daisy, CSK Himachal Pradesh Agricultural University, Hill Agricultural Research and
Extension Centre, Dhaulakuan, Dist. Sirmour-173001, India, bunchy@rediffmail.com
Bashir, Muhammad, Crop Diseases Research Institute, National Agricultural Research Centre
(NARC), Park Road, Islamabad, Pakistan, fodder47@hotmail.com
Baum, Michael, International Center for Agricultural Research in the Dry Areas (ICARDA), PO Box
5466, Aleppo, Syria, m.baum@cgiar.org
Bayaa, Bassam, Faculty of Agriculture, University of Aleppo and International Center for Agricultural Research in the Dry Areas (ICARDA), PO Box 5466, Aleppo, Syria, b.bayaa@cgiar.org
Bishaw, Zewdie, Seed Unit, International Center for Agricultural Research in the Dry Areas
(ICARDA), PO Box 5466, Aleppo, Syria, z.bishaw@cgiar.org
Brand, Jason, Department of Primary Industries Victoria, Horsham, Private Bag 260, Horsham,
Victoria 3401, Australia, jason.brand@dpi.vic.gov.au
vii
viii
Contributors
Buchwaldt, Lone, Agriculture and Agri-Food Canada, 107 Science Place, Saskatoon, SK, S7N 0X2,
Canada, BuchwaldtL@agr.gc.ca
Byrne, Oonagh M., School of Plant Biology/Centre for Legumes in Mediterranean Agriculture
(M080), The University of Western Australia, 35 Stirling Highway, Crawley WA 6009, Australia,
oonagh.byrne@uwa.edu.au
Chandra, S., Indian Institute of Pulses Research (IIPR), Kanpur 208024, Uttar Pradesh, India, mali@
iipr.ernet.in
Chen, Chengci, Central Agricultural Research Center, Montana State University, Moccasin, MT
59462, USA, cchen@montana.edu
Chen, Weidong, United States Department of Agriculture (USDA) Agriculture Research Service
(ARS), Washington State University, Pullman, WA 99164, USA, w-chen@wsu.edu
Coyne, C., Western Regional Plant Introduction Station, 59 Johnson Hall, Pullman, WA, USA,
coynec@wsu.edu
Cubero, J.I., Departamento de Mejora Gentica Vegetal, Instituto de Agricultura Sostenible (CSIC),
Apartado 4084, 14080 Crdoba, Spain, ge1cusaj@uco.es
Davidson, Jenny, South Australian Research and Development Institute, GPO Box 397, Adelaide, SA
5001, Australia, davidson.jenny@saugov.sa.gov.au
Diekmann, Jrgen, International Center for Agricultural Research in the Dry Areas (ICARDA), PO
Box 5466, Aleppo, Syria, J.diekmann@cgiar.org
Erskine, William, Centre for Legumes in Mediterranean Agriculture (CLIMA), The University of
Western Australia, 35 Stirling Highway, Crawley, WA 6009, Australia, werskine@cyllene.uwa.edu.
au
Fernndez-Aparicio, M., Institute for Sustainable Agriculture, CSIC, Apdo 4084, E-14080 Crdoba,
Spain, monfapru@yahoo.com
Ford, Rebecca, BioMarka, Faculty of Land and Food Resources, The University of Melbourne, Victoria,
3010, Australia, rebeccaf@unimelb.edu.au
Fratini, R., Departamento de Biologa Molecular, Universidad de Len, Len, Spain, richard.fratini@
unileon.es
Furman, B.J., United States Department of Agriculture (USDA) Agriculture Research Service (ARS),
Subarctic Agricultural Research Unit, 533 East Fireweed Avenue, Palmer, AK 99645, USA, Bonnie.
Furman@ars.usda.gov
Gangaiah, B., Division of Agronomy, Indian Agricultural Research Institute, New Delhi, India,
bandla_gan@hotmail.com
Grusak, Michael A., United States Department of Agriculture (USDA) Agriculture Research Service
(ARS), Childrens Nutrition Research Center, Department of Pediatrics, Baylor College of Medicine,
1100 Bates Street, Houston, TX 77030, USA, mgrusak@bcm.tmc.edu
Haddad, Atef, International Center for Agricultural Research in the Dry Areas (ICARDA), PO Box
5466, Aleppo, Syria, a.haddad@cgiar.org
Islam, Q.M., Shaqul, Bangladesh Agricultural Research Institute, Joydebpur, Gazipur-1701,
Bangladesh, qazishaq06@yahoo.com
Kharrat, M., National Institute for Agronomic Research (INRAT), Tunis, Tunisia, moha.kharrat@
gmail.com
Kumar, Shiv, Indian Institute of Pulses Research, Kanpur 208024, India, Shiv_iipr@email.com
Kumari, Safaa G., International Center for Agricultural Research in the Dry Areas (ICARDA), PO
Box 5466, Aleppo, Syria, s.kumari@cgiar.org
Larsen, Richard, United States Department of Agriculture (USDA) Agriculture Research Service
(ARS), 24106 N. Bunn Road, Prosser, WA 99350, USA, richard.larsen@ars.usda.gov
Lutfor Rahman, M., Bangladesh Agricultural Research Institute, Gazipur 1701, Bangladesh, Lutfur@
bari.gov.bd
Makkawi, Mohamed, Seed Technology Unit, Dhaid Research Station, Ministry of Environment and
Water, PO Box 12503, Dhaid, Sharjah, United Arab Emirates, mamakk_2000@yahoo.com
Contributors
ix
Makkouk, Khaled M., International Center for Agricultural Research in the Dry Areas (ICARDA),
PO Box 5466, Aleppo, Syria, k.makkouk@cgiar.org
Materne, M., Grains Innovation Park, Department of Primary Industries, Private Bag 260, Horsham,
Victoria 3401, Australia, michael.materne@dpi.vic.gov.au
Matiur Rahman, M., Bangladesh Agricultural Research Institute, Gazipur 1701, Bangladesh,
baridres@citechco.net
McNeil, David L., School of Agricultural Science, University of Tasmania, Hobart, Tasmania,
Australia, david.mcneil@utas.edu.au
Mihov, Miho, Dubroudja Agricultural Institute, General Toshevo, 9520, Bulgaria, mihov43@abv.bg
Muehlbauer, Fred J., United States Department of Agriculture (USDA) Agriculture Research Service
(ARS), 303 Johnson Hall, Washington State University, Pullman, WA 99164-6434, USA, muehlbau@
wsu.edu
Mustafa, Barkat, BioMarka, Faculty of Land and Food Resources, The University of Melbourne,
Victoria, 3010, Australia, b.mustafa@pgrad.unimelb.edu.au
Niane, Abdoul Aziz, Seed Unit, International Center for Agricultural Research in the Dry Areas
(ICARDA), PO Box 5466, Aleppo, Syria, a.niane@cgiar.org
Pala, Mustafa, International Center for Agricultural Research in the Dry Areas (ICARDA), PO Box
5466, Aleppo, Syria, m.pala@cgiar.org
Prez de la Vega, M., Departamento de Biologa Molecular, Universidad de Len, Len, Spain, m.
perez.delavega@unileon.es
Pramanik, S.C., Indian Institute of Pulses Research, Kanpur 208024, Uttar Pradesh, India,
parmanik_03@yahoo.co.in
Quinn, Mark A., Department of Crop and Soil Sciences, Washington State University, Pullman, WA
99164-6420, USA, quinnm@wsu.edu
Raghuvanshi, Rita S., Dean, College of Home Science, G.B. Pant University of Agriculture and
Technology, Pantnagar 263145, India, rita_raghuvanshi@yahoo.com
Rajesh, P.N., United States Department of Agriculture (USDA) Agriculture Research Service (ARS)
and Department of Crop and Soil Sciences, 303 Johnson Hall, Washington State University, Pullman,
WA 99164-6434, USA, pnrajesh@wsu.edu
Redden, B., Australian Temperate Field Crops Collection, Department of Primary Industries, Private
Bag 260, Horsham, Victoria 3401, Australia, Bob.Redden@dpi.vic.gov.au
Regassa, Senait, Oxfam America-Horn of Africa Regional Ofce, Addis Ababa, PO Box 25779 code
1000, Ethiopia, senaitr2@yahoo.com
Rubiales, Diego, Institute for Sustainable Agriculture, Consejo Superior de Investigaciones Cientcas (CSIC), Apdo 4084, E-14080 Crdoba, Spain, ge2ruozd@uco.es
Sabaghpour, S., Dryland Agricultural Research Institute, Kermenshah, Iran, sabaghpour@yahoo.com
Sarker, Ashutosh, International Center for Agricultural Research in the Dry Areas (ICARDA), PO
Box 5466, Aleppo, Syria, a.sarker@cgiar.org
Saxena, Mohan C., A-22/7 DLF City, Phase 1, Gurgaon, Haryana 122002, India, m.saxena@cgiar.
org
Sharma, Balram, Division of Genetics, Indian Agricultural Research Institute, New Delhi 110012,
India, drbrsh@yahoo.co.in
Sharma, S.K., National Bureau of Plant Genetic Resources, Pusa Campus, New Delhi 110012, India,
director@nbpgr.ernet.in
Shivakumar, B.G., Division of Agronomy, Indian Agricultural Research Institute, New Delhi, India,
bgskumar@yahoo.com
Shrestha, R., National Grain Legumes Research Program, Rampur, Nepal, renuka_shrestha@hotmail.
com
Siddique, K.H.M., Institute of Agriculture/Centre for Legumes in Mediterranean Agriculture, The
University of Western Australia, 35 Stirling Highway, Crawley, Western Australia 6009, Australia,
ksiddique@fnas.uwa.edu.au
Contributors
Singh, D.P., Dean, College of Post Graduate Studies, G.B. Pant University of Agriculture and Technology, Pantnagar 263145, India, dhanpalsingh@indiatimes.com
Singh, Guriqbal, Department of Plant Breeding, Genetics and Biotechnology, Punjab Agricultural
University, Ludhiana, Punjab, India, singhguriqbal@rediffmail.com
Singh, K.K., Indian Institute of Pulses Research, Kanpur 208024, Uttar Pradesh, India, singhkk28@
rediffmail.com
Taylor, Paul W.J., BioMarka/Center for Plant Health, School of Land and Environment, The University
of Melbourne, Victoria 3010, Australia, paulwjt@unimelb.edu.au
Tullu, Abebe, Crop Development Centre, University of Saskatchewan, Saskatoon, SK S7N 5A8,
Canada, a.tullu@usask.ca
Turner, David W., School of Plant Biology, Faculty of Natural and Agricultural Sciences, The
University of Western Australia, 35 Stirling Highway, Crawley, WA 6009, Australia, David.Turner@
uwa.edu.au
Turner, Neil C., Centre for Legumes in Mediterranean Agriculture, M080, The University of Western
Australia, 35 Stirling Highway, Crawley, WA 6009, Australia, ncturner@clima.uwa.edu.au
Ujagir, Ram, Department of Entomology, G.B. Pant University of Agriculture and Technology,
Pantnagar 263145, India, ruape@gbpuat.ernet.in
Vandenberg, Albert, Crop Development Centre, University of Saskatchewan, Saskatoon, SK S7N
5A8, Canada, vandenberg@usask.ca
Vishnyakova, M., Leguminous Crops Genetic Resources Department, N.I. Vavilov Institute of Plant
Industry (VIR), 4244, Bolshaya Morskaya Street, 190000, St Petersburg, Russia, m.vishnyakova@
vir.nw.ru
Yadav, N.K., National Grain Legumes Research Program, Rampur, Nepal, nkyadav56@rediffmail.
com
Yadav, S.S., National Agricultural Research Institute (NARI), Kana Aburu Haus PO Box 4415 Lae
411, Morobe Province, Papua New Guinea, shyamsinghyadav@yahoo.com
Yenish, Joseph P., Department of Crop and Soil Sciences, Washington State University, Pullman, WA
99164-6420, USA, yenish@wsu.edu
Foreword
Lentil, its ancient origins notwithstanding, is very much a crop for todays
world which must confront the problems of food security, poverty and
water scarcity, and find sustainable food systems in a changing climate.
Lentil can help address each of these concerns. It has highly digestible, proteinrich grain, fixes nitrogen through association with bacterial rhizobia, uses
water efficiently, and is a vital component of cereal-based systems in marginal environments. Once considered primarily a food for the poor, its virtues are now widely appreciated.
I clearly recall my appointment as the first lentil breeder at the International Center for Agricultural Research in the Dry Areas (ICARDA), and the
excitement of being a young researcher at a young research centre. I contributed to the excellent book Lentils, edited by Colin Webb and Geoffrey
Hawtin, published by CABI in 1981. Twenty-seven years later, it is time to
take stock of the progress made. During this period, the global area under
lentil cultivation has grown by over 70%. Production has increased by
nearly 160%, partly due to the increase in cultivated area, but more importantly because productivity has increased by two-thirds. Clearly, considerable
progress has been made in translating research results into higher yields at
farm level. However, with population growth fuelling an ever-increasing
demand for food, further yield improvements will be needed. It is my earnest
hope that this book, which provides a comprehensive review of current lentil
research, will act as a knowledge platform for further advances.
Mahmoud Solh
Director General, ICARDA
xi
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Introduction
W. Erskine,1 F.J. Muehlbauer,2 A. Sarker3 and B. Sharma4

1The
University of Western Australia, Crawley, Western Australia, Australia;

State University, Pullman, Washington, USA; 3International Center for
Agricultural Research in the Dry Areas (ICARDA), Aleppo, Syria; 4Indian Agricultural
Research Institute, New Delhi, India
2Washington
The lentil (Lens culinaris Medikus subsp. culinaris) is a lens-shaped grain

legume well known as a nutritious food. It grows as an annual bushy leguminous plant typically 2045 cm tall, which produces many small purseshaped pods containing one to two seeds each. The morphology of the crop
is detailed by Saxena (Chapter 4, this volume).
Lentil seed is a rich source of protein, minerals (K, P, Fe, Zn) and vitamins for human nutrition (Bhatty, 1988). Furthermore, because of its high
lysine and tryptophane content, its consumption with wheat or rice provides a balance in essential amino acids for human nutrition. Lentil straw is
also a valued animal feed (Erskine et al., 1990).
Lentils were among the earliest of humankinds plant domesticates
(einkorn and emmer wheats, barley, pea, flax and lentil) and are associated
with the start of the agricultural revolution in the Near East (see Cubero
et al., Chapter 3, this volume). The crop was part of the assemblage of neareastern grains that spread across the Old World. It is now produced across
the dry areas of the globe and, in the Old World, from Bangladesh in the
east to Morocco in the west, and from Russia in the north to Ethiopia in the
south. The adaptation of the crop is discussed by Materne and Siddique
(Chapter 5, this volume).
With these origins it can be no surprise that the crop is well referenced
in early literature. Esau sold his birthright for a mess of pottage made of lentils (Bible Genesis 25: 2934). Lentils are also listed in the Koran (Second
Surah; Al-Baqarah) as one of the products of the earth which the Jews asked
Moses to request from God, following the period in which manna and
quails were the only food available to them. A paste of cooked lentils was
found in Egypt in a 12th-Dynasty tomb at Thebes (240022 bc). The Romans
used lentils widely in soups and Apicius, who in the 1st century wrote the
oldest surviving book on cookery, included a recipe for lenticulata a wellspiced dish of lentil and mussels with honey and vinegar. Its ancient and
CAB International 2009. The Lentil: Botany, Production and Uses
(eds W. Erskine et al.)
W. Erskine et al.
rather more obscure uses include being used as packing material to transport a Middle Kingdom obelisk from Egypt to Rome in the reign of the
Emperor Caligula, and as the preferred feed for the Roman armys information network carrier pigeons (Nygaard and Hawtin, 1981).
Today orange split and dehulled lentil is an important dal eaten with
rice and wheat in South Asia. In the Middle East lentil soup and mujaddarah
(whole lentil with cracked wheat or rice) are popular. Uses of lentil are covered by Raghuvanshi and Singh (Chapter 25, this volume).
Currently lentil is primarily grown in the developing world with a
particular concentration in Asia where India and Turkey are the largest producing countries (see Erskine, Chapter 2, this volume). Global production
of lentil is at c.3.6 million t. In the New World, Canada is the leading producer followed by the USA and Mexico with pockets of production in
Argentina, Chile, Colombia, Ecuador and Peru. In Europe, Spain, France
and Russia are the major lentil producers. Australia has come to dominate
southern hemisphere production since the late 1990s. Internationally the
trade in small-seeded, red cotyledon lentil is dominated by Australia, Canada and Turkey, whereas the market in the large-seeded, green lentil is
held by Canada and the USA. Countries in the Indian subcontinent, West
Asia and North Africa are the major importers of red lentil. Southern Europe
and South America import large-seeded green lentils. Spain is a major
importer of the so-called Spanish brown lentil produced in the US Pacific
Northwest.
Lentil is an important food legume crop component of farming and
food systems of many countries globally. It plays an important role in
human, animal and soil health improvement occupying a unique position
in cereal-based cropping systems. Its ability in nitrogen fixation and carbon
sequestration improves soil nutrient status, which in turn provides sustainability in crop production systems. Lentil is tolerant of different soil types
and of low fertility and this has assured its place as a crop of marginal lands.
Globally lentil productivity was only 560 kg/ha during the period
19611963 and reached 950 kg/ha by 20042006. Despite the increase, these
current yields are still low by comparison with other crops because of the
limited yield potential of lentil landraces, which are also vulnerable to an
array of stresses. Yield limiting factors are lack of seedling vigour, slow leaf
area development, high rate of flower drop, low pod setting, poor dry matter, low harvest index, lack of lodging resistance, low or no response to
inputs, and exposure to various biotic and abiotic stresses. The major abiotic limiting factors to production are low moisture availability (see Shrestha
et al., Chapter 12, this volume) and high temperature stress in spring, and,
at high elevations, cold temperatures in winter. Mineral imbalances like
boron, and salinity and sodicity though localized do cause substantial yield
loss (see Yadav et al., Chapter 13, this volume). Among biotic stresses, lentil
rust, vascular wilt and Ascochyta blight diseases are the most important
fungal pathogens (see Chen et al., Chapter 17, this volume). Additional
constraints to production include agronomic problems of pod dehiscence
and lodging, and suboptimal crop management.
Introduction
Adequate variability for many of the crops genetic constraints exists

within the crop gene pool allowing manipulation through plant breeding
(see Furman et al., Chapter 6; Sharma, Chapter 7; Sarker et al., Chapter 8;
Rahman et al., Chapter 9; and Muehlbauer et al., Chapter 10, all in this volume). However, several other important traits, such as biomass yield, pod
shedding, nitrogen fixation, resistance to pea leaf weevil (Sitona sp.) and
aphids, and the parasitic weed broomrape (Orobanche sp.) are not currently
addressable by breeding because of insufficient genetic variation. Molecular
approaches may be applied to solve some of these intractable problems.
Looking forward, escalating costs to produce inorganic nitrogen fertilizer, reducing availability of water for agriculture, climate change, and an
increasingly nutrition-conscious consumer society collectively give a bright
future for a highly nutritious food produced by a nitrogen-fixing crop like
lentil adapted to the cereal-based farming systems in marginal lands.
References
Bhatty, R.S. (1988) Composition and quality of lentil (Lens culinaris Medik.): a review.
Canadian Institute of Food Science and Technology 21, 144160.
Erskine, W., Rihawe, S. and Capper, B.S. (1990) Variation in lentil straw quality.
Animal Feed Science and Technology 28, 6169.
Nygaard, D.F. and Hawtin, G.C. (1981) Production, trade and uses. In: Webb, C. and
Hawtin, G.C. (eds) Lentils. Commonwealth Agricultural Bureau, Slough, UK,
pp. 713.
Global Production, Supply and Demand

W. Erskine
The University of Western Australia, Crawley, Western Australia, Australia
2.1. Introduction
This chapter surveys global and national production, export and demand/
consumption of lentil. It provides an update on the topic written in 1981 by
Nygaard and Hawtin in Lentils (Nygaard and Hawtin, 1981). The chapter is
based primarily on FAOSTAT the statistical database of the United Nations
Food and Agriculture Organization (FAO, 2008), which provides country-/
global-level estimates of crop production, area harvested, yield, exports,
imports and consumption since 1961. So we can see the big picture. But
crucially it provides no hint of the situation at the sub-national, district and
importantly household level.
2.2. Production, Productivity and Area

Globally lentil ranks sixth in terms of production among the major pulses
after dry bean, pea, chickpea, faba bean and cowpea (Table 2.1). World lentil
production constituted 6% of total dry pulse production in 20032006.
World lentil production has risen steadily by four times (412%) from an
average of 917,000 t in 19611963, when data collection started, to 3,787,000 t
in 20042006 (Fig. 2.1). The growth is clearly in two phases: Phase 1 was
from 1961 to 1979 with a growth rate of b = 25,900 t/year (R2 = 0.85) and
Phase 2 since 1980 with a faster growth of b = 79,100 t/year (R2 = 0.84). The
production gains come from a combination of increases in both area sown
and productivity per hectare. The area harvested in the same time period
has risen steadily from 1.639 million ha in 19611963 to 3.982 million ha in
20042006 with b = 60,200 ha/year (R2 = 0.95) (Fig. 2.1).


Table 2.1.
2008).
Global production as dry grain of major pulses from 2003 to 2006 (Source: FAO,
Pulse
Production (1000 t)
Beans
Peas
Chickpeas
Faba beans
Cowpeasa
Lentils
Pigeon peasa
Other pulsesa
Total pulsesa
only available to 20032004.
4.5
Area harvested (million ha)
4.5
4.0
4.0
b = 60.2; R 2 = 0.95
3.5
3.5
3.0
3.0
2.5
2.5
b = 79.1; R = 0.84
2.0
2.0
1.5
1.5
1.0
1.0
0.5
0.5
b = 25.9; R = 0.85
06
03
20
00
20
20
97
94
19
91
19
88
19
85
19
82
19
79
19
76
19
73
19
70
19
67
19
19
64
19
61
19
Production (million t)
aData
18,791
11,167
7,979
4,452
3,890
3,563
3,185
6,334
59,361
Years
Fig. 2.1. Annual evolution of global lentil production (million t; ) and area harvested
(million ha; ) since 1961 (Source: data from FAO, 2008).
Global lentil productivity was only 560 kg/ha in 19611963, while it

reached 950 kg/ha by 20042006, increasing overall by a healthy rate of
8.6 kg/ha/year (R2 = 0.85) (Fig. 2.2).
Lentil production is concentrated in the developing world, whence 75%
of production emanated in 20022006. The big three lentil producers are
India, followed by Canada and then Turkey, which collectively accounted
for 68% of global production in 20022006 (Fig. 2.3 and Table 2.2).
Most lentils are grown in Asia, where production totalled 2,300,890 t in
20022006 accounting for 61% of lentil production globally (Table 2.2). India
is the biggest lentil producing country in the world, producing 972,100 t of
lentil from 1.492 million ha in 20022006. In Asia the top league of lentil
producers comprises in descending order India, Turkey, Nepal, Syria,
China, Bangladesh and Iran, all of which produce more than 100,000 t/year.
To this list Pakistan may be added as the only other substantial producer.
W. Erskine
1200
y = 8.6111x + 503.7
R 2 = 0.849
Mean yield (kg/ha)
1000
800
600
400
200
0
1961
1966
1971
1976
1981 1986
Year
1991
1996
2001
2006
Fig. 2.2. Annual global mean grain yields (kg/ha) since 1961 (Source: FAO, 2008).
Europe
1%
Other Asia
19%
Turkey
16%
Canada
26%
India
26%
Other North
America
6%
Africa
3%
Australasia
3%
South America
0%
Fig. 2.3. Distribution of global lentil production among India, Canada and Turkey
(the big three producers) and by continent (Source: FAO, 2008).
Other Asian countries listed in Table 2.2 regularly produce less than 5000 t/
year. Over the last two decades lentil production has dropped markedly in
Iraq, Jordan and Lebanon.
Yields in Asia averaged 817 kg/ha, which is significantly below the
world average of 936 kg/ha. So those areas where lentils are most important are those with the lowest yield. Within Asia, productivity is particularly
Table 2.2. Global, continental and national lentil production (1000 t), area harvested
(1000 ha) and mean yield (kg/ha) from 20022006 (Source: FAO, 2008).
Region
Country
Africa
Algeria
Egypt
Eritrea
Ethiopia
Kenya
Madagascar
Malawi
Morocco
Tunisia
Asia
Armenia
Azerbaijan
Bangladesh
China
India
Iran
Iraq
Israel
Jordan
Lebanon
Myanmar
Nepal
Pakistan
Palestine
Syria
Tajikistan
Turkey
Uzbekistan
Yemen
Europe
Bulgaria
Croatia
Cyprus
Czech Republic
France
Greece
Hungary
Iceland
Italy
Macedonia
Russia
Slovakia
Spain
Ukraine
Area (1000 ha)

146.74
1.15
1.03
0.50
89.56
0.37
1.13
1.59
49.60
1.80
2817.36
0.01
1.14
130.20
69.00
1491.50
225.54
2.25
0.10
0.88
0.80
3.64
186.03
38.65
0.84
143.90
0.45
439.90
0.90
9.32
55.15
1.99
0.09
0.01
0.00
10.32
1.47
0.45
1.76
0.10
8.20
0.83
29.83
0.10
Production
(1000 t)
94.56
0.54
1.82
0.03
64.00
0.57
0.73
1.54
24.45
0.88
2300.89
0.02
2.11
118.95
144.23
972.10
113.23
2.00
0.03
0.57
0.80
1.84
159.34
21.90
0.54
159.33
0.40
596.34
0.35
6.81
41.74
2.27
0.12
0.01
0.00
14.78
1.75
0.53
0.03
1.21
0.08
8.64
1.00
11.26
0.06
Productivity
(kg/ha)
644
468
1761
60
715
1520
647
973
493
490
817
2125
1856
914
2090
652
502
889
300
644
1000
504
857
567
643
1107
889
1356
389
731
757
1140
1280
565
333
1432
1188
1189
688
764
1053
1216
378
600
(Continued)
W. Erskine
Table 2.2. continued
Region
Country
Area (1000 ha)
Production
(1000 t)
Productivity
(kg/ha)
Canada
Mexico
USA
887.11
708.20
7.72
171.19
1227.30
985.35
7.36
234.60
1383
1391
953
1370
141.23
139.63
1.60
13.28
1.50
1.16
3.90
3.16
3.56
4060.86
126.95
124.00
2.95
9.08
2.00
0.89
1.10
2.08
3.01
3800.52
899
888
1844
684
1333
769
282
658
846
936
North and Central

America
Australia and
Oceania
Australia
New Zealand
South America
Argentina
Chile
Colombia
Ecuador
Peru
World
low in South Asia with mean yields in Bangladesh, India, Nepal and Pakistan averaging 914, 652, 857, and 567 kg/ha, respectively.
Africa accounts for only 2.4% of lentil production globally and African
production is characterized by a low mean yield of 644 kg/ha. The only
significant producers are Ethiopia with a mean harvested area of 90,000 ha
and Morocco with 50,000 ha. Over the last three decades lentil production
in both Algeria and Egypt, once substantial, has dwindled to insignificance.
Both have become major importers.
Europe accounts for just 1.1% of global lentil production with 42,000 t
produced annually in 20022006. In terms of area harvested the main producers in Europe, in descending order, are Spain, France and Russia.
Lentil production in North America constitutes 32.2% of the total global
picture. Canada is the dominant player producing an average of 985,000 t
with a high average yield of 1391 kg/ha. Canada is now the worlds second
largest lentil producer after India and this emergence into global lentil production has arisen since the 1980s.
In South America total lentil production averaged only 9080 t/year and
is now not significant globally because of decreases in production over the
last two decades in Argentina, Chile and Colombia, where lentil was formerly important.
Australasia has arisen to prominence in lentil production through the
widespread production of the crop in Australia, where the area harvested
was 140,000 ha and production 124,000 t in 20022006. This growth in lentil
production in Australia occurred in the late 1990s.
Looking nationally at the importance of lentils against other pulses, lentil dominates the pulse sector in Nepal and constituted 60% of total dry
pulse production in 20022006. Elsewhere the percentage of lentil against
total pulse production was in descending order: Syria 52%, Bangladesh 35%
(not accounting for grass pea), Turkey 34%, Jordan 20%, Canada 19% and
Iran 16% in 20022006. In India the worlds largest lentil producer lentil
production accounted for only 7% of the total pulse production of 13.4 million t in 20022006.
The seedcoat colour of lentils can be green, tan, grey, brown or black
and the colour is often accompanied by mottling and speckling patterns.
Cotyledons are yellow, red or uncommonly green. Lentil production comprises approximately 70% of red cotyledon small-seeded rounded lentils,
typically 24.5 mm in diameter called red lentils; 25% large-seeded (46 mm
in diameter) more flattened seeds with yellow cotyledons, commonly
known as green lentils; and 5% brown lentils and other types.
2.3. Exports
The annual global export of lentil was 1,140,805 t in 20032005. This comprised 31.7% of production globally. Canada dwarfed other lentil exporting
nations by annually shipping 441,925 t of lentil in 20012005 (Table 2.3). The
top five exporters collectively accounted for 81% of global exports and after
Canada, the other important exporters fall into two groups: (i) the major
developing-world producers India and Turkey; and (ii) middle-ranking
production countries in the developed world Australia and USA, where
the crop is grown primarily as an export commodity.
The export trade is composed of different lentil types: small red lentils
sold as either red splits; red de-hulled but un-split, called football; and
small-seeded entire grains (usually with red, and unusually yellow, cotyledons); and green lentils large-seeded more flattened seeds with yellow
cotyledons. Although reliable data are unavailable on the relative importance of these various sectors, approximately 60% of world trade is in red
lentils, 35% in the greens, and 5% the browns and others.
Table 2.3. Average annual exports (t) of lentil from top
exporting nations from 2001 to 2005 (Source: FAO, 2008).
Country
Exports (t)
Canada
India
Turkey
Australia
USA
Syria
Nepal
China
441,925
140,687
127,669
107,701
105,976
39,470
34,934
25,066
10
W. Erskine
2.4. Demand/Consumption
We have already seen that lentil is primarily grown in the developing world
and especially in Asia. The major lentil-growing nations in the developed
world (Canada, USA and Australia) grow lentil to export it mostly to the
developing world (Table 2.3). So the increasing demand for lentil consumption is one from the developing world spurred by growing numbers of
mouths to feed. There can be little doubt that the main driver for increased
demand and consumption of lentil is the increasing population in Asia.
Turning to the other side of the coin, as 31.7% of the crop is exported so
very importantly 68.3% is eaten locally. Lentil grain has a high level of
protein, and also contains dietary fibre, vitamin B1 and minerals (see Grusak, Chapter 23, this volume). Lentils are mostly mixed with the carbohydrate staples rice and wheat, which results in a complete protein diet (see
Chapter 23). So the crop is making a major contribution to diets, especially
in the Third World of Asia.
Turning to imports (Table 2.4), the situation is much more diversified
than for exports, for which the trade is concentrated among five key players, with the top ten importers spanning only 59% of global trade. Among
importing countries there are several types:
First, there are India and Turkey, both of which are both major producers
and trading nations. They import in order to complete trading contracts
Table 2.4. Imports (t) of lentil into the top-40 importing countries (including food aid)
from 2001 to 2005 (Source: FAO, 2008).
Rank
Country
Imports (t)
Rank
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
Bangladesh
Egypt
Sri Lanka
Algeria
Pakistan
Colombia
Spain
India
Turkey
Italy
Sudan
Mexico
Saudi Arabia
France
Peru
Germany
Morocco
Ecuador
Venezuela
USA
117,306
97,574
93,172
57,288
55,078
53,956
49,990
49,638
31,929
30,397
30,027
28,540
26,029
24,199
21,867
21,228
16,560
14,521
14,519
13,984
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
Country
Ethiopia
UK
Netherlands
Chile
Belgium
Greece
Brazil
Eritrea
Israel
Jordan
Canada
Lebanon
Malaysia
Argentina
Haiti
Kuwait
Iran
South Africa
Portugal
Panama
Imports (t)
13,927
13,793
13,060
12,851
12,377
11,290
10,837
9,322
9,044
7,793
7,516
6,679
6,233
6,041
4,768
4,694
4,586
4,389
4,085
4,052
11
Table 2.5. Extract from FAO lentil consumption data

(kg/capita/year) (Source: FAO, 2008).
Country
Canada
Sri Lanka
Nepal
Syria
Turkey
Eritrea
Lebanon
Algeria
Jordan
Colombia
Iran
Bangladesh
Consumption (kg/capita/year)
6.0
4.5
4.1
3.7
2.9
2.9
2.6
2.3
2.2
2.1
1.5
1.5
when on occasion national production is insufcient to meet export

demands.
Second, there are those countries where historically lentil was grown and
consumed and there remains a residual demand in the diet for lentil
grain. But in these countries production has dropped over the last few
decades. Such countries include Egypt, Algeria, Pakistan and Colombia.
A third but similar group of countries are those which have rapidly
growing populations, a high consumption demand but still maintain
lentil production. This group comprises Mexico and Bangladesh, which
both import from their much larger neighbours.
Fourth, there are European countries such as Spain, Italy and France
where there is a demand for vegetarian food like lentil and this demand
outstrips the local supply.
Finally, there are countries such as Sri Lanka and those of the Middle
East where the crop is not produced but lentil forms part of the diet.
Consumption data from FAO in Table 2.5 confirm the importance of lentil
in the diet in several very poor countries such as Bangladesh, Eritrea, Nepal
and Sri Lanka.
Macro-data give no picture of the consumption of the crop within countries by region and by social group. However, pulses are generally consumed by the poorer sector of the population who cannot afford meat
products as a result of their high price. In Bangladesh it is recognized that
protein deficiency and vitamin A deficiency are major nutritional problems,
with consequences to Bangladeshs health (Rosenberg, 2005). The consumption and use of lentil at the local level has been investigated within nutrition
surveys in Ethiopia (Yetneberk and Wondimu, 1994) and Syria (Mokbel, 1986;
Ghosh, 2004). Lentil is an important dietary item in several often poor
parts of the Third World, contributing to warding off malnutrition through
12
W. Erskine
a balanced diet. Clearly the old adage that lentils are poor mans meat still
remains firmly applicable today.
References
Food and Agriculture Organization (FAO) (2008) FAOSTAT Statistical Database of the
United Nations Food and Agriculture Organization (FAO), Rome. Available at:
http://faostat.fao.org/site/567 (accessed 18 June 2008).
Ghosh, S.A. (2004) Poverty, household food availability and nutritional well-being of
children in north west Syria. PhD thesis, University of Massachusetts, Amherst,
Massachusetts, USA.
Mokbel, M. (1986) Nutritional and dietary patterns in rural Syria: implications for
ICARDA mandate crops. Farming Systems Research Report No. 18. International
Center for Agricultural Research in the Dry Areas (ICARDA), Aleppo, Syria.
Nygaard, D.F. and Hawtin, G.C. (1981) Production, trade and uses. In: Webb, C. and
Hawtin, G.C. (eds) Lentils. Commonwealth Agricultural Bureau, Slough, UK,
pp. 713.
Rosenberg, I.H. (2005) Interdepartmental Committee on Nutrition for National Defense
Surveys in Asia and Africa. Journal of Nutrition 135, 12721275.
Yetneberk, S. and Wondimu, A. (1994) Utilization of cool season food legumes in
Ethiopia. In: Telaye, A., Bejiga, G., Saxena, M.C. and Solh, M.B. (eds) Cool-Season
Food Legumes of Ethiopia. International Center for Agricultural Research in the
Dry Areas (ICARDA), Aleppo, Syria, pp. 6073.
Origin, Phylogeny,
Domestication and Spread
J.I. Cubero,1 M. Prez de la Vega2 and R. Fratini2
1Instituto
de Agricultura Sostenible (CSIC), Crdoba, Spain; 2Universidad de Len,

Len, Spain
3.1. Historical Data

Lentil was one of the first domesticated plant species, its remains being
as old as those of einkorn, emmer, barley and pea (Harlan, 1992). It has been
cultivated for 10,000 years in the most difficult agricultural environments,
being perhaps second only to barley in this sense. The plant was given the
scientific name Lens culinaris in 1787 by Medikus, a German botanist and
physician (for references given in this section, see Cubero, 1981).
Since 3029 bp, at least, they were popular and appreciated in Egypt, being
offered as presents to the gods. According to Athenaeus, Alexandria is full of
lentil dishes. Around 2100 years ago lentils had the same price as wheat, but
later the price declined, as Martial states in one of his Epigrams: receive lentils of the Nile, a present for Pelusium; they are cheaper than spelt. Martial
and also Juvenal, another Roman writer, described a lentil dish eaten by the
poorest, in which lentils were cooked together with the pods.
The Greek philosopher Theophrastus (~2300 bp) mentions a white kind
that is sweeter. The ancient Greek enjoyed lentils very much, especially in
soups. Aristophanes (2400 bp) said: You who dare to insult lentil soup,
sweetest of delicacies. The Greeks also made bread out of lentil flour. In
Rome, Columella (~1950 bp) said that they were too valuable as human food
to be used as animal fodder, even when they were used to feed pigeons; he
placed them only after faba beans among pulses. Romans imported lentils
from Egypt, where Pliny described the growing of lentil plants from seed
and reported the varieties sown at the time; he described two varieties one
rounder and blacker, the other normal in shape. Pliny also mentioned the
medicinal properties of lentils and a variety of ways of boiling or otherwise
cooking lentil seeds for a number of remedies. At the same time, the naturalist
Dioscorides (~1950 bp) mentions lentils in his De Materia Medica, and
Apicius gives details of several lentil recipes.
13
14
J.I. Cubero et al.
In the Middle Ages, lentil was a minor crop. Al Awam, a writer from
southern Spain in the 12th century, mentions two varieties: one of them
large and white that, when soaked in water, does not produce a black tint;
a second being a wild species of very poor quality. The 15th-century Hortus Sanitatis, collecting information from the Dioscorides and other ancient
references, lists some medicinal properties of lentils. In Spain, GabrielAlonso de Herrera writes in 1502 the best ones are the biggest ones they
are large and white and do not produce a black tint in water. He also
mentions that lentils were grown in light and dry soils, in rather cool places
and generally in fallow lands: their culture is not expensive at all since it is
not necessary to plough to sow them. If they are grown it is because in
some years lentils were very well sold.
Tournefort, based on several authors of the 17th century, lists seven species, briefly described as: (i) vulgaris, reddish seed (perhaps var. variabilis
Bar.); (ii) vulgaris, yellowish seed (var. vulgaris Bar.?); (iii) vulgaris, blackish
seed; (iv) maculata, marbled (var. dupuyensis Bar.); (v) major (var. macrosperma Bar.); (vi) hungarica major, perennis (perhaps nummularis Bar.);
and (vii) monanthos (perhaps a type not known in Europe at the present
time). Tourneforts description and drawings are excellent. The only doubtful point is the qualification of perennis given to hungarica.
Miller, in 1741, lists only three species: (i) vulgaris, common lentils;
(ii) major, greater lentils; and (iii) monanthos, lentils with a single flower.
He states that:
there are several varieties of the first and second sorts, which differ from
each other in the colour of their flowers and fruits; but they are accidental
and will often arise from the same seeds these plants are very common
in the warm parts of Europe, and in the Archipelago, where they are food
of the poorer sort of people these plants are one of the least of the pulse
kind they will be grown upon a dry barren soil best.
As in the time of Columella, lentils were also used to feed pigeons.

A rural Cyclopaedia (sic) of the mid-19th century mentions that they
were introduced into England from France during the 15th century, and by
the middle of the 19th century Britain had four varieties that were briefly
described as big, small, red and yellow. Thus, it seems that the European
variety structure was largely the same from the 16th to the 19th century and
probably the same as in the Middle Ages.
If lentils have been maintained by farmers through the ages, it is most
likely because they grow in poor soils, rough climates and harsh conditions
for humans, animals and crops. In many cases, they may be the only source
of protein available to farmers.
3.2. The Genus Lens and the Vicieae

The word Lens was used by Tournefort to name his genus III of his classis X
(De herbis et suffruticihus). Lens had been already used by most of the
many Theatri, Prodomi, Horti and Historiae Plantarum during the
Origin, Phylogeny, Domestication and Spread
15
16th century, but Tournefort was the first to use this word to designate
a specific genus (historical references in Cubero, 1981). After him, Lens
was also considered a genus by Miller, and by Adanson (who separated
Tourneforts Vicia, Ervum and Lens), and by Moench who included his Lens
esculenta (= L. culinaris Medikus) and Lens monantha (= Vicia monanthos) in
his Lens genus.
Linnaeus included it into Cicer and later in Ervum. The Linnean Ervum
was split very soon into Ervum and Lens, and later into Ervilia as well. The
first to use Lens systematically in a modern concept was Godron (1843), who
established the differential characteristics of Vicia, Ervum and Ervilia;
but during the 19th century, most of the authors followed the Linnean
treatment; others preferred Godrons. Boissier included Ervilia and the
non-Lens Ervum species in Vicia, keeping the name Ervum for the actual Lens
(Ervum Boissier is thus a synonym of Lens Miller). Some others included
Lens in Vicia and even in Cicer and Lathyrus. The idea of merging Lens into
Vicia has been kept up to recent times as well as the use of Ervum Boissier
instead of Lens. All the other names, including Ervum sensu Linnaeus, were
abandoned by the end of the 19th century.
There was also confusion concerning the author to whom the name Lens
had to be credited. Even in recent times, well known floras have credited it
sometimes to Adanson, Moench and Tournefort and also to Grenier and
Godron. Finally, the Congress on Botanical Nomenclature decided in 1966
to conserve Miller as the author. Lens Miller is, thus, a nomen conservatum,
having priority over older names (Gunn, 1969).
All the confusion just described has a strong biological basis. The tribe
Vicieae is formed by four closely related genera and many intermediate forms
have been recognized. They are the sources of abundant synonyms as well
as of newly proposed genera. The four genera are Vicia L., Lathyrus L.,
Pisum L. and Lens Miller. Cicer L. was separated in a monotypic tribe,
Cicerae (Kupicha, 1977). Many other genera were included previously in
the tribe (for example, Ervum L., Ervilia L., Faba L., etc.) and the synonyms
are many.
A differential set of characteristics of the four Vicieae genera (Vicia, Lens,
Pisum and Lathyrus) was given by Cubero (1981). Most of the characteristics
overlap to a variable degree. Species of doubtful systematic position, sometimes raised to generic status, are frequent. For most of the morphological
characters there is a continuum from one genus to the others, especially
for Vicia and Lens. In fact, Bueno (1976) studied the genus Vicia including a
macrosperma accession of L. culinaris as a reference. She used a total of 59
characters (42 quantitative and 17 qualitative); the distance of Vicia faba to
the closest species was much greater than that of lentils to many Vicia species, thus, challenging the independence of Lens if the old genus Faba was
included in Vicia.
The debate concerning a genus Lens separated from Vicia is old.
Linnaeus observed: [Lens] only differs from Vicia by the stigma. Boissier
observed that Lens seems to be a Vicia with some Lathyrus characters, in
particular those referring to the style. Lens seems to be one of the extremes
16
J.I. Cubero et al.
of the vicieoid continuum as intermediate forms are found both in Vicia

and in Lens, for example in Vicia section lenticula, Vicia sativa platysperma
and Vicia lunata. Perhaps the best example is Lens montbretii, whose taxonomic position was doubtful (Cubero, 1981), moved to Vicia montbretii
although considered as a lentoid Vicia because of its calyx, style and
flattened seeds (Ladizinsky and Sakar, 1982), although molecular data
clearly distinguish Lens from V. montbretii (Mayer and Bagga, 2002). Vavilov
(1949/50) showed that more or less flattened seeds can be found in all four
genera; he mentioned this character as one good example of his law of
homologous series of variation. The morphological continuum can be also
observed in a parallel response under domestication in lentil (L. culinaris),
common vetch (V. sativa) and grass pea (Lathyrus sativus), as shown by
Erskine et al. (1994).
The confusion arises from the fact that the genera of Vicieae are
members of a young group in active evolution radiating from a single
origin, hence showing a mixture of characters. We denominate a genus a
cluster of forms sharing a greater proportion of characters than other forms,
but the presence of intermediate species among genera as well as of intermediate species within a genus is unavoidable. We try to define our taxa in
the most accurate way, but our words will never contain all the natural
variation.
3.3. Taxonomy of Lens

Morphological and molecular studies
The morphological characteristics of the Lens species as well as synonyms
are given by Cubero (1981). Table 3.1 summarizes the main differential
characters among the species included in Lens. Intermediate forms are not
infrequent, as the different Lens species are also radiating from a common
ancestor.
Thus, it is not surprising that Boissier described his Ervum cyaneum
as a mixture of ervoides, nigricans and orientalis characters. It is easy to find
herbarium specimens successively determined as nigricans and orientalis, or
peduncles in orientalis without and in ervoides with awns, orientalis with
large pods, strongly pubescent forms of culinaris and orientalis from Syria,
Palestine and Turkey, as well as pubescent culinaris from India, and ervoides
with long and large leaflets. Some authors have suggested that nigricans
could have been cultivated in some areas of Turkey, but Ladizinsky (1979)
thought instead that these forms could be culinaris with nigricans stipules.
He also points out that the populations of nigricans reported from orientalis
areas could be orientalis forms with nigricans stipules.
The taxonomy was always confused with a lingering doubt about the
subspecific or specific status of Lens orientalis, and the phylogenetic position
of Lens nigricans within the genus. As L. orientalis is clearly the wild ancestor
of the cultigen (see below), its status has to be considered as subspecific
Culinaris group differential characters (Source: Cubero, 1981; Ladizinsky, 1993, 1997; Ferguson et al., 2000).
Character
Toothed stipules
Leaflets per leaf
Rachis length (mm)
Aristate peduncle or awn
Calyx teeth/corolla ratio
Peduncle/rachis ratio
Pod pubescent
Seed diameter (mm)
ervoides
nigricans
lamottei
odemensis
tomentosus
orientalis
culinaris
No
46
515
Rarely
<1
>1
Yes
23
Yes
68
825
Yes
1
>1
No
2.53.5
Slightly
610
710
Yes
>1
>1
No
2.53.5
Slightly
68
820
Yes
=
>1
No
2.53.5
No
612
720
Yes
>1
>1
No
2.53.5
No
68
525
Yes
<1
>1
Rarely
2.53.5
No
1016
2050
Yes
1
1
Rarely
49
Table 3.1.
17
18
J.I. Cubero et al.
(i.e. L. culinaris ssp. orientalis; Harlan and De Wet, 1971). The classical structure of the genus included L. culinaris, L. orientalis, L. nigricans and Lens
ervoides (Ladizinsky, 1979). In 1984, accessions of nigricans were reclassified
as odemensis and a new taxonomy of the genus was proposed, Lens culinaris
with cultigen subspecies culinaris and wild ssp. orientalis and odemensis, and
Lens nigricans with two subspecies, nigricans and ervoides (Ladizinsky et al.,
1984). However, Ladizinsky later recognized the following taxa: L. culinaris,
with subspecies culinaris and orientalis, Lens odemensis, L. ervoides and L. nigricans (Ladizinsky, 1993). Two new species have been recently added to the
genus, Lens tomentosus (ex L. orientalis; Ladizinsky, 1997) and Lens lamottei
(ex L. nigricans; Van Oss et al., 1997).
Traditional taxonomy was based on morphology; however, morphological
similarities or dissimilarities do not necessarily reflect phylogenetic relationships, sometimes they are the consequence of evolutionary convergence.
For instance, Fratini et al. (2006) using pollen and pistil morphological characteristics, and pollen functioning, found that L. odemensis and L. nigricans
clustered closely to each other and both to L. c. orientalis, next to this cluster
was L. c. culinaris microsperma; L. ervoides was loosely related to all these
taxa, but L. c. culinaris macrosperma was clearly separated from all other
Lens taxa including the microspermas (Table 3.2). Studies are very much
dependent on both the sets of accessions and the characteristics evaluated;
even with the same Lens accessions, different clusters have been obtained
depending on whether quantitative or qualitative characters were scored
(Ahmad et al., 1997).
Ferguson et al. (2000), using morphological as well as molecular markers,
considered both odemensis and tomentosus as subspecies of L. culinaris. The
following classification of the genus Lens was proposed by these authors:
L. culinaris, with four subspecies, namely, culinaris, orientalis, tomentosus
and odemensis, L. ervoides, L. nigricans and L. lamottei. This nomenclature is
currently being used by the National Centre for Biotechnology Information
(NCBI).
Table 3.2 shows different studies from the year 2001 onwards which
have genetically grouped Lens species according to several criteria. Besides
the study by Zimniak-Przybylska et al. (2001), whose phylogenetic nomenclature based on SDS-PAGE coincides with the study of Ferguson et al.
(2000), the rest of the studies using other molecular markers have all clustered tomentosus and odemensis in a group apart from that including culinaris
and orientalis. Recent phylogenetic analyses based upon internal transcribed
spacer (ITS) sequences, molecular markers and convicilin sequences strongly
suggest: (i) the divergent condition of nigricans from the remaining Lens
species; (ii) the close relationship between culinaris and orientalis supporting
their subspecific status; (iii) the relationships between the other taxa are
less clear cut; however (a) tomentosus is the sister taxon to the culinarisorientalis group, (b) lamottei is probably the closest relative to ervoides, and
(c) odemensis is generally joined to the culinarisorientalistomentosus
cluster (Mayer and Bagga, 2002; Sonnante et al., 2003; Durn and Prez de la
Vega, 2004).
Lens species clustered according to molecular markers, hybridization and morphological characteristics (year 2001 onwards).
Clusters
Methoda
Group 1
SDS-PAGE
1
1
1
1
1
1
culinaris
orientalis
tomentosus
odemensis
culinaris
orientalis
1
1
1
1
culinaris
orientalis
culinaris
orientalis
ITS
ITS + cpDNA
ITS
Group 2
Group 3
2 ervoides
2 lamottei
3 nigricans
ZimniakPrzybylska et al.
(2001)
2
2
2
2
2
tomentosus
odemensis
ervoides
lamottei
odemensis
3 nigricans
Mayer and Bagga

(2002)
2
2
2
2
tomentosus
odemensis
lamottei
ervoides
3 ervoides
3 nigricans
Group 4
4 nigricans
Group 5
Group 6
Table 3.2.
Reference
Mayer and Bagga

(2002)
Sonnante et al.
(2003)
19
(Continued)
20
Table 3.2.
continued
Clusters
Methoda
Group 1
Group 2
Group 3
Group 4
Group 5
Group 6
Reference
FISH highly
repeated DNA
RAPD + ISSR
1
1
1
1
1
1
culinaris
orientalis
culinaris
orientalis
culinaris
orientalis
2 tomentosus
3 odemensis
4 lamottei
5 ervoides
6 nigricans
Galasso (2003)
2 tomentosus
3 odemensis
4 lamottei
5 ervoides
6 nigricans
2 odemensis
2 nigricans
3 ervoides
1
1
1
1
culinaris(m)
orientalis
odemensis
nigricans
2 ervoides
3 culinaris(M)
Durn and Prez

de la Vega (2004)
Fratini et al.
(2004); Fratini and
Ruiz (2006)
Fratini et al.
(2006)
Crossability
Morphological
(flower traits)b
aAbbreviations
used: cpDNA, chloroplast DNA; FISH, fluorescent in situ hybridization; ISSR, inter-simple sequence repeats; ITS, internal transcribed spacer;
RAPD, random amplified polymorphic DNA.
b(m), microsperma; (M), macrosperma.
J.I. Cubero et al.
21
In addition, Galasso (2003), by means of in situ hybridization of highly

repeated DNA sequences, found: (i) a typical fluorescent in situ hybridization (FISH) karyotype for each species; (ii) very little variation within species (except in odemensis); (iii) a great similarity between ssp. culinaris and
ssp. orientalis; and (iv) favoured the separation of tomentosus and culinaris
into different species, as well as that of lamottei and nigricans, in both cases
despite their morphological similarity (van Oss et al., 1997). Galasso (2003)
also mentioned that two Syrian tomentosus accessions were more similar in
her study to culinaris than to the other tomentosus accessions, as Ferguson
et al. (2000) had also found between Syrian tomentosus and orientalis. It
would be interesting to find out if this similarity is a result of a misclassification, as Galasso suggests.
As a summary of this section, taxonomic analyses based on morphological and/or biochemical markers ranged from four species in 1979,
namely, L. culinaris, L. orientalis, L. nigricans and L. ervoides; to two species in
1984: L. culinaris (with subspecies culinaris (the cultigen), orientalis and
odemensis) and L. nigricans (with ssp. nigricans and ervoides); again to four species
in 1993, i.e. L. culinaris (ssp. culinaris and orientalis), L. odemensis, L. nigricans and
L. ervoides; to also four species in the year 2000, i.e. L. culinaris (ssp. culinaris,
orientalis, tomentosus and odemensis), L. nigricans, L. ervoides and L. lamottei;
and finally to six species after 2000, which are L. culinaris (ssp. culinaris and
orientalis), L. odemensis, L. tomentosus, L. nigricans, L. ervoides and L. lamottei.
The contradictory results of different studies are a consequence of the
evolutionary process itself. Lens species share many common structural and
biochemical features, and the distances obtained in different studies are a
function of the origin of the accessions involved in them, as well as of the
particular characters and molecular markers chosen. Subtle differences can
place accessions in distinct clusters. It is not a mistake of the experimental
method, rather it is a consequence of a radiating evolution. Fig. 3.1 shows
the geographical distribution of Lens species.
Defining biological species

Crosses among species and subspecies help to clarify the systematics of the
genus by establishing biological barriers to the access to a common gene
pool.
Ladizinsky (1979) made crosses between culinaris (three accessions representing extreme values for some characteristics as, for example, the seed),
orientalis (four) and nigricans (one). The F1 generation of the culinaris orientalis crosses grew normally in which the F2 generation segregated for growth
habit, flower colour, pod dehiscence and seedcoat colour. Meiosis was almost
normal with the presence of a quadrivalent in most of the cells examined
and, in very few cases, a trivalent and one univalent. This result supported
previous proposals considering orientalis as a subspecies of culinaris. The
similarity between them, and even the possible relationship of common
descent, was first pointed out by Barulina (1930) and later by Zohary (1972).
22
O
D
T
L
E
N
L. culinaris ssp. orientalis

L. odemensis
L. tormentosus
L. lamottei
L. ervoides
L. nigricans
0 250 500
J.I. Cubero et al.
Fig. 3.1. Geographical distribution of Lens species.
1000 km
23
According to Ladizinsky (1979), culinaris and orientalis share the same

karyotype, comprising two pairs of metacentrics (one of them with a
secondary constriction), two pairs of large submetacentrics and three
acrocentrics. Lens nigricans has a slightly different karyotype: four pairs of
metacentrics (one of them with a secondary constriction too) and three of
acrocentrics. However, other authors have found different karyotypes than
those described by Ladizinsky. Thus, Balyan et al. (2002) describe the karyotype of nigricans as constituted by one pair of metacentric, three of submetacentric and two acrocentric chromosomes. Different strains of culinaris
microsperma differ in the number of metacentrics, submetacentrics and acrocentrics, and orientalis strains in the position of the NOR locus (proximal or
distal to the centromere) and the number of 5S loci (one or two) (Fernndez
et al., 2005). Likewise, Galasso (2003) described intraspecific karyotype
variations in several Lens species. These results indicate that karyotypic
variation within Lens can be higher than expected and this is important for
the following section.
The first hybrids between culinaris and nigricans also developed
normally. However, later on (Ladizinsky et al., 1984), crosses with other
accessions of nigricans were unviable. Therefore, in 1984, the former
nigricans accession was reclassified as L. odemensis and the members of the
genus Lens were grouped in two biological species, as mentioned above
(Ladizinsky et al., 1984).
The two subspecies of L. culinaris are easily crossed with each other
(Ladizinsky, 1979; Fratini et al., 2004), although fertility of their intersubspecific hybrids depends on the chromosome arrangement of the wild parent
(Ladizinsky, 1979; Ladizinsky et al., 1984). Three crossability groups have
been identified in the wild ssp. orientalis: a common, a unique, and an intermediate. Crosses between members of the common and unique groups
yield aborted seeds which can be rescued by embryo culture; members of
the intermediate group are cross-compatible with the other two groups
(Ladizinsky, 1993; van Oss et al., 1997). Subspecies orientalis is readily
crossed with L. odemensis, the hybrids are vegetatively normal but partially
sterile due to meiotic irregularities resulting from three chromosome
rearrangements between the parental strains (Ladizinsky, 1993).
Lens tomentosus is morphologically closer to L. c. ssp. orientalis than
to any other Lens taxon, nevertheless, they are isolated one from another by
hybrid embryo breakdown, complete sterility and five chromosomal
rearrangements (van Oss et al., 1997), which supports the idea of a specific
status for tomentosus. Likewise, L. tomentosus is reproductively isolated
from L. lamottei and L. odemensis by hybrid-embryo abortion (van Oss et al.,
1997).
Linkage studies have revealed chromosomal rearrangements between
L. culinaris and L. odemensis (Ladizinsky, 1993), which could possibly explain
why certain accessions of odemensis are freely crossed with culinaris (Ladizinsky, 1979) and others need embryo rescue (Fratini and Ruiz, 2006). Lens
odemensis is cross incompatible with nigricans and ervoides due to hybridembryo abortion (Ladizinsky et al., 1984; Ladizinsky, 1993).
24
J.I. Cubero et al.
The morphological differences between L. nigricans and L. lamottei are

limited to stipule shape; however, the two species differ by four reciprocal
translocations and one paracentric inversion, resulting in the complete
sterility of their hybrids (Ladizinsky et al., 1984). Lens nigricans L. ervoides
interspecific hybrids are vegetatively normal but completely sterile
(Ladizinsky, 1993).
Fratini and Ruiz (2006) made extensive crosses between L. c. ssp. culinaris
and L. nigricans, L. ervoides and L. odemensis. Hybrids between the cultigens
and the other species were viable only through embryo rescue; the rates of
adult plants obtained were low: 9% with odemensis and 3% with nigricans and
ervoides. Previously, it had been already shown that crosses between culinaris
and nigricans or ervoides needed embryo rescue to recover interspecific hybrids
(Ladizinsky et al., 1984; Ladizinsky, 1993). Therefore, in view of the above, it
seems that odemensis belongs to the secondary gene pool and nigricans and
ervoides can be classified in the tertiary gene pool (Ladizinsky, 1993).
A summary about the phylogeny of the genus

Hybridization barriers support the idea of six differentiated species. Lens c.
orientalis obviously belongs to the primary gene pool, L. odemensis rather to
the second. However, the success in such crosses (needing embryo rescue
or not) depends on the particular accessions used in the study. Lens nigricans and L. ervoides belong to the tertiary gene pool, but can become part of
the secondary gene pool by means of embryo rescue. Further hybridization
studies are needed to establish whether L. tomentosus and L. lamottei belong
to the secondary or tertiary gene pool.
3.4. Taxonomy of the Cultivated Lentil

Alefeld cited in Cubero (1981) recognized eight subspecies including both
orientalis and nigricans (he used L. esculenta Moench.): (i) schniffspahni
(syn. orientalis); (ii) himalayensis (syn. nigricans); (iii) punctata (syn. culinaris);
(iv) hypochloris; (v) nigra (syn. culinaris); (vi) vulgaris; (vii) nummularia; and
(viii) abyssinica.
His subspecific names were later used by Barulina to name her
varieties. Between Alefeld and Barulina there were no detailed studies,
floras giving only simple varietal names (e.g. var. pilosisima Schur., var.
microsperma Baumg. and var. macrosperma Baumg.). Barulina raised the two
last names to the subspecific status, although molecular evidences indicate
that they are varieties within the subspecies L. c. culinaris. Thus, for instance,
macrosperma and microsperma accessions from Spain are more similar than
microspermas from different European countries (Durn and Prez de la
Vega, 2004).
The most detailed and complete study of the cultivated lentil was
made by Barulina (1930), a disciple and latterly wife of N.I. Vavilov.
25
She considered two subspecies, according to the size of the seed, the main
objective of human selection. Barulina also considered the geographical
distribution of clustered characters, defining regional groups or greges. The
main characters in her study to define subspecies were pods, seeds and
distinct differences in the length of flowers. Secondary characters included
size of leaflets, length of vegetation and height of plants. Important
characters with which to define the groups within the subspecies included
dehiscence, length of calyx teeth, and number of flowers per peduncle.
Varietal identification was realized choosing convenient, non-geographical
and sometimes utilitarian characters. Barulinas description of macrosperma
and microsperma is as follows.
(A) Lens culinaris ssp. macrosperma (Baumg. pro var.) Barulina. Large pods
(1520 7.510.5 mm), generally flat. Large flattened seeds ( 68 mm).
Cotyledons yellow or orange. Flowers large (78 mm long), white with
veins, rarely light blue. Peduncles with two to three flowers. Calyx teeth
long. Large leaflets (1527 410 mm), oval (length:width ratio = 33.5).
Plant height from 2575 cm. No geographical groups. 12 varieties. (B) ssp.
microsperma (Baumg. pro var) Barulina. Pods small or medium (615 3.57
mm), convex. Seed flattened-subglobose (diameter:thickness ratio = 1.53),
small or medium ( 36 mm), diverse in colour and pattern. Flowers small
(57 mm long) from white to violet. Peduncles bearing one to four flowers.
Leaflets small (815 25 mm), elongated, linear or lanceolate (length:width
ratio = 45). Height of plants 1535 cm. 46 varieties grouped in six greges:
europeae, asiaticae, intermediae, subspontanea, aethiopicae and pilosae (see Cubero,
1981 for details).
The geographical distribution of subspecies and greges can be seen in

Fig. 3.2, drawn according to Barulinas own map.
3.5. The Centre of Origin

The three main problems regarding the biological nature of any crop are:
(i) where and when the biological species originated; (ii) where and when
that species became a crop; and (iii) how it has evolved as a crop.
Archaeological data were given by Cubero (1981) and this is summarized in Fig. 3.3. The oldest remains were found in Hacilar (Turkey), Ramad
(Syria), Jarmo (Iraq), Jericho (Palestine), Beidha (Jordan) and Ali Kosh (Iran)
dated around 9000 bp, in aceramic Neolithic layers. Even older remains
were found in Mureybit (Syria), c.10,500 bp, but they were wild lentils.
The oldest Greek lentils were dated around 8000 bp, then Central Europe
(50007000 bp, from classical Neolithic to early Bronze; in both regions
there are some doubtful nigricans seeds). Late arrivals were those of India
(30004000 bp) and Western Europe (France, Germany: 30003500 bp). Egypt
also shows a later arrival (c.5000 bp) than in Greece and Central Europe but
conditions in the Nile Delta are not favourable for preserving agricultural
remains.
26
macrosperma
europeae
asiaticae
intermediae
microsperma
subspontanea
aethiopicae
pilosae
0 250 500
Distribution of L. culinaris greges according to Barulina.
J.I. Cubero et al.
Fig. 3.2.
1000 km
BP
9500
95008000
80006500
6500
0
250
500 km
Fig. 3.3. Archaeological data. Lines indicate, respectively, the main distribution area of the orientalis and the proposed domestication area
within it.
27
28
J.I. Cubero et al.
Concerning the place of origin of lentils as a crop, according to the

archaeological data, the distribution of wild species and the existence of the
remains of both wild and cultivated lentils in the same region, the Near
East is the obvious candidate. Ladizinsky (1999) found the cultivated lentil
to be monomorphic for several characters, but considerable polymorphism
was found to exist in the wild accessions of ssp. orientalis. However, three
accessions from eastern Turkey and northern Syria were shown to share the
above characters with the cultigen and therefore can be regarded to be
members of the genetic stock from which the cultivated lentil was domesticated. Zohary (1999) comes to the conclusion that lentil was domesticated
once or only a few times, this conclusion being based on founder effects
revealed by chromosome and DNA polymorphisms, as well as evidence
from domestication traits and species diversity. Lev-Yadun et al. (2000)
suggested a place close to or overlapping the area where einkorn and
emmer wheats were domesticated in the Fertile Crescent (Fig. 3.3).
Barulina (1930), instead, suggested the eastern border of south-west
Asia as a possible centre of origin of the cultivated lentil, as the region
between Afghanistan, India and Turkistan (i.e. the HimalayaHindu Kush
junction) showed the highest proportion of endemic varieties, a pure
Vavilovian criterion to define centres of origin. She noticed that the area of
distribution of wild lentils did not overlap much with that of the domesticated ones, but she maintained her idea considering that the eastern part of
the orientalis area of distribution reaches Turkmenia.
Today, the abundance in endemic forms of the cultigen as the most
important criterion to locate a centre of origin is taken as a secondary argument (Ethiopia is also rich in endemism and it is not a candidate for centre
of origin). De Candolles ideas were taken up again some time ago (Harlan,
1992). Thus, the overlap between the wild ancestor and the cultigen and
the archaeological remains connecting both are sine qua non conditions to
establish such a centre of origin.
Figures 3.1 and 3.2 show the distribution of wild and cultivated lentils
(Barulinas greges of L. culinaris are used for that purpose). Three groups are
restricted to very concrete areas: pilosae to the Indian subcontinent, aethiopicae to Ethiopia and Yemen, and subspontanea to the Afghan regions closest
to the Indian subcontinent. All three have very small and dark-coloured
seeds, violet flowers, few flowers per peduncle, calyx teeth much shorter
than the corolla, few leaflets per leaf, and dwarf plants. On this common
background, each one of these three greges shows a distinct character:
pilosae a strong pubescence; subspontanea very dehiscent pods, being
purple coloured before maturity; and aethiopicae pods with a characteristic
elongated apex. These particular characteristics are shown together with a
cluster of primitive characteristics closely related to orientalis.
The other three microsperma groups (europeae, asiaticae and intermediae)
and macrosperma are rather cosmopolitan and are clearly intermixed, even
when intermediae has a rather restricted area. Seeds are variable in size,
but in general, they are wider than 4 mm. They have more flowers per
peduncle, more leaflets per leaf, and the calyx teeth are equal to or larger
29
than the corolla. White flowers are common, seeds being diverse in colour.
Figures 3.1 and 3.2 suggest that subspontanea overlaps only with orientalis,
and aethiopicae with ervoides; pilosae does not overlap with any wild lentil.
All other microsperma as well as macrosperma lentils overlap to a greater or
lesser extent with all the known wild lentils. On the other hand, no lentils
have been found in the sites dating back to the seventh millennium bp in
Turkmenia.
The high degree of endemism that exists in the Afghanistan
IndianTurkmenian area is better explained, as in all other species, by an
intense genetic drift, typical of highly diversified environments, coupled
with artificial selection carried out by very diverse human populations,
with drastic genetic fixation and losses providing secondary centres of
diversity.
To sum up, the area from western Turkey to northern Iraq contains not
only all the wild lentils but also lentoid characters such as flattened pods
and seeds present in other species like V. montbretii (a bridge between Vicia
and Lens?) and V. lunata. The known data point to the southern Turkey
northern Syria region as the most likely place of lentil domestication. Some
populations of orientalis were unconsciously subjected to automatic selection here, leading to a new crop, Lens culinaris.
3.6. The Spreading of Lentil Culture

Figure 3.3 shows that lentils follow the spreading of the Neolithic agricultural techniques (Harlan, 1992), being first cultivated in the Fertile Crescent,
then moving to Greece, Central and Western Europe along the Danube, to
the Nile Delta, and eastwards to India. From Greece, lentils found their way
to Central Europe through the Danube. There are lentil archaeological
remains in the Iberian Peninsula since the early Neolithic, the oldest ones
corresponding to the eastern part of the Peninsula. In particular in the cave
de les Cendres, there are remains of several crops (Triticum monococcum,
Triticum dicoccum, Triticum aestivum, barley, pea, grass pea, lentil and faba
bean, i.e. a typical Near East crop complex) dated by 14C to 7540 140 bp.
According to the seed size, the archaeological lentil findings in the Iberian
Peninsula fall within the range of the microsperma sizes (Bux, 1997). Intermediae forms reached Sicily, and asiaticae forms reached Sardinia, Morocco
and Spain (Fig. 3.2), suggesting the arrival in these countries of lentil stocks
from Central Europe, or from the route of the isles.
The lentils probably reached the cradle of Indoeuropean people after
the Greek ancestors split, as suggested by De Candolle (1882) on linguistic
grounds: Greek for lentil is phakos, but lens in Latin, lechja in Illyrian, and
lenzsic in Lithuanian. It is likely that the ancient Greeks took the word from
the aboriginal Mediterranean populations they conquered. Figure 3.2 shows
that central Russia, then Siberia, was more likely reached from the
western coast of the Black Sea or from the Danube valley rather than from
Mesopotamia or Central Asia.
30
J.I. Cubero et al.
The arrival of lentils to the Nile Delta had to be much earlier than the
archaeological remains suggest due to its proximity, both geographical and
cultural, to the Fertile Crescent; but the Delta environment is not helpful in
preserving organic materials. Ethiopia was probably reached from the Arabian coast (at that time it was the Arabia Felix, more humid and fertile than
nowadays) rather than by the Nile. Lentils likely travelled in the Near East
complex (along with barley, wheat, chickpea, faba bean, etc.), established in
the Ethiopian highlands since primitive times, and have been evolving since
then in isolation, producing much endemism that allowed Vavilov to place
the centres of origin of crop plants that only arrived there by farming. Indeed,
aethiopicae exhibits very primitive characters, meaning that lentils probably
also arrived at a very primitive stage of domestication.
Lentils did not reach India before 4000 bp, probably carried by the Indoeuropean invasion (De Candolle, 1882) through Afghanistan, but again there is
the problem of availability of archaeological findings. Primeval introduction
was probably performed by very small samples of a common origin as the
variability found in the Indian subcontinent in the local landraces is very limited, in spite of being the largest lentil-growing region in the world; the asynchrony in flowering of the local pilosae ecotype, probably a consequence of a
long reproductive isolation period, is now being broken by plant breeding
methods in order to widen the genetic base available (Erskine et al., 1998).
The diversity in the centre of origin seems to be well maintained. By
using molecular markers, Ferguson et al. (1998) located areas of high diversity for the wild relatives: L. culinaris ssp. orientalis (south-west Turkey,
north-west and south Syria, and in Jordan), L. odemensis (south Syria), L.
ervoides (coastal border region between Syria and Turkey) and L. nigricans
(west Turkey).
3.7. Evolution of the Cultivated Forms

Lens orientalis is the wild form out of which the cultigen developed. Lens
culinaris forms show, in general, greater height, longer rachis and greater
number of leaflets per leaf, greater leaflet area, greater number of flowers
per peduncle, peduncle shorter or equal to the rachis, higher frequency of
white flowers, and also larger pods and seeds. All these characters are
related to the increase in yield. Seed size is the only character that can
indicate domestication in archaeological remains.
It was mentioned above that there are cultivated groups with some primitive characters; up to the Middle Ages there were blackish and not sweet
varieties, as well as others with normal and rounder seeds. Obviously, they
could be impurities coming from mixtures with some vetches (for example,
Vicia angustifolia or Vicia platysperma), as subsistence farmers did not worry very
much about botanical purity. But these forms could also be mixtures with wild
Lens species or any real primitive landrace.
Figure 3.2 shows a clear regional varietal facies according to Barulinas
study, performed when modern agriculture had not reached these regions.
31
Western lentils have the main characters of big seeds, a high number of
large leaflets, and calyx teeth longer than the corolla. A great discussion is
raised on the origin of some characteristics of the cultigen. Lens odemensis
could have provided the genetic raw material for bigger seeds and all the
correlated characters. Crosses between culinaris and odemensis (then classified as nigricans; Ladizinsky et al., 1984) resulted in a greater development
of tendrils while culinaris shows, although rarely, branched tendrils. Calyx
teeth are also larger in the western lentils, a character that can be correlated
with leaflet area; calyx teeth are shorter than corolla in orientalis but as
long as the corolla in odemensis. The opposite is true for the eastern lentils,
orientalis playing the leading role in crop evolution.
The original stock of pilosae (Indian endemism) could have been originated in south Turkeynorth Syria, where hairy orientalis and culinaris
forms are found. The short calyx of aethiopicae forms could have an ervoides
origin.
The origin of such facies can be attributed to the classical sources of
variation in crop evolution: mutation, migration, selection, genetic drift and
crosses with companion weeds (wild lentils). Figure 3.1 shows that orientalis
is the only one spreading eastwards, ervoides to the south (Ethiopia), and
both nigricans and ervoides to the west. Some L. odemensis accessions can be
readily crossed with the cultigen (Ladizinsky et al., 1984). Lens nigricans and
L. ervoides should be excluded because crosses with the cultigen result in
hybrid embryo abortion (Ladizinsky et al., 1984; Fratini and Ruiz, 2006), but
it cannot be ruled out that these crosses eventually (even if sporadically)
succeed in natural environments, nor that all strains of nigricans and ervoides
behave in a similar way. Thus, if the distribution of wild lentils was the
same as that at the Barulinas time, the regional facies can be explained.
To sum up, lentils were domesticated, in the Near East or, more accurately, in the foothills of the mountains of southern Turkey and northern
Syria. The raw materials were populations of orientalis, but primitive farmers could also have used some other species of the genus, whose similarity
has been shown in the present chapter, in mixed populations rather than in
pure strands. But orientalis and odemensis forms are the most likely candidates to have been companion weeds of the cultigen. However, molecular
marker analyses have indicated that the genetic variability within cultivated
lentils is relatively low (Sonnante et al., 2003; Durn and Prez de la Vega,
2004), which supports the idea that microsperma and macrosperma morphotypes are simple variants for quantitative traits resulting from disruptive
selection (Sonnante et al., 2003). It is difficult to establish how much the wild
relatives have contributed to the cultigen gene pool.
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species and their interspecic hybrids as measured by morphological characters.
Euphytica 94, 101111.
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Balyan, H.S., Houben, A. and Ahne, R. (2002) Karyotype analysis and physical
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Barulina, H. (1930) Lentils of the USSR and other countries. Bulletin of Applied Botany,
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Bueno, M.A. (1976) Taxonoma y Cariologa en el Gnero Vicia. Doctoral thesis,
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Ferguson, M.E., Ford-Lloyd, B.V., Robertson, L.D., Maxted, N. and Newbury, H.J.
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Plant Morphology, Anatomy and

Growth Habit
Mohan C. Saxena*
International Center for Agriculture Research in the Dry Areas (ICARDA),

Aleppo, Syria
*Present address: A-22/7 DLF City, Gurgaon, Haryana, India
4.1. Introduction
The lentil is a slender, softly pubescent, light green, annual herbaceous plant
(Fig. 4.1), generally between 20 and 30 cm tall, although there are cultivars as
short as 15 cm and as tall as 75 cm (Duke, 1981; Muehlbauer et al., 1985). The
plant is indeterminate and it exhibits considerable variation in its growth
habit, with single stem and erect to semi-erect and compact growth to much
branched low bushy forms, mainly depending on genotype although environmental conditions affect the expression of these characters (Saxena and
Hawtin, 1981).
Under optimum environmental conditions, as obtained in late-winter
and early-spring season sowings in the Mediterranean region of West Asia
and North Africa (WANA), lentil plants make fast vegetative and reproductive growth and reach maturity in 75100 days after sowing. In this respect,
lentil, of all cool-season food legume crops, compares well with barley (Hordeum vulgare), which is the fastest growing winter cereal and well adapted
to dry rain-fed conditions in the WANA region. However, in most lentilproducing countries, plant growth of the winter-sown crop is rather slow
in the early vegetative phase because of suboptimum ambient temperatures and it gains momentum only in spring when temperatures rise. As a
result, the winter-sown crop takes 120160 days to reach maturity (Saxena
and Hawtin, 1981), extending to 180 days in some environments of West
Asia. Moisture supply during the reproductive phase of crop growth
much affects the complete realization of the growth and yield potential of
lentil genotypes.
The lentil plant exhibits a wide range of morphological variations in
both its vegetative and its reproductive organs. That is why a major early
attempt to classify cultivated lentils into subspecies by Barulina (1930) was
mainly based on the characteristic morphological features of pods and
34

Plant Morphology, Anatomy and Growth Habit
35
Fig. 4.1. Lentil plant structure: (a) dry seed; (b) moisture-imbibed seed; (c) a newly emerged
seedling showing hypogeal germination; (d) a young seedling showing two bifoliate leaves;
(e) branch with leaves, owers and pods of microsperma lentil.
seeds, length of flower, size of leaflets, and plant height. Lentil cultivars
were grouped in two intergrading clusters of seed sizes: (i) small-seeded
lentils (microsperma) with relatively small and swollen pods (615 3.57.0
mm) and small seeds (36 mm diameter); and (ii) large-seeded lentils (macrosperma) with relatively large flattened pods (1520 7.510.5 mm) and
large seeds ranging in diameter between 6 and 9 mm (Fig. 4.2). It appears
that with domestication there was not only the development of pod indehiscence but also a gradual increase in seed size as testified by the fact that
macrosperma forms appear more recently in archaeological sequence only
in the first millennium bc (Zohary and Hopf, 2000). However these two
groups are freely intercrossable and a complete range of seed sizes, from
microsperma to macrosperma types, is present in the cultivated species (Muelbauer et al., 1985). During the last decades, through broadening of the
genetic base, these two groups of lentils have become indistinguishable,
and the grouping is no longer much used by lentil workers.
36
M.C. Saxena
Fig. 4.2. Variation in pods and seeds of lentil. Top row, macrosperma type lentil:
(a and b) pods; (c) side view and (d) front view of seed. Bottom row, microsperma
type lentil: (e and f) pods; (g) side view and (h) front view of seed.
4.2. Root System

The lentil plant has a slender taproot system with a mass of fibrous lateral
roots. Large genotypic variation has been reported in root growth in terms
of taproot length, number of lateral roots, total root length, root weight
(Sarker et al., 2005) and number of hairs per unit root surface area (Gahoonia et al., 2005). Genotypic differences have also been observed in the rate of
root growth (Gahoonia et al., 2005; Sarker et al., 2005). Because of the importance of root traits in the capture of moisture and inorganic nutrients from
soil, particularly on low fertility and water-holding-capacity soils, positive
association has been observed between rate and amount of root surface
development and the crop yield (Gahoonia et al., 2005; Sarker et al., 2005).
Sarker et al. (2005) attributed the drought tolerance of breeding line ILL
6002, derived from a cross between Canadian cultivar Liard (ILL 4349) and
Argentinean cultivar Precoz (ILL 4605), mainly to its root characteristics of
fast growth and large number of lateral roots.
Interesting variations in pattern of root growth in different ecotypes of
lentils adapted to different types of soils in major lentil production areas in
the Indian subcontinent have been reported by Nezamuddin (1970). The
patterns recognized on the basis of depth and lateral proliferation were: (i)
a much-branched, shallow root system going down only to a depth of about
15 cm; (ii) a slender deep taproot going down to 36 cm depth; and (iii) an
intermediate form (Fig. 4.3). The shallow, profusely branched roots were
found on light alluvial soils with the small-seeded lentil ecotypes with
highly branched shoots. The deep root system was found in the ecotypes
(a)
37
(b)
(c)
Depth (cm)
10
20
30
40
10
20
30
40
50
60
70
Width (cm)
Fig. 4.3. Root system in lentils: (a) shallow; (b) intermediate; and (c) deep (Source: adapted
from Nezamuddin, 1970).
grown on heavy black cotton soils in central India that tend to develop large
cracks on the surface and thus rapidly lose moisture from the top soil layer.
These ecotypes have sparsely branched shoots and relatively large seed
although they belong to microsperma type. The intermediate root system
types are grown in the Punjab and the North-West Frontier Province of
Pakistan.
The taproot and the laterals in the upper soil layer carry numerous
small round or elongated nodules (Fig. 4.4) when the plant grows on a
medium that contains appropriate strains of Rhizobium. Most nodules however become oblong because of the presence of an apical meristem, and several, particularly those on the taproot, become multi-lobed as a result of
bifurcation of the apical meristem. The nodules may start appearing as early
as 15 days after emergence, but the peak growth in their number and mass
occurs when the plant reaches peak vegetative growth, and it starts declining with the onset of flowering, although this could be delayed to some
extent if the soil moisture supply at that time would improve by rain or irrigation (Saxena and Hawtin, 1981). Healthy nodules have a pinkish white
appearance and when cut show pink discoloration of leghaemoglobin. The
nodules provide good medium for the growth of larvae of Sitona weevil
and can therefore be seen bored and eaten by them in fields infested by
Sitona, the insect mainly prevailing in the WANA region (see Ujagir and
Byrne, Chapter 18, this volume). The affected plants show severe nitrogen
deficiency mainly because of curtailed nitrogen fixation.
38
M.C. Saxena
Fig. 4.4. Taproot and laterals showing elongated nodules.
4.3. Stems and Plant Structure

The lentil stems are thin, square and ribbed at the angles and generally herbaceous and weak; particularly at the early vegetative stage, but in several
genotypes they get stronger with advancement in age. The basal part of the
stem becomes woody and lignified as plant growth advances. The degree of
pubescence varies with genotypes, from almost glabrous to very hairy, and
hairs are rather fine and soft. The stems are generally light green in colour
but in several genotypes they may have varying degrees of anthocyanin
pigmentation ranging from only on the basal parts to the whole of the
stems, while in others it could be completely absent.
Depending on genotype and the growth environment, the plant height
may range from 15 to 75 cm. A study by Malhotra et al. (1974), with 47
diverse lentil genotypes grown on sandy loam soil in northern India in a
spacing of 50 cm 10 cm, showed that the plant height ranged from 17 to
55 cm. Tullu et al. (2001), in their 2-year study with a core collection of lentil
germplasm comprising 287 accessions from diverse geographical regions
(Asia, Africa, North America, South America and Europe) and 15 registered
cultivars, grown under tilled and no-till production systems in Washington,
USA, confirmed that plant height at maturity was not only affected by genotypes but also by the environment. The height varied at the no-till site from
15 to 40 cm in 1996 and from 20 to 63 cm in 1997, while the respective ranges
for the conventional tillage site were 1943 cm and 2060 cm. The average
39
was 27 cm in 1996 and 37 cm in 1997. This difference was attributed to better rainfall distribution in 1997 than in 1996. The plant canopy width also
showed large genotypic variation, and, as an average of four environments,
it ranged from 8 to 18 cm at flowering and from 12 to 32 cm at maturity.
Plant height and canopy width were highly correlated with total biomass
and seed yield.
As plant height is highly influenced by environment, assigning an absolute value for the height of a specific cultivar may be of limited value. The
course of growth in plant height is also highly influenced by environment
(Saxena and Hawtin, 1981). The internodal length is rather small in the
lower part of the stem and it becomes progressively bigger in the upper
part up to 75% of the height, and then decreases again. Most of the nodes
are formed by the time of peak flowering. The dry weight accumulation in
the stem follows a sigmoid course and the rate of accumulation is highly
influenced by both genotype and environment. Studies at the International
Center for Agriculture Research in the Dry Areas (ICARDA), Syria, in a
Mediterranean environment, showed that when lentil was sown in winter
the rate of dry matter accumulation in the stem was very slow in the first
90100 days and then it increased sharply with age. In the case of the springsown crop the early slow growth phase was rather short. Improvement in
moisture supply increased dry matter accumulation, late in the season, particularly in the spring-sown crop. Almost 50% of stem dry matter was accumulated after the peak in flowering, indicating that the stem continued to
be an active sink for photosynthates and perhaps other plant nutrients even
during reproductive growth (Saxena and Hawtin, 1981). There is thus scope
for exploiting genotypic differences in remobilizing the nutrients from stem
to seed formation for improving the harvest index in lentil.
Lentil genotypes show large differences in their branching pattern,
although the expression of this trait is also greatly influenced by environment. Nezamuddin (1970) described the branching pattern in lentil genotypes grown in India. The pattern ranges from erect compact, with a narrow
branch angle, to prostrate or spreading where the branch angle is rather
wide and the lower branches remain lying close to the ground. Several
intermediate types are also found. The plant may have only a few or many
primary branches that arise directly from the main stem, and often many
secondary branches arising from primary branches. In some cases there
could be even tertiary branches. Malhotra et al. (1974) reported a range of
1.511.1 primary branches per plant in their study of 47 lines evaluated
under spaced planting. The production of branches is highly affected by
plant population, the number decreasing with increasing plant density
(Wilson and Teare, 1972).
A schematic depiction of different branching patterns and plant structure, observed in the lentil germplasm evaluated at Tel Hadya, Syria, is
shown in Fig. 4.5 (Saxena and Hawtin, 1981). A highly branched bushy pattern was typified by the accession no. 43633; a sparsely branched, tall erect type
by cultivar Laird (ILL 4349); a moderately branched, semi-tall erect plant
structure by the Egyptian cultivar Giza 9 and the Argentinean cultivar
40
M.C. Saxena
60
60
50
50
Height (cm)
40
40
(a)
(b)
(c)
(d)
(e)
30
30
20
20
10
10
0
0
10
20
30
40
50
60
70
80
90
100
110
120
0
130
Spread (cm)
Fig. 4.5. Schematic depiction of variation in branching pattern and plant structure in lentil:
(a) highly branched bushy; (b) sparsely branched, tall erect; (c) moderately branched, semi-tall
erect; (d) moderate to highly branched, semi-tall, subcompact; (e) moderate to highly branched,
short, subcompact.
Precoz (ILL 4605); a moderate to highly branched, semi-tall, subcompact

pattern by Syrian landrace Local Large (ILL 4400); and a moderate to
highly branched, short, subcompact pattern by Syrian landrace Local Small
(ILL 4401).
A ratio of plant height to canopy width is a good measure of plant
structure and it has been used to characterize growth habit in lentil germplasm evaluation (Ferguson and Robertson, 1999).
4.4. Leaves
Lentil leaves are alternate, compound and pinnate with one to eight pairs of
subsessile or sessile, ovate, elliptical or lanceolate leaflets. Each leaf is subtended by a small pair of stipules. The rachis is 45 cm in length and it may
terminate in a bristle or simple (sometimes dichotomous) tendril having a
length that may be similar to that of the rachis, particularly in the upper
leaves (Fig. 4.1). Some genotypes develop tendrils at a fairly early stage of
growth, while most tend to develop the tendrils only on leaves that unroll
just before flowering. At maturity the tendrillous habit keeps the canopy
upright, helping in machine harvest. Some other genotypes remain tendrilless. The colour of the leaves can be yellowish green, light yellow green,
dull green, dark green or dark bluish green. Under low temperature conditions, the leaves may develop purplish discoloration, which disappears as the
temperatures increase. Leaf chlorosis is also common under low temperature
and high moisture conditions on calcareous soils in the Mediterranean
41
region, but the leaves gain their normal green colour as the season advances,
and temperature and moisture conditions become optimum. The leaf pubescence may be absent, slight, medium or dense.
The leaflets of macrosperma types are relatively larger in size (1527 mm
in length, 410 mm in width) than those of microsperma type (815 mm long
and 25 mm broad) and the former are generally oval (length:width ratio =
33.5) and rarely elongated while the latter are elongated, linear or lanceolate (length:width ratio = 45) (Barulina, 1930). Szucs (1972) studied the
relationship between the leaflet width and seed diameter in 49 microsperma
and 45 macrosperma types and reported high positive correlation. The number of leaflets varies among genotypes, and within a genotype with nodal
position. The first two leaves are simple, scale like and largely fused with
two scale-like stipules (Fig. 4.1d). The following two or more leaves are
bifoliate and the subsequent ones are multifoliate. There are no stipels.
There is a small pulvinus at the base of each leaflet which causes the leaflets
to fold up together in the evening or during the day when the plant
experiences moisture stress.
4.5. Flowers and Flowering

The lentil plants are indeterminate and they come to flower after a period of
vegetative growth, the length of which varies considerably between genotypes grown in a single location. The flowering is acropetal; hence lower
nodes may bear pods close to maturity while younger nodes may carry
flowers (Fig. 4.1e). The duration of flowering of a single plant and crop
stands varies enormously; it may range from less than 10 days to longer
than 40 days depending on the genotype and the environment.
The flowers are borne on a single axillary raceme inflorescence, with a
slender peduncle, 25.5 cm long (mostly 34 cm long), on a reproductive
node. The rachis of the inflorescence ends in a filiform apex (Fig. 4.1e). There
are no bracts. The number of peduncles per plant varies with genotype, but
more so by environment. Malhotra et al. (1974) reported a range from 10 to
150 peduncles per plant in their evaluation of 47 lentil genotypes in India.
Each peduncle bears from one to four flowers (sometimes as many as
seven flowers have been found). The flowers are complete, have typical
papilionaceous structure, and are small (49mm long). Their standard petal
is white, pink, purple, light purplish blue, or pale blue (Muehlbauer et al.,
1985). In some genotypes it is white with light purplish blue veins. The
wings and keel are generally white, but wings may have a violet-blue tinge.
The calyx tube divides near its base into five narrow pointed lobes that
curve over and beyond the corolla. The stamens are diadelphous (nine plus
one) with the upper vexillary stamen free. The ovary contains one or two
ovules (sometimes three ovules) and it terminates in a short curved style. The
style is pilose on the inner side and the stigma is slightly swollen and glandular. The flowers are cleistogamous and almost exclusively self-pollinated
although some cross-pollination (less than 0.08%) may occur through thrips
42
M.C. Saxena
and other such insects but not by wind or honeybees (Muehlbauer et al.,
2002). Petals expand at a rate equivalent to about one-quarter of the length
of sepals every 24 hours and most flowers would have been pollinated by
the time petal length exceeds three-quarters that of sepals. Thus, the timing
of emasculation for artificial hybridization in a breeding programme is critical. The opening of all flowers on a single branch takes about 2 weeks to
complete (Nezamuddin, 1970). Flowers open before 10.00 h on cloudless
days but not until 17.00 h when the sky is overcast. At the end of the second
day and on the third day all open flowers close completely and the colour
of the corolla begins to fade. The visual appearance of the pod occurs after
34 days.
4.6. Pods
Lentil pods are oblong, laterally compressed, bulging over the seeds, 620
mm long and 3.511.0 mm wide; rounded to slightly cuneate at the base,
short beaked, smooth, with persistent calyx, and they contain one to three
seeds. As mentioned earlier, the main distinguishing feature between macrosperma and microsperma types is the size and shape of the pods and seeds
(Fig. 4.2). The unripe pods are usually green although in some genotypes
the colour can be purple, violet or spotted. Their lateral expansion is complete before grain filling starts. To start with, the pods are flat and as the
grain filling advances, they become somewhat swollen over the seeds. This
is particularly conspicuous in the microsperma types in which the pod surface becomes convex while in the macrosperma types it remains generally
flat (Fig. 4.2). As the crop reaches maturity, the colour of the pods changes
and, depending on the genotype, the pod wall becomes straw-coloured,
light beige, light brown, dull brown or spotted.
The number of pods per peduncle normally varies from one to four
although up to six have been found. Muehlbauer (1974) studied the genotypic difference in the number of pods per peduncle in 45 space-planted
diverse lentil lines. The frequency of one pod to the peduncle was greatest,
closely followed by that of two pods per peduncle, whereas the frequency
of three pods per peduncle was very low.
The number of pods per plant, which is a very important yield determinant in lentil, varies considerably depending on the genotype as well as the
environment. In an evaluation of Indian lentil genotypes, planted with a
spacing of 50 cm 10 cm, Malhotra et al. (1974) observed a range from 17.2
pods to 216 pods per plant, whereas, Singh and Singh (1969) found more
than 500 pods per plant in a crop planted with 60 cm 30 cm spacing.
4.7. Seeds
Lentil seeds are typically lens shaped (Fig. 4.2). Their diameter ranges from 2
to 9 mm (Barulina, 1930). The large-seeded types generally have a diameter
43
range from 6 to 9 mm, medium size from 5 to 6 mm, and the small from 3 to
5 mm. They may have a globose shape (diameter:thickness ratio ranging
from 1.5 to 2.5) or flattened (diameter:thickness ratio ranging from 2.5 to 4).
The testa colour can be pink, yellow, green, dark green, grey, brown or
black. In some genotypes the testa has dark brown or black spots, speckling
or mottling (Muehlbauer et al., 2002). The seed surface is generally smooth
but in some large-seeded types it may be wrinkled. The hilum is narrowly
elliptical and minute, and its colour is white or dull brown. The cotyledon
colour may be orange, yellow or green, the latter turning yellow after a
period of storage (Kay, 1979; Duke, 1981; Muehlbauer et al., 1985).
The number of seeds per plant is closely correlated with the number of
pods per plant, and is, therefore, an important yield attribute. Muehlbauer
(1974), in his evaluation of 45 diverse lines of lentil, observed a range from
17.6 to 139.6 pods per plant, with a mean of 62.6 pods per plant.
The 100-seed weight may range from 1.07 to 8.55 g depending on genotype. The environmental effect on seed weight is generally not very conspicuous. The 100-seed weight in the range from 1.1 to 4 g is found in the
small-seeded types, while the large-seeded types generally have a range
from 4 to 8.2 g (Barulina, 1930). The evaluation of a core collection of 287
lentil germplasm accessions by Tullu et al. (2001) revealed that the variation
ranged from 1.3 to 7.4 g/100 seeds, with an overall mean of 3.6 g. Also there
was a highly significant variation for seed size between and within yellowand red-cotyledon types, a range from 1.7 to 7.4 g/100 seeds for yellowcotyledon-type accessions and from 1.3 to 5.2 g/100 seeds for red-cotyledon
accessions. The country of origin was a major factor in determining the seed
weight and seed colour in the tested accessions.
The seeds show a 34 week period of dormancy after harvest and the duration varies with genotypes. Non-dormant seeds, when placed in moist soil at
optimum temperature, rapidly imbibe moisture attaining complete imbibition
in 12 h. Small-seeded lentils imbibe water which is 85% of their initial air-dry
weight, while the large-seeded lentils imbibe more than 100% of their initial airdry weight. The germination is hypogeal (Fig. 4.1), and the emergence in the
field occurs in 79 days in the spring-sown crop in the Mediterranean region as
the ambient temperature is around 20C, while the winter-sown crop may take
2530 days to emerge because the temperatures are less than 10C.
4.8. Anatomy
Barulina (1930) has investigated the anatomical characteristics of roots,
stem, leaf, fruit and seeds of lentil and provides illustrations and a short
description of each of these parts.
The transverse section of the taproot of lentil reveals that its epidermal
cells are small and compact and bear root hairs. The cortex comprises multiple layers of thin parenchymatic cells. The central cylinder, bound by
endodermis, has both primary and secondary xylems, bast bundles, and
phloem in between the bast bundles and secondary xylem.
44
M.C. Saxena
The transverse section of the stem of adult lentil plants is typical of

dicotyledonous plants with a small pith in the centre and conspicuous vascular bundles. Primary and secondary xylem tissues are intercepted by
modular rays and are bounded by phloem tissues. In the angular sides of
the stem there are bundles of mechanical fibres in the cortical region. These
contribute to the strength of the stem. In addition, in the transverse sections
of the stem of large-seeded lentils fibro vascular bundles are visible at the
angular parts. The medulla of the stem is often filled with starch grains. The
anatomical structure of the branch is very similar to that of the stem except
that the central part is rather large and filled with parenchymatic tissues.
The epidermal cells of the leaf are covered with a thin cuticle and they
are interspersed with simple lens-shaped stomata. Below a compact layer of
palisade tissues, there are several layers of spongy parenchyma, both containing chloroplasts. In the cross-section of the leaf at the midrib the bundle
of mechanical fibres can be seen. There are also cells filled with calcium
oxalate crystals.
The cross-section of the cover of unripe pods shows an outer and inner
epidermis, the outer one conspicuously lined with cuticle. Immediately
below the outer epidermis are parenchymatic tissues containing chloroplasts. Interspersed are the cells loaded with oxalate crystals. Below the
parenchymatic tissues and above the lower epidermis are mechanical fibre
tissues.
The cross-section of the seed testa shows a conspicuous cuticular layer
followed by a thin light layer just above the long and compact palisade
tissues constituting the epidermis. There is a layer of hypoderma just
below the epidermis and this is followed by several layers of thin-walled
parenchyma.
The cotyledons have a thin layer of epidermis followed by large storage
cells containing starch crystals.
4.9. Growth Habits in Lentil and Plant Types Adapted to Different

Agroecological Conditions
A combination of plant structure and development traits determines the
plant type of a genotype. An ideal plant type for a given environment is one
that provides the best adaptation to that environment (Lal, 2001). Stem
height, number of branches, and angle of branching, which determines the
canopy width, are the key traits that determine the structure of a plant. The
rate of early vegetative growth and ground-cover development (which is a
reflection of early vigour and which affects interception of radiation and
water use efficiency and the size of the vegetative frame of the plant; Silim
et al., 1993), phenological traits in relation to the prevailing environmental
conditions, and the ability of the plants to partition photosynthate and other
nutrients into the seed are some of important developmental and physiological traits to be considered for designing an ideal plant type for a particular agroecological situation.
45
According to Kusmenoglu and Muehlbauer (1998), increased seed yield

in lentil has been obtained through development of cultivars with shorter
vegetative and generative growth periods, greater rates of crop and seed
growth, and a higher seed-partitioning coefficient. This strategy is preferred
for the regions where there is little or no rainfall during the later stages of crop
growth and development (Silim et al., 1993). For irrigated conditions, where no
serious water scarcity is expected during the critical stages of crop growth, a
different growth habit and plant structure might be needed. According to
Solanki et al. (2007), the ideal types of lentil for these conditions should be the
ones that do not revert back to vegetative growth on receiving irrigation at
the time of flowering and are late maturing to be able to make full use of the
long growing season made possible by irrigation. An erect plant type with
shorter internodes and strong stem, so that the crop may not lodge, and bearing pods 1015 cm above the ground surface permitting mechanized harvest,
would be of additional advantage. Cultivar Matilda is a good example of
this type of lentil adapted to the Wimmera region of Victoria, Australia,
where rains do occur late in growing season (Brouwer, 1995).
Several researchers (Solanki et al., 1992, 2007; Lal, 2001; Om Vir and
Gupta, 2002) have proposed plant ideotypes that would adapt to different
agroecological conditions in India. As the conditions in different lentilgrowing areas are vastly different, Lal (2001) has suggested the following
different plant types to cover major production systems: (i) for dry/rain-fed
conditions deep root system with high root volume, profuse branching,
semi-spreading to spreading growth habit to prevent evaporation, low transpiration, pubescent foliage, reduced biomass, high harvest index and short
duration; (ii) for assured moisture conditions erect to semi-erect growth
habit with short internodes, compact plant type with restricted branching,
high biomass production capacity and long duration; (iii) for intercropping
with other crops erect and compact growth habit with shorter internodes
and a capacity to have good photosynthesis at low-light intensity; (v) for
relay cropping/multiple cropping fast germination capacity, high early
vigour, early flowering and responsiveness to improved agronomic inputs.
References
Genetics and Plant Breeding Supplement 40. USSR Institute of Plant Industry of
the Lenin Academy of Agricultural Science Leningrad, USSR, pp. 265.
Brouwer, J.B. (1995) Lens culinaris (lentil) cv. Matilda. Australian Journal of Experimental
Agriculture 35(1), 117.
Duke, J.A. (1981) Handbook of Legumes of World Economic Importance. Plenum
Press, New York, pp. 5257.
Ferguson, M.E. and Robertson, L.D. (1999) Morphological and phenological variation
in wild relatives of lentil. Genetic Resources and Crop Evolution 46, 39.
Gahoonia, T.S., Ali, O., Sarker, A., Rahman, M.M. and Erskine, W. (2005) Root traits,
nutrient uptake, multi-location grain yield and benefit-cost ratio of two lentil (Lens
culinaris, Medik.) varieties. Plant and Soil 272, 153161.
46
M.C. Saxena
Kay, D. (1979) Food Legumes. Tropical Products Institute Crop and Products Digest
No. 3. Tropical Products Institute, London, UK, pp. 4871.
Kusmenoglu, I. and Muehlbauer, F.J. (1998) Genetic variation for biomass and residue
production in lentil (Lens culinaris Medik.). II. Factors determining seed and straw
yield. Crop Science 38, 911915.
Lal, S. (2001) Plant type concept in pulses. Souvenir: National Symposium on Pulses for
Sustainable Agriculture and Nutritional Security, 1719 April, New Delhi. Indian
Society of Pulses Research and Development (ISPRD), Kanpur, India, pp. 5662.
Malhotra, R.S., Singh, K.B. and Singh, J.K. (1974) Genetic variability and genotypeenvironmental interaction studies in lentil. Journal of Research Punjab Agricultural
University 10(1), 1721.
Muehlbauer, F.J. (1974) Seed yield components in lentils. Crop Science 14, 403406.
Muehlbauer, F.J., Cubero, J.I. and Summerfield, R.J. (1985) Lentil (Lens culinaris
Medik.). In: Summerfield, R.J. and Roberts, E.I.I. (eds) Grain Legume Crops. Colins,
London, UK, pp. 266311.
Muehlbauer, F.J., Summerfield, R.J., Kaiser, W.J., Clement, S.L., Boerboom, C.M.,
Welsh-Maddux, M.M. and Short, R.W. (2002) Principles and Practices of Lentil
Production. United States Department of Agriculture (USDA) Agriculture Research
Service (ARS). Available at: http://www.ars.usda.gov/is/np/lentils/lentils.htm
(accessed on 28 February 2008).
Nezamuddin, S. (1970) Miscellaneous, Masur. In: Kachroo, P. (ed.) Pulse Crops of India.
Indian Council of Agricultural Research, Krishi Bhawan, New Delhi, pp. 306313.
Om Vir and Gupta, V.P. (2002) Analysis of relationships of yield factors in microsperma
macrosperma derivatives of lentil. Legume Research 25(1), 1520.
Sarker, A., Erskine, W. and Singh, M. (2005) Variation in shoot and root characteristics
and their association with drought tolerance in lentil landraces. Genetic Resources
and Crop Evolution 52(1), 8795.
Saxena, M.C. and Hawtin, G.C. (1981) Morphology and growth patterns. In: Webb, C.
and Hawtin, G.C. (eds) Lentils. Commonwealth Agricultural Bureau, Slough, UK,
pp. 3952.
Silim, S.N., Saxena, M.C. and Erskine, W. (1993) Adaptation of lentil to Mediterranean
environment. I. Factors affecting yield under drought conditions. Experimental
Agriculture 29, 919.
Singh, K.B. and Singh, S. (1969) Genetic variability and interrelationship studies on
yield and other quantitative characters of lentils. Indian Journal of Agricultural
Science 39, 737741.
Solanki, I.S., Singh, V.P. and Wadia, R.S. (1992) Model plant type in lentil (Lens
culinaris Medik.). Legume Research 15(1), 16.
Solanki, I.S., Yadav, S.S. and Bahl, P.S. (2007) Varietal adaptation, participatory
breeding and plant type. In: Yadav, S.S., McNeil, D.L. and Stevenson, P.C. (eds)
Lentil: an Ancient Crop for Modern Times. Springer, Dordrecht, The Netherlands,
pp. 255274.
Szucs, A. (1972) Study of the cultural value of lentil varieties. Agrobotanika 14,
103124.
Tullu, A., Kusmenoglu, I., McPhee, K.E. and Muehlbauer, F.J. (2001) Characterization
of core collection of lentil germplasm for phenology, morphology, seed and straw
yields. Genetic Resources and Crop Evolution 48, 143152.
Wilson, V.E. and Teare, I.D. (1972) Effect of between and within row spacing on
components of lentil yield. Crop Science 12, 507510.
Zohary, D. and Hopf, M. (2000) Domestication of Plants in Old World, 3rd edn.
Oxford University Press, UK, pp. 328.
Agroecology and Crop Adaptation

M. Materne1 and K.H.M. Siddique2
1Department
2The
of Primary Industries, Horsham, Victoria, Australia;

University of Western Australia, Crawley, Western Australia, Australia
5.1. Introduction
Lentil (Lens culinaris Medikus ssp. culinaris) is one of the worlds oldest cultivated plants. It was domesticated in the Fertile Crescent of the Near East
over 7000 years ago (see Cubero et al., Chapter 3, this volume). Lentil was
introduced into the Indo-Gangetic Plain around 2000 bc and now half of the
worlds current lentil production is in South Asia. In the Americas, lentil
production became widespread in Chile, Argentina and Brazil (Barulina,
1930), prior to expanding into the Palouse region of north-eastern USA in
1916 and western Canada in 1969 (Muehlbauer and McPhee, 2002). In Australia, significant commercial cultivation of lentil only began in 1994.
Lentil has become established in a wide range of agroecological environments but production is limited in tropical areas. The spread of lentil
from the centre of origin has been accompanied by the selection of traits
important for adaptation to environments that can be defined by climate,
soil and their impact on season length, abiotic and biotic stresses.
5.2. Production Environments

Temperature, and the distribution and quantity of rainfall are the main
determinants of where and when lentil is grown around the world. In West
Asia, North Africa and Australia lentil is sown in winter in areas that receive
annual rainfall of 300450 mm. In these regions, low temperatures and radiation restrict vegetative growth during winter, but growth is rapid in spring
when temperatures are rising. Ripening occurs prior to, or early in summer,
when temperatures and evaporation are high and rainfall low. In subtropical regions of Pakistan, India, Nepal and Bangladesh, lentil is also grown as
a winter (rabi) crop but temperatures are higher and crops are grown
47
48
primarily on residual soil moisture from monsoon rains. In high altitude

and/or latitude areas such as central Turkey, USA, Europe and Canada,
where the winters are too cold for lentils to grow reliably, sowing is delayed
until spring. Lentils are grown on stored moisture and/or snow melt supplemented by rainfall during spring and summer when temperatures are
warm and day lengths long. In broad terms lentils have been selected for
adaptation to these three major climatic regions of the world.
Within each growing region, variations in climate, soils, and their interactions affect lentil productivity and quality either directly or indirectly
through their influence on foliar and soil-borne diseases, pests and interactions with rhizobia. While natural selection has been important in establishing crops in traditional lentil-growing areas, the development of new
cultivars and/or cultural practices will improve the adaptation, yield and
quality of lentil in traditional and newer growing regions. For example, late
sowing in the USA and Turkey avoids extremely cold temperatures and the
lentil cultivars grown are adapted to growing in summer. Within a region,
traits may provide benefits for adaptation over large areas such as with water
stress, or smaller areas such as those that occur with local and variable soil
toxicities in Australia. To a large extent, cultural practices have been based
on the use of landraces but this is changing as new varieties, agronomic,
technological, cultural and financial opportunities become available.
5.3. Phenological Adaptation to the Environment

Importance of phenology for adaptation
The clustering of traits that define the phenological adaptation of lentil to
an ecological environment indicates that local environments are important
in the evolution of lentil (Erskine et al., 1989). Matching a crops phenology
to an environment is a key part of improving adaptation and increasing
crop yields. The major constraints to seed production in most environments
are drought and high temperatures, particularly during flowering and seed
growth. Summerfield et al. (1989) reported that, under controlled conditions,
progressively warmer temperatures post-flowering restricted vegetative
growth, accelerated progress towards reproductive maturity and reduced
seed yield in lentil. In temperate and Mediterranean environments, crops are
also at risk from frost during flowering and pod fill.
Flowering time is particularly important for adaptation as it determines
the length of the vegetative phase (sowing to flowering), and determines
the climatic conditions that the crop will be exposed to during reproductive
growth. Lentil genotypes adapted to West Asia and high latitudes flower
too late and produce few pods when grown in southern Asia (Indian subcontinent) because reproductive development begins when conditions are
increasingly hot and dry. Similarly, lentils from high latitudes flower and
mature too late for economic yields when grown in winter in West Asia
and southern Australia. Alternatively, in high latitude countries such as
49
North America, lentils from the Middle East and South Asia flower too
early and biomass production and seed yields are low.
Adaptation to an environment does not necessarily infer optimum
adaptation in any one year due to the variability of climate. For example,
flowering time and the length of the reproductive period were negatively
correlated with seed yield in a very dry year in Syria but not in a wetter
year (Silim et al., 1993). In a similar environment in Australia, dry years are
also associated with an increase in the frequency of frosts and earlier flowering may expose crops to a greater risk of damage. If flowering is too early
in these environments crops cannot produce adequate biomass to sustain
large seed yields in average or higher rainfall years.
Flowering response of lentil to photoperiod and temperature

Understanding the flowering response of lentil has been a key to understanding adaptation. Roberts et al. (1988) found no evidence for a specific
low-temperature response in lentil, while Tyagi and Sharma (1981) reported
a small response. Like most annual winter crops, the timing of phenological
events in lentil is modulated primarily by the responsiveness to photoperiod and temperature (Summerfield et al., 1985, 1996). In lentil, Erskine et al.
(1990) characterized 231 accessions for their flowering response to photoperiod and temperature in glasshouse experiments which were confirmed in
field experiments in Syria and Pakistan (Erskine et al., 1994). In both studies, a large proportion of the variation among accessions for time to first
flower and response to temperature and photoperiod was related to origin.
In particular, sensitivity to photoperiod was dependent on latitude of origin. Accessions from lower latitude countries such as India, Ethiopia and
Egypt have a longer non-responsive phase, are less sensitive to photoperiod
and more sensitive to temperature (Erskine et al., 1994). A lower sensitivity
to photoperiod improved adaptation at lower latitudes by ensuring that
flowering occurs at shorter daylengths and is not delayed past the optimal
and before temperatures rapidly increase.
The dissemination of lentil into new environments has caused selection
of different regionally specific balances between photoperiod and temperature for the control of flowering. For example, in Australia, the most broadly
adapted lentil cultivars were inherently late to flower but had a relatively
large response to temperature and small response to photoperiod (Materne,
2003). Flowering in these genotypes is delayed when days are cooler and
shorter, characteristics of a long growing season, but relatively early at
lower latitudes where winter temperatures are higher and the length of the
growing season is shorter. Inherently early, photoperiod-responsive genotypes, including landraces from Syria, are more specifically adapted to winter sowing in longer season areas of southern Australia. Inherently early,
temperature-responsive but photoperiod-insensitive genotypes, such as Precoz, are specifically adapted to winter sowing in lower latitude short season environments. Inherently late genotypes with only moderate responses
50
to temperature and photoperiod were too late to flower in all winter sowing areas of Australia and are specifically adapted to growing in the warm
long days of summer at higher latitudes. The photothermal model and climatic data have also been used to select genotypes suited to winter sowing
in the highlands of central and eastern Anatolia (Keatinge et al., 1996).
5.4. Impact of Abiotic Stresses on Lentil Adaptation

Water availability
Water availability, in particular rainfall in dryland farming, is a major determinant of grain yield in most crops whether it occurs during the growing
season or prior to sowing and stored in the soil for later use by the crop. In
lentil, total growing season rainfall accounted for 4155% of the total variation in seed yield over many sites and seasons in the West Asia and North
Africa (WANA) region (Erskine and Saxena, 1993); up to 80% of the variation in seed yield over several seasons at one site in Syria (Erskine and El
Ashkar, 1993), and 56% of the total variation in seed yield among 11 diverse
lentil genotypes in south-eastern Australia (Materne, 2003).
Compared to pulses such as faba bean and chickpea, lentil is grown in
drier areas of WANA with as little as 250 mm of annual rainfall. However,
a lack of water is the major limitation to lentil production worldwide and,
in its most severe form, drought results in crops with no economic grain
yield (see Shrestha et al., Chapter 12, this volume). Autumn- or winter-sown
crops in Mediterranean environments are likely to experience intermittent
drought during vegetative growth and terminal drought during reproduction when temperatures are rising and rainfall is decreasing. Spring-sown
crops in Mediterranean environments and winter-sown crops in the semiarid tropics rely on available soil moisture and experience increasing water
stress during the growing season. In South Asia, drought may also occur
during plant establishment if sowing is delayed and plant roots cannot
reach subsoil moisture.
Natural selection and breeding under variable rainfed environments
has resulted in landraces and cultivars that have increased water use efficiency and are responsive to moisture availability. Crops must access and
use as much water as possible to produce biomass and seed while not losing tissues that have been produced using water. Appropriate phenology,
resistance to abiotic diseases and tolerance to abiotic stresses are likely to be
important in increasing water use efficiency in an environment. Various
mechanisms such as high early vigour, and early flowering and maturity
have been proposed for escaping terminal drought. In Syria, the midflowering genotypes ILL 4400 and ILL 4401 had high seed yields in most
seasons (Murinda and Saxena, 1983) but they were intolerant to drought in
another study where early flowering genotypes were highest yielding (Silim
et al., 1993). Similarly, early flowering and maturing genotypes were high
yielding in low rainfall, low yielding environments in Australia but the yields
51
of these genotypes were low compared to the best mid-flowering genotypes

over eight locations and 5 years (Materne, 2003). Although early flowering
is important for drought escape, it is not effective if genotypes have a relatively low mean yield over many seasons and sites, due to an inability to
respond to increasing available soil moisture. Genotypes that were responsive to temperature for flowering were the most broadly adapted in Australia and may avoid drought by flowering at an optimal time in most years
but earlier in drought years when temperatures are often higher (Materne,
2003).
Early sowing has been advocated in many areas to avoid rising temperatures and drought during reproduction, and to maximize yields. However, early sowing may expose crops to increased weed competition, diseases
and abiotic stresses. Thus, addressing constraints to enable early sowing is
another effective method of improving tolerance to terminal drought.
Waterlogging
Lentil does not tolerate waterlogging at germination compared to cereals,
and vegetative growth and roots are severely depressed by waterlogging
after emergence (Jayasundara et al., 1998). Waterlogging can reduce lentil
yields at any time during the growing season especially in lower lying areas
or on poorly structured soils where drainage is impeded. In Nepal, sowing
of lentil after rice is delayed until the soil dries, but in most areas of the
world waterlogging is avoided by not growing the crop in prone areas or
by using drainage systems. The sensitivity of lentil and potential sensitivity
of nitrogen fixation to waterlogging and anaerobic conditions makes irrigation more difficult and its poor growth response explains the low use of
lentil in irrigated cropping systems. Opportunities exist to improve adaptation to waterlogging (Bejiga and Anbessa, 1995).
Temperature
Temperature has been shown to influence the evolution and adaptation of
lentil worldwide and, with water availability, essentially determines the
growing season for lentil. The effects of temperature on flowering have
been discussed. Erskine (1996) also found that among 171 lentil genotypes
from Syria and Turkey, large-seeded (macrosperma) accessions had a longer
reproductive period than small-seeded (microsperma) accessions in Syria. He
concluded that larger seeded accessions were higher yielding in cooler seasons due to a longer seed-filling period, but were lower yielding at higher
temperatures. Worldwide, green lentils are more prominent in colder areas
where lentil is grown in summer and matures when temperatures are
cooler. Conversely, red lentils dominate production in winter-growing areas
where temperatures are increasing during seed maturation. Extremes of
temperature during lentil growth and reproduction can result in more
52
specific and dramatic effects. For example, high temperatures during spring
reduce seed set and cause rapid maturation in West Asia, North Africa and
Australia while cold winters limit production to spring sowing in higher
altitude or latitude regions.
Low temperature
In the USA and Turkey, lentil are traditionally grown in spring as cold winter temperatures kill lentil plants. However, large yield increases have been
achieved by sowing lentil in winter rather than spring if problems associated with a lack of winter hardiness can be overcome (Muehlbauer and
McPhee, 2002). Genotypes able to survive temperatures below freezing during vegetative growth have been selected from field environments where
they survived temperatures as low as 26.8C (Erskine et al., 1981; Hamdi
et al., 1996; Stoilova, 2000; Chen et al., 2006) and in controlled environments
using a cold treatment of 15C (Ali et al., 1999). Cold tolerant genotypes
originated from high elevation areas, indicating that lentil has adapted to
growing at colder temperatures in these areas rather than relying on
avoidance (Hamdi et al., 1996). In the USA, winter hardy types have been
selected and evaluated with a range of agronomic practices to initiate winter sowing of lentil, but reliability of production is still a major issue (Chen
et al., 2006).
In some Mediterranean environments such as in Australia, frost during
the reproductive period can cause major economic losses by killing flowers,
pods and seeds with associated reductions in seed yield and quality. Currently, the only mechanism for limiting frost damage is avoidance by using
later flowering genotypes or delaying sowing. However, the benefits of
these strategies may be small or non-existent in years when frosts occur,
and result in lower yields in other years (Materne, 2003).
High temperatures
Summerfield et al. (1989) reported that under controlled conditions, progressively warmer temperatures post-flowering restricted vegetative growth,
accelerated progress towards reproductive maturity and reduced seed
yield. High temperatures associated with strong winds and dry soils are
especially damaging. Rhizobia are also susceptible to higher temperatures,
particularly when conditions are moist (Malhotra and Saxena, 1993). Earlier
flowering and maturity can avoid high temperatures in winter-sowing environments but may expose crops to a greater risk of frost damage in some
environments and may limit yield potential.
Nutrient toxicities
Lentil are typically grown and adapted to neutral to alkaline soils but yields
are compromised where soils are acidic, sodic, saline or have high levels of
boron. Alleviating such toxicity problems through soil modification is not
53
an economic or practical solution and therefore growing more tolerant cultivars is considered the best approach to overcoming these constraints.
Boron
Boron toxicity is an increasingly recognized problem of agriculture in the
arid areas of West Asia and Australia (Ralph, 1991; Yau and Erskine, 2000;
Hobson et al., 2006). Levels as low as 4 ppm have produced visual toxicity
symptoms on lentil 26 days after sowing compared with barley and oats at
47 and 37 days, respectively (Chauhan and Asthana, 1981).
Hobson et al. (2003) identified lines with tolerance to boron among landraces from Ethiopia (ILL 2024), Afghanistan (ILL 213A, ILL 1818, ILL 1763,
ILL 1796) and the Middle East (ILL 5845), while accessions from Europe
had the least tolerance. These origins of tolerance are consistent with wheat
(Moody et al., 1988), winter barley (Yau, 2002) and field pea (Bagheri et al.,
1994) and indicate where excess boron is potentially a problem. When
grown in a reconstituted core resembling natural high soil boron distribution in Australia, ILL 2024 had no significant reduction in yield where high
subsoil boron (18.2 mg/kg) occurred at a depth of either 30 or 10 cm in the
soil profile compared with reductions of 32 and 91%, respectively, in the
Australian cultivar Cassab (Hobson et al., 2004). While the two tolerant
accessions ILL 2024 and ILL 213A were generally characterized by an ability to partially exclude boron from shoot tissues, ILL 2024 had less leaf toxicity symptoms but ILL 213A maintained better growth at high leaf boron
concentrations (Hobson, 2007).
Boron deficiency has been identified as a limitation to lentil production
on soils in Nepal (Srivastava et al., 1999) and India (Sakal et al., 1988). Germplasm from South Asia exhibit least symptoms of boron deficiency while
those from the Middle East had the most severe symptoms (Srivastava et al.,
2000), supporting the data from boron toxicities studies. Gahoonia et al.
(2005) found lentil lines with different root morphologies better able to
scavenge micronutrients such as boron from the soil and increase yield by
1020%. There is increasing evidence that the same genetic mechanisms are
likely to control tolerance to both boron deficiency and toxicity, predominantly boron exclusion (Yau and Erskine, 2000; Dannel et al., 2002). This
dual control may restrict the adaptability of lentils from South Asia to Australia and West Asia and vice versa.
Salinity
Salinity occurs mainly in arid and semi-arid regions such as in West and
Central Asia and in Australia, and where irrigation has led to salinization in
the Indo-Gangetic Plain of South Asia, West Asia, North Africa, western
USA and Australia (McWilliam, 1986; Saxena et al., 1993; Nuttall et al., 2003).
Legumes are relatively sensitive to salt; lentil is comparatively more sensitive
than field pea and faba bean and similar to chickpea (Saxena et al., 1993).
Variation in the tolerance of lentil to MgSO4, NaCl, Na2SO4 and MgCl2
has been identified (Jana and Slinkard, 1979) but most research has focused
54
on tolerance to NaCl. NaCl tolerant accessions have been identified based

on seedling symptoms: DL 443 and Pant L 406 (Rai et al., 1985), ILL 5845,
ILL 6451, ILL 6788, ILL 6793 and ILL 6796 (Ashraf and Waheed, 1990) and
LG 128 (ILL 3534) (Maher et al., 2003). Importantly, a positive correlation
was observed between salt tolerance at different stages of growth (Ashraf
and Waheed, 1993). Maher et al. (2003) found that tolerant lentil accessions
were unaffected by NaCl concentrations commonly found in Australia but
the growth of local cultivars was severely reduced. High-yielding breeding
lines with improved NaCl tolerance have since been identified and commercialized in Australia (Materne et al., 2006).
The response of crops to salinity is affected by many other environmental factors such as soil water status, relative humidity, temperature and
nutrition (Saxena et al., 1993; Lachaal et al., 2002). For example, lentils that
are slow growing were found to be more sensitive to salt than those growing
rapidly (Lachaal et al., 2002). Breeding for increased vigour and the elimination of other abiotic and biotic stresses may offer potential to indirectly
reduce the effects of salinity.
Sodicity
In lentil, increasing soil sodicity (1025 exchangeable sodium percentage,
ESP) reduced plant height, leaf area, leaf dry weight, total biomass production and seed yield in lentil, and reduced both the nitrate reductase activity
in the leaf and the total concentration of nitrogen (Singh et al., 1993). Gupta
and Sharma (1990) found lentil had a sodicity threshold of 14.0% compared
to a value of 40.2% for wheat. Sodicity also results in poor soil structure
which can inhibit root growth and water penetration, thus limiting the
plants access to water and nutrients or causing transient waterlogging.
Breeding for stronger roots or waterlogging tolerance may overcome some
of the effects of sodicity but research is limited.
Iron and zinc deficiencies

Symptoms of iron deficiency are common in lentil, especially on high pH
soils where iron becomes less available. In Syria, iron deficiency symptoms
were positively correlated with cold susceptibility, which supports observations in other parts of the world and the proposition that wet conditions on
calcareous soils increase the development of iron deficiency symptoms and
together they affect root activity (Erskine et al., 1993). Yield losses of 1825%
were recorded among genotypes susceptible to iron deficiency in India (Ali
et al., 2000). Similarly, in Syria, yield losses of 47% were reported in genotypes with severe iron deficiency symptoms, but no yield loss occurred in
accessions with mild visual symptoms (Erskine et al., 1993). Genotypes from
relatively warmer climates such as in India and Ethiopia are more sensitive
to iron deficiency, while those from the Middle Eastern countries such as
Syria and Turkey have high levels of tolerance (Erskine et al., 1993; Ali et al.,
55
2000). The dramatic symptoms associated with iron deficiency enable the
relatively simple selection of tolerant genotypes in target areas such as the
Middle East and Australia.
Among the cool season pulses, lentil is the most sensitive to zinc deficiency (Ali et al., 2000). In developed countries, iron and zinc deficiencies
are ameliorated through the application of fertilizers, but the use of efficient
cultivars will increasingly become important as fertilizer costs increase.
Poor nitrogen fixation, due to factors affecting the host, the rhizobia and
the symbiosis between the two, can cause nitrogen deficiency and reduced
yield in lentil. In this case the adaptation of both the host and rhizobia and
the symbiotic relationship are important to ensure good nitrogen fixation,
high yield and rotation benefits to cereals. While rhizobia are well adapted
to some soils, their persistence and performance can be relatively poor on
acid or saline soils (Slattery and Pearce, 2002). Rhizobia strains have been
identified that are more tolerant to these soil types and, with a compatible host that is also inherently tolerant to the soil stress, offers opportunities to improve adaptation to acidic or saline soils (Rai and Singh, 1999;
Slattery and Pearce, 2002). However, the ability to implement improved
rhizobia on fertile, alkaline soils that had a long history of host crops and
naturally high levels of rhizobia may be difficult (Slattery and Pearce,
2002).
5.5. Impact of Diseases on Lentil Adaptation

In traditional lentil-growing areas diseases have coevolved over thousands
of years whereas in newer areas diseases often accompany the spread of
lentil, becoming significant only if adapted to the environment. As a result,
quarantine restrictions on plant matter and seed occur in some countries to
limit the introduction and establishment of diseases. Major diseases of lentils are outlined in Chapter 17.
Resistance or tolerance to major disease is a key to the adaptation of
lentil in most areas, especially where cultural controls such as fungicides
are unavailable or uneconomic. In recent years, resistant cultivars have
become available to farmers. For example, Fusarium wilt-resistant cultivars have been released in Syria, and Stemphyllium blight- and rustresistant cultivars in Bangladesh. Although host resistance is an attractive
method of disease control, more virulent pathotypes can arise through
introduction, recombination or mutation and lead to susceptibility of the
crop. This process of coevolution between diseases and host necessitates
ongoing breeding and highlights the importance of understanding the
pathogen population. Diseases also interact with agronomic practices that
change the environmental conditions in which the crop grows and crop
growth, thus impacting on the adaptation of disease and host. For example, in Australia, early sowing increased the prevalence of the diseases
caused by Ascochyta lentis and Botrytis spp. (Knights, 1987; Materne,
2003).
56
5.6. Impact of Management on Lentil Adaptation

Management techniques and lentil genotypes grown have evolved over a
long period of time and are relatively stable in traditional lentil-growing
areas. However, with globalization, lentil germplasm has been distributed
around the world and new methods and tools have become available to
farmers. These can be used to improve yield and profitability if optimal
management practices are used with genotypes adapted to the practices.
Management is increasingly being recognized as a key component in genotype by environment interactions. Agronomic management is discussed in
more detail in Chapter 14.
Impact of time of sowing on lentil adaptation

Lentil is sown after the onset of autumn rains in Mediterranean climates,
after the harvest of summer crops in tropical and subtropical regions and in
spring in colder areas (Muehlbauer et al., 1995). In the USA, Turkey and
New Zealand large yield increases are achieved by sowing lentil in winter
rather than spring, however problems with a lack of winter hardiness and/
or increased incidence of diseases, along with weed control issues, must be
addressed before winter sowing becomes widespread.
The early sowing of lentil often produces the highest seed yield in
most environments. However, a delay in sowing may result in the higher
yield where early sowing increases the incidence and severity of a disease,
insects, weeds or excessive vegetative growth and severe lodging. Early
sowing increases yield potential as it lengthens the vegetative growth
period, but this also increases the period of coincidence of pathogens,
pests and weeds with the crop to increase biotic stress pressure. Therefore
resistance to diseases and pests, suitable control strategies, or changes in
morphology to reduce disease, such as improved lodging resistance to
avoid Botrytis grey mould, may be needed in order to gain the benefits of
early sowing.
Management practices can also impact on the relative yield response of
genotypes and significant interactions between genotype and sowing rate,
application of irrigation, row spacing, and sowing depth can also affect
yield and thus adaptation.
Impact of weed management methods on lentil adaptation

As a result of their slow growth during winter and short stature, lentil competes poorly with weeds and weed control is a major limitation to growing
lentil worldwide, particularly with early sowing. In many traditional lentilgrowing countries weeds are removed by hand, but this is time consuming
and increasingly uneconomical due to high labour costs. Tillage also reduces
57
the impact of weeds but this can delay sowing and increase the risk of erosion and soil degradation. Production in some regions, especially North
America and Australia, is dependent on herbicides for weed control. However, broadleaf-herbicide options are limited and most can cause injury to the
lentil crop. Cultivars with improved tolerance to herbicides such as diflufenican and metribuzin, 4-(4-chloro-o-tolyloxy)butyric acid (MCPB) (Muehlbauer
and Slinkard, 1983), trifluralin (Basler, 1981) or Clearfield Lentil with tolerance to imidazolinone herbicides (Holm et al., 2002), and with good early
vigour offer potential to improve weed control in lentil. Crop topping and
weed wick wiping techniques have also been developed to control difficult
weeds in Australia (Preston, 2002). However, crop topping is more difficult
in later maturing and taller crops and may not be compatible with cultivars
that have improved harvestability. The ability to effectively control weeds
is vital if lentil is to be adapted to high-input continuous-cropping systems
where good weed control is targeted in all crops.
Impact of harvesting methods on lentil adaptation

The large-scale production of lentil in developed countries occurs with
mechanized harvesting systems, yet many traditional lentil-producing
countries are still harvesting by hand. None the less, hand harvesting is
considered a major constraint to lentil production and its high cost has
caused a large decrease in lentil production in Jordan and Syria (Erskine
et al., 1991).
Cultivars adapted to mechanical harvesting have been released in Syria,
Lebanon, Iraq and Turkey and their use, combined with mechanized harvesting, increased net returns to growers (Sarker and Erskine, 2002). Short
height, pod drop, pod dehiscence, lodging and uneven ripening cause seed
losses in crops that are machine harvested (Erskine, 1985; Silim et al., 1989;
Erskine et al., 1991; Ibrahim et al., 1993). The optimal time for harvesting
lentil is also relatively short with plants dropping pods, shattering and
lodging if they are harvested too late. Mechanized harvest is a multidimensional problem that can be solved by using suitable cultivars, methods such
as rolling to ensure a smooth seedbed, and a harvester setup effective for
lentil, including the use of flexi-fronts. Desiccation or swathing also allows
timely harvest especially where temperatures are declining at harvest time
such as in Canada. Breeding can assist harvest mechanization through the
development of taller, non-lodging cultivars that mature evenly and retain
their pods and seeds at maturity. However, in the USA, tall, erect cultivars
had lower yield and competed poorly with weeds compared to shorter
branching types (Muehlbauer et al., 1995). Erskine and Goodrich (1988)
found that lodging increased with plant height and late maturity and therefore tall or late genotypes must have good lodging resistance for machine
harvest. Breeding lines that were erect at harvest are also more prone to
pod-drop due to high winds, highlighting the need to simultaneously breed
for both traits (Materne et al., 2002).
58
Improving adaptation through understanding genotype by environment interactions

in lentil
The key to understanding adaptation is in the interpretation of genotype by
environment interactions (GxE) and how genotypes respond to a changing
environment. It is of particular importance where breeding programmes are
developing cultivars for soils, climates and biotic factors that vary widely
across locations and years. Most of these environmental conditions are a
result of natural variation but some such as soil fertility and weed control
can be affected by management over time. Management practices can also
impact on the relative yield response of genotypes and significant interactions between genotype and sowing rate, sowing time and application of
irrigation have been identified in lentil.
The complexity of adaptation is demonstrated in the significant GxE in
all studies conducted on lentil, even though the potential to identify GxE is
often limited by the number and/or diversity of genotypes, sites or years.
The great challenge is to identify genotypes that are broadly adapted across
a defined region, or genotypes specifically adapted to particular regions,
and the traits that are important for this adaptation. The first indications
and explanation of GxE in lentil occurred when morphological differences
in lentil landraces were related to specific adaptation to a region, for example flowering time (Barulina, 1930; Erskine and Witcombe, 1984; Erskine
et al., 1989). Many other examples of soil, climate and biotic factors have
been reviewed that can explain GxE between diverse regions.
Within a region, GxE can be difficult to explain. Hamdi et al. (1991)
found only small GxE in West Asia but specific adaptation can be identified
from other studies. ILL 4400 and ILL 4401 had a high yield in most years
and sites in Syria (Murinda and Saxena, 1983) but were intolerant to drought
and were lower yielding than earlier flowering genotypes in dry environments (Silim et al., 1993). Under irrigation in Egypt GxE was significant for
yield in the 1 year of testing and were related to differences in soil type and
temperature (Hamdi and Rebeia, 1991). In two separate studies in India,
some lentil genotypes were identified that were stable in yield over 3 years
of testing and others that were better adapted to low- and high-yielding
years (Sharma, 1999; Solanki, 2001). Sharma (1999) proposed early maturity
for stable yields. In Ethiopia, significant GxE was explained by the distribution of seasonal rainfall with later maturing genotypes being favoured when
rainfall occurred over a longer period (Bejiga et al., 1995). In some studies
the highest yielding genotypes were the most unstable for yield and performed poorly in low-yielding environments, but high-yielding genotypes
with average stability in yield were considered to have the greatest economic potential (Bejiga et al., 1995; Chowdhury et al., 1998; Elwafa, 1999;
Sharma, 1999; Solanki, 2001). However, this was not the case in Australia
where the intermediate flowering group of cultivars had the highest and
most stable yields (Materne, 2003). While flowering response explained
much of the variation in grain yield across sites in this study, Indian genotypes had the desired flowering response for broad adaptation but were
59
low yielding across Australia, indicating that other factors were also important for adaptation.
5.7. Conclusion
Lentil has historically been grown as a source of protein to complement
cereals in the diet. However, the adaptation of lentil is increasingly related
to its economic value to farmers who ultimately decide to grow the crop. To
improve adaptation, the key traits that facilitate increased production (level
and reliability of seed production), farming system benefit and price (quality of seed), or reduce cost must be identified, quantified and addressed
through improved production technology and breeding. More specifically,
profitability is a culmination of a species interaction with the environment,
in particular the quantity and distribution of rainfall and temperature which
influences the length of growing season, soil characteristics and biotic
stresses, and cultural practices. Farming system benefits include those
involved with rotation, such as providing a disease break or improving soil
nutrition, weed management, and the timing and simplicity of total farm
operation.
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genetic characterisation of flowering responses to photoperiod and temperature.
Tyagi, M.C. and Sharma, B. (1981) Effect of photoperiod and vernalization on flowering
and maturity in macrosperma lentils. Pulse Crops Newsletter 1(2), 4041.
Yau, S.K. (2002) Comparison of European with West Asian and North African winter
barleys in tolerance to boron toxicity. Euphytica 123, 307314.
Yau, S.K. and Erskine, W. (2000) Diversity of boron-toxicity tolerance in lentil growth
and yield. Genetic Resources and Crop Evolution 47, 5561.
Genetic Resources: Collection,

Characterization, Conservation
and Documentation
B.J. Furman,1* C. Coyne,2 B. Redden,3 S.K. Sharma4

and M. Vishnyakova5
1International
Center for Agriculture Research in the Dry Areas (ICARDA), Aleppo,

Syria; 2Western Regional Plant Introduction Station, Pullman, Washington, USA;
3Department of Primary Industries, Horsham, Victoria, Australia; 4National Bureau of
Plant Genetic Resources, New Delhi, India; 5N.I. Vavilov Institute of Plant Industry
(VIR), St Petersburg, Russia
*Present address: United States Department of Agriculture (USDA) Agriculture
Research Service (ARS), Palmer, Alaska, USA
6.1. Major Ex Situ Collections

Conservation of Lens germplasm is almost entirely ex situ as seed, including
wild and domestic annuals as well as wild perennials. The International
Center for Agricultural Research in Dry Areas (ICARDA) has the global
mandate for research on lentil improvement, and thus houses the world
collection of Lens, which includes around 10,800 accessions. The ICARDA
lentil genetic resources collection includes 8860 accessions of cultivated lentil from more than 70 different countries representing four major geographic
regions (Fig. 6.1), 1373 ICARDA breeding lines, and 583 accessions of six
wild Lens taxa, representing 24 countries (Table 6.1; see Cubero et al., Chapter 3, this volume). The majority of the ICARDA collection (48%) consists of
accessions from Central and West Asia and North Africa, the centre of origin and primary diversity (Zohary and Hopf, 1988; Ferguson and Erskine,
2001; see Cubero et al., Chapter 3, this volume), while South Asia contributed an additional 25%. All members of Lens are self-pollinating diploids
(2n = 2x = 14; Sharma et al., 1996), although relatively high levels of outcrossing have been reported (Erskine and Muehlbauer, 1991).
The lentil accessions in the ICARDA collection have been obtained from
113 ICARDA collection missions (46%), 56 donor institutions (44%) and
ICARDAs breeding programme (10%). Collection missions contributed
1734 cultivated and 374 wild accessions. However, based on the numbers of
64

Genetic Resources
Wild Lens
Cultivated lentil
Fig. 6.1. Distribution of the ICARDA lentil collection locations.

65
66
B.J. Furman et al.

Table 6.1. Wild Lens accessions maintained at ICARDA.
Number of
Taxonomic name
Lens culinaris ssp. odemensis
L. culinaris ssp. orientalis
L. culinaris ssp. tomentosus
Lens ervoides
Lens lamottei
Lens nigricans
Total
Countries represented
4
14
2
15
3
8
46
Accessions
65
268
11
166
10
63
583
wild Lens accessions available, germplasm from North African countries

such as Algeria, Libya, Sudan and Tunisia appear to be under-represented
in the collection, as does germplasm from the new Central and West Asian
republics of the former Soviet Union (Ferguson and Erskine, 2001). Recent
missions to Central Asia and the Caucasus have substantially decreased
geographic gaps from this region, yielding a total of 122 cultivated and 27
wild Lens accessions (K. Street, ICARDA, Syria, personal communication).
Studies on the distribution of genetic variation (Ferguson et al., 1998) suggest that the overall collection priority for wild species should focus on
south-west Turkey, particularly the provinces of Burdur, Isparta and Afyon.
The other major collections worldwide include those at the Australian
Temperate Field Crops Collection (ATFCC) in the Department of Primary
Industries, Victoria, Australia (http://149.144.200.50:8080/QMWebRoot/
SiteMain.jsp) with 5250 accessions, Pullman United States Department of
Agriculture (USDA) Agricultural Research Service (ARS) with 2797 accessions (http://www.ars.grin.gov/), the N.I. Vavilov All-Russian Research
Institute of Plant Industry (VIR) (http://vir.nw.ru/) with 2396 accessions,
and the National Bureau of Plant Genetic Resources, India with 2212 accessions (Dwivedi et al., 2006). In addition, many countries maintain working
collections associated with their national breeding programmes. Such collections are conserved for short- to medium-term utility, and thus storage
conditions may not be as stringent for maintenance of viability as in the
major collections.
The major collections follow the recommended storage guidelines for
genebanks (FAO/IPGRI, 1994), containing a base collection of over 100
seeds at between 18 and 20C for long-term conservation and an active
collection of at least 1000 seeds at 25C for medium-term storage and distribution. Seeds are dried to 15% relative humidity before storage. Seeds are
stored in moisture-proof containers. Vacuum and heat sealing of plasticlined foil envelopes is one of the common genebank procedures for base
collections, allowing seeds to remain viable for up to 100 years given optimal storage conditions. The active collections are generally stored in plastic
screw-top containers for easier access. The active collections should retain
Genetic Resources
67
high viability for at least 30 years, given high initial viability, low seed
moisture and immediate storage postharvest. Optimization of storage conditions is an inexpensive and efficient means of extending seed longevity,
as well as reducing the annual costs for maintenance of high levels of seed
viability in the active collections.
The base collections are necessary to maintain the genetic integrity of
the seed samples of accessions as originally received, as well as provide
very long-term conservation of genetic diversity. They are also utilized to
replenish seed stocks in the active collections when necessary. Regeneration
of the active collections occurs when seed numbers get low or if there is a
risk of low seed viability. Viability tests are therefore carried out regularly
to determine the need for regeneration. In addition, safety duplicate collections for long-term storage in other locations are maintained. For example,
ICARDA maintains safety duplications in both Mexico and India. A third
collection is being prepared for inclusion in the newly developed Arctic
Genebank at Svalbard, Norway. ICARDA similarly maintains the safety
backup collection for the National Board of Plant Genetic Resources, India.
A decision guide for seed regeneration and precautions to maintain genetic
integrity is outlined by Sackville Hamilton and Chorlton (1997).
The major collections make their seed available upon request. ICARDA
distributes approximately 1000 or more accessions/year, free of charge,
upon agreement to the Standard Material Transfer Agreement. The other
institutes have similar guidelines. These collections are utilized for crop
improvement, basic research and augmentation of national collections.
6.2. Core Collections

ICARDA was responsible for creating a reference core (also referred to as
composite) collection for lentil as part of a large-scale programme of the
Consultative Group on International Agricultural Research (CGIAR) Generation Challenge Program that aims to explore the genetic diversity of the
global germplasm collections held by the CGIAR research centres (http://
www.generationcp.org). The global composite collection of 1000 lentil
accessions was established representing the overall genetic diversity and
agroclimatological range of lentil. This collection was compiled from landraces, wild relatives and elite germplasm and cultivars representing the
overall ICARDA collection in both distribution and type. The methodology
combined classical hierarchical cluster analyses using agronomic traits,
and two-step cluster analyses using agroclimatological data linked to the
geographical coordinates of the accessions collection sites (Furman, 2006).
The hierarchical cluster analysis ensured that the level of variation found
in the larger collection was maintained in the composite collection. In addition, 64 accessions of landraces, released cultivars and breeding materials
for their resistances to a number of stresses affecting lentil production were
included in the composite collection upon recommendation of ICARDA
breeders.
68
B.J. Furman et al.
The USDA ARS lentil collection selected a core of 234 accessions based
on country of origin (Simon and Hannan, 1995). Recently, the core was
extended (384 accessions) to add mapping population parents, cultivars and
wild accessions, and a subset of pure lines was created. This pure line subset will be distributed to scientists interested in linkage disequilibrium mapping in lentil.
6.3. In Situ Conservation

There has been no systematic attempt to conserve Lens diversity in situ
using either on-farm techniques or reserves. However, protected areas
throughout the range of the genus undoubtedly contain Lens species. Conservation of Lens is passive in these protected areas (species presence and
genetic diversity is not being actively managed or monitored) and is thus
susceptible to genetic erosion. More active conservation of Lens diversity is
found in a few managed reserves of the eastern Mediterranean (e.g. Ammiad
in eastern Galilee, Israel; in Turkey at Kaz Dag, Aegean Region; and Amanos, Mersin and Ceylanpinar in the south-east). The latter reserve is particularly important as this area is one of the two centres of genetic diversity
for Lens culinaris ssp. orientalis (Ferguson et al., 1998). A second centre was
identified in southern Syria (Ferguson et al., 1998), and Maxted (1995) proposed the establishment of a genetic reserve for Vicieae species in the region
of Mimas, Djebel Druze in southern Syria. Recently this area was designated
as a genetic reserve by the Syrian Scientific Agricultural Research Commission, as part of their Global Environment Facility funded Conservation and
Sustainable Use of Dryland Agrobiodiversity project (Maxted et al., 2003).
Although recent progress has been made towards in situ conservation in the
areas of highest Lens genetic diversity, there remains an urgent need to systematically establish both reserves for the wild species of Lens and on-farm
projects to conserve the ancient landraces of cultivated Lens species.
6.4. Characterization and Evaluation of Collections

The goals of maintaining germplasm collections include preservation of
genetic diversity of the species as well as providing information necessary
for their utilization. As such there are three major activities of any genebank
(Reed et al., 2004):
1. Maintaining genebank inventory using passport information including
accession identiers, taxonomic classication, country of origin and site
data with referencing by latitude, longitude and altitude, whenever possible, dates of acquisition, collector/donor details, synonyms for accessions
acquired from other genebanks, etc.
2. Obtaining and documenting accession characterization data using standard trait descriptors (IBPGR, 1985).
Genetic Resources
69
3. Evaluation of accessions for key agronomic traits to enhance their potential for utilization.
Maintenance of reliable passport information is essential for accurate record
keeping. It is also important to provide information as to the conditions in
which the accessions would be likely to be adapted. Characterization data
describe discrete classifications for morphological traits such as colours of
seed, stems and flowers, as well as standards for quantitative traits such as
flowering time, plant height, 100-seed weight, etc. Full records are very
important to check for duplication of accessions and for quality assurance
on identity. This can be an issue in genebanks with multiple sources of
accessions, as well as large numbers of accessions and species.
Targeted and more efficient utilization of germplasm by plant breeders/researchers can be achieved if the trait characteristics of accessions are
known. These can include the agronomic, disease reaction, yield and quality data of accessions in a particular study, and over different studies for
each accession. For example, ICARDA published the Lentil Germplasm Catalog (Erskine and Witcombe, 1984) providing a listing of 20 trait evaluations
for 4550 lentil accessions. Since its publication, characterization data have
increased for over 4000 additional accessions.
Working with world lentil germplasm that is conserved at ICARDA has
revealed substantial variability among landraces and wild species for economically important traits. Studies on genetic variability have been conducted since the early 1980s among germplasm of diverse origin. Morphological traits were critically examined in variable climatic conditions for use
in breeding and selection programmes (Sarwar et al., 1982; Sindhu and
Mishra, 1982; Erskine and Witcombe, 1984; Erskine et al., 1985, 1989; Erskine
and Choudhary, 1986; Lakhani et al., 1986; Baidya et al., 1988; Ramgiry et al.,
1989; Sarker et al., 2005). These studies revealed responses of flowering to
temperature and photoperiod (Erskine et al., 1990, 1993, 1994), tolerances to
abiotic stresses (Erskine et al., 1981; Summerfield et al., 1985; Erskine and
Goodrich, 1991; Hamdi et al., 1992; Ibrahim et al., 1993; Silim et al., 1993a, b;
Hamdi and Erskine, 1996; Srivastava et al., 2000; Yau and Erskine, 2000;
Malhotra et al., 2004; Sarker et al., 2004, 2005), genetic variability and sources
of resistance to fungal diseases (Khare et al., 1991; Agarwal et al., 1993;
Bayaa et al., 1994, 1995, 1997; Abou-Zeid et al., 1995; Robertson et al., 1996;
Bayaa and Erskine, 1998; Nasir and Bretag, 1998; Malhotra et al., 2004) and
viruses (Makkouk et al., 2001). A number of these accessions possessed
multiple resistances (Robertson et al., 1996). Such desirable variability in the
crop gene pool allows for genetic enhancement through plant breeding.
Unfortunately several other important traits, such as nitrogen fixation, resistance to pea leaf weevil (Sitona sp.), aphids and broomrape (Orobanche sp.)
are not currently addressable by breeding due to lack of sufficient genetic
variation (Sarker and Erskine, 2006).
Evaluation of wild species for resistance to key stresses, including
winter hardiness, drought tolerance and resistance to lentil vascular wilt
and Ascochyta blight, demonstrated sufficient variability for these traits
70
B.J. Furman et al.
(Bayaa et al., 1994, 1995; Hamdi and Erskine, 1996; Hamdi et al., 1996;
Robertson et al., 1996; Nasir and Bretag, 1998). Variation in agronomic characters was also explored within wild germplasm. Results showed that there
was no striking variation within the wild species for seed and straw yields.
Direct selection of wild lentil germplasm for biomass yield under dry conditions is apparently of little value for transfer to cultivated lentil.
The ICARDA composite collection has been evaluated for key morphological and quantitative traits (B. Furman, California, unpublished). In addition, the composite collection has been characterized with 24 simple
sequence repeat (SSR) markers (e.g. Hamwieh et al., 2005) and diversity
parameters such as polymorphic information contents, allelic diversity, etc.
have been determined (publication in preparation). This should allow for
the extraction of core and reference collections for use by breeders and
geneticists. Large-scale genotyping is also underway on the lentil core collection held by the USDA/ARS in Pullman, Washington, USA using 39
mapped SSRs (C. Coyne, Washington, personal communication). The ARS
studies are for fine mapping of disease resistance quantitative trait loci
(QTL) identification (F. Muehlbauer, Washington, personal communication).
6.5. Utilization of Lentil Germplasm

In general, breeders narrow the genetic diversity of their breeding populations in the process of selecting the required trait combinations for improved
varieties (Maxted et al., 2000). Genebanks aim to conserve the original landrace diversity both for current and for unforeseen future needs in breeding, as well as surveying and enhancement of germplasm diversity for key
traits and identification of novel genes in wild relatives. The provision of an
evaluation/passport database covering the major world collections means
that targeted multiple trait searches of most of the available lentil germplasm can now be carried out more efficiently and effectively. Strategies
such as allele pyramiding in key traits and sourcing of novel genes from
wild relatives should now be easier to realize.
The standard practice of genebanks to document passport data on the
origin, synonyms and collection data, with details of source location and
associated agriculture, enables the traceability of accessions along with
details of collection procedure and evolutionary status. In addition, such
data provide information about likely adaptation characteristics in relation
to the physical environment and the local climatic regime. Unless conservation is linked to utilization there is a risk that genebanks may become museums (Maxted et al., 2000). Knowledge of the growth characteristics and
reactions to abiotic and biotic stresses of accessions is thus exceedingly
important.
Each of the major genebanks has their own documentation system
that can be utilized by curators for efficient maintenance and distribution
of seed. In addition, these data can now be outsourced for multiple traits from
combined databases, such as constructed by ATFCC, using International
Genetic Resources
71
Crop Information Systems (ICIS) platforms for digital search and retrieval
across relational databases over countries and years. An example is the
ATFCC International Lentil Information System (ILIS) evaluation database for lentil germplasm, which combines databases of ICARDA, USDA
and ATFCC, enabling a more comprehensive search of the combined
genetic resources over large collections for multiple trait expressions
(http://149.144.200.50:8080/QMWebRoot/SiteMain.jsp).
As far as possible, the internationally standard descriptors (IBPGR,
1985) are used for trait records in ILIS. Additionally ILIS has options for
converting quantitative trait data to a 19 scale based on arithmetic division
of the range of values (outlying values quality checked and filtered out), or
on statistical normalization of data from each location/year set to a mean of
0 (zero) and a range of +/2 standard deviations for 95% of the variation if
distributed as a standard normal distribution. These conversions eliminate
the environmental component from observed values, as well as the genotype environment interaction, to provide genetic comparisons of landraces
for trait expressions. These data can be combined across locations and years,
and across different genebanks, to enable initially simple searching for
accessions with either high or low trait expressions according to breeding
goals. Selected accessions can then be requested online from genebanks,
and the breeder then has a prospective subset of germplasm for validation
in the respective target environments for farmer clients, and choice of
parents for the breeding programme.
The ILIS database allows online users to run searchquery interrogations for germplasm with multiple trait expressions, currently from the
combined ICARDA, USDA and ATFCC collections. This is a very powerful
search engine to assist genebank clients, to add value to the collections, and
to facilitate utilization for crop improvement. It is becoming more necessary
in the current trend for economic rationalization to demonstrate and justify
the expense for genebank operations, in both the short term with current
crop improvement outcomes, and the long term where a genetic insurance
strategy may be very prescient to cope with a looming climate change.
Close linkage of lentil collections with breeders, pathologists and students, for evaluating key traits is becoming a significant genebank activity.
The ATFCC staff actively work with lentil breeders in germplasm enhancement for evaluation of tolerances to salinity, frost, herbicides and disease
(Redden et al., 2006). Similarly, ICARDA has identified sources of resistance
to rust, vascular wilt and Ascochyta blight in the domestic gene pool, and
found additional sources of resistance to vascular wilt and Ascochyta blight,
and greater cold tolerance in the wild progenitor L. culinaris ssp. orientalis
(Ferguson, 2000).
Two important new developments in molecular genetics are its applicability to genebanks. One is the use of molecular markers for DNA fingerprint
characterization of individual accessions. These tools enable the identity of
accessions between collections to be confirmed, detection of duplicates within
collections, and analyses of genetic relatedness among accessions to define the
structure of genetic diversity within a species/genus. This knowledge assists
72
B.J. Furman et al.
breeders seeking allelic diversity for particular traits. The other application
is the capacity of allele mining for various traits using association genetics,
both within species/wild relative collections, and across species with comparative genomics (Spooner et al., 2005).
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food legume germplasm collections. HortScience 30, 907.
Sindhu, J.S. and Mishra, H.O. (1982) Genetic variability in Indian microsperma type
lentil. LENS Newsletter 9, 1011.
Genetic Resources
75
Spooner, D., van Truenen, R. and de Vicente, M.C. (2005) Molecular Markers for
Genebank Management. International Plant Genetics Resources Institute (IPGRI)
Technical Bulletin No. 10. IPGRI, Rome, Italy.
Zohary, D. and Hopf, M. (1988) Domestication of Plants in the Old World. Clarendon
Press, Oxford, UK.
Genetics of Economic Traits

B. Sharma
Indian Agricultural Research Institute, New Delhi, India
7.1. Introduction
Although lentil is among the most prominent grain legumes globally, its
economic position strengthened significantly when it became important in
international trade about two decades ago. This, coupled with the International Center for Agricultural Research in the Dry Areas (ICARDA) focused
attention on the crop as its world mandate, served as a stimulus for
enhanced research in the area of genetics, breeding and biotechnology. This
chapter deals with the genetics of characters showing discrete, identifiable
and contrasting phenotypes. The inheritance and selection for traits displaying continuous variation is covered by Sarker et al. in Chapter 8, Rahman
et al. in Chapter 9 and Muehlbauer et al. in Chapter 10, and gene mapping,
linkage and the use of marker-assisted selection are covered by Ford et al. in
Chapter 11 (all in this volume).
7.2. Induced Mutagenesis

Well-identified traits with contrasting phenotypes in different genetic backgrounds are a prerequisite to embark upon genetic analysis. Attempts to
identify such genetically analysable traits genetically in lentil germplasm
yielded only about three dozen morphological traits for which dominant
and recessive phenotypes were available (Mishra and Sharma, 2003). Clearly
it is essential to discover more genes by induced mutation. A few hundred
mutations is a reasonably good starting base for linkage studies. Some
induced mutations also have breeding value, while many others may serve
as dependable selectable markers and donors for economic traits. As in
other crops, chlorophyll mutations are easily induced in lentil (Sharma and
Sharma, 1981a; Dixit and Dubey, 1986a) and are recessive to the normal
76

77
green phenotype (Vandenberg and Slinkard, 1987, 1989) and have been
reported by Solanki and Sharma (2005) and Ladizinsky (1985). A wide range
of mutations are reported for plant height, growth habit, branching pattern,
stem structure including fasciation, leaf morphology including a leaf mutant
with complex tendril structure, inflorescence, calyx shape, flower structure,
pod characters, glabrous, waxless, short rachis, acute pod, sterility and
seedcoat colour, curled apex, stunted growth, radicle hypertrophy, bilobed
cotyledon leaves, tricotyly, barren apex, and giant seedlings (Sharma and
Sharma, 1978c, d, e, 1979, 1980a; Dixit and Dubey, 1986a; Tyagi and Gupta,
1991; Tripathi and Dubey, 1992; Ramesh and Tyagi, 1999; Vandana et al.,
1999). The mutation frequency can be increased manifoldly by identifying
M1 plants with maximum mutagenic damage in terms of seedling injury
and sterility (Sarker, 1985; Sharma, 1986; Sarker and Sharma, 1988, 1989;
Solanki, 1991; Solanki and Sharma, 2000).
Mutations in dwarfing genes had a great impact in the history of plant
breeding. Such mutations were reported in lentil by Sharma and Sharma
(1981b). Ladizinsky (1997) isolated two spontaneous dwarf mutations (genes
df1 and df2) from a segregating population. Independently induced dwarf
mutations were given different names by their respective authors, for example
compact and bushy (Sharma and Sharma, 1978a, b). Tirdea and Mancas (1986)
isolated several mutations with high amino acid content. Mutations of a physiological nature (Sharma and Sharma, 1980b) such as early mutations (Sharma
and Sharma, 1982) and male sterility (Sharma and Sharma, 1978f) have a
potential for developing varieties and hybrids with better adaptability.
7.3. Inheritance of Morphological Traits

Growth habit
Ladizinsky (1979) reported incomplete dominance of bushy/erect growth
habit and proposed the gene symbol Gh. The gene analysed in this study
appears to be different from the recessive ert gene discovered and mapped
subsequently (Emami and Sharma, 1999a). The erect phenotype denoted by
ert is recessive to the most prevalent growth habit in lentil in which the
plant remains procumbent for about a month and the branches start growing upright thereafter. This growth habit can be called semi-erect or semispreading. The wild relatives of cultivated lentil have prostrate stems for
much longer. The gene Ert has been linked with three other genes: Rdp, Bl
and Gs (Emami and Sharma, 1999a; Kumar et al., 2004a, 2005a).
Red foliage
Several studies have analysed red pigmentation of different plant parts in
lentil: epicotyl (gene Gs for green stem; Ladizinsky, 1979), leaf (gene Bl for
brown leaf; Emami and Sharma, 1996a) and pods (gene Grp for green pod;
78
B. Sharma
Vandenberg and Slinkard, 1989; and Pdp for pod with violet stripes; Havey
and Muehlbauer, 1989). The gene symbol for anthocyanin development on
immature pods was changed to Rdp (red pod) by Emami (1996) with a view
to avoid ambiguity as green pod (for which gene Grp was first proposed)
could be a contrasting phenotype for a variety of colours of the pod wall,
that were dominant in some cases (e.g. yellow pods of chlorophyll mutations, or green plants producing yellow pods as the recessive gene gp in
pea), and recessive in others (against red pod). The GsRdp linkage was
the first to be reported (Ladizinsky, 1979; Zamir and Ladizinsky, 1984;
Havey and Muehlbauer, 1989; Muehlbauer et al., 1989; Weeden et al., 1992)
and was called Linkage Group 1 of lentil.
Light green foliage

The intensity of leaf colour within the normal range of foliage colour has also
been considered to be a major property distinguishing the macrosperma lentils
from microsperma, the former being yellowish green. The variety Precoz is a
well-known example. Genetic analysis (Kumar, 2002) between normal green
and light green genotypes showed segregation of a single gene with normal
green as dominant identified by gene symbol Gl. It was also found linked
with the genes for plant height, pubescence development, and number of
leaflets per leaf in the order PhGlPubHl (Kumar et al., 2005c).
Leaflet shape, size and number

The shape and size of leaflets are traits of taxonomic significance as the macrosperma lentils differ from microsperma by their leaf size and morphology.
Kumar et al. (2005d) identified two contrasting phenotypes for leaf shape:
oval and acute. The trait was reported to be monogenic, oval being dominant over acute. Gene symbol Ol is proposed. It is linked with the genes for
leaflet size (Blf), stipule size (Lst) and pod size (Lpd), and the Globe mutant
(Kumar et al., 2006, unpublished data).
Crosses between genotypes with broad and narrow leaflets showed
monogenic inheritance with incomplete dominance. This trait was identified by gene symbol Blf (Kumar et al., 2004b) and its linkage with the genes
Ol, Lst, Lpd and Glo established (Kumar et al., 2006, unpublished data). The
phenotype with high number of leaflet pairs (more than six in any leaf on a
plant) is monogenically dominant over the one with fewer leaflets (Kumar
et al., 2005c). It has been given the gene symbol Hl.
Tendril formation
The paripinnate compound leaf of lentil terminates with or without an apical unbranched tendril that varies in size from a spur to 5 cm or more in
79
length in different genotypes. Vandenberg and Slinkard (1989) analysed the

inheritance of the presence or absence of tendrils and showed the tendrilless phenotype to be monogenic recessive, and proposed the gene symbol
Tnl. Distortion of segregation may occur due to misclassification of the F2
plants with rudimentary tendrils. Out of nine crosses attempted by Kumar
(2002) F2 segregation fitted well to a 3:1 ratio in only four crosses. Five
crosses had an excess of tendril-less plants. None had an excess of tendrilled
plants. Crosses showing abnormal segregation had Lens 6163 as the male or
female parent. Obviously, Lens 6163 is a tendril-potent genotype with minute tendrils. As a result, some of the tendrilled F2 plants were included in
the tendril-less class. Monogenic inheritance of tendril formation has been
confirmed (Vaillancourt and Slinkard, 1992; Emami, 1996).
Plant pubescence
Pubescence development has been considered an important attribute in lentil taxonomy and is a major criterion to distinguish microsperma from macrosperma lentils. This trait has been treated differently by workers mainly
because the intensity of pubescence varies across plant parts and developmental stages. Generally, maximum expression of pubescence is seen on the
growing tips at the vegetative stage and on the inflorescence. Vandenberg
and Slinkard (1989) analysed the relationship between pubescent and glabrous pods and assigned the gene symbol Glp. Emami (1996) observed that
pubescence on the peduncle is a more reliable trait and redesignated the
gene as Pdp. Kumar (2002) on a closer look concluded that pubescence, when
present, is noticeable on the entire plant, with a variable degree of expression on different organs and gradually disappearing on aged parts. Thus, a
plant has to be either totally pubescent or glabrous. The gene symbol was
therefore again changed to Pub and later shown to be linked to the gene for
light green foliage (Hoque, 2001; Hoque et al., 2002b). With two additional
morphological markers the linkage sequence was extended to PhGl
PubHl (Kumar et al., 2004a, 2005c).
Plant height
Tallness in plants is almost universally shown to be dominant over dwarfness since Mendels times. It is associated with the synthesis or working
efficiency of gibberellins and other growth hormones. Tallness is dominant over dwarfness in lentil also. The gene Ph for plant height, first
reported by Tahir et al. (1994), belongs to the linkage group comprising
eight morphological (Ph, Gl, Pub, Hl, P, Mot, Brt, Pi) and at least 13 isozyme
markers (cf. Kumar et al., 2005c). In addition to this discrete segregation in
the F2, plant height has been found to be quantitatively inherited (Haddad
et al., 1982).
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B. Sharma
Globe mutant
Gupta et al. (1983) induced a globular-shaped ornamental dwarf mutation
in Lens 830 that is recessive to the wild phenotype. These observations were
confirmed by Emami (1996) and Kumar (2002). Its linkage has been established with the genes for leaflet shape (Ol) and size of leaflets (Blf), stipule
size (Lst) and pod size (Lpd) (Kumar et al., 2006, unpublished data).
Stem fasciation
Stem fasciation as a genetic trait has been known in plants since the times of
G. Mendel. Induced mutations for stem fasciation in lentil were variously
reported by Sharma (1977), Sharma and Sharma (1980a), Tyagi and Gupta
(1991) and Solanki (1991), and was always found to be recessive (Sharma
and Sharma, 1977).
Stipule size
The stipules of lentil vary from very small (4 mm long, 1 mm broad) to very
large (11 mm long, 4 mm broad). Crosses between extreme phenotypes
(Kumar, 2002) revealed that stipule size has Mendelian monogenic inheritance with incomplete dominance. Gene symbol Lst was proposed for
stipule size. Its linkage with the genes Ol, Blf, Lpd and Glo for other morphological traits has been demonstrated (Yogesh et al., unpublished data).
Flowering time
Although flowering time is highly influenced by temperature and photoperiod (Summerfield et al., 1985), genetic control of days to flower is well
established in plant species. These two environmental factors have played a
vital role in the adaptation of lentil in spatially isolated parts of the globe
(Erskine et al., 1989, 1990). A model of photothermal response of flowering
was developed on the basis of field evaluation of the ICARDA world collection (Erskine et al., 1994) and this genetic variation has been successfully used
to select lentil genotypes for the West Asian highlands (Keatinge et al., 1996).
Kant and Sharma (1975) observed that all microsperma lines in a germplasm collection were early (flowering in 6080 days) and the remaining macrosperma genotypes originating from the western hemisphere were always
late, producing flowers after 80 days. Some of these macrosperma genotypes
did not produce flowers under the short-day conditions of Indian winter.
Sarker et al. (1998, 1999) concluded monogenic inheritance of flowering
time with earliness being recessive, for which gene symbol Sn was proposed.
Early flowering transgressive segregation in F2 is a consequence of interaction between recessive sn and minor genes for earliness. It was also concluded
81
that flowering time in lentil is determined by this major gene in combination with a polygenic system.
Earliness was shown to be linked to Scp (the gene for seedcoat pattern
that was replaced by the MotSpt locus; Emami, 1996) and Pep (for pubescent peduncle, now Pub) in Linkage Group 5. The recombination fraction
between Sn and Pep was estimated to be 38 8.7.
Another analysis of flowering time in lentil (Emami, 1996) based on 25
crosses between early, medium and late maturing strains in all possible combinations revealed its quantitative nature with continuous variation and
transgressive segregation in the F2 without maternal effect. Interestingly, the
widest range of transgressive segregation was recorded when early maturing
microsperma varieties developed in India (e.g. PKVL-1, Lens 830, Pant L 406,
Pant L 639, etc.) were crossed with the earliest divergent macrosperma variety
Precoz of Latin American origin, as by Tyagi and Sharma (1989). The parent
varieties flowered after 7072 days, but their hybrids threw transgressants
that flowered as early as 4547 days and as late as 120 days. In the same
experiment, crosses among microsperma varieties did not yield transgressive
segregants flowering earlier than 60 days. Clearly, microsperma and macrosperma lentils have different sets of genes controlling flowering time, and all
microsperma genotypes share the same gene pool. Similar transgressive
segregation can be expected for other economically important traits.
Flower colour
Several flower colours have been identified in lentil, namely violet, pink, blue
and purple. The intensity of flower colour varies depending on genotype, age
of flower, and is subject to temperature and sunlight conditions. Red (or pink)
flowers were never observed in a large germplasm collection maintained at
the Division of Genetics, Indian Agricultural Research Institute, Delhi nor in
the ICARDA lentil collection (A. Sarker, personal communication). Probably, the terms violet (Lal and Srivastava, 1975), blue (Ladizinsky, 1979) and
purple (Kumar, 2002) have been used synonymously. Accordingly, different gene symbols have been assigned, for example V for violet and P for
pink (Lal and Srivastava, 1975), W for white (Wilson and Hudson, 1978),
and P for purple (Kumar, 2002). The possibility of multiple alleles to explain
some of the intermediate colours cannot be ruled out. Lal and Srivastava
(1975) and Gill and Malhotra (1980) reported complete monogenic dominance of violet flower over white flower. Kumar et al. (2005b) confirmed
monogenic complete dominance of purple flower over white and reported
its linkage with mottled seed pattern and brown ground colour of testa in
the sequence PMotBrt. This is in agreement with Hoque et al. (2002a).
Number of flowers per peduncle
The number of flowers borne on a peduncle in lentil may have an impact on
its productivity potential. Its genetic analysis is complicated because flower
82
B. Sharma
number per peduncle is inconsistent in expression declining with plant age,

and is highly influenced by the environment. This may cause misclassification of F2 plants. Nevertheless, considering its breeding value Gill and Malhotra (1980) concluded that flower number is a monogenic trait with the
two-flower phenotype dominant over three-flowered. Subsequent studies
of Emami (1996) and Kumar (2002) produced contrary results. The latter
author used the prolificacy potential to classify plants. A plant was treated
as three-flowered or four-flowered if it produced that many flowers even
on a single peduncle. Using this procedure both these studies concluded
that higher flower number per peduncle is dominant.
Pod size
The size of lentil pods varies in a wide range and is often but not completely
associated with seed size. The inheritance of pod size was studied by Kumar
et al. (2005e) who found monogenic inheritance with incomplete dominance
and no maternal effects. This trait was assigned gene symbol Lpd which is a
member of the linkage group LstOlBlfLpdGlo (Kumar et al., 2006,
unpublished data).
Pod dehiscence
Pod dehiscence is a trait of great economic value as it sometimes causes significant losses before or during harvest. Pod dehiscence is a survival trait in
wild species such as Lens orientalis. Among the cultivated lentils also, the
oriental microsperma lentils have a higher degree of dehiscence than their
occidental macrosperma counterparts.
Ladizinsky (1979, 1985) found the dehiscent phenotype to be completely
dominant over indehiscence and assigned gene symbol Pi. The inheritance
of the indehiscence trait was confirmed by Vaillancourt and Slinkard (1992).
Among cultivated lentils, a response to selection for pod indehiscence indicates the presence of quantitative variation for the trait (Erskine, 1985). The
Pi gene was shown to be linked to isozyme locus Gal-1 (Havey and Muehlbauer, 1989) in what was called Linkage Group 1 in the following order:
Pgm-cFkPgd-pMe-2Aat-cGal-1PiGh/Ert (Tahir and Muehlbauer, 1994). Tahir et al. (1993), however, did not show close linkage of Pi to
Gh/Ert and placed it in a different linkage segment and also changed the
numbering of linkage groups. Havey and Muehlbauer (1989) also placed
the Pi and Gh/Ert genes in different linkage segments.
Seedcoat colour
Seedcoat colours have been given several names in the literature, for example black, grey, brown, beige, yellowish grey, tan and green. As observed
83
by Emami (1996), the lack of precision in naming testa colours may be partly
due to the influence of cotyledon pigments on the colour of seedcoat. He
noticed that the pigments of orange, pinkish orange (called brown by Emami
and Sharma, 1996b, c) and green cotyledons are absorbed in the testa tissue
distorting its true colour. Seeds with colourless testa and orange cotyledons
appear pinkish and those with green cotyledons appear green. However,
the seedcoat may have green colour of its own a hypothesis that needs to
be confirmed experimentally. The pigment of yellow cotyledons does not
appear to have such a migratory effect. Therefore it would be desirable to
use only yellow-cotyledon genotypes for genetic analysis of testa colour
where genotypes with different seedcoat colours and patterns are used as
parents. The black testa is obviously dominant or hyperstatic over all other
colours. Care must be taken, especially in the case of black seed colour, to
distinguish black seeds caused by ground colour and due to dense black
spotting. It appears that the so-called tan colour is the colourless situation
although Vandenberg and Slinkard (1990) concluded that tan testa is produced by a dominant gene (Tgc) that in combination with another gene, Ggc
for grey seedcoat, gives rise to brown testa. Recessive homozygotes for both
genes (ggcggctgctgc) produce seeds with green testa. According to this
hypothesis, if the three major genes for testa colour (Blt, Ggc and Tgc) are in
dominant homozygous (also heterozygous) condition, the seeds will appear
black. The migration of green pigment from cotyledons into seedcoat was
never considered in these studies. Absorption of orange, brown (pinkish
yellow) and green pigments by seedcoat from cotyledons can modify its
colour phenotype markedly.
The lentil seedcoat has three (or four) basic colours, that is black (if it is
really different from black spotting), brown, grey and green. Other colours
are a result of either epistatic effects or colour distortions. The green colour
of testa, if such a trait exists, could give other hues in combination with
brown and grey ground colours.
Black seedcoat
With proper care, it is possible to identify genotypes that are black seeded
because of ground colour and not due to black spotting as strains with black
spotting on non-black ground colour are available. The black testa colour
has an interesting mode of inheritance (Emami and Sharma, 2000; Sharma
et al., 2004a). Vandenberg (1987) found one dominant gene with codominant expression for black seedcoat in one cross and two recessive genes for
the same phenotype (i.e. black testa recessive) in two other crosses (Vandenberg and Slinkard, 1990). It is difficult to understand the absence of
black pigment being dominant over its presence. All other studies (Vaillancourt and Slinkard, 1992; Emami, 1996; Emami and Sharma, 2000; Sharma
et al., 2004a) confirmed monogenic inheritance of black testa. These
workers also observed that the F1 plants produce a mixture of black and
non-black (brown, grey, green or colourless) seeds. The numbers of black
and non-black seeds born on the F1 plants do not make a genetic ratio.
84
B. Sharma
Vandenberg (1987) described this phenomenon as codominant expression

of dominant and recessive alleles for black testa. It must be remembered
that codominance is simultaneous expression of the two alleles (phenotypes) of a gene in each cell of the same organism. Genetic analysis shows
that the F2 plants give a perfect ratio of 3 black + mixture : 1 true breeding
non-black. Emami and Sharma (2000) called it a case of dosage effect,
incomplete dominance or lack of penetrance/expressivity and concluded
that the heterozygous plants produce seeds with black and non-black
testa as a result of the quantity of black pigment accumulated in the testa
in the course of development. Apparently, the seeds that develop on the
upper nodes during late reproductive phase run short of time to
accumulate black pigment in sufficient quantity with a single dose of
dominant Blt gene before the plant completes its life cycle. This conclusion is supported by the fact that the dominant homozygotes with two
doses (Blt Blt) produce seeds with black testa only. Gene symbol Blt was
proposed for black seedcoat with new interpretation of its manifestation
and inheritance.
Vandenberg (1987) and Vandenberg and Slinkard (1990) concluded that
black testa colour is controlled by two recessive genes (blsc1 and blsc2) in
one cross but it was inherited monogenically in another. However, Vaillancourt and Slinkard (1992) subsequently demonstrated monogenic inheritance of black testa colour and also concluded that Vandenbergs hypothesis
about two-gene control of black testa trait was caused by an error in scoring
of the F2 phenotypes. Emamis (1996) results were in complete agreement
with those of Vaillancourt and Slinkard (1992). In view of the uncertainties
caused by the Vandenbergs report, Emami and Sharma (2000) considered
his gene symbol Blsc to be invalid and proposed a new gene symbol, Blt, for
black testa with monogenic control.
Brown, grey and green seedcoat
The inheritance of brown seedcoat in lentil was investigated by Ladizinsky
(1979), Vandenberg (1987), Vandenberg and Slinkard (1990), Singh and
Singh (1993), Emami (1996) and Kumar (2002). Ladizinsky (1979) reported
brown testa to be dominant over yellowish grey. In all these studies brown
testa displayed monogenic dominance over grey and tan. Vandenberg and
Slinkard (1990) further demonstrated digenic dominance of brown testa
over green. Both grey and tan were shown to be dominant over green. Consequently, a two-gene hypothesis was proposed: gene Ggc for grey and Tgc
for tan testa. Thus, double dominant GgcTgc genotypes produce brown
seeds while seeds with green testa will be born on recessive homozygous
ggctgc plants. This hypothesis has not been challenged in other studies even
though the authors did not examine the possibility of pigments migrating
from cotyledons into the testa of developing seeds. Kumar et al. (2005b)
however proposed the gene symbol Brt for brown testa, which is now
included in the short gene map PMotBrt (Hoque et al., 2002a; Kumar
et al., 2005b).
85
Tannin in seedcoat
Presence of tannin in lentil seedcoat can be detected only if all other pigments are absent and the seed appears colourless. Vaillancourt et al. (1986)
analysed the inheritance of tannin in the seedcoat. Presence of tannin was
monogenically dominant over its absence, for which gene symbol Tan was
proposed. Presence of tannins (proanthocyanidins) in the seedcoat leads to
its discoloration (browning) during storage (Bezeda, 1980). Although lentil
strains with black testa are generally high in tannins, a strain (PI 211602)
with the highest tannin content of 2.29% of its seedcoat among the germplasm of 87 lines screened did not have black testa.
Seedcoat spotting
Ladizinsky (1979), Vandenberg and Slinkard (1990) and Emami (1996)
observed the presence of spotting on the testa to be dominant over the spotless phenotype. Seedcoat pattern is designated by a generalized gene symbol Scp. Vandenberg and Slinkard (1990) identified two types of spotting,
that is spotting (gene Scps) and dotted (Scpd), and concluded that they are
multiple alleles at the Scp locus. This however was untenable as they also
had true breeding strains with a mixture of both spotted and dotted patterns, which was called the flex phenotype. A closer look on the lentil genotypes in a germplasm collection by Emami (1996) led him to confirm the
two types of seedcoat pattern in lentil, however, both behaving as independent traits, each dominant over the spotless phenotype and giving perfect
monogenic segregation in the F2. One pattern consisted of minute round
spots distributed uniformly over the seedcoat, and the other comprised
patches of irregular shape sparsely spread over the seed. They were named
as mottling and speckling, respectively, and designated by gene symbols
Mot and Spt. Attempts to recombine the Mot and Spt genes indicated tight
linkage with Scp (MotSpt) locus. The Mot gene (synonymous with Scpd) was
placed between the P (purple flower) and Brt (brown testa) genes (see above).
Seed size
Even though the cultivated lentils are divided into microsperma and macrosperma primarily on the basis of differences in seed size, there is continuous
distribution for the trait. Taking seed size as a criterion, typical Indian lentils
(microsperma) form a group of genotypes with 1000-grain weight between
1235 g. Typically, macrosperma lentils generally weigh above 35 g/1000 seeds.
Ladizinsky (1979) crossed L. orientalis and a cultivar of Lens culinaris (both
with seed diameters between 4 and 5 mm) with a cultivated strain having
8 mm seed size (a macrosperma genotype) and observed intermediate seed
size in the F1 with continuous variation in the next generation. Further, none
of the segregants reached the level of the large-seeded parent.
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B. Sharma
Seed size is an economic trait of great significance and deserves the

attention of geneticists and breeders accordingly. It is a special attribute in
lentil consumption and trade. The wholesale price of small- and large-seeded
lentils differs by a large margin depending on the consumer preference and
farmers choice. An important study of lentil seed mass was made by Abbo
et al. (1991) using quantitative trait loci (QTL) analysis to analyse the continuous variation expressed in segregating populations. They showed that
seed weight was under polygenic control with partial dominance for low
seed weight alleles.
Hard seededness
The inheritance of hard seedcoat with poor absorption of water by the testa
appears to be complicated. Ladizinsky (1985) reported hard seedcoat to be a
monogenically inherited trait and designated it with gene symbol Hsc. Vaillancourt and Slinkard (1992) also reported monogenic inheritance but of an
opposite nature. Hard seedcoat was dominant over soft seed in a cross with
Lens ervoides (Ladizinsky, 1985) and in another cross between cultivated lentils (Vaillancourt and Slinkard, 1992). However, in crosses with L. culinaris
ssp. orientalis hard seedcoat was recessive. It is a unique situation that even
among wild relatives hard seed is reported to be dominant in one species
(L. ervoides) and recessive in the other (L. orientalis). According to Ladizinsky
(1985), pod dehiscence (Pi) in L. orientalis is controlled by two complementary
genes, one of which is linked to Hsc for hard seedcoat at 18 map units.
Vaillancourt and Slinkard (1992) also observed that in some crosses
seeds imbibed water but remained dormant for 23 weeks. Thus germination was influenced by the genetically caused hard coat as well as physiological dormancy. Vaillancourt and Slinkard (1992) recorded continuous
variation for germination in such crosses.
The genes for pod indehiscence (pi) and soft seeds that germinate without dormancy (Hsc/hsc) must have played an important role in domestication of lentil as they should have been subject to intensive human selection
in the early history of lentil cultivation.
Cotyledon colour
Genetics of cotyledon colour in lentil presents a fascinating picture. Cotyledon colour has attracted the attention of geneticists and lentil breeders
owing to its market value. The microsperma lentils are predominantly orange
(hence they are called red lentils) and macrosperma lentils usually produce
yellow cotyledons (they are called green lentils in trade jargon because varieties with light-green seedcoats are most preferred and traded).
As cotyledons are sporophytic tissue, their colour can be visualized in
the freshly harvested seeds, and segregation for colour can be recorded in
the seeds harvested from F1 plants. Emami (1996) constructed a simple
87
device to see cotyledon colour in intact seeds (Emami and Sharma, 1996b)
without scratching the seedcoat (as done by Slinkard (1978) and Sinha et al.
(1987)) and then improved the device into a compact seed scanner (Sharma
et al., 2005). The analysis of cotyledon colour has involved crossing genotypes with red and yellow cotyledons (Tschermak-Seysenegg, 1928; Wilson
et al., 1970; Singh, 1978; Slinkard, 1978; Sinha et al., 1987; Emami, 1996;
Kumar, 2002) with orange found to be dominant over yellow. Slinkard
(1978) designated the gene as Yc.
However, in addition to the well-known bright-yellow cotyledon, there
is another yellow phenotype with a pinkish or brownish tinge, which was
called brown so as to distinguish the two variants of yellow cotyledons.
The two yellow phenotypes are distinguishable, preferably against visible
light or under ultraviolet fluorescence (a technique proposed by Wilson
et al. (1970) and Papp (1980) and personal communication). Genetic analysis
(Emami and Sharma, 1996b, c) revealed the independent nature of the two
yellow colours and named their controlling genes as Y and B, respectively.
The double dominant YB leads to orange cotyledons, while double recessive plants (yybb) fail to develop pigment in their cotyledons and remain
green at maturity. The inhibitory gene hypothesis of Slinkard (1978) suggesting involvement of gene i-Yc has not been confirmed. Nevertheless,
Vaillancourt and Slinkard (1993) subsequently demonstrated linkage
between the putative inhibitory gene i-Yc for cotyledon colour and isozyme
markers Aco I, grp and Aat-p. Thus, existence of two genes associated with
yellow cotyledon colour is confirmed. The i-Yc gene proposed by Slinkard
(1978) could be the B gene of Emami and Sharma (1996b, c).
Another gene was discovered in this study (Emami, 1996; Emami and
Sharma, 1999b; Sharma and Emami, 2002), which in the recessive state gives
rise to dark-green cotyledons (much darker than the yybb cotyledons). It was
named as Dg. Sharma and Emami (2002) concluded that while genes Y and B
are individually involved in the synthesis of the yellow and brownish components of the orange complex, respectively, gene Dg acts on their common
precursor at an earlier stage in the synthesis of cotyledon pigments and,
when mutated, blocks its synthesis. Consequently, both the yellow and the
brown pigments are eliminated simultaneously and the cotyledons acquire
deep-green colour. The dgdg (dark-green) situation should be functionally
equivalent to yybb homozygotes which, however, produce light-green cotyledons. This also suggests that Dg is more effective in its function and the two
other genes are leaky in the recessive condition. Thus, Dg has a recessive
epistatic effect on the other two pigment genes and the three genes (Y, B, Dg),
in different combinations, give rise to five F2 phenotypes in the ratio of 27
orange:9 yellow:9 brown:3 light green:16 dark green (Sharma et al., 2004b).
Protein content
Lentil is the richest source of dietary proteins among plants that are consumed without industrial processing. Attempts to analyse the inheritance
88
B. Sharma
of seed protein in lentil (Hamdi et al., 1991; Chauhan and Singh, 1995; Tyagi
and Sharma, 1995) revealed the quantitative nature of this trait. This has
been the pattern in almost all crops. Protein content on a dry weight basis
varied from 20.4 to 28.3% in a sample of 32 germplasm lines and cultivars
(Tyagi and Sharma, 1995). Hamdi et al. (1991) analysed 3663 germplasm
accessions from all lentil-growing regions of the world and recorded a range
of 34.086.0 g/1000-grain weight and 25.729.8% seed protein. Seed protein
content had negligible correlation with grain yield and seed size (r = 0.1
and 0.17, respectively). Protein content is almost universally (especially in
cereals; Watson et al., 1965) negatively correlated with seed size. This principle does not seem to apply in the case of lentil. In fact, Tyagi and Sharma
(1995) observed that protein content in lentil has a positive, although statistically non-significant (r = 0.26), correlation with seed size.
Abiotic stresses
Frost injury
Cold tolerance is an essential requirement when lentils are cultivated during winter or at high altitudes. In recent years, winter hardiness has acquired
special significance in West Asia as winter planting of lentil has become
increasingly popular due to ICARDAs efforts (Sarker et al., 2002; Emami
and Sharma, 2005). Sources of cold resistance have been identified among
germplasm (Erskine et al., 1981; Spaeth and Muehlbauer, 1991), including
wild species (Hamdi et al., 1996).
Eujayl et al. (1999) reported monogenic inheritance of radiation-frost
tolerance and assigned gene symbol Frt. This gene was also tagged with
random amplified polymorphic DNA (RAPD) marker OPS-16750 at 9.1 cM.
Kahraman et al. (2004a, b) concluded that winter hardiness in lentil is a polygenic trait and several QTLs together accounted for 42% of the variation in
their recombinant inbred line populations. Further study of the inheritance
of winter hardiness and discovery of additional QTLs may account for
additional variation and improve prospects for marker-assisted selection.
Drought tolerance
The water requirement of lentil is low. In fact, excessive water supply is
damaging to the crop. This explains the farmers choice of lentil in droughtprone arid regions. Wild relatives of lentil may serve as a good resource to
reduce its water requirement further (Hamdi and Erskine, 1997). A reliable
screening technique for drought tolerance in cowpea in the greenhouse
described by Singh et al. (1999) may be used in lentil as well.
Soil factors
Genetic variation has been found in lentil for response to salinity, nutrient
deficiency and toxicity but no genetic analysis has yet been reported. This
89
variability is covered by Yadav et al. in Chapter 13 (this volume). Briefly

genetic variation in salinity tolerance was reported by Jana and Slinkard
(1979), Ashraf and Waheed (1990, 1993) and Ashraf and Zafar (1997). Variations in response to iron and boron deficiency and boron toxicity have been
found (Erskine et al., 1993; Srivastava et al., 2000; Yau and Erskine, 2000;
Sharma and Sarker, 2008).
Disease resistance
Rust resistance
Rust requires humid conditions for pathogen growth. It is the most important disease in India economically (Singh and Sandhu, 1988) with reported
losses of up to 70% of the lentil crop as a result of rust (Erskine and Sarker,
1997; Negussie et al., 2005).
Precoz, a germplasm accession received from ICARDA in the early
1980s, was found to be universally rust resistant in multilocation trials in
India (B. Sharma, unpublished data, reported in the Workshop Proceedings
of All-India Coordinated Project on Pulses, Kanpur). Most studies on the
inheritance of lentil rust resistance have reported monogenic control, resistance being dominant over susceptibility (Sinha and Yadav, 1989; Singh and
Singh, 1992; Chauhan et al., 1996; Kumar et al., 1997a, b). However, besides
reports of incomplete resistance, duplicate dominant genes controlling resistance have also appeared frequently (Singh and Singh, 1990; Lal et al., 1996;
Kumar et al., 1997b, 2001; Chahota et al., 2002; Mishra, 2006). Only one unconfirmed report suggested rust resistance as a recessive trait in the variety DPL
21. Mishra (2006) also confirmed that the dominant gene for resistance in the
variety Precoz (macrosperma from Latin America) is different from that in
Pant L 4 which is of Indian origin (microsperma). Thus two separate genes
controlling rust resistance have evolved in spatially (may be also temporally)
isolated groups of lentil. The gene symbols proposed are Urf1 in Precoz,
Urf2 in Pant L 4, E 153 and Lens 4147, and the unconfirmed urf3.
Two varieties (Pant L 639 and Pant L 408) developed at Pantnagar
(India) were widely rust resistant when they were commercialized. More
than a decade after their release for cultivation, Chahota et al. (2002) reported
Pant L 639 to be susceptible at Palampur (425 m above mean sea level,
1200 mm rainfall) and Jaacch (428 m above msl, 1500 mm rainfall) in
Himachal Pradesh, where Precoz was still resistant. The inoculum was
multiplied from a single rust colony. It needs to be confirmed whether it is
a case of resistance breakdown or race-specific reaction of the variety Pant L
639. Precoz was crossed with two susceptible varieties (i.e. Pant L 639 and
Lens 259) and with both varieties F2 segregation was in the ratio 15
resistant:1 susceptible at three locations. Lal et al. (1996) had earlier reported
two dominant genes for rust resistance in Precoz at Palampur, when it was
crossed with six susceptible genotypes. This is unequivocal proof that Precoz carries two dominant genes (and not one) for rust resistance.
90
B. Sharma
Wilt resistance
Occurrence of wilt in lentil is frequently associated with moisture stress and
the intensity of disease depends on inoculum saturation in the soil. Therefore it is imperative to have an accurate screening technique for proper
germplasm evaluation and genetic analysis. A highly efficient method of
field screening has been developed at ICARDA where a wilt-sick plot of
large size ensures complete mortality of susceptible genotypes (Bayaa et al.,
1995; Sarker et al., 2004).
Continuous efforts have been made to breed wilt-resistant lentils (Izuqierdo and Morse, 1975). Repeated screening of 500 accessions from the core
collection of lentil germplasm at ICARDA for 3 years yielded 34 strains that
were confirmed for wilt resistance (Sarker et al., 2004). Rigorous screening
through F5F8 led to the identification of 753 resistant lines. Interestingly,
71.6% of wilt-resistant selections were small seeded, whereas only 40.9% of
large-seeded selections were resistant to wilt, suggesting that the gene(s)
for small seed are loosely associated with wilt-resistance genes.
On the basis of a test of allelism, Kamboj et al. (1990) concluded that
wilt resistance in lentil is conferred by five dominant genes, two of which
were duplicate in the variety Pant L 234 and two others complementary in
JL 446 and LP 286. Abbas (1995), however, found only one dominant gene
for wilt resistance in crosses at ICARDA. Eujayl et al. (1998b) also recorded
monogenic inheritance for wilt resistance and designated the gene as Fw. It
was tagged with RAPD marker OPK-15900 with 10.8 cM map distance between
them. This study also established linkage of Fw with the RAPD markers
OP-B17800 and OP-D15500 in coupling phase and OP-C04650 in repulsion
phase. Hamwieh et al. (2005) identified one SSR and another AFLP marker
that linked with Fw at distances of 8.0 and 3.5 cM, respectively.
Ascochyta blight resistance
Ascochyta blight has worldwide occurrence and causes severe damage to
lentil yield and grain quality (Muehlbauer and Chen, 2007). In India it is
mainly confined to the mountainous regions where lentil is cultivated during warm summers with high humidity. Tivoli et al. (2006) listed the sources
of resistance to foliar diseases of lentil, including blight. Searches for sources
of blight resistance have been made at many centres (Ahmed and Morrall,
1996; Ahmed et al., 1996). The germplasm accessions PI 339283 from Canada
(Andrahennadi et al., 1996) and ILL 358 in Australia (Nasir and Bretag, 1996)
have been reported to be the top-ranking blight-resistant genotypes.
Blight resistance was shown to be monogenic recessive in the variety
Laird, and two strains from the ICARDA germplasm collection (ILL 538
and ILL 5684) by Tay and Slinkard (1989). Ahmad et al. (1997) reported two
complementary dominant genes for blight resistance in a cross between L.
ervoides and Lens odemensis. However, only one dominant gene was found
in this study when genotypes of L. culinaris were crossed with each other.
The existence of two complementary dominant genes within the cultivated
lentils was established by Nguen et al. (2001). Ford et al. (1999) mapped two
91
RAPD markers, RV01 and RB18, in flanking positions of the resistance gene
Ral1 (Abr1) 8.0 and 3.5 cM away. Two more RAPD markers (UBC2271290
and OPD-10870) were located in flanking positions of the recessive gene for
resistance, ral2, at 6.4 cM and 10.5 cM distances, respectively, in the variety
Indianhead (Chowdhury et al., 2001).
Anthracnose resistance
Resistance to anthracnose in lentil has been reported to be under the control
of one recessive gene, lct-1, in cultivar Indianhead, and two dominant
genes, designated as LCt-2 (in PI 320937) and LCt-3 (in PI 345629) (Buchwaldt et al., 2003). Two races of the causal organism, Colletotricum truncatum, were isolated and named as Ct1 and Ct0. Anthracnose-resistant
germplasm accessions and the Canadian cultivars CDC Robin, CDC Redberry and CDC Viceroy were resistant to race Ct1 but susceptible to Ct0.
In fact, no genotype resistant to race Ct0 was identified. However, Tullu
et al. (2005) observed that species from the secondary gene pool of Lens (several accessions of L. ervoides, Lens lamottei and Lens nigricans) showed high
to medium levels of resistance to the anthracnose race Ct0. Tullu et al. (2003)
tagged the LCt-2 locus with RAPD markers OPE061250 and UBC704700 in
flanking positions at 6.4 cM and 10.5 cM, respectively.
Stemphylium blight and powdery mildew
These are other locally important diseases of lentil, for which sources of
resistance are available (B. Sharma, Delhi, India, unpublished results). However, their inheritance patterns have not been subjected to genetic analysis.
Virus resistance
Pea seed-borne mosaic virus (PSbMV) is the only prominent disease in lentil, which is transmitted through seed as well as by aphids (Hampton and
Muehlbauer, 1977). Four germplasm strains immune to PSbMV were identified by Haddad et al. (1978): PI 212610, 251786, 297745 and 368648. Their
crosses with the susceptible varieties Tekoa and Precoz segregated into 3
susceptible:1 immune ratio in the F2. Monogenic inheritance and the recessive nature of viral immunity were confirmed in all four crosses. The gene
symbol sbv was proposed to denote the PSbMV resistance in lentil.
Allozyme variation
Allozyme variations had been the markers of choice for mapping in lentil
(Zamir and Ladizinsky, 1984; Tadmor et al., 1987; Havey and Muehlbauer,
1989; Muehlbauer et al., 1989; Vaillancourt and Slinkard, 1992, 1993; Tahir
et al., 1993) before DNA markers became available (Eujayl et al., 1997, 1998a;
Rubeena et al., 2003). Short linkage sequences were reported on the basis of
linkages among allozymes used in different crosses (Gulati et al., 2002).
92
B. Sharma
Cytogenetic studies
Knowledge of cytological structure helps in planning breeding strategies,
particularly in crosses involving allopolyploid species and wild relatives. The
possibility of successful hybridization between different Lens species and
subspecies has been demonstrated by several studies. Chromosomal behaviour in distant hybridization will be a matter of concern in lentil breeding.
Several attempts have been made to define the standard karyotype of
lentil but frequently reported contradictory results. The total chromosome
length of 39.31 m was reported in the variety Pant L 639 by Gupta and
Singh (1981) and 31.77 m in Type 36 by Dixit and Dubey (1986b). The total
chromatin length varied from 28.2 to 72.3 m in different microsperma cultivars of L. culinaris ssp. culinaris (Dixit and Dubey, 1986b). Sindhu et al. (1983)
reported that the total chromosome length varies from 21.47 m in L.
ervoides to 30.4 m in L. orientalis. The pooled length of L. culinaris chromosomes was 27.43 m. The lowest chromatin length of 16.9 m was reported
by Lavania and Lavania (1983). It has also been demonstrated that L. orientalis and L. culinaris contain higher amounts of DNA in their genomes than
L. ervoides. This observation corresponds with their chromatin length.
The chromosome structure has also been shown to vary in different
species and among genotypes of the same species. According to Sindhu
et al. (1983), the chromosome complement in the somatic tissues of L. culinaris, L. orientalis and L. nigricans includes three sub-metacentric (one of
them with a secondary constriction), one metacentric and three acrocentric
chromosome pairs. This can be taken as the most widespread karyotype of
lentil. However, there are several variations. Lens ervoides has four submetacentric chromosomes (one of which is satellited) and three acrocentric
chromosomes. The variety Pant L 639 carries four metacentric, three submetacentric and no acrocentric chromosomes (Gupta and Singh, 1981) while
L. ervoides has three acrocentric and four sub-metracentric chromosomes
(Sindhu et al., 1983). Metacentric chromosomes are absent in L. ervoides. At
the other extreme was variety Lens 3847 in which all the seven chromosomes are sub-metacentric (Lavania and Lavania, 1983). All three types of
chromosomes (three acrocentric, one metacentric, three sub-metacentric) are
present in L. culinaris (Sindhu et al., 1983).
The secondary constriction is located on the fifth chromosome of L.
ervoides and on the fourth chromosome of other species (Gupta and Singh,
1981). However, inconsistent reports have appeared about the number of
satellited chromosomes in cultivated lentil. One satellited chromosome was
reported by Bhattacharjee (1951), Sharma and Mukhopadhyaya (1962) and
Sinha and Acharya (1974) but no satellited chromosome was found in germplasm accessions carrying EC (i.e. exotic collection) numbers (Sinha and
Acharya, 1974), and the varieties Pant L 639 (Gupta and Singh, 1981) and
Type 36 (Dixit and Dubey, 1986b) of Indian origin.
In conclusion, it seems difficult to define a standard karyotype for Lens
species, subspecies and further lower taxa. There could be karyotypic
groups, which need to be identified.
93
Two cytological aberrations in the form of chromosomal breaks have

been identified and the break points were successfully used as markers in
gene mapping (Tahir et al., 1993). Chromosomal interchanges have been
reported in lentil and mapped (Tadmor et al., 1987; Gupta et al., 1999). Gupta
et al. (1985) identified a spontaneous trisomic in the progeny of an interchange heterozygote. A triploid plant of lentil was described by Gupta et al.
(1984). Tetraploids were created experimentally by Gupta and Singh (1982).
Chromosomal eliminations and translocations lead to distorted segregation ratios in crosses of ssp. odemensis with other subspecies (Vaillancourt
and Slinkard, 1992). Goshen et al. (1982) identified three reciprocal translocations in ssp. culinaris ssp. odemensis crosses. The progenies of intersubspecific hybrids frequently have more abnormal segregation than intraspecific
crosses.
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Genetic Enhancement for Yield and

Yield Stability
A. Sarker,1 A. Aydogan,2 S. Chandra,3 M. Kharrat4 and S. Sabaghpour5

1International
Center for Agriculture Research in the Dry Areas (ICARDA), Aleppo,

Syria; 2Central Research Institute for Field Crops (CRIFC), Ankara, Turkey;
3Indian Institute of Pulses Research (IIPR), Kanpur, Uttar Pradesh, India; 4National
Institute for Agronomic Research (INRAT), Tunis, Tunisia; 5Dryland Agricultural
Research Institute, Kermenshah, Iran
8.1. Introduction
Lentil (Lens culinaris Medikus subsp. culinaris) is one of the ancient crops of
world agriculture as evident from its occurrence in archaeological excavations. Cultivated lentil originated from L. culinaris Medikus subsp. orientalis
in the Near East arc and Asia Minor, and considerable diversity in forms
and morphological variants are available in these areas (Zohary, 1972; Zohary
and Hopf, 1973; Williams et al., 1974; Ladizinsky, 1979). It is a self-pollinated,
annual diploid (2n = 2x = 14) species, is a short, slender annual legume that
was among early-domesticates in the Fertile Crescent of the Near East. Since
domestication, lentil has become an important staple pulse crop traditionally grown in West Asia, the Indian subcontinent, East and North Africa,
and to a lesser extent, in Central Asia, the Caucasus and southern Europe.
In the recent past, lentil is also cultivated in South and North America and
in Oceania. It plays an important role in human and animal nutrition and in
soil health improvement. Its cultivation enriches soil nutrient status by adding nitrogen, carbon and organic matter, which promotes sustainable crop
production systems. Lentil is one of the important pulses for crop intensification in West and Central Asia, and diversification in South Asia (Sarker
et al., 2004b).
Lentil seed is a rich source of protein, minerals (K, P, Fe, Zn) and vitamins for human nutrition (Bhatty, 1988; Savage, 1988), and the straw is a
valued animal feed (Erskine et al., 1990a). Furthermore, because of its high
lysine and tryptophan content, its consumption with wheat/rice provides a
balance of essential amino acids for human nutrition. Lentil is predominantly eaten in the Indian subcontinent as boiled or fried dal. It has a consistency like soup and is usually eaten with unleavened bread (roti). Boiled
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Genetic Enhancement for Yield and Yield Stability
103
rice is also served as a staple with lentil dal. Khichri is made from a mixture
of splitted/dehulled lentil and cracked wheat or rice. In West Asia and
North Africa, Mujaddarah, made of whole lentil and immature wheat seed,
is a popular dish. Of course, lentil soup is popular all over. Also, lentil may
be deep-fried and eaten as snack, or combined with cereal flour in the preparation of such foods as bread and cake. Lentil, like other pulse crops, provides nutritional security to low-income consumers in many developing
countries. Because of its multifaceted importance, various national programmes and the International Center for Agricultural Research in the Dry
Areas (ICARDA) are engaged in research and development to improve
yield through improved agronomic practices and genetic enhancement.
World lentil production has tripled from 1.1 million t in 1971 to 3.84
million t in 2006, and this has been accompanied by increased yields from
611 to 898 kg/ha, respectively (FAO, 2007). Three top-ranking countries,
namely, India, Canada and Turkey have increased their productivity and
production. Although the area planted to lentil in Turkey has declined in
recent years, lentil area has increased greatly in India, Canada, Australia
and Ethiopia. However, area expansion and productivity has increased in
the developed world most particularly in Canada and the USA, compared
to the traditional lentil-growing countries.
8.2. Historical Perspectives

The domestication of lentil occurred, together with that of emmer and
einkorn wheat, barley, pea, chickpea, bitter vetch and flax, during the Neolithic Agricultural Revolution, which is thought to have taken place in the
eastern Mediterranean around the 8th and 7th millennia bc (Zohary and
Hopf, 1973). Lentil spread rapidly with that of Neolithic agriculture to the
Nile Valley, Europe and Central Asia. It was part of the Harappan crop
assemblage in the Indian subcontinent between 2250 and 1750 bc (Zohary
and Hopf, 1993). After 1500 ad, the Spanish introduced lentil to South
America via Chile (Solh and Erskine, 1984). More recently it has been cultivated in Mexico, Canada, the USA, New Zealand and Australia.
Although lentil was cultivated as early as 8000 bc in the Middle East
(Hansen and Renfrew, 1978), it remained an under-exploited and underresearched crop until recently. Systematic research for its improvement
started at some national institutions and at ICARDA, Syria, during the last
three decades. Under the Consultative Group of International Agricultural
Research (CGIAR), ICARDA has the world mandate for lentil improvement.
The lentil improvement programme at ICARDA is closely linked with the
national agricultural research systems and with advanced research institutes in the world to address this mission.
The breeding objectives at ICARDA and in national programmes are
targeted to yield improvement and its stability and to address specific needs
of various agroecological regions. A world collection of wild and cultivated
lentil germplasm is maintained at ICARDA and has been instrumental in
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A. Sarker et al.
the development of improved genetic stocks suitable to different environmental niches. Except for a few traits, sufficient variability for important
economic characters including stress resistance is present in the germplasm
for use in breeding programmes. Following a bulk-pedigree method, the
lentil breeding programme at ICARDA constructs new genotypes to deliver
to the national programmes through the International Nursery Network.
8.3. Constraints to Production Addressable by Breeding

Average lentil yields are low because of the limited yield potential of landraces, which are also vulnerable to an array of stresses. The yield limiting
factors are lack of seedling vigour, slow leaf area development, high rate of
flower drop, low pod setting, poor dry matter, low harvest index, lack of lodging resistance, low or no response to inputs, and various biotic and abiotic
stresses. The major abiotic limiting factors to lentil production are low moisture availability and high temperature stress in spring, and, at high elevations,
cold temperatures in winter. Mineral imbalances like boron, iron, salinity and
sodicity though localized do cause substantial yield loss. Among biotic stresses,
rust, vascular wilt and Ascochyta blight diseases caused by Uromyces viciaefabae (Pers.) Schroet., Fusarium oxysporum f. sp. lentis Schlecht and Ascochyta
fabae Speg. f. sp. lentis, respectively, are globally important fungal pathogens
of lentil (Agarwal et al., 1993; Bayaa and Erskine, 1998). Other diseases such
as Botrytis blight (Botrytis cinerea Pers.), Stemphylium blight (Stemphylium
botryosum Wallr.), collar rot (Sclerotium rolfsii Sacc.), root rot (Rhizoctonia solani
Kuhn) and stem rot (Sclerotinia sclerotiorum (Lib.) de Bary) appear locally, but
cause substantial yield loss (Sarker et al., 2004a). Additional constraints to
production include agronomic problems of pod dehiscence and lodging, and
inadequate crop management practices by growers.
Adequate variability for many of the important traits exists within the
crop gene pool allowing manipulation through plant breeding. However,
several other important traits, such as biomass yield, pod shedding, nitrogen fixation and aphids, and the parasitic broomrape (Orobanche sp.) are not
currently addressable by breeding because of insufficient genetic variation
within cultivated germplasm and crossable wild species.
8.4. Genetic Resources and Variation in Key Traits

The international lentil breeding programme is built upon the foundation
of the germplasm collections and their efficient use. A large number of
germplasm accessions are maintained at ICARDA; the National Bureau of
Plant Genetic Resources, India; the Vavilov Institute of Plant Industry, Russia; and at the United States Department of Agriculture (USDA), USA. The
ICARDA collection is by far the largest and comprises 10,282 cultivated accessions from 72 countries and 583 wild species accessions from 24 countries.
Among cultivated species conserved at ICARDA, c.17% are breeding lines,
developed at ICARDA through cross breeding.
105
Marked variation among the characters for use in breeding and selection
programmes has been reported for various morphological characters (Sindhu
and Mishra, 1982; Erskine and Witcombe, 1984; Erskine et al., 1989; Ramgiry
et al., 1989), responses in flowering to temperature and photoperiod (Summerfield et al., 1985; Erskine et al., 1990b, 1994a), winter hardiness (Erskine et al.,
1981; Sarker and Erskine, 2006), iron-deficiency chlorosis (Erskine et al., 1993),
boron imbalances (Srivastava et al., 2000; Yau and Erskine, 2000) and drought
tolerance (Hamdi et al., 1996; Sarker et al., 2001). Germplasm and new breeding
lines resistant to fungal diseases (Bayaa and Erskine, 1998; Sarker et al., 2004a)
and viruses (Makkouk et al., 2001) are available at ICARDA.
Wild relatives also have shown considerable variability for morphological traits (Robertson and Erskine, 1997), winter hardiness (Hamdi et al.,
1996), Fusarium wilt and Ascochyta blight resistance (Bayaa et al., 1994,
1995), anthracnose resistance (Tullu et al., 2005) and susceptibility to Sitona
weevil (Mustafa et al., 2008).
8.5. Breeding Strategies and Approaches

From the onset of the breeding programme at ICARDA, a multidisciplinary
approach involving breeders, plant protectionists, agronomists, postharvest
technologists, socio-economists has been followed to develop and transfer
improved varieties/technologies to farmers. The improvement programme
also follows a decentralized breeding strategy to develop cultivars for shortseason environments of southerly latitude countries and winter-hardy
cultivars for the highlands. Among diseases, rust screening is done at hot
spots in Bangladesh, India and Ethiopia, and Ascochyta blight evaluation in
Pakistan.
Recently, the lentil breeding programme has adopted Participatory
Varietal Selection (PVS) to enhance adoption of improved varieties as indicated by Witcombe and Joshi (1996). Although over the past two decades a
number of lentil cultivars have been released in many countries, their adoption by farmers has been slow for several reasons: poor adaptation to varying environments and vulnerability of pests and diseases, inadequacy of the
seed distribution system, lack of extension services and inattention to consumers tastes. In this context, it is imperative that farmers are involved in
the development of high-yielding lentil cultivars and production technology, right from the start. With their native wisdom, first-hand knowledge
of the crop and cropping systems and other local conditions, farmers can be
effective collaborators in this endeavour.
8.6. Major Agroecology and Specific Breeding Objectives

Knowledge of the patterns of variation in the world germplasm collection is
the key to understanding factors affecting lentil adaptation. The geographic
distribution of variation of landraces in the world lentil collection for
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A. Sarker et al.
morphological characters, temperature and photoperiod effects on flowering,

winter hardiness, iron-deficiency chlorosis and boron imbalances collectively
illustrate the specificity of adaptation in lentil. Additional information on the
specificity of adaptation within the crop has come from collaborative yield
trials of common entries evaluated at widely differing locations.
Understanding of genotypes and environmental factors, the local constraints to production and consumer requirements of different geographic
areas for seed as food and straw as feed, has been a guide to the international
breeding programme at ICARDA for developing new genetic materials for a
series of separate, but finely geographically-targeted streams, linked closely
to national breeding programmes (Robertson and Erskine, 1997).
The major target agroecological regions of production of lentil are: (i)
South Asia, East Africa and Yemen; (ii) low to medium elevation areas in
the Mediterranean; and (iii) the high elevation area of West Asia and North
Africa. These regions correspond to the maturity groups of early, medium
and late maturity (Table 8.1). Within each of these major regions there are
specific target areas. In addition, lentil improvement activities have recently
been extended to the Central Asia and the Caucasus (CAC) region where
the initial thrust is to study lentil adaptation to the prevailing agroclimatic
conditions. For Latin America, large-seed, yellow-cotyledon lentils, suitable for mechanical harvest with resistance to rust and Ascochyta blight
are preferred.
The lentil breeding programme generally uses parents of diverse origin
with known traits with the aim to combine gene(s) that contribute to yield
and resistance to major biotic and abiotic stresses. Wide crosses among
cultigens are also done by manipulating planting dates and providing 18 h
extended light period to the parents to attain synchrony in flowering. In
addition, crosses are made to study the inheritance pattern of specific trait(s)
and to develop recombinant populations for biotechnological research.
More than 250 crosses are commissioned at ICARDA every year. At present
the programme uses breeding lines in hybridization, developed through
crossing multiple parents, thus allowing recombination of desirable genes
in newly constructed genotypes for yield improvement and stability over
years and across production zones.
8.7. Breeding Methodologies

In the early stages of lentil cultivar development, most of the cultivars
released have been derived from pure line selection within heterogeneous
landraces (Muehlbauer, 1992). Due to increased efforts in lentil breeding
nationally and internationally, new lentil cultivars/genotypes are now
being developed through cross breeding. The methods of breeding lentil
are similar to those utilized in breeding other self-pollinated crops that
include pure line selection or hybridization followed by the bulk method,
the pedigree method, the single seed descent, or some modification of these
procedures. At ICARDA a bulk-pedigree method is used, where single plant
Region
A. South Asia and East Africa
B. Mediterranean low to
medium elevation
Key traits for recombination

1. India, Pakistan, Nepal
and Ethiopia
2. Bangladesh
3. Yemen
1. 300400 mm annual rainfall
2. <300 mm annual rainfall
3. Morocco
4. Egypt
C. High elevation
D. Central Asia and the Caucasus
E. Latin America
1. Anatolian highlands
2. North African highlands
Seed yield, early maturity, resistance to root diseases, rust

and Ascochyta blight
Seed yield, extra-earliness and combined resistance to rust
and Stemphylium blight diseases
Early maturity, Ascochyta blight resistance
Biomass (seed + straw), attributes for mechanical harvest and
wilt resistance
Biomass, drought escape through earliness and wilt resistance
Biomass, attributes for mechanical harvest and combined
resistance to rust, Ascochyta blight and wilt
Seed yield, response to irrigation, earliness and Fusarium wilt
resistance
Biomass, winter hardiness and Ascochyta blight resistance
Seed yield, low level of winter hardiness and rust resistance
Seed yield, large-seed, good standing ability
Large-seed, resistance to rust and Ascochyta blight
Table 8.1. Target agroecological regions of lentil production and key breeding aims.
107
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A. Sarker et al.
selection is done from F4 bulks to develop F5 progeny. A scheme of breeding methodology followed at ICARDA is shown below.
Hybridization (Parent A Parent B): at Tel Hadya, ICARDA HQ (year 1)
Conrmation for hybridity (F1) at summer nursery, Lebanon (year 1)
F2 bulks grown at Tel Hadya, ICARDA HQ (year 2)
F3 grown at summer nursery in Lebanon (year 2)
F4 grown at Tel Hadya and single plant selection performed (year 3)
F5 families evaluated for agronomic traits and Fusarium wilt reaction in wiltsick plot at Tel Hadya (year 4)
F6 progenies are evaluated for yield, agronomic traits and Fusarium wilt at
Tel Hadya (year 5)
F7 xed lines are evaluated in preliminary yield trials at three contrasting
locations: Breda, Tel Hadya (Syria) and Terbol (Lebanon)
F8 advanced yield trials at these three locations except those bred for southerly latitude countries
F9 incorporation in international nurseries
Mutation breeding programmes are underway at the Bangladesh Institute of Nuclear Agriculture (BINA), Bangladesh; the Indian Agricultural
Research Institute (IARI), India; and the National Institute of Agricultural
Biology (NIAB), Pakistan. Enormous variability has been created for economic traits. Generally, gamma rays are used to create macromutations and
EMS (ethyl methane sulfonate), NEU (nitroso ethyl urea) and SA (sodium
azide) are useful to generate micromutations in lentil.
8.8. Breeding Achievements

Broadening the genetic base in South Asia
About 50% of the worlds lentil is grown in South Asia (FAO, 2007). The
region grows only small-seeded red lentil, pilosae type, an endemic group
with a narrow genetic variability, short stature and early maturity. For a long
time, lentil improvement programmes in South Asia were handicapped due
to lack of access to sufficient genetic variability for improvement because of
lack of overlap in flowering with introduced lentils. Most materials of West
Asian origin, when grown in short-season environments of South Asia,
come to flower when the South Asian landraces are maturing (Ceccarelli
et al., 1994). This observation prompted research on the environmental factors that control flowering in lentil. Summerfield et al. (1985) concluded that
temperature and photoperiod modulate flowering in lentil. In a comprehensive study with a broad spectrum of lentil germplasm, Erskine et al. (1994b)
found that germplasm of Indian origin are more sensitive to temperature
and less responsive to photoperiod than germplasm from West Asia.
To widen the narrow genetic base of lentil in South Asia, in an early
attempt, Precoz, a bold-seeded, early-maturing germplasm of Argentine
origin and other early maturing exotic germplasm was introduced from
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ICARDA and utilized in breeding programmes in the region (Erskine et al.,

1998). This led to the development of improved cultivars and a yield jump
in Bangladesh (Barimasur-2 and Barimasur-4) (Sarker et al., 1999a, b),
Nepal (Shekher and Shital) and Pakistan (Manshera-98, Shiraz-96,
Masoor-93 and Masoor-2002). Precoz when evaluated under both Indian
and Syrian environments was shown to have a major gene Sn for early
flowering (Sarker et al., 1999c).
As a spin-off to research towards broadening the genetic base and based
on findings of quantitative responses to photothermal effects, Summerfield
et al. (1985) proposed a model for flowering in lentil. According to the
model, rates of progress towards flowering (i.e. 1/f, the reciprocal of the
time to first flower, f in all genotypes vernalized or not) were linear functions of both mean temperature, t, and photoperiod, p, with no interactions
between the two terms. So, over a wide range of conditions covering the
photothermal regimes experienced by the lentil crop worldwide, time to
flowering was described by the equation:
1/f = a + bt + cp
where a, b and c are constants which differ between genotypes and the value
of which provide a sound basis for screening germplasm for sensitivity to
temperature and photoperiod. Although these two environmental factors
affect the same phenological event, Summerfield et al. (1985) suggested that
the responses are under separate genetic control.
Identification and exploitation of resistance to abiotic stresses

Winter hardiness
In the highlands of central Anatolia of Turkey, Iran, Afghanistan and Baluchistan province of Pakistan, lentil production can be increased significantly by shifting sowing from spring to early spring or winter planting.
The winter crop allows optimum vegetative growth and higher water use
efficiency which leads to higher yield. There is a potential to replace about
400,000 ha of spring lentil with a winter lentil in the highlands of West Asia
(Sakar et al., 1988).
Spring-sown lentils in the highlands frequently suffer from terminal
drought, which can be avoided by early sowing; this also allows taller canopy development suitable for machine harvest. Cultivars with winter hardiness (such as Kafkas, Cifci and Uzbek) are now being cultivated in
Turkey. Several additional lines are identified for future release (LC 9978057,
LC 9977006, LC 9977116, LC 9978013, ILL 759, ILL 1878, ILL 4400, ILL 7155,
ILL 8146, ILL 8611 and ILL 9832). In Iran lines with winter hardiness, early
growth vigour and rapid ground cover (ILL 662, ILL 857, ILL 975, ILL 1878)
are under on-farm evaluation. Selection at the Arid Zone Research Center,
Baluchistan, Pakistan resulted in the release of cultivar ShirAZ-96, based
on ICARDA germplasm for winter cultivation (Sarker and Erskine, 2006).
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A. Sarker et al.
The focus in winter/autumn sowing is to translate the research results into

production gains on-farm; in this regard weed control is critical. Production
increases from early sowing are only possible in combination with winter
hardiness of the cultivars.
Drought tolerance
Tolerance to moisture stress is a key trait in rain-fed lentil production.
Drought is the major abiotic stress in many countries of the world (Johansen
et al., 1994). In the Mediterranean environment of West Asia and North
Africa, lentil generally suffers from terminal drought (Sarker et al., 2001). In
South Asia, the crop is grown on conserved soil moisture, and occasionally
faces intermittent drought. Drought-tolerance research, particularly development of screening techniques for the identification of drought-tolerant
genotypes, became a key research issue in ICARDAs crop improvement
strategy (Malhotra et al., 2004). Drought-tolerance mechanisms, such as
escape, dehydration avoidance and dehydration tolerance have been used to
select drought-tolerant genotypes. Traits like early seedling establishment,
early growth vigour and rapid ground coverage, high biomass development,
early flowering and maturity were taken into consideration to select droughttolerant genotypes. Selection in drought-prone sites (at the Syrian experimental station Breda and farmers fields where rainfall <280 mm) is the key
to success in identification of drought-tolerant genotypes. Seedling shoot
and root traits such as taproot length and lateral root number are important
traits for drought tolerance (Sarker et al., 2005). The resulting droughttolerant lines are made available to national programmes through the Lentil
International Drought Tolerant Nursery (LIDTN) (Malhotra et al., 2004).
Salinity tolerance
Soil salinity is a major obstacle to crop production in arid and semi-arid
regions of the world. Like other field crops, lentil often suffers from soil
salinity. The crop is very sensitive to salinity. Nevertheless, some genetic
variation in response to salinity has been reported (Ashraf and Waheed,
1990; Ameen, 1999).
Mineral imbalances
Soil mineral imbalances sometimes pose a serious threat to lentil production locally. Boron (B) toxicity occurs primarily in some arid areas in alkaline soils and is reported from Australia, India, Pakistan, Iraq, Peru and
Turkey. Screening has revealed significant variation in B toxicity with tolerance reported in lentil (Yau and Erskine, 2000). By contrast, B deficiency is a
problem in other regions, such as the eastern Terai plain of Nepal, eastern
India and northern Bangladesh, where the soil is leached. Landraces from
Nepal and Bangladesh were shown to be tolerant to B deficiency, and accessions of West Asian origin were highly inefficient in B-deficient soil (Srivastava et al., 2000). Among B-deficiency tolerant lines, ILL 2580, ILL 5888, ILL
111
8009 and ILL 8010 showed higher potential in the B-deficient Chitwan
region of Nepal and in the northern region of Bangladesh.
Gene mining and utilization to combat biotic stresses
Diseases
Among biotic stresses, diseases, caused by various fungal pathogens, are
the most devastating yield-limiting factors of lentil. Of them, Fusarium wilt,
rust and Ascochyta blight are globally important diseases over a wide range
of environments and are addressed by the international breeding programme at ICARDA.
Screening techniques for these diseases have been developed and
sources of resistance have been identified. Fusarium wilt-resistant lines
have been identified through rigorous screening in a >15-year-old wilt-sick
plot at ICARDA, Tel Hadya. Systematic screening of all new germplasm
and breeding lines is carried out. Recently a total of 34 new sources of resistance were identified from 1500 accessions of diverse origin and included in
the breeding programme (Sarker et al., 2004a). Rust-resistant lines have been
reported from India, Bangladesh and Ethiopia, and some are released as
varieties (Agarwal et al., 1993; Bakr, 1993; Bejiga et al., 1998). Lentil accessions with a high level of resistance have been identified for foliar as well as
for seed infection of Ascochyta blight and are being used in a breeding programme. Sources of resistance to minor diseases and viruses are also
reported (Agarwal et al., 1993; Bakr, 1993; Makkouk et al., 2001; Tullu et al.,
2005). Three disease nurseries, namely, the Lentil International Rust Nursery (LIRN), the Lentil International Fusarium Wilt Nursery (LIFWN) and
the Lentil International Ascochyta Blight Nursery (LIABN) are made available to national programmes. The next goal is to ensure multiple disease
resistance over specific production zones.
Construction of suitable plant type for mechanical harvest
Lentil production in Australia, Canada and the USA has been completely
mechanized since cultivation began. However, the availability of combine
harvesters, tall plant stature and large field sizes contrast with the situation
in the Mediterranean basin. Lentil cultivation in West Asia and North Africa
has been threatened due to rising cost of agricultural labour with hand harvesting accounting for approximately 47% of the total cost of production.
Therefore to reduce costs, it is essential that lentil harvest be mechanized.
To address this constraint, ICARDA has developed economic machine harvest systems for lentil cultivation involving cultivars with improved standing ability, a flattened seedbed and the use of cutter bars. The Center has
developed and promoted with NARS a lentil production package that includes
mechanization and the use of improved cultivars with good standing
ability. Such cultivars include Idlib-2, Idlib-3 and Idlib-4 in Syria, Hala
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A. Sarker et al.
and Rachayya in Lebanon, IPA-98 in Iraq, Saliana and Kef in Tunisia,

and Firat-87 and Sayran-96 in Turkey. On average, mechanical harvesting combined with improved cultivars having good standing ability reduces
harvest costs by 1720% (ICARDA, 2001).
Development of micronutrient-dense cultivars
Currently, there has been interest at national and international levels to
enhance the nutrient content in our food. One approach is to develop nutrient-dense cultivars in staple foods. In this endeavour, the first step is to
characterize the genetic diversity for nutrient content that exists in current
cultivars and genotypes, and use this information in breeding. In preliminary screening of 1645 lentil accessions comprised of breeding lines, landraces and released cultivars, iron (Fe) content varied from 41 to 132 mg/
kg and zinc (Zn) content ranged from 22 to 78 mg/kg, which suggest scope
of improvement in these traits through plant breeding (ICARDA, 2007). Some
of the identified cultivars are under fast-tracking to disseminate to farmers in
Ethiopia, Bangladesh, Nepal, Syria, Turkey, Portugal and Morocco (Table 8.2).
The most commonly grown improved cultivars are also being crossed with
micronutrient-dense accessions to enrich the cultivars with higher micronutrients. The high content lines are being delivered to national programmes
through the International Nursery Network. High content high content
lines are being crossed to develop transgressive segregants with even higher
contents. For example, parents with an Fe content >80 mg/kg are being
used to develop cultivars with an Fe content of >120 mg/kg. Similarly, high
Zn content accessions are included in a crossing programme to develop
cultivars with >85 mg/kg of Zn. Nutrient-dense lentil cultivars could provide marketing opportunities for producers, thereby increasing sales. An
additional benefit to farmers is that under some soil conditions (e.g. low
mineral availability), nutrient-dense cultivars have the potential to improve
seedling establishment and resistance to certain diseases.
Varietal releases by national programmes
As we stated earlier, ICARDA supplies national breeding programmes
through the International Nursery Network with a range of genetically
fixed materials and segregating populations. These are for use in selection
programmes according to the national programmes specific needs. On the
basis of yield, phenological adaptation, agronomically desirable traits, resistance to prevailing stresses, quality aspects, farmers and consumers preference, etc., national scientists identify and select promising lines/single
plants for further evaluation and eventual release for commercial cultivation. In this endeavor, to date 104 lentil cultivars have been registered by 31
countries emanating from ICARDA-supplied genetic materials for high
yield, disease resistance, drought tolerance, suitability to harvest mechanization and seed traits (Table 8.3).
Cultivars with high concentration of Fe and Zn.
Cultivar
Country
Alemaya
Idlib-2
Idlib-3
Idlib-4
Barimasur-4
Barimasur-5
Barimasur-6
Abda
Ethiopia
Syria
Syria
Syria
Bangladesh
Bangladesh
Bangladesh
Morocco
Fe (mg/kg)
Zn (mg/kg)
82.3
73.2
72.3
68.5
86.2
86.0
85.2
69.7
66.0
52.3
51.2
62.4
51.1
59.0
66.1
56.3
Cultivar
Country
Fe (mg/kg)
Meyveci-2001
Beleza
Shital
Shekhar
Khajurah-1
Khajurah-2
Sisir
Alemaya
Turkey
Portugal
Nepal
Nepal
Nepal
Nepal
Nepal
Ethiopia
64.5
74.0
75.7
83.4
78.8
100.7
98.0
82.0
Zn (mg/kg)
53.4
56.6
59.0
56.0
58.0
59.0
64.0
66.0
Table 8.2.
113
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Table 8.3.
Lentil varieties released by national programmes that emanated from ICARDA-supplied genetic materials.
Region
Country
Asia
Bangladesh, India, Nepal, Pakistan, China,

Afghanistan, Iran, Iraq, Syria, Lebanon,
Jordan, Yemen, Turkey
Ethiopia, Egypt, Morocco, Libya, Tunisia,
Algeria, Lesotho, Sudan
Argentina, Canada, Ecuador, USA
Africa
The Americas
Oceania
Europe
Central Asia and
the Caucasus
Australia, New Zealand

Portugal
Georgia, Uzbekistan, Azerbaijan
No. of
varieties
53
30
7
10
2
2
Traits
High yield; wilt, rust, Stemphylium blight and
Ascochyta blight resistance; better standing ability;
high biomass; early maturity; winter hardiness
High yield; wilt and rust resistance; early maturity;
resistance to lodging; winter hardiness
High yield; rust and Ascochyta blight resistance;
good standing ability
High yield; Ascochyta blight resistance; erect canopy
High yield; erect; large seed; tall
High yield; large seed; tall
A. Sarker et al.
115
20
18
16
14
Variety
12
10
13
11
9
10
9
7
6
2
18
17
3
1
Landrace Crosses Landrace Crosses Landrace Crosses Landrace Crosses Landrace Crosses Landrace Crosses
19801984
19851989
19901994
19951999
20002004
20052007
Year of release
Fig. 8.1. Trend of cultivar releases from landraces and cross breeding.
Of these, 44 varieties were directly released from germplasm in the

early phase of lentil improvement and 60 cultivars have been released from
breeding lines in the recent past. There is a clear trend of increasing development of new cultivars through cross breeding (Fig. 8.1). The breeding
lines were developed involving multiple parents of diverse origin. A yield
potential up to 3.42 t/ha has been observed from large plot trials in Ethiopia. Cultivars developed through hybridization have wide adaptation and
stable yields as a result of the buffering effect of genes aggregated from
diverse multiple parents.
8.9. Evolution of the International Breeding Programme

The lentil breeding strategy at ICARDA has changed with time. Initially, in
Stage 1, variation in the germplasm collection was directly exploited. Selection was made among and within locally adapted landraces. These selections were distributed to national programmes through the International
Nursery Network to test for local adaptation. As a result many cultivars
released by national programmes are actually selections from landraces in
the ICARDA germplasm collection (Robertson et al., 1996). These Stage-1
registrations emphasize first the value of direct exploitation of landraces
and secondly the under-exploited nature of lentil germplasm.
The particular combinations of characters required for specific regions
were often not found on the shelf in the germplasm collection. Consequently,
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A. Sarker et al.
ICARDA started hybridization and selection from segregating populations

from crosses made at ICARDA to produce fixed lines as Stage 2 material.
Such selections were then distributed after multiplication to the national
programmes for selection in their respective agroclimatic zones. This has
resulted in the release of a number of cultivars in different regions.
However, lentil lines developed from selection at ICARDA in West Asia
are mostly limited to adaptation to the home region. As a result, the breeding
programme has decentralized to work closely with national programmes. For
the other regions, at Stage 3, crosses are agreed upon with national cooperators and made at ICARDA, Tel Hadya, Syria and then country-specific segregating populations shipped to national cooperators for local selection and
release as cultivars. Approximately 250 such crosses are made annually at
ICARDA. Selections made by national programmes are then fed back into the
International Nursery Network for wider distribution.
In Stage 4, the national programmes directly use ICARDA-derived material in their own hybridization programme and selections are made locally.
8.10. Future Directions

Exciting gains in sustainable production arise from the integration of a
change in agronomic practice with a high-yielding new cultivar. Several
such prospects exist for lentil and are given below.
In South Asia, large areas are left fallow over winter following the
harvest of rainy-season paddy rice. Farmers need early-maturing, diseaseresistant varieties with late sowing potential for such situations or when
land becomes available for lentil planting after the monsoon floodwater
has subsided. When such extra-early maturing cultivars with combined
resistance to diseases will be available, the prospect will open of a major
expansion of lentil production in the post-rice system in the Indian
subcontinent.
In some areas around the Mediterranean, such as in south Syria, lentil
production has ceased primarily because of severe yield losses caused by
Sitona weevil. With the identification of sources of resistance to this insect in
wild species, most particularly in L. culinaris ssp. orientalis, it may be possible
to construct lentil genotypes resistant to this pest (Mustafa et al., 2008).
In the highlands of West Asia, still a vast majority of lentil is currently
spring sown. However, adoption and expansion of sowing in winter, coupled
with the use of a winter-hardy cultivar will give substantial yield advantages.
Although agronomic constraints to winter-sowing lentil in the highlands
require further research, substantial yield increase can be expected in the
future to augment the area and production in this agroecological zone.
Molecular tagging of genes responsible for resistance to biotic and abiotic stresses needs to be done to assist the breeding programme. Further,
research on functional genomics to address some of the key problems
should be initiated. Development of transgenics for drought tolerance using
dreb1A gene (Liu et al., 1998) and to control Orobanche through incorporation
117
of the Bar gene (Thompson et al., 1987) and other notorious weeds need to
be exploited.
In-depth physiological research to meet challenges due to the effect of
climate change is warranted to assist breeding programmes.
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Breeding for Short Season Environments
M. Matiur Rahman,1 Ashutosh Sarker,2 Shiv Kumar,3 Asghar Ali,4

N.K. Yadav5 and M. Lutfor Rahman1
1Bangladesh
Agricultural Research Institute, Gazipur, Bangladesh; 2International

Center for Agricultural Research in the Dry Areas (ICARDA), Aleppo, Syria; 3Indian
Institute of Pulses Research, Kanpur, India; 4National Agricultural Research Centre,
Islamabad, Pakistan; 5National Grain Legumes Research Program, Rampur, Nepal
9.1. Introduction
Lentil (Lens culinaris ssp. culinaris Medik.) from South Asia contributes 34% to
global production. The crop was introduced to the region from West Asia
through Afghanistan around 2000 bc (Cubero, 1981) and is grown extensively
under varied agroclimatic conditions. South Asian lentils have limited genetic
variation for agromorphological traits (Erskine et al., 1989, 1998; Erskine and
Saxena, 1993) and this is an apparent result of a bottleneck during introduction of the germplasm to the region. Natural selection and farmer selection
for adaptation and local preferences have had a profound effect on seed traits,
pubescence, size of vegetative organs, phenology, plant height and pod shape
as well as flowering responses. Lentils from South Asia are exclusively of the
pilosae ecotype which is characterized by dense pubescence on vegetative
organs (Barulina, 1930) and rudimentary tendrils (Vandenberg and Slinkard,
1989). The main driver behind this differentiation was adaptation to match the
prevailing climatic factors, particularly photoperiod and temperature (Erskine
et al., 1989). Cultivated lentils have been described as belonging to two seed
groups: macrosperma (69 mm diameter) and microsperma (25 mm). There are
strong preferences for small seed types with red cotyledons among consumers in the South Asian countries while preference for the large-seeded lentil
with yellow cotyledons is limited to a few pockets in North Pakistan and
India. Central India is one region where large-seeded cultivars of microsperma
lentil are preferred over small-seeded ones.
9.2. Present Status

In South Asia, the lentil is extensively grown in India, Nepal, Bangladesh
and Pakistan where 1.83 million ha are cultivated with total production of
121
122
M. Matiur Rahman et al.
1.30 million t and average productivity of 699 kg/ha accounting for 46% of
the global area and 34% of global production (see Erskine, Chapter 2 this
volume). Although during the past two and a half decades (19802006) production has increased in the region, the production increase was 53% during 19811991, 23% during 19912001 and <2% since the beginning of this
century.
Analysis of each country for the period 19812006 showed that India
and Nepal experienced positive growth in area and production of lentil,
whereas it was negative in Bangladesh and Pakistan. In Nepal, lentil cultivation has recently been extended in the hill and mountain regions with
substantial gain in productivity (Shrestha et al., 2005). Presently, India
accounts for 76% and 80% of the total South Asian production and area,
respectively, in the region, followed by Nepal (12%, 10%), Bangladesh
(9.2%, 7.7%) and Pakistan (1.5%, 2.3%).
9.3. Production Base

Lentil is grown in South Asia as a post-rainy season crop mostly under rainfed conditions in areas where scanty rainfall is frequently observed. The
lentil crop season is characterized by mild and relatively short winters during the vegetative phase and high temperatures with receding soil moisture
during the reproductive phase. As a result, the crop experiences terminal
drought and forced maturity. A substantial area of South Asia lentil is
grown as the only legume in the winter season in rotation with summer
crops such as rice, maize, sorghum, cotton and jute. However, it is also
grown as an intercrop with wheat, barley, mustard, linseed, grasspea or
field pea. In Bangladesh and the north-eastern states of India and Nepal,
lentil is relay cropped with rice in which the lentil seeds are sown in the
standing rice crop about 15 days before its harvest under the utera system
or immediately after the harvest of rice under the paira system. In both cropping systems, lentil is grown predominantly on residual soil moisture.
9.4. Production Constraints

The average yield of lentil in South Asia (699 kg/ha) is 24% lower than the
world average (1053 kg/ha) (FAO, 2007). The reasons for low yield are
occurrence of various biotic, abiotic and edaphic factors at different growth
stages (Ali et al., 2000; Haqqani et al., 2000; Matiur Rahman et al., 2000; Pandey et al., 2000). Being a cool-season crop, lentil is grown on conserved soil
moisture in the post-rainy season and generally, no rainfall except occasional scattered showers occur during winter in the region. As a result, soil
water deficit during crop establishment and in the post-flowering period
emerges as the major yield constraint along with the rising temperature.
Salinity and deficiencies of soil nutrients are also observed locally in the
region. For example, boron deficiency has been identified as a limiting
123
factor in some parts of Nepal. Lentil is also sensitive to zinc and iron deficiency and poor nitrogen fixation. The crop often encounters pod dehiscence and lodging as well under the conditions prevalent in the region.
Vascular wilt caused by Fusarium oxysporum Schlect. f. sp. lentis Vasudeva
and Srinivasan and rust caused by Uromyces viciae-fabae (Pers.) de Bary are
widespread diseases inflicting serious yield losses across the countries.
Stemphylium blight caused by Stemphylium botrysum Wallr. in Bangladesh
and Nepal, Ascochyta blight (Ascochyta lentis Bond and Vassil) and Botrytis
grey mould (Botrytis cinerea Pers. Ex Fr.) in Pakistan, and collar rot (Sclerotium rolfsii Sacc.), root rot (Rhizoctonia solani Kuhn) and stem rot (Sclerotinia
sclerotiorum) in India and Nepal are also of economic importance. Among
insects, aphid (Aphis craccivora Koch) is an important field pest while
bruchids (Bruchus spp. and Callosobruchus spp.) cause serious grain losses
during storage. These stresses appear individually and/or in combination
with varying intensity depending on the location and year, thus, causing
great fluctuation in lentil production in the region. Hence, development of
resistant varieties against major diseases and realizing the yield potential of
existing commercial cultivars have been given priority in the past while
there have been limited efforts towards breaking yield barriers in lentil in
South Asia.
9.5. Timing of Flowering

The onset of flowering is an important phenological event for the short
period of the vegetative phase, and determines the potential of the crop
under the climatic conditions to which the crop will be exposed during
reproductive growth. The optimum flowering response differs between
regions and is simple to measure within a target region (Erskine et al.,
1994a). Being a quantitative long day plant, wide genetic variation for flowering response to photoperiod and temperature has been reported in the
global germplasm repository at the International Center for Agricultural
Research in the Dry Areas (ICARDA) (Erskine et al., 1990). Presently, the
understanding of the genetic control of flowering is limited. However, the
available results show that early flowering is largely determined by a single
recessive gene sn in addition to some polygenes (Sarker et al., 1999). This
results in transgressive segregation in the progeny of crosses between early
and late flowering parents (Tyagi and Sharma, 1989; B. Sharma, Chapter 7
this volume).
Sowing time has a profound effect on flowering pattern, productivity
and success of the crop as it causes considerable change in the plant environment with respect to temperature, photoperiod and availability of soil
moisture (Ramakrishna et al., 2000). The optimum planting time of lentil in
Pakistan, northern India, Nepal and Bangladesh is the last week of October
to mid-November depending on the location and soil moisture (Ahad Mia
and Matiur Rahman, 1993; Ali et al., 1993; Neupane and Bharati, 1993) while
in central India and Pakistan highlands (>
_1000m altitude), lentil is planted
124
from September to mid-October (Ali et al., 1991, 1993). Delay in planting

progressively shortens the growing period as a result of the rapid rise in
temperature accompanied by low moisture availability which leads to inadequate biomass production, poor pod filling and forced maturity (Erskine
et al., 1994b; Ali et al., 2000).
Optimum temperature for lentil growth and development ranges from
15 to 25C (Clarke et al., 2005). Crop duration is extended by cool weather
(Summerfield, 1981). The lentil season in South Asia coincides with a sharp
rise in temperature (>30C) by the end of February in central India, West
Bengal, Bangladesh and early April in northern India, Nepal and Pakistan.
The rising temperatures coupled with receding soil moisture push the
plants into forced maturity. Thus, the crop duration in this region varies
from 100 to 140 days depending on location. Therefore, cultivars with high
yield potential should have a crop duration of around 100 days in central
India, 110 days in Bangladesh and 140 days in the cooler areas of the region.
9.6. History of Improvement

Lentil breeding has a relatively short history of organized research as compared to other major crops such as maize and soybean. The earliest record
of organized research in South Asia goes back to 1943 when a lentil breeding programme was taken up under the State Department of Agriculture in
different provinces of undivided India. This received a further boost with
the establishment of the All India Coordinated Pulses Improvement Project
in 1967. With the creation of ICARDA in 1977, lentil improvement programmes in South Asian countries received valuable assistance by having
access to global germplasm, yield nurseries and basic information. This collaboration has resulted in the development of improved cultivars and widening of the genetic base. Over the years, these countries have established
sound lentil breeding programmes. The lentil breeding programme in Nepal
began in 1977 and systematic genetic improvement started in 1985 with the
establishment of the National Grain Legume Research Program (Bharati
and Neupane, 1991). The Bangladesh Agricultural Research Institute (BARI)
launched a pulse research programme in 1991. In Pakistan, the National
Agricultural Research Centre in Islamabad has major responsibility for lentil improvement. The work initiated in 1981 under the National Coordinated
Research Programme on pulses.
9.7. Germplasm
Collection of genetic resources in India started in the early 1960s under a
Rockefeller Foundation programme which was funded by PL 480 assistance
and extended to other South Asian countries in 1977 under the Arid Lands
Agricultural Development Program (ALAD). The global lentil germplasm
repository of 8748 landrace accessions and 479 wild accessions maintained
125
at ICARDA contains 86 accessions from Bangladesh, 2118 from India, 475

from Nepal and 297 from Pakistan (Sarker et al., 2005). Among the national
genebanks, India and Bangladesh hold 2238 and 2045 accessions, respectively. In addition, a large number of introductions in the form of exotic
lines and yield nurseries have also been accumulated to enrich the local
gene pool. About 7533 accessions in India and 2500 in Bangladesh have
been introduced through ICARDA. Studies on quantitative agromorphological variation in germplasm from 13 countries revealed that the Indian
lentils have limited variation. India and Pakistan lie among the countries
with the lowest quantitative agromorphological variation. The group was
characterized by early flowering and maturity, low biological yield, short
stature, lowest pod height and small seeds (Erskine et al., 1989). Evaluation
of 456 germplasm accessions showed large variability in the indigenous collection in India (Table 9.1).
9.8. Breeding Objectives

The major target of the current breeding programmes in South Asian countries is to match crop duration with the target environments coupled with
resistance to major diseases and improved seed size. Genotypes which
mature relatively early are better adapted to variable environments. Thus,
short duration cultivars with rapid grain fill may avoid substantial terminal
stress (Summerfield, 1981). It is, therefore, essential to breed short duration
cultivars combining resistance to key biotic and abiotic stresses (Saxena
et al., 2005). Extra earliness combined with resistance to rust and Stemphylium blight is important for the short season environment of Bangladesh
and the eastern and central regions of India while genotypes resistant to
root diseases and rust are crucial for Nepal and central India. Medium
duration cultivars are also widely cultivated in Nepal. Ascochyta blight is
Table 9.1.
Mean and range for various traits in Indian lentil germplasm.
Trait
Days to 50% owering
Days to maturity
Plant height (cm)
Primary branches
Pod length (cm)
Pod width (cm)
Pods per peduncle
Peduncles per plant
Pods per plant
Yield per plant (g)
100-grain weight (g)
Seeds per pod
Seed diameter (mm)
Mean
Minimum
Maximum
127.5
179.6
31.4
2.9
1.1
0.5
2.0
36.7
55.0
12.0
2.90
2.02
0.44
108
154
17
1
0.9
0.4
1
11
14
0.4
1.6
1
0.35
199
202
57
7
1.7
0.9
3
110
180
31.3
8.5
3
0.67
126
important in the highlands of north India and Pakistan. Adequate variability

for these traits exists within lentil genetic resources allowing for manipulation through breeding (Erskine et al., 1989).
9.9. Breeding Strategy

Efficient screening techniques depend on the ability to reproduce the most
probable conditions of stress in the target environment (Wery et al., 1994). It
requires defining the most frequently occurring stress in the crop cycle and
reproducing it in conditions where screening of a large number of genotypes can be made. These two steps are essential for the reproducibility of
screening. The next step is to define the plant traits required in the target
environment. As mentioned earlier, terminal heat stress is frequently associated with soil moisture deficiency at the reproductive stage. Therefore,
germplasm must be planted at the appropriate time at the screening location so as to ensure that the stress coincides with the critical crop stages.
Since seed yield is not a reliable criterion for selection under stress conditions (Muehlbauer et al., 1985), it is suggested that indirect selection be practised through selecting traits that have high heritability and that are strongly
associated with yield. Although one cannot expect any single trait to determine yield under drought in view of the variability of stress and complexity
of yield, early growth vigour has shown strong correlation with biomass
and seed yield in lentil and is of practical value in predicting final yield
under a short duration environment (Erskine and Saxena, 1993). The early
flowering and maturing cultivars of lentil display a typical drought-avoidance
strategy at the reproductive stage when high temperature and water deficit
induce rapid senescence and early maturity (Erskine et al., 1994a; Shrestha
et al., 2006a). For biotic stresses, particularly diseases, field screening at hot
spots, as well as pot screening or glasshouse screening under controlled
conditions, has been standardized and is being employed for identification
of resistant donors (Khare et al., 1993; Gaur and Chaturvedi, 2004) and
inheritance studies. Field screening in sick plots of large size has been perfected with remarkable success at ICARDA.
Under short-duration rainfed environments, the superior performance
of lentil genotypes from South Asia and derivatives from crosses between
South and West Asian lentils has been associated with rapid canopy cover,
early phenology and high harvest index (Shrestha et al., 2005). Higher grain
yield in the derivatives from crosses between the South and West Asian
genotypes is attributed to larger seed size introgressed from the West Asian
genotypes in the typical short duration background of South Asian genotypes (Shrestha et al., 2006a). Therefore, genotypes with rapid ground cover,
early phenology, a prolonged flowering and podding period, leading to
increased dry matter production, more pods, high harvest index, efficient
water use and large seeds should be selected for the short duration environment of South Asia (Shrestha et al., 2005). Crossing of superior parental lines
from the South and West Asian germplasms, early generation selection, and
127
evaluation under a range of environments could be the preferred breeding

strategy for further improvement towards adaptation and increased yield
of this important legume crop in South Asia (Shrestha et al., 2005).
As lentils have an indeterminate growth habit, water deficit at the
reproductive stage affects both vegetative and reproductive development
by reducing leaf area, dry matter production, number of branches and, consequently, the number of flowers, pods and seeds. Under water deficit conditions, the number of pods and seeds and seed size are most important traits
related to seed yield in lentil (Shrestha et al., 2006b). Correlations between
traits must be given careful consideration and interpretation by plant breeders. Number of pods per plant, seeds per pod, secondary branches per plant,
plant height and straw yield are reported to have significant positive correlation with grain yield in lentil (Erskine, 1983; Singh and Singh, 1991; Pandey
et al., 1992). Thus, selection for increasing seed yield would not adversely
affect straw yield. Muehlbauer et al. (1985) concluded that branching pattern
and number of fruits to reach maturity are the most important characters that
contribute positively to yield. Therefore, early maturing plants having early
growth vigour and a greater number of pods should be selected for short season environments, as they will be able to escape terminal stresses. In South
Asia, small-seeded and semi-spreading cultivars are mostly grown under
normal and late sown situations, and yields are stable because the plants are
able to fill the available space by initiating lateral branches and compensating
for poor emergence. Early flowering combined with early growth vigour,
large seeds and cold tolerance are some of the desirable attributes of new
plant types. For late sown conditions, semi-spreading growth, early maturity
and large seeds are important traits (Singh, 1997).
9.10. Creation of Variability

The scope of selection for desirable genotypes depends on the extent of exploitable genetic variability. Some of the promising traits in the indigenous gene
pool of South Asia are early maturity and more pods per plant and per cluster
(Bhag Singh and Rana, 1993). Extreme specificity of adaptation limits the scope
of direct introduction of exotic germplasm in the region. Sharma et al. (1993)
reported that most of the Indian lentils (microsperma type) are early maturing
and the Mediterranean macrosperma lentils are late maturing. Podding potential was higher in small-seeded genotypes than large-seeded ones and smallseeded types have better stability. This variability should be exploited to
widen the genetic base through hybridization. A wide range of variation,
including transgressive segregation, was obtained both for crop duration and
for seed size from a cross between indigenous microsperma and exotic macrosperma (Precoz) types, resulting in selection of extra short duration and
erect genotypes (gene Ert) with large seeds and tolerance to key stresses
(Sharma et al., 1993; Tufail et al., 1993; Emami and Sharma, 1999). Therefore,
introgression of exotic genes from diverse origins into locally adapted cultivars should be the approach for widening the genetic variability.
128
9.11. Breeding Methods

The methods of breeding lentil are similar to those utilized to breed other
self-pollinated crops and include pure line selection, back crossing and
hybridization, followed by bulk pedigree, single seed descent or some modification of these selection procedures. The bulk-pedigree method has been
the preferred breeding method. Porta-Puglia et al. (1993) described different
breeding methodologies for single and multiple stress resistance in coolseason food legumes. They suggested pedigree selection for disease resistance and the bulk-pedigree method for abiotic stresses. ICARDA follows
the bulk-pedigree method in which the crosses are advanced as bulks from
F2 to F4 and the selected plants are carried forward as plant progenies from
F5 onwards. Although many cultivars have been released in South Asia targeting specific traits and environments, not all are suitable for the short
duration environment. Therefore, only short duration cultivars released
after 1980 (100140 days) are listed in Table 9.2. So far, seven high-yielding
disease-resistant cultivars have been released for cultivation in Bangladesh,
29 in India, five in Nepal and seven in Pakistan.
Pure line selection

In the early stages of lentil breeding, most of the cultivars released were derived
from selection within the heterogeneous landraces. Most of the expansion in
the range of adaptation has resulted from empirical selection of superior performance of individual genotypes in new climate zones. A large number of
landraces collected from different regions were evaluated and based on multilocation testing for yield and disease reaction, and superior genotypes with
wide adaptability were commercialized. Some of the popular cultivars developed through this method are: L 9-12, T 36, BR 25, C 31, JL 1, Pant L 406, Pant L
639, Lens 830, Lens 4076, B 77, Vipasha, VL 1, K 75, JL 3 and VL 4 in India;
Barimasur 1 in Bangladesh; Shital in Nepal; and Masoor 85 in Pakistan.
Cultivars found popular in one country/region are often introduced
and acclimatized in similar conditions prevailing in another country through
ICARDA and if found suitable, are released as cultivars. Among the lentil
cultivars released through introductions from ICARDA are: Vipasha and
VL 507 in India; Mansehra 89 and Shiraz 96 in Pakistan; and Simal,
Sikhar, Khajura Masuro 1 and Khajura Masuro 2 in Nepal.
Recombination breeding
Recombination breeding, or the selectioncrossingselection cycle, consists of
controlled crossing between parents of choice followed by pedigree selection
or its various modifications in the segregating generations for the targeted
trait(s). The lentil breeding programmes generally use parents of diverse
origin with known traits with the aim to combine genes that contribute to
Country
Cultivar
Year
Pedigree
Days to
maturity
100-grain
weight (g)
Grain yield
(t/ha)
Bangladesh
Barimasur 1
Barimasur 2
Barimasur 3
Barimasur 4
Barimasur 6
Barimasur 7
Binamasur 1
Pant L 234
Asha
Pant L 639
Ranjan
LL 56
Malika
Arun
LL 147
Sapna
Jawahar Lentil 1
Pant Lentil 4
Lens 4076
Priya
Pusa Vaibhav
Garima
Narendra Masur 1
Sheri
Subrata
1991
1993
1996
1996
2006
2006
2002
1980
1980
1981
1982
1983
1986
1986
1986
1991
1991
1993
1993
1995
1996
1996
1997
1997
1998
Local selection
F3 ICARDA
Local cross
F3 ICARDA
F3 ICARDA
F3 ICARDA
Mutation
Selection from P 230
Selection from Jorhat local
L 9-12 T 8
Mutant of B 77
L 9-12 L 32-1
Local selection
Mutant of BR 25
PL 284-67 NP 21
L 9-12 JLS 2
Local selection
UPL 175 (PL 184 P 285)
PL 234 PL 639
PL 406 L 4076
(L 3875 P 4) PKVL 1
Pusa 2 NO. 4
Precoz L 9-12
JLS 1 LG 171
JLS 2 T 36
110115
110115
100105
115120
105110
105110
125130
135140
120125
140145
120125
150155
130135
125130
140145
135140
120125
140145
130135
130135
130135
135140
120130
130135
120125
1.51.6
1.21.3
2.22.3
1.71.8
1.92.0
2.22.3
1.51.6
2.3
1.5
1.7
1.7
1.9
2.7
2.5
1.8
2.7
2.5
1.7
2.8
2.7
1.8
2.6
2.6
3.4
2.8
1.51.8
1.52.0
1.72.0
2.02.2
1.82.0
2.02.5
1.52.0
2.12.5
1.71.9
2.12.3
2.02.1
1.61.8
2.12.4
2.12.3
1.92.1
2.02.3
1.51.7
2.12.4
2.12.3
2.12.4
2.32.6
2.02.2
2.02.4
2.32.5
1.82.1
India
129
(Continued)
Table 9.2. Short duration cultivars released in South Asia.
130
Country
Pakistan
Nepal
Cultivar
Year
Pedigree
Days to
maturity
100-grain
weight (g)
Grain yield
(t/ha)
JL 3
Noori
KLS 218
HUL 57
IPL 406
Masoor 85
Mansehra 89
Masoor 93
Shiraz 96
NIAB Masoor 02
Masoor 04
NIAB Masoor 06
Simal
Sikhar
Khajura Masuro 1
Khajura Masuro 2
Shital
1999
2000
2005
2005
2007
1985
1989
1993
1996
2002
2004
2006
1989
1989
1999
1999
2004
Local selection
K 75 PL 639
KLS133 L9362
Mutant of HUL 1
DPL35 EC157634/382
Local selection
Precoz
18-12 ILL 4400
ILL 5865
Precoz Masoor-85
ICARDA selection
Mutant of ILL 2580
LG 7 (ILL 3512)
ILL 4404
ILL 3746 (LG 198)
Pant L 639
ILL 2580
115120
110120
120125
121
125130
120130
90110
110120
140160
90110
100120
90110
140143
140143
125128
130134
130133
3.0
2.7
1.9
1.8
3.9
1.8
4.05.0
2.5
2.8
2.53.0
2.5
2.2
1.41.8
1.31.6
1.31.7
1.41.9
1.82.0
2.02.2
2.12.3
1.41.6
1.41.5
1.71.9
1.51.7
2.12.5
1.92.1
1.82.2
1.51.8
1.92.1
1.42.1
2.12.5
2.53.0
1.52.0
2.12.5
1.82.0
131
yield and resistance to major biotic and abiotic stresses. Crosses between genotypes from South Asia and West Asia which greatly differ in their photoperiod requirements are made either by raising them under long-day conditions
during summer in hilly areas or by providing 18 h of extended light to attain
synchrony in flowering. The first cultivar emanating from recombination
breeding in the region was Pant L 639 in 1981. This was followed by a spate of
high-yielding disease-resistant cultivars from such efforts (Table 9.2). However, the genetic base of these cultivars remained narrow as South Asian
improvement programmes have not used genes from exotic sources on a wide
scale. This bottleneck for lentil improvement in the region has recently been
broken (Erskine et al., 1998) with the introduction of exotic early flowering
materials into the Indian subcontinent. For example, flowering of Precoz, an
early maturing Argentinean landrace, is well synchronized with the local
South Asian germplasm. Studies showed that Precoz has a major gene sn for
earliness and produces early and extra early transgressive segregants that can
fit well in the short growing periods of South Asia (Sarker et al., 1999). Precoz has been used extensively to incorporate such traits as large-seed size,
disease (mainly rust) resistance and early vigour in the background of locally
adapted cultivars since 1985. Precoz was first received in India in the
1982/83 cropping season when its extra-earliness among macrosperma lentils
was noted. It must be noted that a great majority of macrosperma genotypes
received from the western hemisphere are extremely late under the conditions of the Indian subcontinent. Some of them do not flower even after 140
days from sowing when the crop matures under temperature extremes above
40C. Crosses between Precoz and local lentils have resulted in the development of a wide spectrum of extra bold and extra early cultivars in India (NDL
1 and DPL 58) and Pakistan (NIAB Masoor 02).
Mutation breeding
Mutation breeding has definite merit for simply inherited traits and where
suitable and efficient screening procedures are available to identify economically useful mutants from sufficiently large populations of mutagenized
material. There are a few reports of mutation breeding in lentil. Experience
with lentil mutagenesis has shown that the macrosperma cultivars are more
mutable than the microsperma genotypes. Mutation breeding has been successful in developing lentil cultivars with specific traits. So far, seven mutant
cultivars including three cultivars each in India (Ranjan, Arun and HUL
57) and Bangladesh (Bina Masur 1, Bina Masur 2 and Bina Masur 3) and
one in Pakistan (NIAB Masoor 2006) have been reported.
9.12. Future Perspectives

The productivity of the present cultivars is quite low and needs to be
improved through change in the existing plant type and insulation against
132
multiple stresses. Introgression of economically valuable traits through

wide crosses holds promise for a number of important traits, particularly
tolerance to abiotic and biotic stresses. This requires comprehensive screening of the available germplasm in hot spots and under controlled conditions
for directed improvement in lentil. Exciting gains in sustainable production
and expansion of the crops to new niches can be made from incorporation
of desired traits with changes in agronomic practices. Several such prospects exist for lentil in South Asia. The diseases such as rust, wilt and Ascochyta blight are key problems; however, lentil could be put into production
in the large areas that are left fallow during winter after the harvest of late
paddy. For this potential use, super-early photo-thermo-insensitive cultivars that can be grown successfully under late sown situations are needed.
Once super-early-maturing cultivars with multiple disease resistance are
available, a major breakthrough in lentil production in the rice-based cropping systems in the Indian subcontinent could be realized.
Until recently, a major limitation to genomics-enabled lentil improvement has been unavailability of locus-specific PCR-based codominant markers and precise mapping and tagging of useful genes (Ford et al., 2007).
However, recent developments in simple sequence repeat (SSR) marker
technology for lentil at ICARDA (Hamwieh et al., 2005) indicates progress
in this area and possibly will lead to further development. Genes for resistance to biotic and abiotic stresses need to be tagged with molecular markers for effective use in breeding. Some of the major genes or quantitative
trait loci (QTLs) imparting resistance to Fusarium wilt and Ascochyta blight
have been tagged (Eujayl et al., 1998; Ford et al., 1999; Chowdhury et al.,
2001). Further research on functional genomics to address some of the key
problems should be initiated. This requires availability of appropriate mapping populations, and identification and validation of trait-associated markers across different environments (Ford et al., 2007). On the technological
front, lentil needs a major breakthrough in yield levels through morphophysiological changes in plant type, and dissection of abiotic stresses into
its components. Lentil germplasm has shown tremendous genetic variation
for protein and micronutrient (iron and zinc) contents. This provides scope
for biofortification of high-yielding varieties for augmenting nutritional values of the crop.
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10
Improvement in Developed Countries
Fred J. Muehlbauer,1 Miho Mihov,2 Albert Vandenberg,3

Abebe Tullu3 and Michael Materne4
1Washington
State University, Pullman, Washington, USA; 2Dubroudja Agricultural

Institute, General Toshevo, Bulgaria; 3University of Saskatchewan, Saskatoon, Canada;
4Department of Primary Industries, Horsham, Victoria, Australia
10.1. Introduction
Cultivated lentil (Lens culinaris Medikus subsp. culinaris) had its origin in
the Near East arc where the crop was domesticated an estimated 8000 years
ago (Zohary, 1972; Ladizinsky, 1979). The lentil crop is now grown successfully on all continents except Antarctica. Lentil is adapted to most soils and
climates, but is especially important in arid and semi-arid regions. Lentil is
a quantitative long day plant and flowers in progressively longer days. It is
grown as a winter crop in the Mediterranean region, South Asia, Australia
and parts of South America. Though winter-hardy germplasm is available,
the crop is primarily spring sown in most highland regions (>850 m above
sea level) of West Asia, North Africa and in North America.
Most of the worlds lentil crop is grown in the developing countries of
West Asia, North Africa, East Africa and South Asia, where it is an important human food for local populations; while in developed countries production is targeted to specific international markets and speciality types.
Demand for the large green type with yellow cotyledons, typical of Brewer,
Laird and other similar cultivars produced in Canada and the northern
plains states of the USA, has driven production; however, increased international market demand for small red-cotyledon types has led to recent
increased production of that type in Canada, USA and Australia. Several
other types have importance for niche markets, for example the Pardina
preferred in Spain and several other types such as Eston. Domestic use of
lentil in developed countries has remained rather minimal despite extensive
domestic marketing efforts to increase consumption.
Priorities and breeding goals usually differ between regions depending
on critical location-specific problems and special considerations related to
farmers needs and consumer demands. In developing countries particularly in the West Asia and North Africa region, the major breeding goals
137
138
F.J. Muehlbauer et al.
are development of cultivars with higher yields, resistance to major diseases,

suitability for mechanical harvesting and improved residue yields because
of the value placed on lentil straw as animal feed. By contrast, improved disease resistance, reduced tendencies to lodge, reduced pod and seed shatter,
increased seed yield and improved seed quality traits are principal breeding
goals in developed countries. Techniques of breeding lentil are similar to
those used for other self-pollinating crops with some modifications.
10.2. Types of Cultivars

A wide range of lentil cultivars is used throughout the world with the small
diameter red-cotyledon type accounting for most of the production followed by the large and small yellow-cotyledon types. Green lentils range
from 4 to 8 mm in diameter and cultivars can be divided into three market
classes: large, medium and small. The typical large green lentil has lightgreen seedcoats with little or no mottling and cotyledons are yellow. There
are no truly green-cotyledon lentils in the market at the present time
although genetic stocks with green (light and deep) cotyledons are known
(see Sharma, Chapter 7, this volume). Lentils with yellow cotyledons and virtually spotless testa are called green lentil, a term that is a common market
jargon in developed countries. This expression is not used in the major lentilproducing and -consuming region of South Asia. In Canada, the large green
type typical of Laird and more recently CDC Glamis, CDC Sovereign,
CDC Grandora, CDC Plato, CDC Greenland and most recently the imidazolinone-tolerant cultivar CDC Improve have dominated production. While
in the USA, Brewer, Merrit, Pennell and Riveland are the predominant
cultivars. Large green lentils are marketed as whole seeds and so uniformity
of size, shape and colour is important for marketing. In Australia, the trend
has been towards production of the small red type for export to South Asia
and the Middle East. Lentil production in Australia began with the cultivars
Digger, Northfield (these two being the most significant), Matilda, Cobber, Aldinga, Callisto and Kye. Now these cultivars have been replaced
with Nugget and more recently Nipper. US and Canadian production is
also targeted to South Asia; however, considerable tonnage is exported to
countries of southern Europe, particularly Spain, Italy and Greece.
The medium category of green lentils include Richlea, CDC Vantage,
CDC Meteor and the imidazolinone-tolerant CDC Impress. The small
category includes Eston, Pardina and CDC Milestone and more recently
CDC Viceroy. Green lentils at harvest have yellow cotyledons with green
seedcoats that progressively turn brown with age. Browning due to ageing
will however occur only in the lentil strains having tannins in their seedcoat. Important export markets for medium and small green lentils exist in
Algeria, Italy, Greece, Morocco and Spain, as well as South America.
Canadian production of small red lentils has become equal to that of
green lentils, starting with the cultivars Redwing and Crimson, then CDC
Robin and CDC Blaze and more recently CDC Rouleau, CDC Rosetown
139
and CDC Redberry. The most recent cultivars, including CDC Imperial,
CDC Impact, CDC Impala and CDC Maxim, are tolerant to imidazolinone herbicides. US and Canadian red lentil production is targeted to South
Asia; however, considerable tonnage is exported to Mediterranean countries, Egypt and Turkey. As with green lentils, cultivars of red lentils can be
separated into small, medium and large sizes, although the size of each
group is smaller than for green lentils. This is because the so-called green
lentils owe their origin to the microsperma subspecies and the red lentils are
descendants of the microsperma subspecies which are also called Indian lentils (pilosae). Each country has preference for different sized red lentils based
predominantly on the types traditionally grown and consumed. At one
extreme Bangladesh prefers very small red lentils and Sri Lanka prefers
large red lentils. Red lentil cultivars typically have brown or grey to black
seedcoats resulting from various degrees of black spotting or mottling (see
Sharma, Chapter 7, this volume) with red cotyledons but there is small production of red lentils with pale green seedcoats. The shape of red lentils is
an important factor in the process of decortication (removal of the seedcoats) with round and thick seeds being preferred to flat and thin seeds.
There is a huge market for decorticated red lentils in South Asia where they
are sold under the name of Malka Masoor.
Cultivars such as Pardina, important in Spain, and Naslada, important in Bulgaria, and CDC LeMay, the mottled French green type, also represent distinct lentil cultivars. Black seeded (Beluga in the USA, or
Indianhead in Canada) and white-seeded zero tannin cultivars are produced on a limited scale for speciality markets. Cultivars representing each
of these quality types have been released for the local conditions in North
America, Australia and elsewhere, including Bulgaria.
10.3. Genetic Resources Utilization, Interspecific Hybridization and

Introgression
Substantial genetic resources for use in lentil breeding are maintained at the
International Center for Agricultural Research in the Dry Areas (ICARDA),
Aleppo, Syria, and also by national programmes, particularly the USA,
Canada, Australia, Russia, India and a number of other important repositories (Table 10.1). These genetic resources comprise landraces, breeding lines,
cultivars and related wild species, and represent significant genetic variation that is readily available on request. In North America and Australia,
landraces have been widely used in breeding programmes for resistance to
diseases such as Ascochyta blight and Botrytis grey mould and tolerance to
abiotic stresses such as boron toxicity and excess salinity.
Resistance to anthracnose found in Lens ervoides germplasm is being
exploited in Canada by introgressing the resistance genes into cultivated
backgrounds (Tullu et al., 2006). This successful use of L. ervoides holds promise as a source of genes for resistance to other diseases and possibly for plant
habit, biomass production and other important agronomic and marketing
140
Table10.1. Major genebanks and the estimated number of accessions of lentil as of 2005.
Country
Abbreviation
Institution
Australia
ATFCC
2
3
4
5
6
7
8
9
10
11
12
Bangladesh
Bulgaria
Georgia
Greece
Israel
India
Spain
Italy
Canada
New Zealand
Pakistan
NBPGR
IPGR
GASU
GGB
IGB
NBPGR, ARI
SIA
ISCI
PGR
NPGS
NBPGR
13
Poland
NCPGR
14
15
16
Portugal
Russia
USA
ENMP
VIR
USDA, ARS
17
18
Syria
Slovakia
ICARDA
RIPP
19
20
21
Turkey
Czech Republic
Hungary
AARI
AGRITC
ABI
The Australian Temperate Field Crops Collection, Victorian Institute for

Dryland Agriculture (VIDA), Australia
National Bank for Plant Genetic Resources Dacca, Bangladesh
Institute for Plant Genetic Resources, Sadova, Bulgaria
Georgia Agrarian State University, Tbilisi, Georgia
Greek Gene Bank, Thessaloniki, Greece
Israel Gene Bank for Agricultural Crops, Volcani Center, Bet-Dagan, Israel
National Bureau of Plant Genetic Resources, New Delhi, India
Servicio de Investigacion Agraria, Valladolid, Spain
Instituto Sperimentale per le Colture Industriali, Bologna, Italy
Plant Gene Resources, Ottawa, Canada
National Plant Germplasm System, Wellington, New Zealand
National Bank for Plant Genetic Resources, Islamabad,
Pakistan
National Centre for Plant Genetic Resources, Radzikow,
Poland
Estacao Nacional de Melhoramento de Plants, Elvas, Portugal
Vavilov Institute of Plant Industry, St Petersburg, Russia
United States Department of Agriculture, Agriculture Research Service,
Pullman, Washington, USA
International Center for Agricultural Research in Dry Areas, Aleppo, Syria
Research Institute of Plant Production, Genebank of the Slovak Republic,
Piestany, Slovak Republic
Aegean Agricultural Research Institute, Menemen-Izmir, Turkey
AGRITEC Research, Breeding and Services Ltd, Sumperk, Czech Republic
Institute for Agrobotany, Tapioszele, Hungary
No. of
accessions
3,841
1,518
532
36
97
204
8,482
603
348
2,345
678
2,576
4
215
3,338
2,484
10,113
279
615
80
866
No.
141
traits. Further exploitation of L. ervoides and the other wild Lens species
seems warranted.
10.4. Crossing Techniques in Lentil

Healthy plants with well-developed flowers are important for successful
lentil crossing. Female flowers selected for emasculation should have corolla
petals that are unopened and have reached approximately three-quarters
the length of the sepals, although this will vary depending on the parental
line (Plate 1A). The cleistogamous self-pollinating flowers are small and
care must be taken to avoid injury during emasculation. For emasculation,
the petals are carefully folded back (Plate 1C) to expose the anthers, which
are carefully removed using sharply pointed forceps (Plate 1D). More
advanced flowers with very recently dehisced anthers are chosen as a
source of pollen (Plate 1B) and can be identified quickly with some practice.
The emasculated flower (Plate 1E) is pollinated immediately. A common
technique is to use the pollen-laden stigma of the male parent as a brush to
carefully apply the pollen to the stigma of the female flower (Plate 1F). Pollen should have a bright yellow to orange appearance and should be visible
on the surface of the stigma of the female flower (Plate 1F). After pollination, the petals are carefully returned to their original positions covering the
ovary and stigma. Early to mid-morning is generally most appropriate for
lentil crossing. For further information on lentil crossing techniques and
appropriate environmental conditions see reviews by Muehlbauer et al.
(1980, 1985) and Muehlbauer and Slinkard, (1981).
Rather simple equipment is needed for lentil crossing and includes
magnifying glasses, a pair of sharply pointed forceps, a vial of 95% alcohol,
and small tags. Viewing of flower parts can be improved with visors that
provide 5 to 10 magnification. Forceps to be used for emasculation and
pollination are immersed in the alcohol between crosses to prevent contamination by foreign pollen. Small tags are used to record the parents, dates
and the identity of the person making the cross. Some crossing programmes
use colour-coded string to label the crossed flowers. Developing pods of
crossed flowers become visible in 34 days.
Several morphological markers are used to confirm hybridity of the
crossed seeds. For example red versus yellow cotyledon colour has often
been used as a genetic marker to easily identify hybrid seeds, but only when
the yellow-cotyledon parent is used as female. Likewise, the gs gene for
stem coloration is also a useful marker to identify F1 hybrids at seedling
stage where the male and female parents have purple and green stems,
respectively. Molecular markers are also an obvious approach to identify F1
hybrids and can be used effectively to eliminate selfed seeds.
Where large seed-size differences exist between parental lines, it is
advisable to use the large-seeded parental line as the female parent. Such
genotypes also have larger flowers that are easy to handle. This makes
emasculation easier and also reduces the chances for pod shatter. The larger
142
seeded parent with corresponding pod size can accommodate the large hybrid
seeds within their pod walls while the pods of the smaller seeded parent often
cannot. Some breeders use the smaller seeded parent as female because two
hybrid seeds are often set in the pods of the smaller seeded parent. When crosspollination is successful, ovary development is rapid and the developing hybrid
seeds are conspicuous in the pods. Harvesting of the crossed pods is always
done by hand when they turn yellow-brown. Harvested pods should be
allowed to dry completely in envelopes or small muslin cloth bags before the
seeds are removed. Hybrid seeds can be kept temporarily at room temperature; however, for long-term storage, lentil breeding material should be stored
at about 10C with 30% relative humidity or in a freezer at 20C.
Crossing blocks vary with the preferences of the breeder and numerous
layouts have been described that are effective in placing the female and
male parents in close proximity. When crosses are made in controlled environments, pot-grown plants are used and can be placed in close vicinity for
ease of crossing. Crossing plans in controlled environment generally include
several plantings of the parental material at predetermined intervals to
ensure synchrony in flowering of parents differing in phenology. This also
ensures availability of flowers over a longer period.
10.5. Methods Used in Lentil Breeding

Pure line selection either within local landraces or introduced germplasm
has been an important means of developing lentil cultivars with uniformity
and adaptation to local conditions. However, lentil improvement programmes established in developed countries in the past three to four
decades have been largely based on hybridization and selection. ICARDA,
USA, Bulgaria, Canada and later Australia have developed breeding programmes that rely on selection within populations derived from crosses
involving germplasm accessions with specific traits needed to solve important production or quality problems. Breeding lines and advanced material
from those programmes have been released as improved cultivars and have
found their way into breeding programmes of many developing countries.
The methods of breeding lentil are similar to those followed in the
breeding of other self-pollinated crops and include combinations of bulk,
pedigree, and single-seed descent procedures. SSD and mutation breeding
has been employed for specific purposes. The use of doubled haploids in
lentil is not yet possible, but recent progress made in developing electroporation techniques to induce doubled haploids in field pea (Ochatt et al.,
2007) and chickpea (Grewal et al., 2008) may soon be adapted to lentil.
In Canada, unadapted germplasm is introduced into the breeding
stream using a process of systematic introduction of new germplasm, or
SINGing. Since most lentil germplasm is completely unadapted to the long
day environment of the Canadian prairies, it has become a standard practice to use multi-parent crosses to dilute the genetic contribution of unadapted
parents. Three cycles of crossing are used to produce large numbers of F1
143
seeds for field screening. Using this approach it is possible to produce threeway crosses and five-way crosses in which a 50% genetic contribution to the
cross is made by a parent with the best agronomic performance. During the
crossing cycles it is possible to use gamete selection (Singh, 1994) to select
for seedcoat and cotyledon characters.
Bulk population
Bulk population, often with some modification, has been the preferred
method for lentil breeding because of the ease of application and because of
inherent difficulties when applying other methods to lentil. Bulk population breeding of lentil is simple, requires minimal record keeping, and is
not labour intensive. The simplicity and relatively low cost of the bulk
method makes it attractive to most lentil breeders and also provides sufficient seed of segregating populations for evaluation and selection under
diverse environments. This aspect is especially important for traits such as
winter hardiness, where natural selection is relied upon to identify the most
winter-hardy plants. However, caution is needed when using the system in
lentil since good seed set on individual plants within a bulk population may
not necessarily ensure retention of desirable genotypes during generation
advancement. Therefore careful monitoring of bulk populations and their
composition is important. Mass selection for seed size, shape and colour is
easily applied to bulk populations during generation advancement.
Slinkard et al. (2000) described the recombinant-derived family (RDF),
also known as the F2-derived family, method for lentil breeding whereby
early generation selection for yield is practised in F2:4, F2:5 and F2:6 families
to eliminate inferior crosses and inferior F2-derived families. Additional
screening of F2 populations and F2-derived F3 families in rows or microplots in disease nurseries can be integrated into the selection process, provided that sufficient numbers of plants produce adequate quantities of seed.
Regional trials are conducted on the best F2:7 and F2:8 families. Results of
those trials determine which F2-derived families are refined by selection for
phenotypic traits and considered for release as improved cultivars. Advantages of the RDF method include: yield testing in early generations leading
to rapid elimination of low-yielding crosses and families; and a shortened
time period from the initial cross to the final cultivar release.
Another advantage is that selection can be based on phenotypic characters
such as canopy architecture, pod distribution and lodging as early as the F3.
This favours selection for mechanical harvesting systems. It is also possible to
apply mass selection for specific seed diameter and thickness during generation advancement. Also, there is some benefit to be derived from the fact that
the resulting cultivars have a degree of heterogeneity that may improve adaptation to the environment leading to more stable yields. The heterogeneity
maintained in the resulting cultivars may be a disadvantage of the RDF
method and prevent application for Plant Variety Protection (PVP). Although
PVP has generated little interest among lentil breeders in developed countries
144
this is likely to change in Australia where Plant Breeder Rights (PBR) can play
an important role in the effective collection of end-point royalties.
The US lentil breeding programme uses a modified bulk method
whereby the populations are advanced in bulk to the F4 followed by visual
selection for desired plant and seed types. Bulks are retained in the breeding programme for advancement to the F5 and F6 with selection in each
generation for desired types. Populations of selected bulks are grown in the
glasshouse during the off-season and the resulting plants are sown into
individual plant rows in the field for further selection. Selected lines are
then evaluated for agronomic traits, disease resistance, seed quality traits
and yield. Promising selections are tested widely and depending on performance, released as improved cultivars.
Breeding methodology in Australia is also based on a bulk population
method with single pod selection at F2 and F3 and single plant selection at
F4. The F2, F3 and F4 populations are grown in the field during the growing
season and Ascochyta blight-infested lentil stubble is spread over the plots
as a source of inoculum to screen for resistance to this disease. Natural
selection for Ascochyta blight resistance is followed by mass selection for
freedom from Ascochyta blight blemishes using an electronic colour sorter.
Machine harvesting is purposely delayed to shift the population towards
reduced pod drop and seed shatter. With delayed harvest, those genotypes
in the population that are prone to shatter will have lost their seeds while
those that resist shatter will retain their seeds and be harvested, thus shifting the population towards shatter-resistant genotypes. Mass selection for
seed shape (roundness) is accomplished using sieves and selection for seed
colour is accomplished using the electronic colour sorter.
Seed from pods of F2 and F3 plants is grown over summer in tubes in
the glasshouse. Progenies from seed of single pods (selected from F2 and F3
plants) and F4 plants are sown in 5 m rows (approximately 8000) in the field
during the winter season. Selection among those progeny rows is based on
resistance to Ascochyta blight, plant height, resistance to lodging, pod drop
and shattering, vigour/biomass, and timing of flowering and maturity.
Seed from selected rows is assessed visually for seed colour, size, shape and
freedom from Ascochyta blight blemishes.
Selected lines are evaluated for yield, quality and harvestability across
the target range of environments. Stage 1 trials are unreplicated and located
at three sites, stage 2 trials have two replications and eight sites and stage 3
trials have three replications and eight sites. Row-by-column designs are
used and trials are spatially analysed. Screening for the two major diseases,
Ascochyta blight and Botrytis grey mould are done for entries in all stages
of testing in the field and controlled environments, respectively. Breeding
lines are also screened for tolerance to the major herbicides in the field and
toxicity to boron and salinity (NaCl) using soil-based media in a controlled
environment. Physical, milling and cooking qualities of the lines are assessed
in the laboratory. All potential new lentil cultivars are evaluated for response
to commonly used agronomic practices and a management package made
available to farmers to maximize the benefits of new cultivars.
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Promising selections are subjected to pedigree seed production in collaboration with PB Seeds, the commercial seed partner for the breeding
programme. The breeding programmes major focus is on small, medium
and medium-large red lentils and large green lentils; however, small and
medium green lentils and other niche types are also produced. Simple and
complex crosses are made in the glasshouse during the growing season,
which enables crosses to be made using field pollen of elite lines.
Pedigree selection
The pedigree selection breeding method is not particularly suited to lentil
breeding because of plant architecture and plasticity, which allows the plants
to occupy available space. The response of lentil plants to available space is a
disadvantage in selection because performance as individual widely spaced
plants often is entirely different from performance in more dense stands of the
same genotype. Selection based on yields of individual plants has been ineffective for lentil breeding (Erskine et al., 1990). However, selection of individual
plants for highly heritable traits such as seed size, flowering time and height is
likely to be successful. Population sizes for effective use of the pedigree method
for lentil breeding will depend on the genetics and heritability of the traits
under consideration and resources available to the breeding programme.
Single seed descent (SSD)
SSD is well suited to rapid generation advance in lentil breeding provided
there is a facility (glasshouses or off-season nurseries) for growing the populations several times in a year. The primary advantages of the SSD method
are the maintenance of genetic variation during generation advance and the
minimal space required for the method. Haddad and Muehlbauer (1981) found
more genetic variation was maintained by SSD compared to the same populations advanced by the bulk method. In their study, three SSD-derived populations had 10, 9, and 13% more erect lines, respectively, when compared with
the same hybrid populations advanced by the standard bulk method.
SSD has been used effectively to develop recombinant inbred lines
(RILs) for use in genetic linkage analyses and development of genetic maps
of the lentil genome (Eujayl et al., 1998; Kahraman et al., 2004; Hamwieh
et al., 2005). Development of RIL populations is currently underway for
mapping of the genes for resistance to rust, Stemphyllium blight, Ascochyta
blight and Sclerotinia white mould using the SSD method.
Mutation breeding
Mutation breeding merits consideration in a lentil breeding programme
where a desired trait may not exist in available germplasm. The approach is
feasible where suitable screening methods are available and can be employed
146
to evaluate large populations of mutagenized plants. Mutation breeding has

the promise of improvement of important traits in lentil without altering the
genetic background. While this may not always be true, other breeding techniques such as the backcross method may be used to return the genetic background to the desired type. Mutations have been important for genetic mapping
and genomic approaches towards identification of important gene functions.
A number of effective physical and chemical mutagens are available
including X-rays, gamma-rays, fast and slow neutrons, ethyl methane sulfonate (EMS), ethyl nitrous-urea (ENU), methyl nitrous-urea (MNU) and
sodium azide (NaN3). Effectiveness of chemical mutagens depends on
mutagen concentration and treatment conditions and the ability to identify
micro-mutations in the mutagenized material.
Eight lentil cultivars developed through induced mutations using irradiation have been registered in India, Bulgaria and Canada (Bhatia et al.,
2001; Mihov et al., 2001; Toker et al., 2007). Cultivars developed by mutagenesis reportedly vary for vegetative growth period, plant height, seed size and
colour. An interesting dwarf mutant was isolated from mutagen treatment of
Tadjikskaya (Mihov et al., 2001). The dwarf mutant has significantly shorter
internodes and increased branching. The mutant may be of interest to
researchers working on morphological development and comparative genomics. Mutants for flowering traits, plant growth habit and seed quality variations
have been described and may have similar interest in studies of genomics
and morphological development as well as value for breeding purposes.
EMS treatment was used in Canada to identify a lentil line with tolerance to imidazolinone herbicides. The trait was patented (http://www.
patentstorm.us/patents/7232942-description.html) and licensed for use in
Clearfield lentil cultivars. The trait has been transferred to cultivars in all
market classes, resulting in the release of a series of herbicide-tolerant cultivars including CDC Imperial, CDC Impact, CDC Improve, CDC Impala,
CDC Impress and CDC Maxim. Clearfield lentil cultivars are now widely
available in Canada.
The backcross method
Backcross breeding provides predictable results and is generally used for
simply inherited traits with high heritability, for example herbicide tolerance. Backcrossing of the genes for resistance to diseases such as Fusarium
wilt and viruses are prime candidates for improvement of lentil by backcross breeding. The brief history of lentil breeding often means that breeding programmes are targeting improvements in many traits simultaneously
rather than incorporating a single gene into an elite cultivar.
Population improvement and germplasm enhancement
Population improvement has not had an important role in lentil breeding
primarily because of the difficulties in making the necessary numbers of
147
crosses and the lack of an efficient male sterility/restorer system. However,

a form of cyclical recurrent selection may be warranted in order to concentrate minor genes for important quantitatively inherited traits. The process
requires a careful choice of parents, intercrossing to develop a heterogeneous population of heterozygous plants, screening for the trait of interest
and intercrossing of selected individuals to form the base population for the
next cycle of selection. The process is repeated for several cycles and during
the progression through the successive cycles mean performance is evaluated for the amount of genetic variation available for effective selection. The
resulting improved population can then be used in the breeding programme
to develop trait-specific cultivars. The method has been used successfully in
Canada to improve resistance to anthracnose. Each year, multi-parent
hybrids are used to develop F2-derived Fs micro-plots which are screened in
a field disease nursery that is inoculated with anthracnose-infested lentil
debris. Single plants from the most resistant micro-plot in the top 510 resistant populations are selected as parents to begin a new cycle of selection.
Breeders have become increasingly aware of the need to better coordinate germplasm enhancement within regional breeding programmes. The
Australian lentil breeding programme has developed a formalized approach
to incorporate germplasm enhancement using traditional and biotechnological approaches. With this approach it is hoped to locate and incorporate
new characteristics into improved cultivars rapidly. Through this mechanism the breeding programme is well positioned to collaborate with and
provide adapted backgrounds for new biotechnological advances that may
occur (e.g. genetic modification, doubled haploids, etc.) and implement the
technology. It is also expected that advanced breeding techniques and
germplasm enhancement successfully employed in other pulses will be
extended to lentil. For example developments in tissue culture of peas may
open avenues for use in lentils. Identification of the DNA sequence with
specific functions in the model species may aid in locating and cloning
genes with the same or similar function in lentils. A series of projects that use
sequence-based approaches to develop marker-based strategies for increasing genetic gain in lentil have started in Canada, the USA and Australia.
10.6. Selection Procedures for Special Traits

Traits needed for mechanized and extensive production in developed countries include predictable flowering times, tall plants with upright growth
habit and minimal pod and seed shattering. In traditional lentil-growing
regions, optimal flowering response is represented in indigenous landraces.
However, in new production areas optimal flowering response and yield
potential are determined during germplasm assessment. Flowering time in
lentil has been reviewed (Roberts et al., 1988) indicating that progress towards
flowering is predictable and depends on temperature and photoperiod. Where
major changes in crop agronomy were introduced (e.g. tolerance of diseases)
there is also a need to re-establish optimal flowering physiology under the
148
altered circumstances. In Australia, selection for genotypes that are responsive to temperature but not photoperiod for flowering is done by measuring flowering times at two key sites. Important cultivars with broad rather
than specific adaptation in new production areas include Crimson and
Eston in the USA, and Digger and Nugget in Australia.
Tall upright growth habit with resistance to lodging is an important
trait under selection in developed countries. Upright habit is essential for
adaptation to machine harvesting while reducing the potential for the pods
to come in contact with soil surfaces causing staining, seed decay and
reduced product quality. Plant height index as a selection tool has been
established by the US breeding programme. According to the procedure,
the ratio of canopy height at maturity relative to canopy height in the full
flowering stage is calculated and used as a selectable trait. Selections with
ratios close to 1.0 are considered to have good resistance to lodging while
lower ratios indicate less lodging resistance. The cultivars Pennell and Merrit were selected for their resistance to lodging using the plant height index.
In Canada, a selection technique for lodging resistance is under development.
It is based on using a roller to flatten plots just prior to flowering, then visually assessing recovery after 3 days. The cultivar CDC Redberry is highly
resistant to lodging in Canada, resulting in its quick adoption by lentil growers. In Australia, genotypes with good lodging resistance are more prone to
pod drop caused by strong winds at maturity and improved resistance to
pod drop is a key breeding aim.
Fungicides are often available for the economic control of diseases; nevertheless, genetic resistance is a major focus towards improving reliability
of yield and increase profitability. Identification of resistant/tolerant germplasm, determining the inheritance of the trait and formulating selection
procedures for use in breeding are the primary approaches used in breeding. Of particular importance in Canada and Australia is the prevalence of
Ascochyta blight caused by Ascochyta lentis, which has devastated production and product quality in those countries. Of nearly equal importance in
Canada is anthracnose caused by Colletotricum truncatum; however, resistance has been found in L. ervoides and is being transferred to cultivated
backgrounds.
Foliar fungal diseases are particularly devastating in cool and humid environments typical of the winter growing season in Australia, the cool and wet
spring conditions in Canada, and the northern plains states of the USA. The
cultivar Nipper released in Australia has good resistance to grey mould and
Ascochyta blight. Progress in finding germplasm sources with resistance or tolerance to these foliar fungal diseases has been reviewed by Tivoli et al. (2006).
Most traits needed for efficient lentil production are selected under normal field conditions using well-designed nurseries with suitable checks.
Reportedly, selection for yield under variable rainfed conditions has increased
water use efficiency through an increased response to moisture availability
(Erskine et al., 1993; Materne and McNeil, 2007). However, breeding has
increasingly focused on addressing abiotic and biotic constraints, particularly
disease. In the USA and Turkey (Central Anatolia), large yield increases
149
have been achieved by sowing lentil in winter rather than spring using
genotypes tolerant to cold temperatures (Kusmenoglu and Aydin, 1995;
Hamdi et al., 1996; Muehlbauer and McPhee, 2002). The cultivar Morton was
developed in the USA for autumn sowing into standing cereal stubble. The
standing stubble helps with winter survival by trapping snow and providing some protection from freezing temperatures and desiccating winds.
Although generally adapted to alkaline soils, lentil growth can be
affected by hostile subsoil factors such as high pH, toxic levels of boron and
salinity and sodicity. Although variation in tolerance to these factors has
been identified, breeding efforts to target these stresses has been, to our
knowledge, relatively limited. Breeding lines with improved tolerance to
boron, derived from ILL 2024, have been developed in Australia and based
on controlled environment experiments could improve yields by up to 91%
in the target regions (Hobson et al., 2006). Similarly, lines with improved
tolerance to NaCl have been developed and are soon to be released in Australia. These lines have great potential as they are the highest yielding
entries in advanced yield trials (Materne and McNeil, 2007). In the breeding
programme, boron screening involves growing plants in soil that is high in
boron and rating symptom expression, while for NaCl screening, plants are
grown in sand media to which saline water is added after emergence.
ICARDA has the world mandate for lentil improvement and the Center
has been instrumental in developing high-yielding cultivars with greater total
biomass yield, drought tolerance and resistance to disease. Since 1980, more
than 100 lentil cultivars that can be traced back to ICARDA-generated material
have been released by various national programmes. The released cultivars
represent improvement for resistance to pod shattering, lodging, greater plant
height, enhanced yield potential and improved adaptation. As a result large
production increases have been realized in areas of Turkey, South Asia and
other countries of West Asia and North Africa. Developments in lentil germplasm have also benefited the developed countries where the material is routinely used in breeding programmes to broaden the genetic base for traits
under selection.
10.7. Trait Selection for Domestic Markets

For most domestic markets in developed countries, large-seeded lentils with
yellow cotyledons and light-green seedcoats are preferred; however, there
is increased interest in small red-cotyledon cultivars. Many of the traits
needed for domestic markets in developed countries are the same as those
needed for export markets.
Mass selection for physical seed characteristics is done by hand picking,
using sieves or mechanization using small-scale equipment such as gravity
tables or electronic colour sorters. Since most of the lentil crop in developed
countries is exported, trait selection is export-market driven. The important
traits under selection are mostly cosmetic and concern outward physical
appearance, minimal mechanical damage to the seeds during harvesting
150
and processing, short cooking times and good visual appearance after cooking. Recent work on the genetics of green seedcoat colour in Canada showed
that this trait had medium to high heritability (Davey, 2007).
Selection in breeding programmes for market traits use, methods developed by individual breeding programmes with most based on the procedures
developed at ICARDA. Procedures used for determining percentage water
uptake, time for cooking and qualities after cooking have been developed at
ICARDA (Erskine et al., 1985). A small portable decortication machine is used
to decorticate small red lentils to determine percentage yield during the process and also to assess colour of the resulting product. Breeding for processing
and cooking quality will become increasingly important as markets and consumers have more choice and become more sophisticated in their preferences.
10.8. Trait Selection for International Markets

Breeding programmes in developed countries monitor their breeding material to ensure that new cultivars have quality traits that are acceptable to
international markets. In the case of small red-cotyledon cultivars, consideration is given to the yield of the split product and the depth of red coloration of the cotyledons after decortication. Quality traits such as protein and
amino acid concentrations are a lesser concern.
Breeders in North America are concerned with large-seeded green lentils in the size range of 78 mm in diameter and with yellow cotyledons and
light-green seedcoats that lack mottling. While large green lentils have been
the primary focus of breeding in North America, within the past decade
other types particularly the small-seeded red-cotyledon type, have taken on
greater importance for export. In addition to the large green and small red
types, medium-sized yellow lentils typical of the cultivar Eston have
gained some importance while the small brown type with yellow cotyledons typical of Pardina have been widely grown in the USA primarily for
export to Spain, and small marbled types with yellow cotyledons typical of
CDC LeMay have been grown for export primarily to France. In contrast
to North America, lentil production in Australia is predominantly of the
red type but the first widely adapted green lentil Boomer has recently been
released for production and to develop export opportunities.
For the smaller red-cotyledon types, seed thickness is most important
since there is a direct correlation between decortication yield and seed thickness with the thicker seeds generally having higher decortication yields and
less cotyledon chipping. Diameter is also important as markets range from
those that prefer larger splits to those that prefer small round lentils that
can be decorticated without splitting (the so-called footballs and referred
to in India as Malka Masoor).
At ICARDA and in Canada, efforts are underway to understand the
heritability of the content of micronutrients such as iron and zinc in lentil.
In many countries, lentils are regarded as important sources of micronutrients,
and research of this type will expand in future. Once heritability estimates
151
are established, appropriate breeding strategies for biofortification can be

devised. In future, this information may lead to the development of breeding programmes with specific objectives for micronutrient content.
10.9. Linkages and Networks with International Programmes

With ICARDA having the world mandate for improving lentil it is important that national lentil research programmes and particularly the breeding
programmes have close linkages with the Center. Based on close collaboration, large and sustainable increases in lentil production have been achieved
in Canada, the USA, Australia, Turkey and South Asia. Collaboration with
ICARDA has formed the basis of breeding in developed countries and currently strong ties of collaboration exist there. The most valued result of linkages of ICARDA with breeding programmes in developed countries has
been the germplasm exchanges for mutual benefit and discussion of problem areas leading to research collaboration.
ICARDA maintains the most extensive world collection of lentil germplasm that includes the largest number of accessions of wild species and
also the largest collection of landraces and improved germplasm. This resource
has been used extensively for breeding and genetic investigations. Linkage
projects with ICARDA have been able to map the quantitative trait loci
(QTL) for tolerance to cold and winter injury in lentil and to foster the introduction of winter germplasm into the USA, Turkey and Central Asia. Other
productive linkage projects have led to the first genetic linkage map of the
lentil genome (Hamwieh et al., 2005) and the mapping of genes for resistance to Fusarium wilt. An ongoing linkage project between ICARDA and
the USA is working towards understanding the genetics of resistance to
rust and Stemphylium blight in lentil and placing the relevant resistance
genes on the lentil genetic map.
An effort is currently underway in Canada in collaboration with the
USA and Australia to identify genetic stocks and hybrid populations that
can be used for in-depth genomics research on lentil which is expected to
improve our understanding of the lentil genome and how it compares to
other cool-season food legumes and to the model legume species, Medicago
truncatula and Lotus japonicus.
10.10. Summary and Conclusions

Lentil breeding in developed countries has focused on developing cultivars
to meet international market demands, increasing yields through improved
disease resistance and improving quality traits. Systematic germplasm collection in the centre of origin for the wild species has provided a means to
broaden the genetic base for breeding and selection. The expanded lentil
germplasm pool is being exploited for resistance to disease, for example
resistance to anthracnose found in L. ervoides. Additional exploitation of the
152
wild species may be forthcoming for resistance/tolerance to other biotic

stresses and especially abiotic stresses such as heat and drought.
Progress has been made in recent years towards understanding the
genetics of important traits in lentil and location of the genes on the lentil
genetic map. While lentil genomics is currently far behind that of the major
legume crops such as pea, chickpea, dry bean and soybean, efforts are underway in several labs in developed countries towards marker development
and genetic mapping. Markers closely linked to genes of interest are being
sought for such traits as resistance to Fusarium wilt, Stemphyllium blight,
rust, Ascochyta blight, anthracnose, winter hardiness, resistance to drought
and a number of other important traits.
Lentil research programmes have not identified improved nitrogen fixation as an important priority either by management or genetic manipulation.
With increased costs of nitrogen fertilizers this aspect of lentil research may
acquire increased importance. Current estimates of nitrogen fixation by lentil
crops are few and indicate only nominal contributions to soil nitrogen status.
Excellent cooperation has been developed among lentil breeders in developed countries and exchanges of breeding materials have taken place on a
regular basis as well as exchanges of information at regional, national and
international conferences and workshops.
In most countries breeding programmes are funded by local government and much of the advisory, cultivar-testing and seed-distribution roles
are also controlled by government agencies. However, in developed countries private investment is important. In Canada, the USA and Australia,
farmers invest in lentil research, including breeding, through research levies collected on production. In these and other countries private companies
have increased investment in the cultivar release process by undertaking
the multiplication of new cultivars, distribution of seed and, in the case of
Australia, collecting royalties for investment back into agricultural research.
In Canada, the Saskatchewan Pulse Growers Organization supplies operational funds for the lentil breeding programme and in return, receives the
rights to commercialize all lentil breeding lines produced by the breeding
programme. In the USA, the USA Dry Pea and Lentil Council collects an
assessment on the first sale of the lentil crop with the funds collected being
allocated to marketing and research programmes.
References
Bhatia, C.R., Maluszynski, M., Nichterlein, K. and van Zanten, L. (2001) Grain legume
cultivars derived from induced mutations and mutations affecting nodulation.
Mutation Breeding Review 13, 144.
Davey, B.F. (2007) Green seed coat colour retention in lentil (Lens culinaris). MSc.
thesis, University of Saskatchewan, Saskatoon, Saskatchewan, Canada.
Erskine, W., Williams, P.C. and Nakhoul, H. (1985) Genetic and environmental
variation in the seed size, protein, yield and cooking quality of lentils. Field
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Erskine, W., Isawi, J. and Masoud, K. (1990) Single plant selection for yield in lentil.
(1993) Current and future strategies in breeding lentil for resistance to biotic and
abiotic stresses. Euphytica 73, 127135.
Eujayl, I., Baum, M., Powell, W., Erskine, W. and Pehu, E. (1998) A genetic linkage
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Hamdi, A., Kusmenoglu, I. and Erskine, W. (1996) Sources of winter hardiness in wild
markers and the localization of fusarium vascular wilt resistance. Theoretical and
Hobson, K., Armstrong, R. and Nicolas, M. (2006) Response of lentil (Lens culinaris)
germplasm to high concentrations of soil boron. Euphytica 151, 371382.
Kahraman, A., Kusmenoglu, I., Aydin, N., Aydogan, A., Erskine, W. and Muehlbauer,
F.J. (2004) QTL mapping of winter hardiness genes in lentil. Crop Science 44,
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Kusmenoglu, I. and Aydin, N. (1995) The current status of lentil germplasm exploitation
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Ladizinsky, G. (1979) The origin of lentil and its wild genepool. Euphytica 28, 179187.
Materne, M. and McNeil, D.L. (2007) Breeding methods and achievements. In: Yadav,
S.S., McNeil, D.L. and Stevenson, P.C. (eds) Lentil: an Ancient Crop for Modern
Times. Springer, Dordrecht, The Netherlands, pp. 241253.
Mihov, M., Stoyanova, M. and Mehandjiev, A. (2001) Some results of application of
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Muehlbauer, F.J. and McPhee, K.E. (2002) Future of North American lentil. In:
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Slough, UK, pp. 6990.
Muehlbauer, F.J., Slinkard, A.E. and Wilson, V.E. (1980) Lentil. In: Fehr, W.R. and
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Muehlbauer, F.J., Cubero, J.I. and Summerfield, R.J. (1985) Lentil (Lens culinaris Medic.).
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Ochatt, S., Pech, C., Conreux, C. and Jacas, L. (2007) Towards the efficient production
of androgenetic doubled haploid plants from pea (Pisum sativum L.) anthers. In:
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Roberts, E.H., Summerfield, R.J., Ellis, R.H. and Stewart, K.A. (1988) Photothermal time
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Tullu, A., Buchwaldt, L., Lulsdorf, M., Banniza, S., Barlow, B., Slinkard, A.E., Sarkar,
A., Taran, B., Warkentin, T. and Vandenberg, A. (2006) Sources of resistance to
anthracnose (C. truncatum) in wild Lens species. Genetic Resources and Crop
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culinaris. Economic Botany 26, 236332.
11
Rebecca Ford,1 Barkat Mustafa,1 Michael Baum2 and P.N. Rajesh3

1The
University of Melbourne, Victoria, Australia; 2International Center for

Agricultural Research in the Dry Areas (ICARDA), Aleppo, Syria; 3Washington State
University, Pullman, Washington, USA
11.1. Introduction
Advances have been made towards understanding the lentil genome and
the development and application of molecular markers to advance breeding
strategies. However, the routine application of markers to lentil breeding
programmes worldwide is limited, most likely because of lack of resources
for broad validation and implementation. Meanwhile, rapid biotechnological advances are fast transforming the field of lentil molecular genetics into
an integrative science resulting from the fusion of statistics, computer science, mathematics, genomics and biology. The molecular maps, genomics
approaches and the availability of gene sequences from lentil as well as the
full genomes of the model organisms are beginning to illuminate the complex and intertwined nature of responses to both biotic and abiotic stimuli.
In particular, the utilization of recently available gene sequences in lentil is
playing a key role in broadening our understanding of the complexity of
the biological systems they underpin.
11.2. Genomics and Gene Identification

One method to identify functionally associated genes is global gene expression profiling, usually performed at the mRNA level. Essentially, this aims to
identify RNAs that are present in a specific tissue sample at a particular time
and in response to a particular stimulus. Gene expression is the cellular process that transcribes the genetic information of the DNA into short-lived
mRNA and from mRNA to proteins, which ultimately determines the functionality of the gene. Therefore, characterizing the RNA population under a
particular environmental and/or developmental condition may provide a
window to understand the dynamic function of genes as well as their mutual
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role in a particular regulatory system. Gene expression profiling holds tremendous promise for dissecting the regulatory mechanisms and transcriptional networks that are involved in defence responses to pathogen attack or
physiological responses to abiotic stress such as drought, cold or salinity.
Various approaches have been developed for analysing differential
gene expression. These include: cDNA-amplified fragment length polymorphism (cDNA-AFLP) (Bachem et al., 1998), suppression subtractive hybridization (SSH) (Diatchenko et al., 1996), and serial analysis of gene expression
(SAGE) (Velculescu et al., 1995), differential display (Liang and Pardee,
1992; Walsh et al., 1992), massively parallel signature sequencing (MPSS)
and microarray technology (Schena et al., 1995). Of these, microarray technology has become the method of choice for large-scale systemic analysis of
differential gene expression profiling. Microarrays comprise DNA probes
immobilized to a solid substrate (Shalon et al., 1996; DeRisi et al., 1997). To
these probes, fluorescently labelled nucleic acid molecules (typically cDNA
prepared by reverse-transcription or mRNA) are hybridized. Implemented
in the context of well-designed experiments, microarrays may provide highthroughput and simultaneous analysis of mRNA for hundreds or thousands
of genes. This method is semi-quantitative, sensitive to low abundance transcripts that are represented on a given array and has been successfully used
to study plant responses to various biotic and abiotic factors in Arabidopsis
thaliana (Reymond et al., 2000; Cheong et al., 2002; Seki et al., 2002), Medicago
truncatula (Fedorova et al., 2002; Kster et al., 2004), soybean (Glycine max)
(Maguire et al., 2002; Thibaud-Nissen et al., 2003) and chickpea (Cicer arietinum) (Coram and Pang, 2005; Mantri et al., 2007).
Two major types of microarrays are used: cDNA microarrays and oligonucleotide microarrays. cDNA arrays comprise complementary DNA
(cDNA) probes of ~3005000 nucleotides, each representing a gene, that are
immobilized to a solid surface such as a glass slide, nitrocellulose or nylon
membrane. The probes are usually PCR products generated from cDNA
libraries or from genomic DNA. Oligonucleotide microarrays or GeneChips
comprise 2080 nucleotide probes synthesized either in situ (on-chip), by
photo-lithography (Fodor et al., 1991) or by conventional synthesis followed
by on-chip immobilization. On the chip, a gene is represented by a set of
oligos, thus conferring greater specificity. Following hybridization of the
probe (usually mRNA or cDNA), GeneChip arrays employ a single channel
detection system. In general, the oligonucleotide arrays require extensive
sequence data and computational input to produce the gene-specific oligonucleotide probes, and the manufacture and data analyses of arrays require
proprietary technology. Indeed, the cost of oligonucleotide microarray fabrication and usage is substantially higher than for cDNA microarrays.
Alternatively, cDNA microarrays may be constructed using existing
cDNA libraries, thus fabrication is dependent upon the availability of a suitable
cDNA clone collection. They may also be used in two-colour co-hybridization
experiments that allow direct comparison of transcript abundance among two
mRNA populations of interest (treatment and control) on the same slide.
This approach helps to remove experimental variability introduced during
157
array fabrication and individual hybridizations. A cDNA microarray system measures the relative increase or decrease in mRNA between different
treatments or tissues of interest.
A prerequisite for designing a cDNA microarray is the existence of a
suitable cDNA library or expressed sequence tag (EST)-enriched library that
has been constructed to contain genes related to the biological condition
under investigation. A search of GenBank for lentil sequences (conducted
on 06/10/07) returned no ESTs and only 485 genome survey sequences
(GSS). In comparison, there were 1960 ESTs and 48,339 GSS from chickpea,
1333 ESTs and 50,208 GSS from cowpea (Vigna unguiculata), and 87,189 ESTs
from field pea (Pisum sativum). Thus, lentil may be regarded as an orphan
crop when it comes to gene discovery and functional genomics studies.
However, considerable genome sequence and ESTs are available for the
model legume species Medicago truncatula, which will benefit lentil through
comparative and syntenic approaches (Zhu et al., 2005; Phan et al., 2007).
Medicago truncatula cDNA microarrays (commercially available or custom
made) may thus be used to study gene expression in lentil.
Although M. truncatula microarrays have not yet been used to study
gene expression in lentil, a multi-species cDNA microarray (PulseChip) was
used to identify homologous genes differentially regulated in lentil in
response to the important fungal pathogen Ascochyta lentis (B. Mustafa, 2007,
unpublished results). The PulseChip comprised ESTs from cDNA libraries
enriched for genes involved in the reactions between chickpea and Ascochyta
rabiei, and between Lathyrus sativus and Mycosphaerella pinodes. Differential
gene transcript profiles were assessed among the resistant (ILL 7537) and
susceptible (ILL 6002) lentil genotypes at 6, 24, 48, 72 and 96 h after inoculation (hai) with A. lentis (AL4 isolate). The non-redundant differentially
expressed genes for each accession and time point were hierarchically clustered using Euclidean metrics. In total, 25 differentially expressed sequences
were up-regulated and 56 were down-regulated in ILL 7537 and 26 were upregulated and 44 were down-regulated in ILL 6002. Several candidate
defence genes were characterized from lentil including; a b-1, 3-glucanase, a
pathogenesis-related protein from the Bet v I family, a pea disease resistance
response protein 230 (DRR230-a), a disease resistance response protein
(DRRG49-C), a PR4 type gene and a gene encoding an antimicrobial SNAKIN2
protein, all of which have been fully gene sequenced (Ford et al., 2007). Several transcription factors were also recovered at 6 hai and future aims will be
to further biologically characterize these and earlier responses, to gain a
comprehensive understanding of the key pathogen recognition and defence
pathways to A. lentis in lentil. Also, the full-length gene sequences will be
used in transgenic studies to further characterize function.
11.3. Genetic Manipulation through Transformation

Commonly, the particle bombardment and the Agrobacterium tumefaciens
infection methods have been used to introduce genes with novel functions
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not readily available in the gene pool of that species. With the explosion of
sequence information available in the databases, transformation systems
have also become useful tools to study gene function via RNA interference
knockout, T-DNA insertion or transforming a genotype lacking a particular gene. Thus a robust, reproducible and efficient transformation system
combined with a protocol to regenerate complete fertile plants from transformed cells is essential to fully study plant gene functions.
Following the initial report of shoot regeneration (Bajaj and Dhanju,
1979) from apical meristems, shoot regeneration was achieved routinely
with various explants such as apical meristems (Bajaj and Dhanju, 1979),
stem nodes (Polanco et al., 1988; Singh and Raghuvanshi, 1989; Ahmad et al.,
1997), cotyledonary node (Warkentin and McHughen, 1992; Sarker et al.,
2003a), epicotyls (Williams and McHughen, 1986), decapitated embryo,
embryo axis and immature seeds (Polanco and Ruiz, 2001) as well as cotyledonary petioles (Khawar and zcan, 2002). The induction of functional
roots on in vitro-developed shoots has been the most challenging part of
lentil micropropagation. To date, no reliable and reproducible efficient rooting protocol is available. The difficulty to induce roots is thought to be associated with the use of cytokinins to obtain multiple shots from the initial
explants (Mohamed et al., 1992; Sarker et al., 2003b). Among the several
studies conducted on root induction from shoots, Fratini and Ruiz (2003)
reported 95% rooting efficiency from nodal segments cultured in an inverted
orientation in media with 5 M indole acetic acid (IAA) and 1 M kinetin
(KN). Sarker et al. (2003b) reported 30% rooting efficiency on Murashige
and Skoog (MS) medium supplemented with 25 mg/l indole butyric acid
(IBA). More recently Newell et al. (2006) obtained 100% rooting efficiency
on nodal micro-cuttings placed inverted in a mixture of sphagnum peat,
coarse river sand and perlite at a 0.5:2:2 ratio, and concluded that the
improved rooting efficiency was due to greater aeration.
To date, transformation of lentil has been reported through A. tumefaciens-mediated gene transfer (Warkentin and McHughen, 1992; Lurquin et al.,
1998; Sarker et al., 2003a) and biolistic transformation including electroporation (Chowrira et al., 1996) and particle bombardment (Gulati et al., 2002;
Mahmoudian et al., 2002). Warkentin and McHughen (1992) first reported
the susceptibility of lentil to A. tumefaciens and later evaluated a number of
explant types including: shoot apices, epicotyl, root, cotyledons and cotyledonary nodes. All explants showed transient b-glucuronidase (GUS) expression at the wound sites except cotyledonary nodes, which were subsequently
transformed by Sarker et al. (2003b). ktem et al. (1999) reported the first
transient and stable chimeric transgene expression on cotyledonary lentil
nodes using particle bombardment. Gulati et al. (2002) reported the regeneration of the first fertile transgenic lentil plants on MS medium with 4.4
M benzyladenine (BA), 5.2 M gibberellic acid (GA3) and chlorsulfuron
(5 nM for 28 days and 2.5 nM for the rest of the culture period), followed by
micrografting and transplantation in soil.
Most recently, Khatib et al. (2007) have developed herbicide-resistant
lentil through A. tumefaciens-mediated transformation. This was achieved
159
with a plasmid construct pCGP1258, harbouring the bar gene conferring

resistance to the herbicide glufosinate ammonium that was transformed
using A. tumefaciens strain AgL0. Three lentil lines, ILL 5582, ILL 5883 and
ILL 5588, were used and a high selection pressure of 20 mg/l of glufosinate
was applied to the explants for 18 weeks. Survival shoots were subsequently
grafted onto non-transgenic rootstock and plantlets were transferred to soil
and acclimatized. The presence of the transgene was confirmed by PCR and
the gene function was confirmed via herbicide application.
11.4. Molecular Screening of Existing Germplasm Resources

The International Center for Agricultural Research in the Dry Areas
(ICARDA) is participating in the genetic diversity analysis of the global lentil germplasm collections held by the Consultative Group of International
Agricultural Researchs (CGIAR) research centres. As a part of Subprogram
1, of the CGIARs Generation Challenge Program, a project will identify a
composite collection of germplasm for individual crops and characterize
each composite set using anonymous molecular markers. The overall goal
of this project is to study diversity across given genera and identify genes
for resistance to biotic and abiotic stresses that can be used in crop improvement programmes. ICARDA is responsible for creating the composite collection for lentil and analysing these accessions for diversity using microsatellite
markers. ICARDA has the global mandate for lentil improvement and holds
the largest global collection of genetic resources with >11,000 accessions.
From this collection, a global composite collection of 1000 lentil accessions
landraces, wild relatives, elite germplasm and cultivars has been established
at ICARDA (Furman, 2006). These will be analysed for molecular diversity
at 30 simple sequence repeat (SSR) loci to: (i) obtain detailed information on
baseline levels of genetic variability within the composite collection of lentil; (ii) determine geographic patterns in the genetic structure of the composite collection; and (iii) provide genome-wide marker information for
future analysis of stress functional and comparative genomics in the Challenge Program.
11.5. Molecular Markers: New Resources

Most recently, many resources have gone towards the development of microsatellite and gene-specific type markers in lentil, preferentially for the
development of markers for use in marker-assisted trait selection. Microsatellite or SSR markers are generally codominant, unilocus, multi-allelic and
species specific. Produced from primers designed to the flanking sequence
of mainly di- and trinucleotide repeats, they have become a marker of choice
in many species to gain understanding of genetic relationships, evolutionary insights and for molecular mapping.
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The successful isolation of microsatellite markers involves several distinct steps:

1. Constructing and screening the small insert genomic library with SSR
motifs.
2. Sequencing the positive clones.
3. Designing primers that can amplify SSR loci.
4. Determining polymorphic SSR primers.
At each stage, there is the potential to lose microsatellite loci, and thus the number of loci that will finally constitute the working primer set will be a fraction of
the original number of clones sequenced. Therefore, the development of microsatellite markers requires a considerable amount of laboratory effort.
Constructing a small insert genomic library is advantageous for microsatellite cloning for two reasons. First, a small insert library can contain a
small array of microsatellites that are more stable in Escherichia coli. Second,
positive clones can be sequenced without subcloning (Weising et al., 1998).
For the library construction, various restriction enzymes, such as PstI, TaqI,
AluI, RsaI, MobI and Sau3AI, have been used in different plant species (Jarret and Bowen, 1994; Rder et al., 1995; Winter et al., 1999). In general, restriction enzymes that are four-base-pair cutters are superior to six-base-pair
cutters for acquiring a large number of small genomic fragments that represent the entire genome (Weising et al., 1998). However, depending on the
genomic sequence, different four-base-pair cutters may have different efficiencies with regard to cloning microsatellites.
A Sau3AI genomic library was constructed from the cultivated accession ILL 5588 and screened with (GT)10, (GA)10, (GC)10, (GAA)8, (TA)10 and
(TAA) probes. Dinucleotide repeats were observed more frequently than
trinucleotide repeats or other motifs (Table 11.1). The microsatellite motifs
were classified as perfect, imperfect, compound perfect or compound
imperfect repeats according to the modified classification of Weber (1990).
The simple/perfect repeats were predominant (56.8%), followed by compound/perfect (16.1%). The compound/imperfect (12.7%) occurred least
often (Table 11.1). Among the perfect repeats, (CA/GT)n motifs were the
most abundant, comprising 24.2% of the isolated clones, followed by (AT/
TA)n repeats (8.9%). Most recently, 126 new SSR markers were generated
using a magnetic bead capture method at Washington State University by
the United States Department of Agriculture (USDA) Agriculture Research
Service (ARS) under the direction of Weidong Chen and P.N. Rajesh.
EST-based intron-targeted amplified polymorphic (ITAP) markers have
also recently been developed from close relative species and applied to lentil. ESTs were compared from phylogenetically distant M. truncatula, Lupinus albus and G. max species to produce 500 ITAP markers that could be
applied to related temperate legume species such as lentil (Phan et al., 2007).
Also, 126 M. truncatula cross-species markers were used to generate comparative genetic maps of lentil (Lens culinaris Medik.) and white lupin (L.
albus Linn.) and macrosyntenic relationships between lentil and field pea
were observed (Phan et al., 2007).
161
Table 11.1. Frequency of microsatellite motif type observed in the lentil genomic library
(Source: M. Baum, 2007, unpublished results).
Type
Microsatellite motif
Simple
Perfect
CA/GT
CG/GC
CT/GA
CTT/GAA
AT/TA
ATT/TAA
Others types
Total
Imperfect
CA/GT
CG/GC
CT/GA
CTT/GAA
AT/TA
ATT/TAA
Other types
Total
Compound
Perfect
Imperfect
Total
Overall total
Number
Occurence (%)
57
2
7
3
21
7
37
24.2
0.8
3.0
1.3
8.9
3.0
15.7
134
56.8
21
0
1
0
3
1
8
8.9
0.0
0.4
0.0
1.3
0.4
3.4
34
14.4
38
30
68
16.1
12.7
28.8
236
100
11.6. Genome and Gene Mapping

In general, whole genome analysis is intended for the following four purposes: (i) development of molecular markers such as restriction fragment
length polymorphism (RFLP), EST, resistant gene analogue (RGA) and single nucleotide polymorphism (SNP); (ii) genetic mapping of the molecular
markers and traits using suitable mapping populations; (iii) gene discovery;
and (iv) sequencing. Furthermore, development of whole genome maps is
not only useful to identify molecular markers linked to the traits of interest
but also for genome characterization such as to determine the recombination rate at different parts of the genome, to classify the genome into gene
rich and gene poor regions and to discover the repetitive element abundant
regions.
Since lentil has little genomic sequence available, the initial genetic
mapping efforts have been initiated with arbitrary markers such as random
amplified polymorphic DNA (RAPD), AFLP and inter simple sequence
repeat (ISSR) markers which are designed to scan the entire genome and
amplify multiple bands at low stringent conditions. Although such random
markers are preferred for initial mapping analysis, they are often not consistent or reproducible and the results vary from lab to lab. However, the
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newly developed gene/locus specific markers are reproducible and represent definite genomic regions. Although a few agronomically important
traits are governed by single genes, most are quantitative in nature and
governed by quantitative trait loci (QTL), which are influenced by environmental factors. Since the expression of a QTL is likely to vary among populations and environments, their genomic location and effect must be
determined for a specific genetic background and environment (Bagge et al.,
2007). In such cases, mapping arbitrary markers will be less informative
than mapping ESTs in association with traits of interest. In addition, if the
genes are already available for phenotypic traits, designing function-associated
markers will have tremendous application in marker-assisted breeding.
These markers are essentially polymorphic sites within the genes that contribute directly to phenotypic trait variation (Bagge et al., 2007). Hence, in
order to effectively apply genomics in breeding, the availability of genomic
tools and extensive genomic information is required including genome linkage maps that are of usable density, built with robust and reproducible markers that may be transferable for integration across genetic backgrounds. A
summary of the available comprehensive linkage maps and the types of
markers and crosses used in their construction is provided (Table 11.2).
Although genetic mapping (linkage analysis) began in lentil in 1984
(Zamir and Ladizinsky, 1984), DNA-based molecular markers were not
used until 1989. Havey and Muehlbauer (1989) mapped RFLP markers generated from a genomic library using an F2 population derived from an interspecific cross (Lens orientalis L. culinaris). In spite of the advantage of
developing a linkage map rapidly, an F2 population is not a permanent
resource for long-term mapping studies and limits the number and type of
phenotypes that may be assessed. Eujayl et al. (1998a) generated the first linkage map using recombinant inbred lines (RILs) with 177 markers (89 RAPD,
79 AFLP, six RFLP and three morphological markers). Of late, there have
been significant efforts to map single locus specific markers. As mentioned
Table 11.2. Published genetic linkage maps for lentil; mapping populations and types of
markers mapped.
Population mapped
Marker types mappeda
Citation
Interspecic F2
Inter-subspecic RILb
Intraspecic F2
Intraspecic RIL
Inter-subspecic F2
Inter-subspecic RIL
Intraspecic RIL
RFLP, isozymes, morphological

RFLP, RAPD, AFLP
RAPD, ISSR, RGA
RAPD, ISSR, AFLP
RAPD, ISSR, AFLP, SSR
AFLP, SSR
SSR, ITAP
Havey and Muehlbauer (1989)

Eujayl et al. (1998a)
Rubeena et al. (2003)
Kahraman et al. (2004)
Durn et al. (2004)
Hamwieh et al. (2005)
Phan et al. (2007)
a AFLP, amplied fragment length polymorphism; ISSR, inter simple sequence repeat; ITAP,
intron-targeted amplied polymorphic; RAPD, random amplied polymorphic DNA; RFLP, restriction
fragment length polymorphism; RGA, resistant gene analogue; SSR, simple sequence repeat.
b RIL, recombinant inbred line.
163
previously, Phan et al. (2007) recently constructed the first linkage map
using sequence-based and gene-specific markers. P.N. Rajesh (2007, unpublished results) mapped PCR-based RFLP markers, cross-species gene specific (CSS) markers and SSR markers from other closely related species such
as pea and M. truncatula using an intraspecific F2 population derived from
crossing Pardina Pennell. Such cross-species markers will help to establish macrosynteny between lentil and other legumes. In addition, SNPs
were discovered from 16,158 bp CSS and 4270 bp RFLP sequences. One SNP
was detected every 311 bp in CSS and every 251 bp in RFLP sequences
between the two cultivars. Also, one SNP was detected every 281 bp in CSS
between the parental lines, ILL 5588 and L692-16-1(s), used for map construction by Hamwieh et al. (2005).
At ICARDA an F2 population of a wider cross, L92-013 (ILL 5588 (L.
culinaris) L692-16-1(s) (L. orientalis)) was used to develop RILs with an
increased chance for polymorphism among parents and segregating progeny by advancing individual F2 plants to F6 using single seed descent (SSD)
(Eujayl et al., 1998a). Using a logarithmic odds (LOD) score of 4.0 and maximum distance of 25 cM, a total of 177 markers (89 RAPD, 79 AFLP, six RFLP
and three morphological markers) were mapped in seven major linkage
groups covering 1073 cM. Marker distances ranged from 0.3 cM to 23.4 cM
with an average spacing of 6 cM. RAPD markers were more evenly distributed among linkage groups than AFLP markers, which tended to cluster
particularly in linkage group 2. The morphological markers, pod indehiscence (Pi), seedcoat pattern (Scp) and flower colour (W) were also mapped.
Building on this, another map comprising 283 markers distributed over 14
linkage groups was constructed, including 39 microsatellite markers
(Table 11.3) and five morphological markers of seedcoat pattern (Scp),
flower colour (W), pod indehiscence (Pi), Fusarium wilt resistance (Fw) and
radiation frost tolerance locus (Rf). The map was constructed using 86
recombinant inbred lines derived from the cross ILL 5588 L 692-16-1(s),
which had been previously used for the lentil linkage map (Eujayl et al.,
1997, 1998a, b, 1999). The new map spanned 750.5 cM (Kosambi function)
with an average distance between two markers of 2.7 cM. The Fusarium
vascular wilt resistance was localized on LG 6, and this resistance gene was
flanked by microsatellite marker SSR59-2B and AFLP marker p17m30710 by
a distance of 8.0 cM and 3.5 cM, respectively. Further analysis for the association between these markers and Fw was confirmed. However, only
SSR59-2B was closely linked with Fw at the estimated linkage distance 19.7
cM (Fig. 11.1; Hamwieh et al., 2005).
11.7. Trait Mapping, QTL Analysis and Marker-assisted Breeding

In lentil, several morphological markers exist such as colour of the cotyledon (Yc), anthocyanin in stem (Gs), seedcoat pattern or spotting (Scp), pod
dehiscence-indehiscence (Pi), early flowering (Sn) and ground colour of the
seed (Ggc). These were mapped onto the lentil genome (Eujayl et al., 1998a;
164
R. Ford et al.
Table 11.3. Distribution of SSR markers within the integrated linkage map of lentil (Source:
Hamwieh et al., 2005).
Linkage group
Length of the linkage

group (cM)
Total no. of
markers
No. of SSR
markers
1
2
3
4
5
6
7
8
9
10
11
12
13
14
93.3
63.5
171.9
90.1
69.6
98.3
47.4
41.6
26.5
13.4
8.5
3.5
7.7
15.2
50
47
41
45
19
35
21
10
3
3
2
2
3
2
12
4
4
5
2
4
1
5
1
0
0
0
0
1
Total
750.5
283
39
Durn et al., 2004). These traits exhibited monogenic dominant mode of

inheritance and hence are qualitative in nature (Durn et al., 2004). Other
mapped qualitatively inherited traits include; Fusarium vascular wilt resistance (Fw) (Eujayl et al., 1998b), anthracnose disease resistance (Lct-2) (Tullu
et al., 2003) and seedling frost tolerance (Frt) (Eujayl et al., 1999).
To date, quantitatively inherited traits have been mapped by Durn
et al. (2004) who detected five QTL for height of the first ramification, three
for plant height, five for flowering, seven for pod dehiscence, one for shoot
number and one for seed diameter. Five and four QTL were identified for
winter survival and winter injury, respectively, and were mapped using a
population of 106 RILs derived from WA8649090 Precoz (Kahraman et al.,
2004). In this study, the experiments were conducted in multiple locations
and only one of five QTL was expressed in all environments. Primary mapping of Ascochyta blight resistance using an F2 population derived from
ILL 7537 ILL 6002 identified three QTL (QTL-1, QTL-2 and QTL-3) accounting for 47% (QTL-1 and QTL-2) and 10% (QTL-3) of disease variance (Shaika,
2004). Identification and mapping of QTL conferring resistance to Stemphylium blight, rust and white mould fungal diseases using recombinant inbred
line populations are underway (W. Chen, Washington State, 2007, personal
communication), as is mapping of physical seed quality traits such as size,
shape and colour, as well as resistance to Ascochyta blight at maturity
growth stages (R. Ford, 2007, unpublished results).
To date, most quantitative traits have been mapped using F2 populations (<100). Since quantitative traits are influenced by both genetic and
165
Fig. 11.1. A genetic linkage map of Lens sp. based on microsatellite, AFLP, RAPD and
morphological markers. The marker names beginning with an asterisk are those markers
mapped by Eujayl et al. (1998b). The values on the left side of the individual linkage groups
represent distance in centimorgans calculated using the Kosambi mapping function
(Source: reproduced from Hamwieh et al., 2005).
environmental effects, RILs or near isogenic lines (NILs) would be more

suitable populations to accurately dissect their components.
One of the primary objectives of associating markers with phenotypic
traits is to expedite crop breeding through marker-assisted selection (MAS).
166
R. Ford et al.
Ideally, the genes controlling a trait of interest are the perfect marker for
MAS. However, this is often made difficult because cloning of the genes is
labour intensive and time consuming. Alternatively, marker(s) that are
tightly linked to and flanking a gene locus that conditions a sizable portion
of the genetic variation accounting for the trait may be selected for with the
premise that the associated chromosomal region contains the functional
gene(s).
Often, genetically linked markers to traits of interest are identified by
coarse mapping and these have limited use in MAS because of the distance
and hence chance of recombination between marker and actual gene locus.
Therefore, genomic regions where the trait is mapped should be characterized at high resolution (since recombination rates may vary at different
genomic regions) and be validated across genetic backgrounds, in order to
determine their utility in MAS. Also, physical characterization of genomic
regions of interest will facilitate cloning of the gene to develop direct markers
(candidate genes) and/or physically closer markers to the gene, increasing
the reliability for MAS. Marker types most useful for MAS should be locus
specific, highly reproducible and easy to discern. These include sequence
tagged site (STS), sequence characterized amplified region (SCAR) or allele
specific amplified primer (ASAP), specific polymorphic locus amplification
test (SPLAT) and PCR-based RFLP markers. When locus specific markers are
not size polymorphic among the parental lines used in the breeding programmes, sequence discriminative methods are required. These include SNP,
cleaved amplified polymorphic site (CAPS) and derived CAPS (dCAPS)
markers. Meanwhile, there are several markers available for different traits
that have the potential for use in MAS and gene pyramiding (Table 11.4).
These include SCARW19 and SCARB18 linked to and flanking the AbR1 A.
lentis resistance loci (Nguyen et al., 2001; Taran et al., 2003). These enabled
successful pyramiding of the AbR1 and ral2 A. lentis resistance loci together
with the LCt2 Colletotrichum truncatum (anthracnose) resistance loci (Taran
et al., 2003).
11.8. The Future of Lensomics

To better target genes that are functionally associated with traits of interest
for breeding purposes, several approaches are predicted for the future.
These include the use of emerging genomic sequence for the development
of gene family specific degenerate primers. This approach has already been
taken, to identify novel defence-related genes within various resistance genes
families. RGAs are a family of sequences associated with pathogen defence
but with unknown function at the time of isolation. These have been isolated
and characterized from lentil using degenerate oligonucleotide primers
(Yaish et al., 2004; R. Ford, 2007, unpublished results), and recently associated
with defence to A. lentis (B. Mustafa, 2007, unpublished results). Yaish et al.
(2004) isolated 32 RGA sequences from ILL 5588 using degenerate primers
designed to conserve motifs in the NBS domain of NBSLRR resistance
167
Table 11.4. Molecular markers closely associated with desirable lentil breeding traits for use
in marker-assisted selection.
Trait mapped
Associated molecular
markers
Fusarium wilt resistance (Fw)

Ascochyta blight resistance (AbR1)
Ascochyta blight resistance (ral 2)
Ascochyta blight resistance
(mapped as a QTL)
Anthracnose resistance (Lct 2)
Frost tolerance (Frt)
Winter hardiness
Fusarium wilt resistance (Fw)
OPK15
RV01, RB18, SCARW19
UBC227, OPD-10
C-TTA/M-AC (QTL1 and
QTL2), M20 (QTL3)
OPE06, UBC704
OPS-16
UBC808-12
SSR59-2B, p17m30710
Citation
Eujayl et al. (1998a)
Ford et al. (1999)
Chowdhury et al. (2001)
Rubeena et al. (2003)
Tullu et al. (2003)
Eujayl et al. (1999)
Kahraman et al. (2004)
Hamwieh et al. (2005)
proteins. Predicted amino acid sequences showed they contained typical

conserved motifs present in the majority of known plant NBSLRR genes
and that they phylogenetically belonged to the TIR-NBS-LRR subclass.
R. Ford (2007, unpublished results) isolated a further five TIR-NBS-LRR
and one Pto kinase type RGA sequences from ILL 7537. Genetic mapping of
RGA markers will potentially identify the genomic regions where the genes
are located that functionally regulate signal transduction and/or defence
pathways. This will also facilitate map-based cloning of the resistance genes.
Other major gene families of interest should become the targets for future
isolation and trait association mapping. These may underpin the control of
expression of other breeding goals (such as disease resistance, abiotic tolerance and physical attributes) and include those thought to act as transcription factors and in key biochemical pathways (i.e. Myb genes). With the
advancement in functional genomics, expression QTL (eQTL) can be identified for the traits of interest by coupling global genome gene expression
profiling and suitable genetic materials and analysis. Since eQTL affect the
expression of the genes for the trait of interest, the markers linked to this
eQTL will have enormous reliability in MAS compared to the markers identified by traditional QTL analysis.
Other biotechnology approaches such as proteomics and metabolomics
will no doubt begin to emerge for lentil, in order to discover and better
characterize the functional mechanisms behind the breeding targets. Ultimately, and with sufficient funding, the combination of lentil omics data
will enable the precise formulation of superior and high-yielding genotypes,
adapted to a multitude of environments and market preferences.
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12
Breeding and Management to Minimize

the Effects of Drought and Improve Water Use
Efficiency
R. Shrestha,1 K.H.M. Siddique,2 David W. Turner2 and Neil C. Turner2
1National
Grain Legumes Research Program, Chitwan, Rampur, Nepal; 2The University of Western Australia, Crawley, Western Australia, Australia
12.1. Introduction
Lentil (Lens culinaris Medikus subsp. culinaris) is an important cool-season
grain legume crop, mainly grown in South Asia, West Asia and North
Africa. In recent years the area under lentil production has also expanded
in Canada, Australia and the USA. It is grown as a winter crop either on
stored soil moisture after summer rainfall in subtropical regions or on limited rainfall in Mediterranean-type environments. Usually growth is restricted
by cool temperatures during vegetative growth in winter and by high temperatures and terminal drought in spring and early summer (Yusuf et al.,
1979; Shah et al., 1987; Erskine and Saxena, 1993; Shrestha, 1996). This chapter reviews the effects of water deficits on growth and yield of lentil and
explores the genetic and agronomic options to minimize the effects of
drought on dry matter (DM) production and seed yields.
12.2. Effects of Water Deficits

Dry matter production
With an adequate water supply, DM in lentil increases rapidly over time,
while water deficits reduce the accumulation of above-ground DM at maturity by up to 3261% (Ashraf et al., 1992; Turay et al., 1992; Shrestha et al.,
2006b). Water deficits reduce the number of leaves and nodes, leaf area, and
increase the rate of leaf senescence (Shrestha, 2005). Lentil genotypes vary
significantly in the above-ground DM they produce under water deficits
(Ashraf et al., 1992; Shrestha et al., 2006b). As lentil has an indeterminate
growth habit, growth continues after flowering, and the imposition of water
deficits in the reproductive phase reduces plant height, leaf area and foliage
172

Breeding and Management to Improve Water Use Efciency
173
DM as well as reproductive structures. There are genotypic differences in

the partitioning of DM to seeds, pod wall and foliage (leaf, stem and flowers) with water deficits, with the proportion of DM in the seeds affected by
water shortage in some but not all genotypes (Shrestha, 2005).
Flower and pod formation

In indeterminate legume species, the total number of seeds produced is a
function of the number of nodes, the number of flowers per node and the
proportion of flowers that develop into mature pods (Egli, 1998). In lentil,
water deficits reduce the total number of flowers produced by up to 45%
and reduce the flowering duration by 12 days compared with adequately
watered lentils. However, a mild water deficit during early flowering that
was relieved during pod development stimulated flower production by
2839% in some genotypes (Shrestha et al., 2006b).
Terminal drought significantly reduces the total numbers of filled pods
and seeds produced and increases the number of single-seeded pods compared with double-seeded pods (Shrestha, 2005). A severe water deficit
imposed during podding stopped the production of new flowers and pods
and reduced the final pod and seed numbers by 6070% compared with
adequately watered lentils (Fig. 12.1), while seed abortion (empty pods)
increased by 5575% (Shrestha, 2005).
300
200
100
Seeds/plant
Pods/plant
150
100
50
0
105
120
135
150
105
120
135
150
Time after sowing (days)
Fig. 12.1. Changes in the number of pods and seeds per plant over time in West
Asian (circle, line ), crossbred (triangle, line ) and South Asian (square, line
) lentil genotypes grown under adequately watered (solid symbols and line) and
water-deficit (open symbols and dotted line) conditions. Lines are the fitted curves.
Bars indicate one SE of the fitted values at maturity where SE larger than the symbol
(Source: adapted from Shrestha, 2005).
174
R. Shrestha et al.
Seed development
The dry weight accumulation of an individual seed follows a sigmoid
growth pattern, with an initial lag phase, followed by a rapid (linear)
increase in DM to a maximum and the final phase where DM accumulation
rate reaches zero at physiological maturity (Fig. 12.2). Final seed size (weight
per seed) depends on how fast (seed growth rate, SGR) and how long (seed
fill duration, SFD) seed growth continues. In lentil, the SGR ranged from 1.2
to 2.5 mg/seed/day and there was a positive correlation between SGR and
seed size (Shrestha et al., 2006a).
Water deficits at the end of embryo cell division (i.e. during the lag
phase of early seed filling) affect the number of cotyledonary cells, while
'Cassab'
60
ILL 7979
'Simal'
(a)
(b)
(c)
(d)
(e)
(f)
50
40
30
Dry weight (mg/seed)
20
10
0
60
50
40
30
20
10
0
0
20
40
60
20
40
60
20
40
60
Time after flowering (days)
Fig. 12.2. Changes in seed size with time after flowering in West Asian (Cassab), crossbred
(ILL 7979) and South Asian (Simal) lentil genotypes grown under adequately watered (a, b, c)
and water-deficit (d, e, f) conditions. The lines are the predicted values using a linear mixed
model with cubic spline up to 50 days after commencement of flowering. Flowering dates: {,
6476 days after sowing (DAS); , 7988 DAS; , 121130 DAS; , 133146 DAS; and ,
150164 DAS (Source: adapted from Shrestha et al., 2006a).
175
water deficits during the linear phase of seed development affect cell expansion and DM accumulation (Munier-Jolain and Ney, 1998; Saini and Westgate, 2000). Water deficits also reduce the final seed size by shortening SFD
as a result of a reduction in assimilate supply rather than inhibition of SGR
(Egli, 1998). A water deficit during early pod development or seed filling
alters both the source and the sink strength and maintains the flux of assimilates to the developing embryo without affecting seed size or SGR (Egli,
1998). Moreover, the ability of plants to mobilize storage reserves to the
seed (Turner et al., 2005) and the ability of developing seeds to maintain a
favourable water status at low water potential helps to maintain SGR, seed
size and prolong the SFD under water deficits (Westgate and Grant, 1989).
In lentil, a water deficit during the reproductive phase reduced both the
source (leaf area and/or assimilates) and the sink strength (abortion of flowers, pods and seeds resulting in reduced pod and seed numbers) thus maintaining SGR and the final size of the seeds that remained. The lack of a
significant effect of water deficit on SFD in lentil may be related to its relatively small seed size (Shrestha et al., 2006a).
Seed yield and harvest index

In lentil, there is a positive association between seed yield and the number of
reproductive nodes, total number of flowers, number of pods, number of
seeds, total DM, and harvest index (HI) (Shrestha, 2005). However, neither
seed size nor seeds per pod were correlated with seed yield. A transient water
deficit during flowering increased seed yield by 1041% and HI by 1821%
in some genotypes due to increases in the numbers of pods (2666%) and
seeds (2230%) (Shrestha et al., 2006b), whereas terminal drought reduces
seed yield by reducing pod and seed numbers (Shrestha et al., 2006a).
Nitrogen fixation
Symbiotic N2 fixation in lentil ranges from 69 to 154 kg N/ha (Kurdali et al.,
1997; Schulz et al., 1999; Schmidtke et al., 2004) where the amounts of N2
taken up from the soil ranged from 58 to 61 kg N/ha (Schmidtke et al.,
2004). Under favourable conditions about 5880% of N2 is derived from the
atmosphere (Saxena and Silim, 1990; Badarneh and Ghawi, 1994; Kurdali
et al., 1997). Water deficits affect N2 fixation either through reduced carbon
supply to the nodules (Subbarao et al., 1995) or reduced flow of carbon into
nodules via the phloem (Serraj et al., 1999) or through a direct effect on the
activity of the nodules (Durand et al., 1987; Subbarao et al., 1995). Water deficits decrease the nitrogenase activity of nodules (Durand et al., 1987; Serraj
et al., 1999) through increasing the resistance of oxygen diffusion to the
bacteroids (Durand et al., 1987), thereby decreasing the number of effective
nodules, nodule dry weight, number of bacteroids and leghaemoglobin
content (Rathore et al., 1992).
176
R. Shrestha et al.
12.3. Crop Management to Improve Water Use Efficiency

Yield (Y) in water-limited environments is a function of water use or evapotranspiration (ET), the efficiency of conversion of the water to DM (water
use efficiency, WUE) and the conversion of vegetative DM to a harvestable
yield (HI) (Passioura, 1977):
Y = ET WUE HI
ET has two components, crop transpiration and soil evaporation. Losses by
runoff and deep drainage below the root zone are considered minimal in
crops such as lentil grown in low-rainfall Mediterranean environments or
on residual soil water. However, soil evaporation can be high and minimizing the proportion of ET lost as soil evaporation is essential to maximize
WUE and yield. Thus to maximize yields in water-limited environments,
agronomic practices to maximize crop transpiration, WUE and HI, and
minimize water loss by soil evaporation are required.
Water use, dry matter production and seed yield
In dryland environments crop water use or ET is correlated with seasonal
rainfall and its distribution, with greater losses in wet seasons than in dry
seasons (Zhang et al., 2000). Soil evaporation is greatest during the early
stages of crop growth, when the canopy cover is small. Early planting coupled with early vigorous growth reduce soil evaporation. In a glasshouse
study 85% of the variation in water use in lentil was explained by leaf area
and 70% by DM accumulation at flowering (Shrestha, 2005). Seed and straw
yields showed a linear increase with the amount of water applied by supplemental irrigation (Zhang et al., 2000; Shrestha, 2005). It is therefore not
surprising that total seasonal rainfall accounted for about 40% of the variation in mean seed yield in Mediterranean environments, with up to 80% of
the variance in mean seed yield accounted for at a single site (Erskine and
El Ashkar, 1993).
Similarly, at a Mediterranean site, Sarker et al. (2003) reported 77% of
the variation in seed and straw yields were due to seasonal rainfall and
temperature, with a positive correlation between seed and straw DM and
total seasonal rainfall. However, in Western Australia, with a Mediterraneantype climate, seed yield was not correlated with total water use or with
water use before flowering (Leport et al., 1998), but was positively correlated with post-flowering water use (Siddique et al., 2001).
There are a range of crop management practices such as minimum tillage, appropriate fertilizer use, improved weed/disease/insect management, timely planting, narrow row width and crop rotations that contribute
to high grain yield by increasing rainfall use efficiency of dryland crops
(Siddique et al., 1998a; Debaeke and Aboudrare, 2004; Turner, 2004; Sarker
and Erskine, 2006). These crop management strategies in a rainfed environment increase water stored in the soil at sowing, increase soil water extraction,
177
reduce water losses by soil evaporation, runoff, through-flow, deep drainage, optimize the seasonal water use pattern between pre- and post-anthesis
use, and improve the tolerance of a water deficit and recovery from it
(Debaeke and Aboudrare, 2004; Turner, 2004). Relay cropping of lentil in
rice fields (broadcasting of seed 12 weeks before the rice harvest) is a common practice in South Asia in ensuring the use of residual soil water
(NGLIP, 1988). Early sowing, selection of varieties with rapid canopy coverage, adequate plant population, optimum fertilizer application and close
spacing help to cover the soil surface, suppress weed growth and thereby
increase WUE (Cooper et al., 1987). Sowing date is the major factor influencing yield and yield components in lentil (Krarup, 1984; Gray and de-Delgado,
1986; Karim et al., 1987; Mohamed, 1988; Singh et al., 1990; Aziz, 1992;
Shoaib, 1992; Varshney, 1992; Shrestha, 1996; Siddique et al., 1998b). Siddique et al. (1998b) showed that in the short-season Western Australian
Mediterranean-type environment delaying sowing reduced yields by up to
30 kg/ha/day at some sites, while a seed yield reduction of 15 kg/ha/day
was reported in the warm-temperate environment of the Kathmandu Valley (Shrestha, 1996). Crop water use decreased with later planting and water
use in the post-flowering period was particularly reduced by a delay in
planting. Under rainfed environments in Nepal and Jordan early-sown lentils produced greater yield as a result of adequate vegetative biomass, long
grain-filling periods and greater number of pods (Neupane and Shrestha,
1991; PAC, 1994; Shrestha, 1996; Rahman et al., 2002). In West Asia and
North Africa early-sown lentil had 2025% higher seed yield compared
with the late-sown lentil (cited in van Duivenbooden et al., 2000).
Water use efficiency

Rainfall received during the winter months in the Mediterranean environments of West Asia and Australia, usually occurs in frequent small showers
and hence soil evaporation may constitute 3050% of ET (160353 mm)
(Table 12.1). In South Asia, where lentils are grown on stored soil water (ET
ranged from 134 to 194 mm), the surface soil is dry and hence soil evaporation is low relative to Mediterranean environments. For example, at Tel
Hadya in northern Syria with its Mediterranean climate, soil evaporation
(80 mm) contributed 28% of the seasonal ET in lentil (Zhang et al., 2000).
The overall WUE values in this study ranged from 3.6 to 30.3 kg DM/ha/
mm and from 0.6 to 20.0 kg seed yield (grain)/ha/mm under rainfed conditions. Supplemental irrigation had the values to 2.829.6 kg DM/ha/mm
and 1.011.2 kg grain/ha/mm, depending upon genotype, growing environment and water use (Table 12.1). Siddique et al. (1998a) showed that lentil could achieve a WUE of 20 kg/ha/mm in south-western Australia, but
in the majority of situations the values of WUE were below 15 kg/ha/mm
(Table 12.1). One of the reasons for the lower WUE was a delay in planting
(Siddique et al., 1998a). Analysis of seed yields and water use using the
French and Schultz (1984) method and a soil evaporation (intercept) of
178
R. Shrestha et al.
3500
7000
(b)
3000
6000
2500
5000
2000
4000
1500
3000
1000
2000
500
1000
Total dry matter (kg/ha)
Seed yield (kg/ha)
(a)
0
0
100
200
300
400 0
100
200
300
400
Total water use (mm)
Fig. 12.3. Relationship between (a) seed yield and (b) above-ground DM and
cumulative water use for selected lentil genotypes from West Asia (), South Asia ()
and crossbreds () (Shrestha et al., 2005). Data from studies in India () (Sharma and
Prasad, 1984), Tel Hadya, Syria ({) (Silim et al., 1987; Zhang et al., 2000), Breda,
Syria () (Silim et al., 1993b) and south-west Western Australia () are also included.
Lines are based on an intercept of 115 mm and water use efficiencies of (a) 15 ( ),
10 (....) and 5 () kg grain/ha/mm, and (b) 40 kg DM/ha/mm (Source: Siddique
et al., 1998a, 2001).
115 mm (Siddique et al., 1998a, 2001) showed that in rainfed environments

of Nepal the WUE of the small-seeded lentils (South Asian and crossbreds)
were similar at 1015 kg grain/ha/mm to those in Western Australia, India
and Syria, while the large-seeded lentils (West Asian) had WUE for grain of
less than 5 kg/ha/mm (Fig. 12.3a). The WUE for DM of the small-seeded
genotypes was about 40 kg DM/ha/mm, whereas the large-seeded genotype had only half of this value (Fig. 12.3b).
Siddique et al. (1998b) studied the effect of sowing rate (20120 kg/ha)
on growth and seed yield of lentil at 13 sites over three seasons in the cropping regions of south-western Australia. On average, the profitability of
lentil crops in south-western Australia can be maximized with a plant density of about 150 plants/m2. It is likely that higher plant densities (up to 230
plants/m2) are optimal when growing conditions are unfavourable and the
growth of individual plants is limited (e.g. in low rainfall environments,
through delayed sowing or with early maturing cultivars). Lentil crops
sown at high densities may also be better able to compete with weeds, be
less prone to aphids and virus attack, help reduce soil evaporation and are
likely to be taller and hence easier to harvest than low-density crops. However, Krarup (1984), Gray and de-Delgado (1986), Karim et al. (1987),
Mohamed (1988), Neupane and Shrestha (1991), Shrestha (1996) and Rahman et al. (2002) reported no effect of seed rates on seed yield and yield
components, which may be due to the small range of seed rates studied (for
example, 3060 kg of seed/ha by Shrestha, 1996).
Crop water use (mm)
WUEDM (kg/ha/mm)
Region
Irrigated
Rainfed
Soil depth
(m)
West Asia
267445
283
277318
160353
213326
93
8795
195321
1.8
1.8
1.0
10.612.3
7.913.8
352678
a183388
b115228
269312
231249
179266
174273
222339c
182
134
159179
169
194 d278
1.7
1.4
0.4
1.7
1.2
1.2
1.2
0.6
0.6
2.84.1
10.112.3
29.6
Australia
South Asia
WUEGRAIN (kg/ha/mm)
Irrigated
Rainfed
Irrigated
Rainfed
9.218.1
14.222.6
5.58.8
5.59.8
6.614.9
5.2
13.530.3
8.513.1
c3.66.2
7.3
24.226.2
8.218.9
3.48.1
4.35.3
2.46.3
2.14.5
3.99.1
0.93.1
0.63.4
1.06.0
10.7
3.820
2.47.2
c1.32.4
4.0
7.312.5
2.3
10.4
1.87.0
1.02.0
5.310.0
9.111.2
4.65.4
7.610.6
Reference
Zhang et al. (2000)
Silim et al. (1987)
Saxena and Silim (1990)
Silim et al. (1993b)
Badarneh and Ghawi (1994)
Sarker et al. (2003)
Siddique et al. (1998a)
Siddique et al. (2001)
Shrestha (2005)
Leport et al. (1998)
Yusuf et al. (1979)
Saraf and Baitha (1985)
Sharma and Prasad (1984)
Lal et al. (1988)
Shrestha et al. (2005)
Table 12.1. Crop water use (evapotranspiration (ET), mm), water use efciency (WUE, kg/ha/mm) for total above-ground DM (WUEDM)
and seed yield (WUEGRAIN) of lentil grown under supplemental irrigation and rainfed environments in West Asia, South Asia and Australia
(Source: adapted from Shrestha, 2005).
a Growing
season rainfall + irrigation.

water extraction + irrigation.
c Intermittent water decit at owering/podding.
d Long season West Asian genotype that used the late-season rainfall.
b Soil
179
180
R. Shrestha et al.
As lentil is a poor competitor, weed species flourish and compete for

water, nutrients and light (Walsh et al., 2004), leading to reduced seed yields
(Kayan and Adak, 2006). Use of selective herbicides, mechanical and or
hand weeding generally improve yield and WUE of lentil crops in dryland
environments. Increased fertilizer use and deep ripping resulting in excessive vegetative growth may lead to an increase in ET and hence greater
water deficit at flowering and in the post-flowering period when crops
grow on stored soil water (Turner, 2004). In lentil, excess nitrogen (with
adequate soil moisture) results in excessive vegetative growth reducing
seed yield, however, high rates of nitrogen fertilizers are seldom used in
lentil as it is a N2 fixing legume.
Harvest index
The ratio of grain yield to total above-ground DM, the HI, is determined by
the DM accumulated and the amount of water available for seed development after podding. In lentil, HI decreases with delay in sowing, and this is
associated with less water availability after flowering (Siddique et al., 1998b).
Thus, early planting and good early establishment by seed priming will
increase the HI.
12.4. Mechanisms of Drought Resistance and their Use in Breeding

Drought resistance is the ability of a genotype to produce higher yield than
another when subjected to periodic water deficits (Turner, 1979). Two
frameworks for the adaptation of plants to drought have been proposed
(Turner, 2003), namely a drought-resistance and a yield-component framework. The drought-resistance framework deals with the specific morphological, physiological and biochemical characteristics that enable plants to
adapt to low water availability, while the yield-component framework
focuses on yield under water-limited conditions and the components of
yield, namely water use, WUE and HI (Turner, 2003). For survival and
improved yields under rainfed conditions, plants usually possess more than
one adaptive mechanism (Khanna-Chopra, 1982). While the yield-component
framework has been helpful in analysing the management practices to improve
yields in water-limited environments, we adopt the drought-resistance framework for considering the traits that assist in improving yields of lentil in
water-limited environments.
Drought-resistance mechanisms have been divided into three categories
(e.g. by Turner, 2003):
1.
2.
3.
Drought escape.
Dehydration avoidance.
Dehydration tolerance.
181
Drought escape
The mechanism identified as important in drought escape is the ability of a
plant to complete its life cycle before the commencement of severe water
deficits (Boyer, 1996). This involves rapid phenological development such
as rapid germination and seedling establishment, early flowering, early
podding, early maturity, and phenological plasticity to take advantage of
longer and cooler seasons (Turner et al., 2001). In lentil, flower initiation is
affected by temperature, photoperiod and vernalization (Summerfield et al.,
1984; Erskine et al., 1989a, b; McKenzie and Hill, 1989). Lentil is predominantly a long-day (1516 h photoperiod) plant (Saxena and Wassimi, 1984),
but genotypes vary considerably from neutral to acute sensitivity in their
responsiveness to photoperiod and temperature (Summerfield et al., 1985b),
and many genotypes show a quantitative vernalization requirement (Summerfield et al., 1985a). Genotypes originating from lower latitudes (Ethiopia,
Sudan, Egypt and India) have a shorter critical photoperiod than genotypes
from higher latitudes (USSR, Turkey), and the magnitude of photoperiodic
response depends on temperature differences (Saxena and Wassimi, 1984;
Erskine, 1997). In the Mediterranean environments of North Africa and
West Asia where lentil is sown in March/April (spring), vegetative growth
coincides with progressively lengthening days and warmer temperature,
and flowering occurs at a photoperiod of 11.612.0 h. In South Asia, northern Argentina and Australia lentil is sown in autumn and vegetative growth
coincides with shortening days and cooler temperatures (Sinha, 1977;
Erskine and Hawtin, 1983; Erskine et al., 1994a; Shrestha et al., 2005) and
flowering occurs at a photoperiod of about 10.611.0 h (Erskine, 1983; Waldia et al., 1988; Shrestha et al., 2005). Mediterranean/West Asian genotypes
flower late when grown under South Asian conditions and as a result, the
duration of flowering and pod filling is truncated by rising temperatures
and decreasing rainfall in spring and early summer, resulting in poor DM
accumulation, very few pods and poor seed yield (Erskine and Hawtin,
1983; Shrestha et al., 2005). In contrast, genotypes originating from South
Asia and Ethiopia, when sown in South Asia, flower and pod early and
thereby escape drought in rainfed environments (Hamdi et al., 1992). Differences in phenological development contribute 4560% of the variation in
seed yield in South Asia (Silim et al., 1993a; Erskine et al., 1994b; Siddique
et al., 2003). Therefore, the appropriate drought strategy for lentil in environments where water deficits and high temperatures at the reproductive
stage induce senescence and early maturity is drought escape (Erskine et al.,
1994b; Thomson et al., 1997; Zhang et al., 2000; Siddique et al., 2003; Shrestha
et al., 2006a).
Initiation of flowering can be unaffected, advanced or delayed in lentil
depending on the severity of water deficits and the crop growth habit (van
Kessel, 1994). In the case of mild water deficits, advancement in the time of
flowering may be related to higher plant temperature as a consequence of
reduced transpiration. While early flowering and rapid phenological
development give higher seed yields and higher yield stability than late
182
R. Shrestha et al.
flowering under terminal drought, seed yields may be reduced when there
is an unpredictable intermittent drought (Siddique et al., 2003). Thus droughtinduced early maturity is advantageous in dry seasons, but high yield is
often associated with longer growth duration, late flowering and greater
water use (Blum, 1997). The ability of plants to adjust the duration of different growth phases in response to the availability of soil water during the
growing season, that is developmental plasticity (Turner, 1979; Subbarao
et al., 1995), enables a plant to produce higher yields when the growing
period is longer and is an important mechanism in unpredictable climates
and under favourable soil water regimes. While lentil is an indeterminate
plant, this quality varies with genotypes. In lentil a period of water deficit
enhances pod fill and hence the length of pod filling varies depending on
the degree and duration of water shortage, particularly in late-maturing
genotypes (van Kessel, 1994). The indeterminate nature of some lentil genotypes is a beneficial trait that contributes to developmental plasticity.
Dehydration avoidance
Both morphological and physiological mechanisms contribute to dehydration avoidance.
Morphological mechanisms
Morphological modifications such as reduction in leaf size, shape and numbers, leaf orientation (change in leaf angle) and leaf rolling are mechanisms
that postpone dehydration because radiation absorption and water loss are
reduced (Kramer, 1980; Turner et al., 2001). With increasing water deficits,
the rate of leaf production decreases (Mwanamwenge et al., 1997; Shrestha,
2005) and leaves became smaller (Mwanamwenge et al., 1997). In lentil, yellow cotyledons appear to be associated with high DM production (Tullu
et al., 2001) and light-coloured foliage at the vegetative stage is associated
with vigorous early growth and establishment (Silim et al., 1993a). Increased
stomatal and cuticular resistance, by increased pubescence, and waxiness
help to reduce water loss. Lentil genotypes with greater wax deposition on
the leaf surfaces showed considerable tolerance to water deficits (Ashraf
et al., 1992). Under rainfed environments, anthocyanin pigment production
in genotypes from South Asia and crossbreds between South Asian and
West Asian genotypes is a mechanism that enhances cold tolerance (Summerfield et al., 1985a). Leaf hairs reduce gas exchange, increase leaf reflectance and reduce leaf heating (Monneveux and Belhassen, 1997). Leaf drop
and rapid senescence of the physiologically older leaves are also adaptive
mechanisms for reducing water loss (Monneveux and Belhassen, 1997).
Deep rooting and high root density are common morphological mechanisms to improve water uptake by extracting water from greater depth
(Turner, 1986). Legumes have lower root length densities (RLD) than cereals when grown in similar soils (Gregory, 1986). Lentil has greater RLD in
183
the upper 030 cm soil layer than in the deeper soil layers (60160 cm)
(Sharma and Prasad, 1984; Silim et al., 1993b; Adak and Biesantz, 1997;
Shrestha et al., 2005), and the rooting depth of lentil under rainfed conditions is about 0.8 m (Gregory, 1986; Siddique et al., 2001). A large vigorous
root system is a major feature of drought resistance under water deficits
when water is available deep in the soil profile and is replenished each year
(Ludlow and Muchow, 1988). Dehydration-avoiding species usually have
deep rooting ability (Boyer, 1996), but increased rooting depth and RLD do
not always reflect the ability of a genotype to extract more soil water if the
wet profile is shallow (Ludlow and Muchow, 1988). The ratio of root-toshoot DM in lentil increased by 14100% compared with well-watered lentil
when water deficits were imposed at the reproductive stage (Shrestha,
2005). The increase in the root-to-shoot ratio was the result of the relatively
greater reduction in shoot growth than in root growth rather than an
increase in absolute root weight (Shrestha, 2005). Sarker et al. (2005) reported
significant variation in taproot length, lateral root number, total root length
and total root weight among lentil genotypes of diverse origins. Genotypes
with a long taproot and more lateral roots have increased drought tolerance
and are being used for breeding high-yielding cultivars under water-limited
conditions (Sarker and Erskine, 2005; Sarker et al., 2005). Lentil genotypes
from South Asia have the highest RLD (at pod filling) followed by the crossbreds (South Asia West Asia), and the lowest in the West Asian genotypes
when grown in the rainfed environment of Nepal (Shrestha et al., 2005).
Physiological mechanisms
Low stomatal conductance is an efficient adaptive response to short periods
of water deficit (Franca et al., 2000). Stomatal closure allows plants to maintain tissue turgour and volume for normal metabolic function by maintaining water uptake and reducing water loss (Kramer, 1980; Turner et al., 2001).
There is considerable variation in the sensitivity of stomata to increasing
water deficits among grain legumes (Leport et al., 1998; Shrestha, 2005). In
lentil, stomatal conductance under well-watered conditions varied from 169
to 400 mmol/m2/s but decreased markedly to 19100 mmol/m2/s as the
leaf water potential decreased below 2.5 MPa (Leport et al., 1998; Shrestha,
2005). No differences among genotypes in the stomatal response to water
deficits were observed (Shrestha et al., 2006b) and stomatal conductance has
not been used as a selection criterion in lentil.
Differences in stomatal conductance will influence the rate of photosynthesis (PN), but water deficits can affect the rate of photosynthesis by affecting the photosynthetic capacity of the leaf as well as stomatal conductance
and can influence the remobilization of carbon to the grain in assimilate
redistribution. With an adequate water supply, leaf PN in lentil ranged from
16.6 to 17.0 mol/m2/s during flowering in both the field and the glasshouse (Leport et al., 1998; Shrestha, 2005) and 8 mol/m2/s during podding
(Shrestha, 2005). Leaf PN was reduced by 2253% when water was withheld
at flowering and podding, and reached very low values when the leaf water
184
R. Shrestha et al.
potential fell to 2.5 MPa (Shrestha, 2005) to 3.0 MPa (Leport et al., 1998).
However, no differences among genotypes were observed in the sensitivity
of photosynthesis to water deficits (Shrestha, 2005) and breeders have not
focused on selecting lines with differences in photosynthesis in lentil.
When water deficits are severe, particularly under terminal drought
where the current PN is impaired during seed filling, remobilization of stem
reserves to the grain can be important (Turner et al., 2005). Therefore, yield
stability under terminal water deficits depends on the ability of crops to
mobilize assimilates, accumulated before and immediately post-flowering, to
the seed (Turner et al., 2001). Genotypes vary in the proportion of pre-flowering
assimilate transferred to grain. In late-maturing lentils, stem reserves act as a
source of carbon for seed filling, while PN is the major source of assimilates in
early maturing genotypes (Clements et al., 1997). In general, grain legumes
show poor remobilization of assimilates produced before flowering to the
seed compared with cereals. This is because of the greater demand for
assimilates for N2 fixation and through competition for assimilates between
vegetative and reproductive organs as a result of their indeterminate growth
habit. Nodules respire about 40% of the carbon assimilated photosynthetically at the vegetative stage (Salon et al., 2001), but in lentil, higher concentrations of nitrogen in the leaves at flowering (3641 mg/g) compared with
maturity (1822 mg/g) suggests that there may be remobilization of nitrogen from the leaf to the seed during pod filling (Whitehead et al., 2000). In
lentil, low soil water almost halved N2 fixation compared with an adequate
water supply (Saxena and Silim, 1990) and genetic variation in N2 fixation
among legume cultivars with response to water deficits has been reported
(Serraj et al., 1999). Also, the genotypic differences in the growth and survival of rhizobial strains (Rhizobium leguminosarum) to increasing soil water
deficits (1.5 MPa) have been reported by Athar (1998), but both the genetic
variation in N2 fixation and the tolerance of water deficits by rhizobia have
to be exploited by breeders and agronomists.
Osmotic adjustment (OA) is an important physiological mechanism by
which plants synthesize and accumulate solutes in cells in response to water
deficits in the soil and/or plant (Turner and Jones, 1980). This net accumulation of solutes lowers the osmotic potential or increases the osmotic pressure of the cells and draws water into the cells and tissues and hence
contributes to the maintenance of turgour, stomatal conductance, photosynthesis and plant growth at progressively lower leaf water potentials (Turner
and Jones, 1980; Subbarao et al., 1995). In a rainfed environment in Western
Australia, lentil showed a high degree of OA (0.6 MPa) with decreasing leaf
water potential (Leport et al., 1998; Turner et al., 2001) and has been shown
to range from 0.0 to 1.8 MPa in a range of genotypes and soil water potentials (Ashraf et al., 1992; Clements et al., 1997; Shrestha, 2005). In a glasshouse study lentil genotypes showed variation in OA during flowering, but
this variation was a result of the varying degrees of soil water deficit among
the genotypes (Fig. 12.4). The values of osmotic potential were lower at
podding than flowering, but no solute accumulation occurred during the
drying cycle at podding (Shrestha, 2005). The solutes that accumulate through
185
Osmotic adjusment (MPa)
2.5
2.0
1.5
1.0
0.5
0.0
30
25
20
15
10
5
Cumulative leaf water potential (MPa)
Fig. 12.4. Relationship between osmotic adjustment (calculated as the difference in

osmotic potential at 100% relative water content between adequately watered and
droughted plants) and cumulative predawn leaf water potential (r2 = 0.62; highly
significant at P = 0.01 level) of six lentil genotypes: Cassab ({), ILL 6829 (), ILL
7979 (), ILL 7982 (), Khajura 2 () and Simal () when water was withheld at
flowering (Source: Shrestha, 2005).
OA are reportedly transported from the senescing leaves to the developing

seeds when the rate of leaf photosynthesis reaches zero (Leport et al., 2003).
Positive correlations between OA and seed yield have been reported for
some species, but no correlation between OA and seed yield or DM have
been observed in lentil (Francis and Siddique, 2001) and no selection for OA
has been conducted in breeding lentil for water-limited environments.
In those rainfed environments in which the crop grows on residual soil
water, one of the mechanisms that increases the efficiency of water use is
increased transpiration efficiency (TE). TE is the transpiration per unit DM
or grain yield (Fischer, 1981) or, at the level of leaf, the moles of carbon (C)
or CO2 fixed per mole of water lost by transpiration (Arnon, 1992). Measurement of the instantaneous TE of the leaf is possible with portable gas
exchange equipment, but an integrated measure of TE is difficult and labour
intensive, requiring the measurement of plant transpiration and soil evaporation and the changes in DM over the same period. Carbon isotope discrimination (CID) can be used as an indirect and rapid tool to measure TE
and has been used in wheat to develop high TE cultivars of wheat for waterlimited environments (Richards, 2006). In lentil, CID values showed significant variation among genotypes, plant parts and age (17.8% in seed to 22.9%
in the leaf at flowering) under adequate water supply, and its values differed
significantly with response to watering regimes (Matus et al., 1995a, b). Lentils also had lower discrimination values under water deficit than in wellwatered conditions. Johnson et al. (1995) reported a negative correlation
between CID and TE in lentil. However, Matus et al. (1996) and Turner et al.
186
R. Shrestha et al.
(2007) found no correlation between CID and TE in different genotypes of

lentil grown under adequately watered conditions and concluded that CID
is not a useful selection tool for TE in lentil. Selection for high TE by any
technique has not been undertaken in lentil breeding programmes.
Dehydration tolerance
Dehydration tolerance is the ability of plant membranes to withstand
mechanical injury/degradation, or the ability of membranes and cytoplasm
to withstand protein denaturation and where cells continue metabolism at
low leaf water status (Turner et al., 2001). Most crop plants die as dehydration reaches a critical level (Turner et al., 2001). Crop species and genotypes vary in the lethal water potential (lowest water potential experienced
by the last viable leaf) (Flower and Ludlow, 1987). In lentil the mechanism
of dehydration tolerance or the level of critical water potential needs further investigation.
12.5. Conclusion
Lentil is an important source of protein and other nutrients essential for
human health, its straw is a valuable feed for livestock. The crop is primarily grown in rainfed conditions, where it is frequently subjected to moisture
deficit conditions. In South Asia, lentil is grown during the winter months
on residual soil water after the harvest of rice, while in West Asia and in
other Mediterranean-type environments such as Australia it is usually sown
during the wet winter months, but in both cases a reducing soil water supply coupled with increasing temperatures during reproductive development in spring results in terminal drought and a limitation to yield.
Rapid phenological development allows the crop to complete its life
cycle before the onset of severe terminal drought, and therefore is an important trait for crop adaptation in water-limited environments. Lentils exhibit
drought escape whereby terminal drought induces early maturity and
senescence. Lentil has an indeterminate growth habit and water deficits
during the vegetative stage reduce canopy development, net photosynthesis and hence assimilate supply. Water deficits during the reproductive
period reduce the reproductive sinks to compensate for the reduced assimilate supply. Mild water deficits, however, promote flower production and
therefore increase seed yield in some lentil genotypes. Although physiological parameters such as leaf photosynthesis, stomatal conductance and OA
are affected by water deficits, no direct correlation between these traits and
seed yield has been observed. Also, as there was no genotypic variation in
rates of leaf photosynthesis and stomatal conductance, and as variation in
OA among genotypes was related to varying degrees of drying, breeding
for increased yields using these physiological traits has not been pursued
to date.
187
The major impact of water deficits in lentil is on reproductive processes.

Water deficits restrict crop growth, reduce flower production by almost half
and reduce the number of fruiting nodes with a significant increase in the
proportion of fruiting nodes bearing a single pod. Pod and seed numbers
are further reduced as a result of increased abortion of flowers and pods.
The reduction in seed number as a result of reduced flower production and
increased flower/pod abscission under water deficits are similar to those
observed in other legume species. However, water deficits do affect the
number of seeds per pod, suggesting that the reduction in seed number balanced the reduced supply of assimilates and helped to maintain seed size.
Seed size and SGR in lentil are conserved under terminal drought, as in
other grain legumes. Reduced pod and seed numbers are a compensatory
mechanism in response to reduced assimilate supply induced by water
deficits, and so the SGR and seed size are maintained. As seed size in lentil
is the primary determinant of quality and price in the marketplace especially for green lentil, the ability to maintain seed size under water deficits
is of particular benefit as the majority of lentils are grown in rainfed environments.
Agronomic options to improve yields in water-limited environments
centre around early planting in Mediterranean-type environments and
rapid establishment in the post-rainy season where lentils are grown on
residual soil water. Seed priming has been shown to benefit yields by
increasing plant establishment when lentils are sown just prior to, or immediately after, harvest of a previous rice crop. Minimum tillage with stubble
retention will also lead to improved crop establishment, reduced soil evaporation loss and greater WUE.
Lentil genotypes (pilosae, microsperma type) from South Asia flower and
mature early so that the crop matures before there is an abrupt temperature
increase and soil water recedes. The large-seeded genotypes from Chile,
Greece, Iran, Iraq, Jordan, Syria, Tunisia and Turkey show a greater degree
of cold tolerance than small-seeded lentils. In South Asia to date, breeding
has concentrated on selection of large-seeded types and yield improvement
through early flowering, an increase in DM and disease resistance. Specific
crosses have been made to incorporate the drought-resistance traits from the
South Asian genotypes, specifically early flowering, with the high yield and
high DM of the West Asian genotypes. This hybridization and population
improvement helps to broaden the genetic base of commercial genotypes.
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Turner, N.C. (1979) Drought resistance and adaptation to water deficits in crop plants.
In: Mussell, H. and Staples, R.C. (eds) Stress Physiology in Crop Plants. John Wiley
and Son, New York, pp. 343367.
Turner, N.C. (1986) Adaptation to water deficits a changing perspective. Australian
Journal of Plant Physiology 13, 175190.
Turner, N.C. (2003) Drought resistance: a comparison of two research frameworks. In:
Saxena, N.P. (ed.) Management of Agricultural Drought: Agronomic and Genetic
Options. Science Publishers Inc., Enfield, New Hampshire, USA, pp. 89102.
Turner, N.C. (2004) Agronomic options for improving rainfall-use efficiency of crops
in dryland farming systems. Journal of Experimental Botany 55, 24132425.
Turner, N.C. and Jones, M.M. (1980) Turgor maintenance by osmotic adjustment: a
review and evaluation. In: Turner, N.C. and Kramer, P.J. (eds) Adaptation of Plants to
Water and High Temperature Stress. John Wiley and Sons, New York, pp. 87103.
Turner, N.C., Wright, G.C. and Siddique, K.H.M. (2001) Adaptation of grain legumes
(pulses) to water-limited environments. Advances in Agronomy 71, 193231.
Turner, N.C., Davies, S.L., Plummer, J.A. and Siddique, K.H.M. (2005) Seed filling in
grain legumes (pulses) under water deficits with emphasis on chickpea (Cicer
arietinum L.). Advances in Agronomy 87, 211250.
Turner, N.C., Palta, J.A., Shrestha, R., Ludwig, C., Siddique, K.H.M. and Turner, D.W.
(2007) Carbon isotope discrimination is not correlated with transpiration efficiency
in three cool-season grain legumes (pulses). Journal of Integrative Plant Biology
49, 14781483.
van Duivenbooden, N.M., Pala, C.S. and Bielders, C.L. (2000) Cropping systems and
crop complementarity in dryland agriculture: a review. Netherlands Journal of
van Kessel, C. (1994) Seasonal accumulation and partitioning of nitrogen by lentil.
Plant and Soil 164, 6976.
Varshney, J.G. (1992) Effect of sowing dates and row spacing on the yield of lentil
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Waldia, R.S., Singh, V.P. and Kharb, R.P.S. (1988) Stability of seed yield of some lentil
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Walsh, M., Siddique, K.H.M., Hashem, A. and Seymour, M. (2004) Weed management
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127134.
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efficiency of chickpea and lentil in a Mediterranean environment. Australian
13
Soil Nutrient Management
S.S. Yadav,1 David L. McNeil,2 Mitchell Andrews,3 Chengci Chen,4

Jason Brand,5 Guriqbal Singh,6 B.G. Shivakumar7 and B. Gangaiah7
1National
Agricultural Research Institute (NARI), Papua New Guinea; 2University of

Tasmania, Hobart, Tasmania, Australia; 3University of Sunderland, Sunderland, UK;
4Central Agricultural Research Center, Montana State University, Moccasin, Montana,
USA; 5Department of Primary Industries, Horsham, Victoria, Australia; 6 Punjab
Agricultural University, Ludhiana, Punjab, India; 7Indian Agricultural Research
Institute, New Delhi, India
13.1. Introduction
If soil fertility is non-limiting and the crop is relatively free from weeds and
diseases through all growth stages, then lentil growth is greatly dependent
on weather conditions. In this situation, physiological processes respond to
changes in air and soil temperature, solar radiation, moisture availability
and wind speed. The importance of weather inputs is highlighted by the
inputs into a lentil crop growth model (LENMOD) developed and calibrated in experiments carried out on a silt loam soil at Lincoln, Canterbury,
New Zealand (43.38S, 172.30E, 11 m above sea level). The model was
created using the cultivar Titore, a small-seeded, red variety of lentil
(McKenzie and Hill, 1989; McKenzie et al., 1994). LENMOD assumes soil
fertility is non-limiting and requires the input of daily values of maximum
and minimum air temperature, solar radiation, precipitation, potential
evapo-transpiration and day length. With only these inputs, LENMOD has
accurately predicted lentil growth in a wide variety of situations (Andrews
et al., 2001).
However, in many situations soil fertility is, or may become, limiting
and long- and short-term availability, and addition and removal of nutrients can become a major limitation on lentil production. Under these circumstances, there is a need to determine the biological and economic
consequences of the available options for dealing with limitations affecting
yield and productivity. The options available are varied and include addition of nutrients in organic or inorganic forms, biological additions (e.g.
rhizobia), crop modifications and soil management and ameliorations. The
economics and effectiveness of these treatments will interact with the variety, availability of products, yield expectations, state of local knowledge,
194

195
growing system, local economics and destiny of the crop. This chapter will
consider all these management options and interactions as they apply to
both high productivity and subsistence cropping of lentils.
13.2. Nutrient Requirements of Lentil

Nutrient removal by the lentil crop
The magnitude of nutrient removal from the soil by a lentil crop is dependent on nutrient availability, crop growth and development, and ultimately
yield. Traditionally, lentil is grown for subsistence or local consumption on
marginal lands on stored soil moisture with no inorganic nutrient input.
Under these conditions, lentil yields are relatively low, for example, average
yields in India are 0.7 t/ha (FAOSTAT, 2006). However, in several countries,
in particular Canada and the USA, over the past 25 years, lentil production
has increased greatly in high fertility soils with adequate soil moisture.
Under these conditions, yields of 3 t/ha can be achieved (Andrews et al.,
2001; FAOSTAT, 2006). The concentrations of essential and beneficial nutrient elements reported in lentil seed are shown in Table 13.1, along with
their role in lentil growth and development. Table 13.2 indicates the removal
of nutrients by high- and low-yielding lentil crops based on the range of
concentrations given in Table 13.1.
Generally, lentils obtain the major proportion of their N requirements
via N2 fixation and on average (but with a high degree of variability) their
residues contribute around 20 kg N/ha to the soil N pool (McNeil and
Materne, 2007). However, there will be a net removal of all other nutrients
from the system on harvest of lentil seed. The magnitude of this loss will
depend on the concentrations of nutrients in the seed and seed yield. For
some nutrients, for example Co, losses are small and are likely to be replenished naturally. However, in other cases, losses will be substantial, especially under high productivity. For example, P loss will be in the range
2.947.25 g/kg of seed (Table 13.1) and soil P levels could eventually limit
plant growth if P fertilizer is not applied. Therefore in many situations there
will be effects of nutrient addition and the response of the lentil crop to different nutrient fertilizers will now be considered.
Nutrient requirements of the lentil plant
With respect to macronutrients, there have been reports of positive
responses to addition of N, P, K, S, Mg and Ca although their effects are not
consistent. Generally, lentil can fix adequate amounts of N for growth, and
addition of N fertilizer does not increase yield (McNeil and Materne, 2007).
However, lentil, as other grain legumes, can take up and assimilate N from
the soil prior to nodulation to an extent that affects growth (Andrews et al.,
1992). If sown into soil with extremely low available N or cold, wet soil,
addition of 1025 kg N/ha at sowing can stimulate early growth, nodulation
196
S.S. Yadav et al.
Table 13.1. Concentrations of essential and benecial nutrient elements in lentil seed, the
role of these elements and the recommendation in relation to their application to lentil crops
(Sources: Andrews et al., 2001; Raven et al., 2004; Urbano et al., 2007; Yadav et al., 2007).
Nutrient
Concentration
(g/kg)a
37.248.8
2.423.7
Ca
Mg
0.422.10
0.492.20
2.947.25
1.00
Cl
NAd
Fe
0.0650.133
0.0090.011
Mn
Zn
0.0120.054
0.0250.062
Cu
0.0090.032
Mo
0.0070.013
Ni
Na
Si
Co
0.0010.003
0.110.79
NA
< 0.001
Roleb
Recommendationc
Nucleic acids, proteins,

cofactors
Cofactor for many enzymes,
osmoticum
Signalling, osmoticum
Cofactor for many enzymes,
osmoticum
Nucleic acids, cofactors,
regulation
Amino acids, lipids,
osmoticum
Cofactor for photosystem II,
osmoticum
Cofactor for many redox
reactions
Cofactor in cell walls
1025 kg N/ha under specic

conditions
2274 kg K/ha under specic
conditions
Not recommended
Not recommended
Cofactor for photosystem II

Enzyme cofactor, e.g. carbonic
anhydrase
Enzyme cofactor,
e.g. cytochrome oxidase
Enzyme cofactor including
nitrogenase
Enzyme cofactor, e.g. urease
Osmoticum
Mechanical, leaf posture
Vitamin B12, synthesized by
rhizobia
20 kg P/ha under specic

conditions
20 kg S/ha under specic
conditions
Not recommended
As per need under specic
conditions
0.51.1 kg B/ha under
specic conditions
Not recommended
26 kg Zn/ha under specic
conditions
Not recommended
935 g Mo/ha under specic
conditions
Not recommended
Not recommended
Not recommended
Not recommended
aTaken
from Andrews et al. (2001), Urbano et al. (2007) and Yadav et al. (2007).
from Raven et al. (2004).
cReferences given in text.
dNA, not available.
bModied
and N2 fixation (McKenzie et al., 2007). Application of higher doses of N

fertilizer or addition of N to fertile soils is likely to have a negative effect on
nodulation and N2 fixation (Turay et al., 1991; Cash et al., 2001).
The literature shows a wide variation in the response of lentils to application of P. McKenzie et al. (2007) emphasized this point by presenting
results from trials in different countries which range from no response to a
60% increase in yield with additional P up to around 20 kg/ha. In central
Montana of the USA, triple super-phosphate was applied to a clay-loam soil
197
Table 13.2. Removal of essential and benecial nutrients in lentil seed, based on maximum
and minimum seed concentrations (cf. Table 13.1) and unlimited (3 t/ha) and limited (0.7 t/ha)
production environments.
Nutrient removal in seed at 3 t/ha
crop (kg/ha)
Nutrient removal in seed at 0.7 t/ha

crop (kg/ha)
Nutrient
Minimum
Maximum
Minimum
Maximum
N
K
Ca
Mg
P
S
Fe
B
Mn
Zn
Cu
Mo
Ni
Na
Co
111.600
7.200
1.260
1.470
8.820
3.000
0.195
0.027
0.036
0.075
0.027
0.021
0.003
0.330
0.003
146.400
71.100
6.300
6.600
21.750
3.000
0.399
0.033
0.162
0.186
0.096
0.039
0.009
2.370
0.003
26.040
1.680
0.294
0.343
2.058
0.700
0.046
0.006
0.008
0.018
0.006
0.005
0.001
0.077
0.001
34.160
16.590
1.470
1.540
5.075
0.700
0.093
0.008
0.038
0.043
0.022
0.009
0.002
0.553
0.001
containing 14 ppm extractable Olsen-P. Significant yield increase occurred

only once in the 2-year study with the lentil variety Vantage at the rate of
14.7 kg P/ha compared to the unfertilized control. However, the increase of
forage yield due to P application was much more apparent than that of
grain yield (Wen et al., 2008). There are regions of Bangladesh, India, Pakistan and other South Asian countries, where application of around 20 kg P/
ha is likely to increase lentil yield (Ali et al., 2000). Also, in high production
systems, additional P at around 20 kg/ha could be beneficial especially if
soil P concentration (sodium acetate extract) is less than 4 ppm (Muehlbauer
et al., 1995; Mahler, 2007). Similarly, there are areas in South Asia where
lentil yield is likely to respond to addition of 20 kg S/ha (Ali et al., 2000)
and the general recommendation under improved practice is to add around
20 kg S/ha especially if soil tests give SO4_S concentrations of less than 10
ppm (Muehlbauer et al., 1995; Mahler, 2007).
There are few reports in the literature of lentil yield responding to additional K, Mg or Ca (Ali et al., 2000; McKenzie et al., 2007). Nevertheless, in
the USA, Muehlbauer et al. (1995) recommended application of 22 kg K/ha
to lentil based on a sandy or eroded soil while Mahler (2007) recommended
application of 56 kg K/ha if sodium-acetate-extracted soil K is 5070 ppm
and 74 kg K/ha if soil K is less than 50 ppm. Addition of K may improve
lentil cooking quality (Wassimi et al., 1978).
The literature indicates that Zn, Mo and B are the micronutrients most
likely to limit lentil growth in the major lentil-growing areas while Mn and
198
S.S. Yadav et al.
Fe may limit lentil growth locally and lentil yields may increase with seed
treatments or foliar sprays of ferrous sulfate (Ali et al., 2000). There are a
number of reports of a yield response of lentil to additional Zn in the range
26 kg/ha (Sharma et al., 1993; Jain et al., 1995; Srivastava et al., 1999). Lentils are particularly susceptible to Zn deficiency when grown after paddy
crop as Zn deficiency is common in these soils (Saxena, 1981). The critical
limit for available Zn ranges from about 0.5 to 1.81 ppm depending on the
extractant used (Saxena, 1981; Mahler, 2007). Mahler (2007) recommended
application of 6 kg Zn/ha when Zn soil test levels were less than 0.6 ppm.
Also, Shuknesha (1977) reported that soil P and Zn can interact such that
when the P : Zn ratio is greater than about 400, Zn may become limiting.
Molybdenum is a component of the nitrogenase enzyme and is thus a
key micronutrient for legumes. Seed treatment with Mo has been reported to
increase seed yields in Bulgaria, India and the USA (Golubev and Lugovskikh, 1974; Ali et al., 2000; Mahler, 2007). Application of 935 g Mo/ha is
recommended in the USA for soils of pH < 5.7 as a routine treatment, or every
third time lentils are grown in a soil (Muehlbauer et al., 1995; Mahler, 2007).
Boron deficiency has been reported to limit lentil growth in soils in
India, Nepal and the USA (Sakal et al., 1988; Srivastava et al., 1999; Mahler,
2007). For example, application of 0.5 kg B/ha increased yield from 0.1 to
1.5 t/ha in a trial in Nepal (Srivastava et al., 1999). Mahler (2007) recommended application of 1.12 kg B/ha to soils testing less than 0.5 ppm B.
Considerable variation in tolerance to B deficiency exists within lentil germplasm and it is possible to select genotypes for low B soils (Sakal et al., 1988;
Srivastava et al., 2000).
13.3. Nutrient Management involving Chemical, Organic and

Biological Sources
Adequate nutrition is essential for optimum crop growth and grain yield
and quality of lentil. Table 13.2 indicates the minimum quantities of nutrients that will need to be replaced to maintain the nutrient status of the soil.
However, response to applied nutrients, in terms of biomass production
and grain yield, will only occur where the availability of a nutrient in soil is
below the critical levels to maintain adequate growth. Nutritional response
will also depend on the yield potential of the crop which will tend to be
lower in water limited situations, that is a crop with a potential of 3 t/ha
grain yield will require more nutrients than a crop with 0.7 t/ha potential.
Some nutrients need to be replaced regularly, while others almost never
because of high natural concentrations and inputs from residues and biological processes. Availability of a nutrient can be influenced by many soil factors
such as pH, water status, buffering capacity and clay content (Marschner,
1993). In addition, nutrient availability can often vary throughout a cropping
season as a result of changes in soil temperature, water status and depletion
from crop usage.
199
Improved nutrition can improve the tolerance of the crop to disease and
other biotic and abiotic stresses (Marschner, 1993). Conversely, where nutrition promotes excessive vegetative growth, it can create conditions conducive to fungal disease. Excessive vegetative growth can also place a crop at
greater risk of yield loss from terminal drought, particularly in rainfed cropping regions, as the vigorous growth consumes moisture reserves that could
be utilized during grain fill.
Macro- and micronutrients essential to lentil crop growth are similar to
most other crop species. However, ultimately, for maximum yield and quality most nutrients will act synergistically, in that the full response to the
addition of one nutrient will only be observed when there are adequate
concentrations of other nutrients.
Macronutrients
As lentils have the ability to fix N, in most cases, a lentil crop can be grown
without additional N when adequate levels of other nutrients and Rhizobium bacteria are available, particularly those that are critical for nodulation
and N2 fixation (see Quinn, Chapter 15, this volume). Additional N up to 20
kg/ha can be beneficial (Kumar et al., 1993; Carter and Materne, 1997; McKenzie et al., 2007). However, high levels of N can reduce nodulation and N2
fixation, but generally do not result in significant yield loss (Bremer et al.,
1989; Singh et al., 1997). Phosphorus is one of the most critical nutrients for
optimum crop growth and grain yield in lentil. Several studies which have
been mentioned earlier also have indicated that in low P soils, application
of approximately 1020 kg P/ha will generally maximize yields (Kumar
et al., 1993; Carter and Materne, 1997; Ali et al., 2000). Low P can also reduce
nodulation and N2 fixation processes (Gupta and Sharma, 1989; R. Norton,
Horsham, 2003, personal communication). Gupta and Sharma (1989) also
reported increases in seed protein with addition of P fertilizer. Similar to N
and P, positive grain yield and biomass response to K, S, Mg and Ca have
been observed (Singh and Chauhan, 1987, 2005; Srinivasarao et al., 2003;
Kiss et al., 2004). Increased K and Mg have also been shown to have benefits
for N2 fixation (Srinivasarao et al., 2003; Kiss et al., 2004).
Micronutrients
Additional micronutrient application can have significant effects on crop growth
and yield. Increases in biomass and/or yield have been recorded for Zn
(Khurana et al., 1998), Mn (Brennan and Bolland, 2003), Fe (Singh et al., 1985),
Mo (Biswapati et al., 1998) and B (Srivastava et al., 2000) when soil availability
was limited. Increased levels of grain protein have also been measured with
additional Zn (Dawood and El-Far, 1994) and Mn (Brennan and Bolland, 2003).
Nutrients to maximize crop growth and yield can be supplied through
both biological (e.g. bio- or organic fertilizers and crop residues) and inorganic chemical methods as discussed below.
200
S.S. Yadav et al.
Organic sources
Farmyard manure (FYM) conserves N during the initial phase of the crop
cycle, reduces N loss and provides better synchronization of N availability
and crop N demand during the latter part of the annual crop cycle (Ghoshal, 2002). Application of 10 t FYM/ha increased the grain yield of lentil
considerably (Sinha and Sakal, 1993). Even the lower dose of 5 t FYM/ha
has been reported to increase the grain yield of lentil by 12.7% (Sekhon et al.,
2008), 17.9% (Singh et al., 2003) and 20.2% (Singh and Sekhon, 2006). FYM
not only provides macro- and micronutrients to the crop but also improves
physical and biological properties of the soil, which help in improving the
grain yield. However, application of FYM to lentil is not a common practice
among farmers.
Chemical fertilizers
A range of solid and liquid chemical fertilizers are used on lentils. In most
cases, the solid chemical fertilizers are in a granulated form and applied at,
or near, sowing. Most of the fertilizers used currently are referred to as
high analysis, meaning that the fertilizer has been processed to contain
accurate amounts of prescribed nutrients. Two of the most common high
analysis fertilizers used in Australia are Grain Legume Super and MonoAmmonium Phosphate having NPKS ratio of 0:15:0:7 and 10:22:0:0,
respectively (Day et al., 2006). These are often applied with added Zn concentrations of up to 5%. Low analysis products such as super-phosphate
can also be used to provide P. While these products contain a known
amount of P, they have not been highly refined and may contain many
additional macro- and micronutrients beneficial for growth. Generally, with
the solid, granular fertilizers, the focus is applying sufficient P to maintain
growth. One risk with solid, granular fertilizers is that they are in a highly
concentrated form, and if placed too close to the seed, they can be toxic.
Hence, in many mechanized regions of the world, technologies have been
developed to place the fertilizer below or to the side of the seed.
Liquid fertilizers can also be used at sowing, but are most commonly
used on a crop to correct nutrient deficiencies that appear during the season. As liquids can often be taken up through the leaves, rapid responses to
their application (e.g. crop colour change) can be observed compared with
the granular fertilizers. Similar to the high analysis granular fertilizers, they
generally contain accurate concentrations of nutrients.
Crop residues
Crop residues are generally considered as waste. However, they contain
large amounts of nutrients and are also a good source of organic matter.
Wheat residue applied in combination with fertilizer to a rice-lentil rotation
201
increased the activity of soil organic matter, thereby enhancing the nutrient
supply rate, which in turn resulted in increased crop growth and better
grain yields (Singh, 1995). However, crop residues are not always available
for use in nutrient management as they have alternate uses as animal feed
especially in developing countries. In some countries such as India (especially northern India), rice straw from the combine-harvested crop is burnt,
which not only results in loss of nutrients but also causes environmental
pollution. The use of rice straw as an effective nutrient management option
needs to be evaluated and, if found suitable, should be popularized.
13.4. Management of Toxic Elements in Lentil

pH Management and pH interactions
Lentil grows well on slightly acidic (5.56.5 pH) to moderately alkaline
(7.59.0 pH) soils. However, it shows best production performance at neutral pH. Lentil performance is severely hampered in soils with acidic (<5.0)
and high (>9.0) pH. Soil pH influences nutrient availability, making some
nutrients deficient while others highly available to the point of becoming
toxic for growing plants. The effects of soil pH and nutrient availability and
their management are discussed below with respect to lentil.
Soil acidity
Soil acidity leads to deficiency of P, K, Ca, Mo and B. It can also lead to toxicity of Al, Fe and Mn. Molybdenum deficiency leads to hampered nodulation and symbiotic N2 fixation of legumes. Owing to the above effects, lentil
productivity was reduced by 71% in acidic soil as compared to normal soil
(Dwivedi, 1996). To alleviate the adverse effects of acidity, liming of soils is
advocated to bring the soil pH to around 6.0. A decrease in exchangeable
Al, Fe and Mn (as a result of their precipitation as carbonates, oxides and
hydroxides, respectively) coupled with an increase in pH, exchangeable Ca
and Mg and available nutrients with application of 6 t lime/ha (Table 13.3)
resulted in increased lentil seed yield from 325 to 1125 kg/ha (Dwivedi,
1996).
The increased availability of P at higher soil pH was ascribed to reduced
P fixation while the increased N availability was a result of the accelerated
decomposition of organic matter coupled with increased biological activity.
Liming increases the availability of soil reserve Mo by increasing pH, hence,
the response to Mo fertilization in the presence of lime would be minimal.
Besides lime, P and Mo fertilization were found effective in alleviating the
Mo deficiency for lentil in acidic soils in the descending order of Mo > P >
lime (Mandal et al., 1998). The interaction effects of P (at medium levels of
25 kg/ha) and Mo fertilization on dry matter production and Mo uptake by
lentil were found to be positive (Mandal et al., 1998). This was possibly
202
S.S. Yadav et al.
Table 13.3. Effect of liming on soil pH, yield and nutrient availability for lentil in an acidic soil
(Source: Dwivedi, 1996).
Lime dose
(t/ha)
0.0
1.5
3.0
4.5
6.0
CDb P = 0.05
aOC,
bCD,
Exchangeable cations
Cmol (p+) (kg/ha)
Available nutrients (kg/ha)
pH
Yield
(kg/ha)
Al
Fe
Mn
Ca
Mg
P2O5
K 2O
OCa
4.6
4.8
5.3
5.9
6.7
4.6
325
475
615
700
1125
685
1.90
1.70
1.40
1.00
0.50
0.25
0.09
0.06
0.05
0.03
0.01
0.02
0.06
0.05
0.03
0.02
0.01
0.01
1.2
1.8
2.4
3.2
4.5
0.5
0.9
1.2
1.8
2.2
2.5
0.8
185
210
225
250
280
15
4.6
5.7
8.3
10.5
13.6
2.5
410
435
465
440
425
18
0.75
0.90
1.02
1.05
1.16
Organic carbon.
Critical difference.
because of alleviation of Al toxicity owing to its precipitation as aluminophosphates by P fertilization (Haynes and Ludecke, 1981). Rhizobium becomes
ineffective in acidic soils; hence, pelleting of Rhizobium-inoculated seeds
with CaCO3 enhanced lentil performance (Sarkar and Pal, 1986).
The mobility of surface-applied lime is low; thus, it has little impact in
situations where subsoil acidity is a problem. Gypsum application is useful
in ameliorating subsoil acidity by precipitation of active Al as Al2(SO4)3 and
thus increasing the Ca:Al ratio of soil. Fly ash, owing to its alkaline nature,
is also effective in ameliorating soil acidity. Similarly, phosphate rocks with
low free carbonate content can also act as liming materials for acidic soils.
Although the increases in soil pH are low with rock phosphate application,
the decrease in exchangeable Al is significant with a corresponding increase
in exchangeable Ca of soils.
Salinity and alkalinity

Salinity
Lentil is one of the most salinity sensitive crops and when it is grown on
residual moisture in the post-rainy season (after rice or fallowing in the
rainy season in South Asia) it is prone to salinity stress as salts accumulate
in the soil solution and are then precipitated towards the soil surface as
the soil dries out. Lentil yield and its biological N contribution get reduced
by 90100% (Hoorn et al., 2001; Katerji et al., 2003) at an electrical conductivity (ECe) of 3.1. At over 0.8 ECe in saline soils, deficiency of K and Ca
(Finck, 1977) coupled with the toxicity of Na sulfates and chlorides results
in reduced nodulation of legumes (Bhardwaj, 1975). This results in poor
yield performance. Salinity-induced nodulation inhibition in lentil was
ascribed to inhibition of root hair expansion leading to root curling (Sprent
and Zahran, 1988).
203
An increase in NaCl concentration in soil resulted in increased uptake

of Na and Cl, and reduced dry weight, K concentration and K:Na ratio of
lentil. The decrease in K:Na ratio indicates the antagonism of Na on K
(Turan et al., 2007). Hence, K fertilization is effective in counteracting the
effects of Na at low to moderate levels of salinity. However, under high
salinity situations, K fertilization may be futile in excluding Na from plants.
The high chloride (Cl1) content of saline soils competitively inhibits phosphate (PO43) uptake of plants (Zhukovskaya, 1973).
Selection of salinity-tolerant genotypes is the most effective way of
overcoming the salinity impacts (Maher et al., 2003). Application of Ca was
found to reduce the Na:Ca ratio and the deleterious effects of NaCl salinity
on germination and seedling growth of lentil were reduced at a 14:1 ratio
(Astaraei and Forouzan, 2000). Inoculation of lentil seeds with Rhizobium in
a saline soil with an ECe of 6 dS/m increased dry matter production by
22.9% (Ahmad et al., 1986). Inoculation also increased the N and P contents
of plants. Seed pelleting of Rhizobium-inoculated seeds with CaSO4 enhanced
lentil performance in a saline soil (Poi, 2005).
Alkalinity
Alkaline soils are characterized by the occurrence of excess Na that
adversely affects the physical properties of soil under waterlogging and
nutrient availability to plants. In sodic soil (pH >8.5), N availability (owing
to high volatilization losses) as well as Fe is reduced. The reduced Fe availability is due to high CaCO3 (Singh et al., 1985) and production of bicarbonate with decomposing organic matter that increases the available phosphate
which reduces Fe availability to the plant (Brown et al., 1959). Zinc availability is reduced as a result of increased Zn fixation (Gupta et al., 1987).
Manganese is also less available (Brennan and Bolland, 2003). Sodic soils
can also lead to toxic concentrations of P (Chabra et al., 1981; Gupta et al.,
1987), B and Mo.
The application of gypsum reduces the available P (Chabra et al., 1981)
by lowering pH and ECe. Gypsum was more efficient in reducing these
parameters than pyrite (Misra et al., 2007). Gypsum application also decreases
the toxicity of Mo to plants owing to antagonistic effects of sulfate on the
molybdate ion. Similarly, gypsum aids in the transformation of B from highly
soluble sodium metaborate to relatively insoluble calcium metaborate (Gupta,
1979): this can result in substantial increases in lentil yield. Zinc fertilization
(20 ppm) in alkaline soils enhanced nodulation and biomass production of
lentil. Zinc induces alkalinity tolerance in lentil owing to reduced Na uptake
and increased K, Ca and Mg uptake resulting in a narrowing down of the
Na:K and Na:(Ca + Mg) ratios (Misra and Tiwari, 1998).
In calcareous soils, the critical limit of available Fe was estimated as 6.95
and 74.5 ppm in soil and lentil plant, respectively. In soils with <6.95 ppm
available Fe, lentil yield increase varied from 13.31 to 53.97% with application of 10 ppm Fe (Sakal et al., 1984). Though Fe deficiency is not seen as a
limitation on lentil production in Mediterranean climates where it originated,
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significant increases (14108%) in straw yield were reported with Fe fertilization (Erskine et al., 1993; Zaiter and Ghalayani, 1994). This can be of
immense value, as straw is highly important in these areas.
It is observed that though germination was not hampered up to a level
of exchangeable sodium percentage (ESP) 30, lentil plants failed to survive
beyond 25 ESP (Ashraf and Waheed, 1990). A decrease of 50% in biomass
production of lentil was recorded at 26 ESP (Das and Mehrotra, 1971). The
seed yield of lentil was reduced from 1.7 to 0.2 t/ha with an increase in ESP
from 8.5 to 36.0. The K, Ca, N and Mg uptake of lentil and crude protein
content of seeds also decreased at higher levels of ESP (Singh, 1988).
Toxicities
In specific environments, a number of other elements can reach toxic levels
sufficient to reduce lentil growth. In a wide variety of situations globally,
excess B can limit growth of lentil. In particular, B toxicity is a problem in
some arid areas of West Asia and Australia (Yau and Erskine, 2000; McNeil
and Materne, 2007). The variation in lentil germplasm to B toxicity can be
used to select B tolerant lines (Hobson et al., 2006). Lentil may also be grown
in soils contaminated with heavy metals (e.g. As) either naturally or by
human input. Under these circumstances, plant growth or safety for consumption may be affected. A variety of solutions have been proposed, such
as inoculation with arbuscular mycorrhizal fungi (Ahmed et al., 2006),
breeding for altered uptake or modifying water management.
13.5. Strategies for Optimizing Nutrient Use Efficiency in Lentil

Nutrients are required to enhance productivity and/or improve the quality
of the produce. However, the nutrient use efficiency of lentil is very low. The
nutrients can either be lost from the system and cause environmental pollution or are not effectively utilized by the crop. This results in increased cost of
production and lower profits for farmers. The nutrient requirements of lentil
will depend greatly on soil status and yield potential of the crop. There is a
need to increase the nutrient use efficiency using the following strategies.
Diagnostic tools for nutrient status

Before raising a crop, the soil should be analysed to assess the status of
macro- and micronutrients so that the crop is fertilized as per its need. The
response to applied P, for example, depends upon the initial P status in the
soil. In low and medium P soils, lentil responded significantly to 40 kg
P2O5/ha, whereas in high P soils, the response was significant only up to 20
kg P2O5/ha (Dhillon and Vig, 1996). It was also found that organic C content was a superior index to Olsen-P for assessing P availability to lentil.
205
Plant samples can also be analysed to assess the nutrient needs of the
crop. For lentil, the critical P concentrations in shoots, leaves and grains are
reported to be 0.28%, 0.33% and 0.26%, respectively (Rashid and Bughio,
1993). Similarly, leaf analysis could be used to assess the requirements for S
fertilization (Huang et al., 1992).
Time and method of fertilizer application in relation to environment

Phosphorus and K are generally applied at the time of sowing the crops.
However, they may also be applied by foliar application. Foliar spray with
solutions of calcium superphosphate at 11.9 kg/ha or a mixture of 23.8 kg
calcium superphosphate + 11.9 kg K2SO4/ha resulted in significant increases
in yield and yield components of lentil (Zahran et al., 1998).
Drought conditions decrease the Zn uptake by lentil (Gulser et al., 2004).
Soil application of 12.5 kg ZnSO4/ha provided the highest grain yield of lentil compared with soil application of 6.25 or 25 kg ZnSO4/ha or foliar application of Zn (Azad et al., 1993). However, Gangwar and Singh (1994) reported
that when Zn was applied through seed coating as ZnO or to soil or by foliar
application as ZnSO4, the grain yields from the three application methods
were in the order seed coating > foliar application > soil application.
Integrated use of different nutrient sources

To obtain optimum yields, the crop needs various nutrients in balanced doses,
which could be applied through different sources. The application of 18 kg N
+ 46 kg P2O5 + 20 kg K2O + 25 kg ZnSO4/ha in combination provided the
highest nodulation (Jain et al., 1995) and grain yield of lentil (Sharma et al.,
1993) in comparison with a range of less complex nutrient treatments. Yadav
et al. (2008) reported higher grain yield and N uptake by lentil with the combined inoculation of Rhizobium and Azospirillum brasilense. The combination
of phosphobacterial inoculation and 35 kg P2O5/ha provided higher water
use efficiency, total P uptake and net returns compared with no P application
or 35 kg P2O5/ha alone (Venkateswarlu and Ahlawat, 1993).
In the absence of Rhizobium inoculation, application of 12.5 kg N + 40
kg P2O5/ha is recommended (PAU, 2006). However, when seed is inoculated with Rhizobium, then 12.5 kg N + 20 kg P2O5/ha is sufficient, thus
offering an opportunity to save 20 kg P2O5/ha (Zarei et al., 2006).
13.6. Economics of Nutrient Management in Lentil

Socio-economic factors
Under all farming conditions, economic considerations (the cost of inputs
and expected returns from the output) should be taken into account before
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S.S. Yadav et al.
deciding the nature of nutrient management. When the expenditure on a

nutrient is not sufficiently compensated by way of increased yield and monetary returns, application of nutrients in the form of manures or fertilizers is
not warranted. Only when there is reasonable assurance that the crop yield
will be increased sufficiently to compensate for the costs of nutrients, does
the possibility exist of going for optimum doses of nutrients.
13.7. Economics of Application of Nutrients and Biofertilizers

While many experiments have looked at the impact of different nutrients
on the productivity of lentil, much less work has been reported on the economics of lentil nutrient management. A brief account of the reports on the
economics of the application of nutrients and biofertilizers to lentil is now
given.
Due to ability of lentil to fix atmospheric N2, little attention has been
given to the economics of N application. However, use of Rhizobium inoculants for managing N nutrition has been considered. In soils lacking adequate rhizobia, additional gross return, net return and net return per rupee
of investment of Rs2521, Rs2496 and Rs0.57, respectively, were obtained
after Rhizobium inoculation (Arpana et al., 2002).
Phosphorus availability can be an important factor determining the success of many legumes including lentil. Besides inorganic sources, P solubilizing bacteria and vesicular arbuscular mycorrhizae fungi have been utilized to
enhance the availability of P and increase the productivity and monetary
returns of lentil. Phosphobacterial inoculation alone or with 35 kg P2O5/ha
gave highest water use efficiency and net returns in a multi-treatment
experiment in Delhi, India (Venkateswarlu and Ahlawat, 1993). In an experiment in Pakistan, application of P at the rate of 50 kg P2O5/ha gave a return
of Rs9.0 for each rupee invested in fertilizer (Khan et al., 2006). (The approximate exchange rate is US$1 = Indian Rs42.)
Potassium is the third most important element needed for healthy
growth and productivity of lentil. It is often adequate in many lentil-growing
areas. However, lentil can show economic responses to application of K.
In Uttar Pradesh, India, application of 40 kg K2O/ha produced an additional yield of 273 kg/ha, a net return of US$65 and a benefit:cost ratio of
11 to 1 (IPNI, 2007). There are also reports that application of 2030 kg S/
ha would be sufficient to realize higher economic returns from lentil and
in some areas, increased monetary returns were observed up to 50 kg S/
ha (Shivakumar et al., 1995). Application of some micronutrients (Fe, Mo
and B) has had a positive effect on the growth and development of lentil
in many areas but their economics has not been studied in detail. However, in the B deficient areas of eastern India, an 8% increase in benefit:cost
ratio was observed in lentil when boronated NPK was used (Dibyendu
et al., 2006).
The economics of a few combinations of nutrients has been studied. In
central India, the heart of the lentil belt, application of 18 kg N + 46 kg
207
P2O5 + 20 kg K2O + 20 kg S/ha was observed to give the highest net

returns when compared to any individual or other combined applications
(Gwal et al., 1995). Sharma (1996) reported significantly higher grain yield,
net return and benefit:cost ratio with the application of 20 kg N + 50 kg
P2O5/ha along with Rhizobium seed inoculation compared to individual
applications or other combinations. Shah et al. (2000) reported that a 40 kg
N + 80 kg P/ha combination gave a net return of US$189 as compared to
the unfertilized control and there was an income of US$7 for each dollar
invested on fertilizers. The net return per rupee of investment with application of chemical N:P:K fertilizer at 20:40:20 kg/ha (Rs2.70) or 5 t FYM/
ha (Rs2.69) was superior over the control (Rs2.41) (Arpana et al., 2002).
As lentil is cultivated in many cropping systems, both sequential and
mixed cropping, quite often nutrient studies have been carried out for the
system rather than for the individual crops. Since it is difficult to separate
individual crop requirements in systems, it is useful to have a general
understanding of the system requirements rather than the component crop
requirements. The fertilizer treatment of 80 kg N + 30 kg K/ha to rice and
10 kg N + 60 kg P2O5/ha to utera lentil (relay seeding of lentil before rice
harvest) in a ricelentil cropping system, resulted in a net return of
Rs12,215/ha. The total net income was more than doubled by applying fertilizer to both crops under residual soil moisture (Dwivedi and Sharma,
2005). As lentil is an important component of many farming systems, nutrient management for the whole system will be more useful than for individual components. Further, the economics of nutrient management for all
nutrients taken together rather than for individual nutrient applications is
needed because of potential antagonistic or synergistic effects.
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Turan, M.A., Turkmen, N. and Taban, N. (2007) Effect of NaCl on stomatal resistance
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Turay, K.K., Andrews, M. and McKenzie, B.A. (1991) Effects of starter nitrogen on
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14
Cropping Systems and Production

Agronomy
Masood Ali,1 K.K. Singh,1 S.C. Pramanik1 and Mohamed Omar Ali2
1Indian
Institute of Pulses Research, Kanpur, Uttar Pradesh, India; 2Bangladesh

Agriculture Research Institute, Ishurdi, Bangladesh
14.1. Introduction
Lentil (Lens culinaris Medikus subsp. culinaris) was among the first crops
domesticated in West Asia and introduced into the Indo-Gangetic Plains
around 2000 bc (Cubero, 1981). It has become an important food legume
crop in the farming and food systems of many countries like West Asia, the
Indian subcontinent, Ethiopia and North Africa and to a lesser extent in
southern Europe. Its seed is a rich source of protein, minerals and vitamins
for human nutrition, and the straw is a valued animal feed (see Grusak,
Chapter 23, this volume; Erskine et al., 1990). Furthermore, as a result of its
high lysine and tryptophan content, its consumption with cereal provides
balanced nutrition. It has the potential to cover the risk generally encountered in dryland agriculture (Kumvanshi et al., 2004). Its ability to sequester
nitrogen and carbon improves soil nutrient status, which in turn provides
sustainability to production systems.
Lentil is produced in over 48 countries worldwide though the major
lentil-producing region is South Asia (see Erskine, Chapter 2, this volume).
In India, cultivation extends from the warm regions of Madhya Pradesh and
Maharashtra to the cold areas of Ladakh in Kashmir at an altitude of over
3500 m above sea level. The crop is thus widely adapted ecologically, primarily for rainfed systems and grown by a wide range of farmers from small to
large. Lentil growers in different countries face the challenges of relatively
similar biotic and abiotic stresses which are responsible for low productivity.
14.2. Lentil in Cropping Systems

Lentil is grown as a winter crop largely under rainfed conditions in residual/
conserved moisture. However, in the northern regions of India, like Punjab,
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Haryana and western Uttar Pradesh, it is also irrigated. Depending upon

annual precipitation and distribution, ground water availability, and demands
of the local cropping cycle, lentil is grown in different cropping systems as a
monocrop or as a component of double cropping, mixed cropping and relay
cropping.
In Bangladesh, lentil is mostly grown in the upland rice (aus)/jute (Corchorus capsularis L.) fallow-lentil cropping pattern, and is usually sown by midNovember. It is also broadcast in aman rice 710 days before harvest to capitalize
on residual moisture. These practices ensure profitable use of rice fallows. More
than 85% of the lentil area in Bangladesh is concentrated in the medium-high
topography lands of Jessore, Faridpur, Kushtia and Rajshahi districts.
The inclusion of lentil in various cropping systems benefits the companion crop or succeeding crop by improving the physical and chemical
properties of soil as a result of biological nitrogen fixation and other rotational effects (Ahlawat et al., 1981; Prakash et al., 1986; George and Prasad,
1989; Ali, 1994; Shah et al., 2003; Wisal, 2003). Experiments conducted in the
temperate ecosystem of Kashmir, India showed that nutrient status and the
balance of nutrients were significantly influenced by the cropping system.
The rice-lentil system increased the nutrient status of soil (Singh and Sofi,
2007). Results from two different experiments conducted in dry farming
areas of Central Anatolia (Haymana-Ankara and Gzl-Konya) to detect
the effect of fallow and winter lentil and different tillage systems on wheat
yields proved that sowing of winter lentil was more profitable instead of
leaving land fallow before wheat. Results showed that protein content of
wheat was increased by 2.122.15% after lentil in Haymana and Gzl (Eser
and Adak, 1999).
Monocropping
Monocropping lentil is a common practice in India in the heavy textured
soils of the Bundelkhand region of Uttar Pradesh and Madhya Pradesh,
where rainfall does not support a good kharif (monsoon) crop. Hence, fields
are left fallow during the kharif season and lentil is sown on the conserved
moisture. It is also practised in heavy rainfall areas (>1000 mm) of Bihar,
west Bengal, Orissa and eastern Uttar Pradesh in the areas of the Diara lands
and Tal lands. In these areas, frequent floods and poor drainage do not
allow raising a kharif crop. Thus lentil is sown as monocrop after the floodwater recedes. Diara lands are highly fertile due to the deposition of silt
during flood and hence a bumper crop of lentil is harvested.
In West Asia and North Africa (WANA) lentil is sown in rainfed cropping areas that receive from 300 to 450 mm annual average rainfall. At the
lower end of this rainfall spectrum it is taken in rotation with barley,
whereas from 350 to 450 mm the predominant cereal is durum wheat and
less commonly bread wheat. Temperatures are moderate in winter and relatively hot in summer leading to low and high evapotranspiration in winter
and summer, respectively.
Cropping Systems and Production Agronomy
215
Double cropping
Lentil is recognized as being important for crop diversification in South
Asia (Sarkar et al., 2004) and consequently its area is expanding. Rice-lentil
is a common production system practised in the lowlands of India, Pakistan, Bangladesh and Nepal. In India double cropping with lentil is generally practised after the harvest of kharif crops. The common crop rotations
are: rice-lentil; maize-lentil; cotton-lentil; pearl millet-lentil; sorghum-lentil;
and groundnut-lentil.
In Bangladesh, on the medium-high topography lands of Kushtia,
Rajbari, Magura and Jessore districts, which have sandy loam soils, lentil is
sown in a broadcast aus paddy rice/jute-fallow-lentil cropping system,
whereas on the medium-low topography, which has clay loam soil, lentil is
grown after the receding floodwater. The common cropping system in this
area is jute-fallow-lentil while the low-lying areas of Faridpur, Kushtia,
Rajbari and Natore, which have heavy clay soils, the broadcast aman (long
season) rice-lentil-fallow system is adopted.
Mixed cropping
In mixed cropping in South Asia, seeds of the component crops are mixed
and broadcast. Mixed cropping is practised in rainfed areas under moisture
stress conditions as an insurance against adverse biotic and abiotic factors
as well as being an economical and efficient use of farm inputs. Reduction
in seed yield of lentil was greater in mixed cropping with wheat in an irrigated environment (Ahlawat and Sharma, 1985). Popular crops for mixed
cropping systems with lentil are Indian mustard, barley and chickpea. In
Turkey, the mixture of 7080% lentil + 2030% wheat gave the highest landequivalent ratio for straw and grain yields (Cftc and lker, 2005).
Intercropping
Under this system, lentil is grown along with other crops in a definite row
ratio/planting geometry. The common intercrops with lentil are wheat, barley, Indian mustard and linseed in South Asia. For mixed/intercropping an
optimum seeding and planting configuration is very important to achieve
high total productivity. In various experiments, lentil + mustard (5:1 or 2:1
row ratio) and lentil + linseed (4:2 or 5:1 row ratio) showed higher yield than
growing lentil alone (Sarkar and Pal, 2005; Sekhon et al., 2007; Yadav and Tripathi, 2007). Singh and Rana (2006) reported that a paired row (30/90 cm) of
Indian mustard + lentil (two rows) intercropping system proved productive
as it gave 370 kg/ha extra yield of lentil without significantly affecting the
seed yield of mustard. Lentil is also intercropped with winter wheat and barley; however, lentil + wheat/barley intercropping is not always productive
and profitable (Gangasaran and Giri, 1985; Prakash et al., 1986). In Bangladesh,
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M. Ali et al.
Ahmed et al. (1987) found that the most profitable seeding ratio for wheat
and lentil was 100:50. Rehman and Shamsuddin (1981) reported that maximum total productivity, land equivalent ratio (LER) and net return under
wheat/lentil intercropping was obtained when 30% of the wheat seed rate
was sown between lentil rows spaced 30 cm apart. Experimental results on
the intercropping of lentil with linseed at different planting configurations
also revealed the advantage of intercropping over lentil grown as a sole
crop (Miah and Rahman, 1993). In the North Eastern Plains of India, western Terai of Nepal and a large area of Bangladesh, lentil is also intercropped
successfully with autumn sugarcane (Srivastava, 1975; Giri, 2005). In autumn,
sugarcane-lentil intercropping revealed higher cane equivalent yield than
sugarcane alone. In this system, sugarcane is planted in the first week of
October and two rows of lentil may be sown 30 cm apart between two rows
of sugarcane. Since lentil is a legume crop, there is no need for applying
nitrogen fertilizer. The lentil crop can be harvested by the end of March or
first week of April. Fertilizer is applied to the sugarcane after the lentil harvest. This system provides 15001600 kg/ha of additional lentil yield with
no adverse effect on cane yield. Lentil is rarely intercropped outside South
Asia.
Relay cropping
Relay planting of lentil in rice is practised in South Asia (Saxena, 1981; Ali
et al., 1993). Under relay cropping, lentil is broadcast into the standing rice
crop (paira or utera cropping) 1015 days before rice harvest. Relay cropping
with rice is generally practised in Chattishgarh and the eastern region
of India and eastern Terai of Nepal. In sandy loam soil of the mediumtopography areas of Jessore, Chuadanga, Magura, Meherpur, Kushtia,
Rajshahi and the Pabna district of Bangladesh farmers also grow lentil in
relay cropping with rice.
This practice takes advantage of the residual moisture remaining after
rice, gains time and economizes the inputs and ultimately is more productive than normal sowing (Chakraborty et al., 1976; Roysharma et al., 1984;
Gupta and Bhowmick, 2005).
14.3. Production Agronomy

Climate
Lentil requires a cold climate. It is sown as a winter season crop in South
Asia and lowland WANA. It is cultivated in a wide range of climatic conditions. The adaptation of lentil to different agroclimatic conditions is covered
in more detail by Materne and Siddique (see Chapter 5, this volume). It can
be grown successfully up to 3500 m above sea level. It is generally grown
on moisture conserved during the rainy season with no inorganic nutrient
217
input in South Asia. Under these conditions, lentil yields are relatively low,
for example, the average yield in India is 0.7 t/ha (see Erskine, Chapter 2,
this volume). However, in several countries, particularly Canada and the
USA, over the past 25 years, lentil production has increased greatly due to
better crop management (Andrews et al., 2001).
Soil
Lentil is adapted to various soil types, from sand to clay loam, provided
there is good internal drainage. Lentil does not tolerate flooding or waterlogged conditions, and does best on deep, sandy loam soils that are moderate in fertility. Good drainage is required, because even short periods of
exposure to waterlogging or flooded field conditions results in wilting and
drying of plants (because excess moisture causes a lack of oxygen supply to
roots, preventing root respiration). A soil pH near 7.0 is best for lentil production. Lentil performance is severely hampered in soils with acidic pH
(<5) and alkaline pH (>9). Soil pH influences nutrient availability, making
some nutrients deficient while other nutrients become highly available and
toxic to growing plants. A decrease in exchangeable Al, Fe and Mn (due to
their precipitation as carbonates, oxides and hydroxides, respectively) coupled with an increase in pH, exchangeable Ca, Mg and available nutrients
with application of 6 t lime/ha resulted in increased lentil seed yield from
325 to 1125 kg/ha (Dwivedi, 1996). In India lentil is grown on a variety of
soils such as light loams and alluvial soils of the Punjab and Uttar Pradesh
to black cotton soils of Madhya Pradesh. This crop is also suited to poorer
types of soils, low-lying situations such as in paddy fields and even to soils
of moderate alkalinity.
Cultivars
The genotype plays an important role in realizing high productivity. Since
genotype environment interactions are significant, choice of a genotype
depends on prevailing agroclimatic conditions, cropping systems, farmers
choice and local market preferences, etc. Lentil genotypes show significant
variation with respect to their productivity, stability and adaptability. Lentil varieties that are suitable for different agroclimatic conditions and cropping systems have been discussed by Sarker et al. (Chapter 8, this volume),
Rahman et al. (Chapter 9, this volume) and Muelhbauer et al. (Chapter 10,
this volume).
Seedbed preparation/land preparation

Land preparation varies according to the soil type, climate and preceding
crop in the field. Tillage options and seedbed preparation particularly in
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M. Ali et al.
North America, Australia and the WANA region are covered by Diekmann
and Al-Saleh (Chapter 16, this volume). A firm, smooth seedbed with most
of the previous crop residues incorporated is best for lentil. Uneven surface,
large clods, stones or protruding crop residue can interfere with seed placement and complicate later swathing and combining. The soil for the cultivation of lentil should be made friable and weed free so that seeding could be
done at a uniform depth. For optimum germination and crop establishment
in South Asia, a temperature range of 1525C is desirable. On heavy soils,
one deep ploughing followed by two to three cross harrowing, especially
under rainfed conditions, ensures proper conservation of moisture in the
soil and better germination and growth of the crop. In developed countries,
a heavy roller is used to smooth the soil surface after planting and to ensure
good soil-to-seed contact for uniform germination and emergence. This
practice also provides a smooth surface to facilitate harvesting either by
direct combining or swathing (Muehlbauer et al., 1998).
Work on tillage management of rice-based sequential cropping systems
at the Indian Institute of Pulses Research, Kanpur, showed that relay planting of lentil in the standing crop of rice gave low yields as compared with
crop grown with cross harrowing after the harvest of rice (Kumar and Ali,
1995). In light soils, less tillage is needed to prepare an ideal seedbed. In
rainfed areas, where moisture is conserved during the preceding monsoon,
land preparation should not be done using a soil-turning plough to ensure
minimum loss of water. Tomar and Singh (1991) reported higher uptake of
N and P by lentil with one ploughing as compared with zero tillage on
sandy loam soils of Agra, Uttar Pradesh. After harrowing, the field should
be given a gentle slope and nicely compacted using a wooden plank. There
should be proper moisture in the soil at the time of sowing for good germination of seeds and to support the plant stand.
Direct seeding (no-till) of lentil into standing stubble left after cereal
(wheat or barley) harvest is becoming an option in developed countries
where soil erosion is a problem. Direct seeding is recommended in the USA
for autumn-sown lentil as a means of conserving soil moisture and to provide some surface protection to reduce winter injury for the developing lentil plants (Chen et al., 2006).
Sowing time
Since lentil is grown in diverse types of agroecological zones in different
cropping systems, its planting time varies accordingly. The time of sowing
is very important in relation to plant growth and phenological development
to realize the full grain yield. Timely planting reduces incidence of rust
(Chaudhary and De, 2005), aphids and Stemphylium (Hossain et al., 2006;
Ihasanul Huq and Khan Nowsher Ali, 2007). Delayed sowing of lentil reduced
the incidence of soil-borne pathogens, collar rot and stunt (Agrawal et al.,
1976; Singh and Dhingra, 1980). However, in Bangladesh delayed planting
has been reported to increase disease and insect infestation and hamper
219
growth (Khan and Miah, 1986). Lentil is generally grown after the rainy
season on conserved soil moisture. Due to short and mild winters in India,
Bangladesh and Ethiopia, the life cycle of lentil is limited to 100120 days.
In South Asia, sowing of lentil begins in mid-October and continues to early
December; however, the optimum seeding time is before 15 November
(Sekhon et al., 1986; Singh and Ram, 1986; Ahmad and Motior, 1993; Ali
et al., 1993; Neupane and Bharati, 1993; Singh et al., 2005; Ihasanul Huq and
Khan Nowsher Ali, 2007). In Dholi and Bihar, India, a crop sown on 25
October outyielded crops sown on 4 November and 4 December. There was
a drastic reduction in yield and yield attributes of lentil when sown on 4
December (Kumar et al., 2005). However, planting can be delayed in the
foothills and eastern parts of Punjab to the end of November without much
loss in yield (Saxena and Yadav, 1976; Dhingra et al., 1983). In the central
zone of India where moisture is limiting, lentil is sown in mid-October. In
Assam and adjoining states, sowing is recommended in the first or second
week of October. Late planting leads to less vegetative growth and forced
maturity and consequently the grain filling is affected. Delay in planting
causes reduction in yield but the magnitude of reduction is large after 15
November (Ali et al., 1993; Nandan et al., 2007). In relay planting, broadcasting of seed depends on the timing of the end of the monsoon cycle and the
maturity of the rice crop.
In the Andean region of Latin America, sowing of lentil is generally
done in the early winter period following the onset of the rains. The lentil is
sensitive to weed competition. Throughout its range under winter rainfed
conditions in the lowland Mediterranean region, sowing is generally
delayed to allow cultivation after the early rains for weed control. So early
sowing with adequate provision for weed control generally results in major
yield gains. In such areas lentil is normally planted at the beginning of the
winter season with an appropriate sowing time being up to the middle of
December (Saxena, 1981). In southern Australia, where a dryland Mediterranean-type climate is prevalent, early sowing (late April to early May)
allowed a longer period for vegetative and reproductive growth, rapid canopy development, greater absorption of photosynthetically active radiation
(PAR), more water use, and, hence, greater dry matter production, seed
yield and water use efficiency than delayed sowing (Siddique et al., 1998).
In cold-prone highland areas of West Asia (above 859 m above sea level)
and in parts of the USA (see below), the crop is spring sown. Spring-sown
lentils in the highlands frequently suffer from terminal drought. However,
shifting sowing from spring to early spring or winter sowing using winterhardy cultivars can increase lentil production significantly. Winter cropping
allows optimum vegetative growth and the development of higher yield
potential (frequently >50% of spring-sown yields) and provides higher
water use efficiency from winter precipitation, providing the cultivar is
winter hardy. There is the potential to replace about 400,000 ha of spring
lentil with a winter crop in the highlands of West Asia (Sakar et al., 1988).
Research on autumn sowing of lentil in these highlands including modelling sowing time from climatic data is summarized in Keatinge et al. (1996).
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M. Ali et al.
Cultivars with winter hardiness are now being cultivated in Turkey and in
Iran such lines are under on-farm evaluation. The focus in winter/autumn
sowing is to translate the research results into production gains; in this
regard weed control is critical. In Argentina and Chile, sowing is done from
May to early July, and the lentils are harvested in November and December. Due to low temperatures that cause winter killing of most lentil cultivars, lentil crops in Canada and the USA are planted in early spring. In the
states of Idaho, Oregon and Washington in the USA the crop is planted in
mid- to late April; while in the northern plains states of North Dakota and
Montana and the western provinces of Canada, the crop is usually planted
in early to mid-May depending on conditions.
Plant population, spacing and methods of planting

Response of lentil to plant densities has been variable, depending upon
genotype, planting time and growing conditions. Seeding rate varies from
region to region and with seed size. In South Asia, a seed rate of 3040 kg/
ha is recommended (Zaman and Miah, 1989). In West Asian countries, a
higher seed rate of 100120 kg/ha produces higher yield, the optimum
plant stand being 275300 plants/m2 (Silim et al., 1990). In Alberta, Canada
107 seeds/m2, in the states of Washington Idaho, 215 seeds/m2, (Muehlbauer et al., 1998) and in North Dakota, 172220 seeds/m2 was optimum
(Eriksmoen et al., 2008). Bukhtiar et al. (1991) reported that in lentil the seed
rate varied according to seed size. The small-seeded cultivar L9-6 produced
a higher yield with a seeding rate of 30 kg/ha while large-seeded cultivar
AARIL 355 gave more yield with a 40 kg/ha seed rate. In Bangladesh a
seed rate of 3035 kg/ha with a plant density of 250 plants/m2 has been
found optimum (Rahman and Miah, 1989). A seed rate of 4045 kg/ha for
large-seeded lentil and 30 kg/ha for small seeded have been reported as
optimum by other workers (Khare et al., 1991; Singh and Singh, 2002).
A reduction in seed yield because of delayed planting could be minimized to a certain extent by relatively closer spacing and use of a higher
seed rate. A seed rate of 40 and 60 kg/ha gave the same yield when planted
on 1 December but when planted on 15 and 30 December a significantly
higher yield was obtained with a 60 kg/ha seeding rate (Ahlawat et al.,
1982). Nandan et al. (2007) found the higher seed rate of 60 kg/ha optimum
over 40 kg in Dholi, Bihar, India. Under utera conditions, a higher seed rate
is generally used as all seeds broadcast in a standing rice field do not germinate and the seedlings are damaged. In West Bengal, a seed rate of 7080
kg/ha gave a higher yield under utera conditions (Gupta and Bhowmick,
2005; Anonymous, 2006). Seed rates of 40 kg/ha and 60 kg/ha were found
to be optimum under conditions at Kanpur and Dholi, Bihar, respectively
(Kumar et al., 2005).
Raised-bed planting has emerged as a promising management technique. At Kanpur, India, raised-bed planting increased grain yield significantly, reduced irrigation requirements, improved branching and podding,
221
and increased water use efficiency over traditional flat-bed planting (Singh,
2006). In agriculturally mechanized countries, lentil is planted using grain
drills, but elsewhere it is still hand broadcast (Gowda and Kaul, 1982) or
seeding is carried out behind a desi (country) plough with a porah (the funnel attached behind the wooden plough used to sow seeds at the appropriate depth). Planting in rows promotes a uniform plant stand, allows for ease
of weeding and other cultural operations and improves productivity. Lentil
seeds should be sown at about 34 cm deep in rows 20 cm apart (Singh
et al., 2005). Bidirectional sowing in a row rectangularity of 22.5 30 cm
gave higher yield than unidirectional sowing in north India (Kler and
Dhillon, 1982).
Fertilizer management
Lentils are generally grown under very low input conditions in otherwise
inherently low productive soils and consequently yields are low. Saxena
(1981) reported that a lentil crop yielding 2 t/ha removed 100 kg N, 28 kg
P2O5 and 78 kg K2O and essential nutrients from the soil. Studies on the
relative contribution of production inputs under different agroecological
conditions in India revealed that in the central zone, fertilizer was the premier input followed by weed management (Ali et al., 1993). Judicious and
balanced use of nutrients helps the plant to grow vigorously and give
maximum yield besides developing resistance against biotic and abiotic
stresses. Thus it is imperative that the required nutrients are applied to
realize higher productivity. Information is reviewed on soil nutrient
management by Yadav et al. (Chapter 13, this volume) and on biological
nitrogen fixation and soil health improvement by Quinn (Chapter 15, this
volume).
Seed priming
Seed priming is another technology through which substantial yield
improvement can be achieved. Seed priming is a technique in which seed is
soaked in water overnight prior to sowing. This is generally done to hasten
germination, especially in cool temperatures and dry soils. Priming of seeds
advances germination by inducing biochemical changes in the seed. Seed
priming (immersing seeds in water for 8 h) prior to planting increased
yields by 2938% in different agroecological conditions (Ali et al., 2003). Ali
et al. (2005) reported that seed priming increased seed yield from 30 to 37%
in Bangladesh. Priming improved plant stands, enhanced early vigour and
resulted in early maturity, reduced disease and increased yield. Seed priming is quite beneficial in population management especially under rice-relay
cropping systems. Experiments conducted during 20052006 in India revealed
that sowing sprouted seeds was beneficial.
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M. Ali et al.
Water management
Crop water requirements depend on soil type, slope of the field, date of
sowing, crop duration, temperature and other factors. Field estimates of the
consumptive use of water have been made for different parts of Egypt and
North India. El-Gibadi and Badawi (1978), using the Blaney and Criddle
formula, indicated that the consumptive use of water in lower, middle and
upper Egypt was 365, 364 and 391 mm, respectively.
Most lentil crops in India are grown on conserved soil moisture as a
monocrop or on residual moisture (after rice harvest) under rainfed conditions. Moisture stress is usually the single most important bottleneck to
achieve the potential grain yields in the Indian subcontinent. Aspects of
drought physiology and the optimization of water use efficiency are to be
found in Shrestha et al. (Chapter 12, this volume). Different workers reported
water requirements in the range of 155483 mm in India (Tickoo et al., 2005).
Saraf and Baitha (1979) computed the water requirements of lentil by following the actual soil moisture depletion on a sandy loam soil of North
India. Water requirements of timely sown lentil ranged from 155 mm with
one irrigation to 214 mm with four irrigations. About 67% of the total soil
moisture depletion occurred from the first 30 cm soil layer and 25% from
second 30 cm layer.
On sandy loam soils with a low water-holding capacity, a positive
response to between one and three irrigations has been recorded on lighttextured soils (Ahlawat and Rana, 2005). Flowering is the most critical
period for irrigation. The highest grain yield of 1547 kg/ha was obtained
when irrigation was scheduled at an irrigation water (depth):cumulative
pan evaporation (IW:CPE) ratio of 0.6 coupled with 70 kg P2O5/ha, which
was 70% higher than the control at the same level of irrigation (Venkateswarlu and Ahlawat, 1993). In black clay soils of Jabalpur and sandy
loam soils of Pusa and Bikramganj, the optimum IW:CPE ratio was observed
to be 0.6 requiring two to three irrigations to produce maximum yields.
However, the irrigation water requirement as well as the total water use
was highest at Jabalpur. At Pantnagar, Ludhiana and Delhi locations in
India, two irrigations of 67 cm depth at different critical stages were adequate (Prihar and Sandhu, 1987). An experiment conducted in the Mediterranean region at the main station at the International Center for Agricultural
Research in the Dry Areas (ICARDA) indicated grain and biomass yield of
lentil increased with increased supplemental irrigation at the reproductive
stage (Oweis et al., 2004). The response to irrigation depended on available
soil moisture and rainfall during the crop growth period. Experiments at
Alberta, Canada indicated that lentil yields increased significantly by supplemental irrigation during drought years in comparison to wet years (Agriculture and Agri-Food Canada, 2008).
Raised-bed planting economizes water as compared to flat-bed planting. At Kanpur, India, lentil planted on raised beds saved 2025% irrigation
water and increased the grain yield (Pramanik et al., 2007). Lentil is also
highly susceptible to over-watering. Temporary waterlogging can cause
223
severe damage. Therefore, only light irrigation is advocated and excess

water should be removed by drainage.
Weed management
Lentils are poor competitors against weeds because the plants are slender
and neither branch sufficiently nor grow tall enough to suppress weeds.
Weeds cause heavy losses to the lentil crop as they rob the soil of its nutrients and moisture. The magnitude of loss depends upon weed species and
their intensity, growing conditions, soil fertility and soil moisture. A full
treatment of weed management is to be found in Yenish et al. (Chapter 20,
this volume).
Effect of integrated management

Integrated management for fertilizers, manures, weeding and inoculum
application has tremendous effect on lentil production. Ali et al. (2003)
reported that a combination of cow dung at 3 t/ha, chemical fertilizers at
20:40:20 kg/ha of NPK, and one hand-weeding at 30 days after emergence
was the optimum management practice for economically viable production.
Miah and Rahman (1993) also reported that proper tillage accompanied by
the application of chemical fertilizers, weeding and disease control can
significantly increase lentil productivity.
Harvesting
Lentil should be harvested when plants begin to turn yellow and pods are
yellow-brown to brown. The mature crop should be harvested at the earliest date so as to avoid shattering of pods and loss of seeds. It is desirable to
harvest the crop during the morning hours to avoid breakage of plants and
shattering losses. The crop should be cut at 57.5 cm from the ground level
to minimize breakage. The crop is harvested by hand pulling, cutting with
sickle bars or by direct combining, depending on field conditions and availability of equipment. A detailed account of the mechanization of harvest is
given by Diekmann and Al-Saleh (Chapter 16, this volume). The handharvested crop is spread on the threshing floor for further drying and run
under tractor/bullocks for threshing. The threshed crop is cleaned manually or by winnowing. Where the crop is to be cut using sickle bars or
swathers, it is advisable to do this operation at higher moisture content of
1820% to avoid yield losses. Where direct combining is practised, the
plants should be sufficiently dry for ease of operation and the seeds should
be at or below 12% moisture content. In some cases in the USA and Canada,
the crop is sprayed with a desiccant to promote uniform maturity and to
ease the combining operation.
224
M. Ali et al.
After harvest, the seed, at a moisture content of 910%, should be stored

in airtight containers. For protection against bruchids in storage, the seeds
may be treated with certain oils (mustard/castor/neem/mahua (Maduca
indica)/coconut/groundnut/sesame) at 912 ml/kg or mixed with inert
material (charcoal powder or lime) or plant product (annola seed powder
or neem leaf powder/asafoetida/neem seed powder) at 912 g/kg seed to
avoid losses resulting from storage grain pests (Anonymous, 2006).
Drying and storage

Lentil seed after harvest should be adjusted to moisture contents of 910%
to prevent damage from mould-causing and storage pests. Lentil can be
dried in heated air dryers, but the maximum temperature should not exceed
43C to avoid cracking of seedcoats. Natural air drying has advantages over
heated air, but it is necessary to have a properly designed system. The
design must ensure good airflow through the seed. This requires that thinner layers of the seed must be used in this process. Further information on
the postharvest processing of lentil can be found in Vandenberg (Chapter
24, this volume).
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C
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emasculation.
Continued
1 Continued
F
Plate 1. Lentil crossing technique showing (D) removal of the anthers; (E) an emasculated female flower showing
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C
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Continued
3 Continued
F
Plate 3. (D) mosaic symptoms on lentil leaves caused by Broad bean stain virus (left) as compared to a healthy
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B
Plate 4. (A) Detection of Bean leaf roll virus by tissue-blot immunoassay (TBIA) in infected lentil stem blot (right)
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D
B
Plate 5. (A) Patch of dodder-infested lentil plants; (B) detail of lentil plants infected by dodder; (C) closer detail of
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Continued
5 Continued
Plate 5. (F) Orobanche aegyptiaca plant infecting lentil; (G) detail of germinated broomrape seed contacting lentil
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15
Biological Nitrogen Fixation and Soil

Health Improvement
Mark A. Quinn
Washington State University, Pullman, Washington, USA
15.1. The Importance of Biological Nitrogen Fixation and Legumes

Legumes have always been a critical component of agroecosystems throughout the world because of their ability to convert atmospheric N into usable
plant protein, their ability to grow in N-poor soils, and their contribution
to the pool of soil N that can be used by succeeding crops. The approximately 250 million ha of legumes grown in the world fix about 90 trillion g
of N each year (Graham and Vance, 2000). It has been estimated that nitrogenfixing legumes save approximately US$710 billion on N fertilizer each year
(Herridge and Rose, 2000; Hardarson et al., 2003). The fixed N is used
directly for plant growth and maintenance, and provides an excellent source
of protein for humans and livestock.
Six main genera of bacteria are involved in the symbiotic relationship
with legumes: Rhizobium, Bradyrhizobium, Sinorhizobium, Azorhizobium,
Mesorhizobium and Allorhizobium, and are collectively known as rhizobia.
There is often taxonomic specificity between species of legumes and rhizobia. For example, Rhizobium leguminosarum bv. viciae is the symbiont of all
Lens species, but also forms associations with other members of the Vicieae
(i.e. Vicia, Pisum, Lathyrus). The strength of the symbiotic relationship that
determines the degree of nodulation and nitrogen fixation is affected by
numerous factors, including legume and bacteria genotype, abiotic conditions, and interactions with other organisms. Many of these factors are
under the control of growers of cultivated legumes.
Legume cultivation is particularly important in N-poor soils where they
can reduce the need for costly inorganic fertilizers. Nitrogen deficiencies are
common in tropical and subtropical soils that have mineral fractions with a
low capacity to retain nutrients and water (Boddey et al., 1997). Not only
does nitrogen fixation allow the growth of legumes in these N-poor soils,
but also it can indirectly increase soil N levels and, thus, support the growth
229
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of other crops grown with the legumes or the succeeding crops after
legumes. Nitrogen-fixing legumes are an important component of healthy
soil to provide sustainable crop production systems.
15.2. Nodulation and Nitrogen Fixation in Lentil

At regional and global scales, the contribution of nitrogen fixation to crops
is substantial. Lupwayi and Kennedy (2007) estimated that in 1974, about
171 million kg N2 was fixed by field pea, lentil, dry bean and chickpea crops
in the Canadian prairies, and about 40 million kg N2 was fixed by grain
legumes in the USA. This represented about 7% of total fertilizer N used by
farmers. At a global level, crop legumes may fix about 90 trillion g N/year
(Graham and Vance, 2000), with lentils accounting for about 0.35 trillion g
N/year (McNeil and Materne, 2007).
At a local scale, nitrogen fixation by lentils is also substantial (Table
15.1). Unkovich and Pate (2000) reviewed various methods and problems
with estimating the amount of biological nitrogen fixation by crops, and
found a high degree of variation in recorded estimates within individual
crops, between years and between locations. Estimates of the percentage of
plant N derived from fixation in lentils ranged from 28 to 87% with an average of 63%. Walley et al. (2007) summarized results from 38 field experiments
with lentils conducted in Canada and reported a range of 988% N derived
from fixation, with a median value of 60%. Other grain legumes have similar
ranges (Peoples and Craswell, 1992; Unkovich and Pate, 2000; Walley et al.,
2007). For lentils and other grain legumes, the theoretical proportion of N
derived from fixation ranges from 0 if soil N is high and/or if soils contain
no effective rhizobia, to 100% if soil N if very low and soils contain compatible strains of rhizobia. The empirical values reported in Table 15.1 for individual field experiments reflect this range. Researchers have examined and
linked a variety of factors responsible for the broad range of estimates,
including lack of effective strains of native rhizobia, lentil genotype, use of
inoculants, soil type, existing soil N levels, crop phenology and type of cropping system (Table 15.1). The exact role of these factors on the contribution
of nodulation and nitrogen fixation to soil health will be discussed below.
Published estimates of the total amount of N fixed by lentil range from
0 to 192 kg N/ha (Table 15.1), with an average value of about 80 kg N/ha in
shoots and roots. This estimate is similar to quantities fixed by chickpea
and dry bean (Unkovich and Pate, 2000). In contrast, up to 327450 kg N/
ha is fixed by lupin, fababean and soybean. The lentil root system takes up
about 2025% of fixed N in the plant, or about 22 kg N/ha in the roots and
nodules (Unkovich and Pate, 2000). In comparison, chickpea, lupin and
lucerne invest more than 40% of fixed N in their root systems. The amount
of N sequestered in nodulated roots is important because this is a source of
N available for subsequent crops, along with any surface residue incorporated into the soil. Unless used as a green manure (see Table 15.1), most of
the fixed N is removed as grain, leaving only a small percentage in the soil.
Estimates of nitrogen xation in lentil.
Location
N xed (%)
Total N xed
(kg/ha)
N added to soil
(kg/ha)
Pakistan
948
948
Pakistan
Jordan
5685
8083
4685
1633
Syria
Syria
Syria
5868
093
5569
111154
1882
1055
Syria, France
Egypt
6375
5255
88147
127139
Canada
7287
129192
Canada
072
081
Canada
076
0105
Canada
5577
1014
Canada
Canada
USA
Australia
6272
86
61
149
67110
169
59a
070
169a
Australia
6090
60110
aAfter
Variability factors
References
Rhizobia strain; plant genotype;

uninoculated control
Year; incorporation of residue
Plant genotype; plant phenology
Year; added N
Year; location; intensity of
management
Year; location, soil insecticide
Rhizobia strain; uninoculated
control
Year; plant genotype; uninoculated
control
Year; plant genotype; location;
location; uninoculated control
Intercropped with ax versus
monocrop; added N
Conventional tillage versus no-till
End of growing season estimate
Year; location
One crop tested; used as a green
manure
Year
Hafeez et al. (2000)

Shah et al. (2003)
Badarneh and Gwawi (1994)
Kurdali et al. (1997)
McNeill et al. (1996)
Keatinge et al. (1988)
Beck et al. (1991)
Moawad et al. (1998)
Rennie and Dubetz (1986)
Bremer et al. (1988)
Biological Nitrogen Fixation and Soil Health Improvement
Table 15.1.
Bremer et al. (1990)

Cowell et al. (1989)
Matus et al. (1997)
van Kessel (1994)
Smith et al. (1987)
Rochester et al. (1998)
Peoples et al. (2001)
grain removal.
231
232
M.A. Quinn
Ghosh et al. (2007) concluded that the carryover of N from grain legumes
for succeeding crops (e.g. sorghum, pearl millet, maize, castor) in dry lands,
and marginal and sub-marginal lands ranged from 35 to 120 kg N/ha. The
carryover for lentils is probably at the lower end of the range, estimated as
45 kg N/ha by McNeil and Materne (2007) and 23 kg N/ha by van Kessel
and Hartley (2000). In comparison to other grain legumes (i.e. common
bean, chickpea), lentils are likely to have positive N balances (Walley et al.,
2007). As discussed below, the carryover N is just one aspect of soil health
provided by lentils.
15.3. Factors that Affect the Contribution of Lentils to Soil Health

The exact contribution of lentils to soil health depends on numerous abiotic,
biotic and production factors that affect nitrogen fixation and the decomposition of lentil roots and surface debris (Table 15.2). Abiotic factors, such as
soil temperature, soil pH, soil salinity and soil nutrients affect rhizobia, the
plant, and/or the rhizobia-plant symbiosis. Biotic factors that affect the
contribution of lentils to soil health includes both intrinsic and extrinsic factors. Intrinsic biotic factors include lentil genotype, strain of rhizobia and
the plant genotyperhizobia strain interaction. Extrinsic biotic factors are
organisms that affect the plant or rhizobia, such as plant pathogens, weeds,
rhizobial competitors and beneficial soil microbes (e.g. mychorrizas, growthpromoting bacteria). Agronomic factors that affect N inputs into soil include
fertilizer regimes, tillage and planting methods, irrigation, mixed and intercropping, crop rotations, and use of green manures.
Abiotic soil factors

Saline soils constitute approximately 19.5% of irrigated land and 2.1% of
dryland agriculture in the world (FAO, 2000), and are a significant challenge to growing lentils and other grain legumes. Rai and Singh (1999) studied the effects of salt stress on lentils and rhizobia and found that saline
soils inhibited nodulation and activities of nitrogenases, glutamine synthetase
and NADH-dependent glutamate synthase in R. leguminosarum. They were
able to identify two rhizobia strains and four lentil genotypes that were more
tolerant of saline soils, exhibiting greater nodulation, nitrogen fixation, total
plant N and plant biomass compared to indigenous strains. Moawad and
Beck (1991) evaluated 229 isolates of R. leguminosarum from lentil-growing
regions of West Asia-North Africa and found considerable variation in salt
and heat tolerance.
Considerable progress has been made in the selection of legumes and
effective rhizobia for acidic soils (Giller and Cadisch, 1995), which comprise
about 1.6 billion ha in the world (Graham and Vance, 2000). Slattery et al.
(2004) conducted a survey of the effectiveness of rhizobia communities at
50 sites in southern Australia and found that nitrogen-fixing effectiveness
Factors that affect the contribution of the rhizobia-lentil symbiosis to soil health.
Factor
Variables
Affect on the rhizobia-lentil symbiosis
Abiotic factors
Temperature
pH
Salinity
Soil N
Soil phosphorus
Susceptible to extremes; adapted to cool soil temperatures

Susceptible to extremes; prefer slightly alkaline to neutral soils
Susceptible to high salinity
Too much N inhibits nodulation and nitrogen xation; adapted to low N conditions
Limiting factor for nitrogen xation; uptake can be enhanced by mycorrhizas and P-solubilizing
bacteria
Zn, Bo and Mo deciencies affect nodulation and nitrogen xation
Interact with rhizobia strains to affect nodulation and nitrogen xation
High degree of variability in competitiveness, response to abiotic conditions, nodule initiation
and nitrogen xation
Numerous strains compete for nodule initiation sites; inoculant strains must be competitive with
indigenous strains
Pathogens and herbivorous insects can reduce nodulation and nitrogen xation by directly
feeding on the root system; foliar pests indirectly affect nodulation and nitrogen xation by
reducing photosynthate production; weeds compete for P and micronutrients and indirectly
affect nitrogen xation via their effect on foliage growth
Very little known about the effects of bacteria-feeding protozoa and microinvertebrates on
rhizobia populations; involved in nutrient cycling
Can increase P uptake and nitrogen xation
Intrinsic biotic
factors
Extrinsic biotic
factors
Micronutrients
Lentil genotype
Rhizobia strain
Intra-strain competition
Plant pests
Soil microinvertebrates
Crop production
factors
Mychorrhizas; Psolubilizing bacteria

Inoculants
Fertilizers
Tillage
Intercropping
Crop rotation
233
Green manure
Essential in rhizobia-decient soils; must be competitive with indigenous strains and adapted to
abiotic conditions; must be compatible with lentil genotypes
N fertilizers reduce nitrogen xation
Reduces soil organic matter; variable affects on nodulation and nitrogen xation
Little or no direct transfer of N to non-legumes; increased nitrogen xation when non-legume
uses soil N; increases soil pool of N
Viable alternative to fallow periods and continuous cereal production; often have a positive N
effect on soils
Provides abundant N as a green manure; can increase organic matter in comparison to fallow;
prevents soil erosion
Table 15.2.
234
M.A. Quinn
was related to soil pH and location. Only 54% of the sites had sufficient
resident populations of R. leguminosarum bv. viciae for effective nodulation
of lentils. Low populations (<10 rhizobia/g soil) of R. leguminosarum bv.
viciae were found in acidic soils. Evans (2005) also found that strains of R.
leguminosarum vary considerably in their ability to nodulate lentil, fix N and
contribute to plant growth in acidic soils. High alkalinity also reduces nodulation, nodule biomass and leghaemoglobin content of lentil nodules
(Misra et al., 2002).
Existing soil N and P contents affect nodulation, nitrogen fixation and
growth of lentils and other legumes. Phosphorus deficiencies are frequently
cited as limiting nitrogen fixation (Giller and Cadisch, 1995) and approximately 33% of arable land is deficient in P, a common problem in tropical
and subtropical soils (Graham and Vance, 2000). Arpana et al. (2002) looked
at the effect of inoculation, fertilizers and irrigation on uptake of nutrients
by lentils grown in Bihar, India over a 2-year period. They found that inoculated lentils increased the availability of N, P and K, when fertilized and
irrigated, compared with uninoculated controls. Researchers have reported
059% increases in lentil yield in response to added P (McKenzie et al.,
2007). Inoculation of lentils and added P has been shown to increase root
biomass, number of nodules, plant growth and nutrients (Shah et al., 2002;
Balyan and Singh, 2005).
Soil micronutrients also affect the contribution of nodulation and nitrogen fixation to soil health (Wani et al., 1995). Prasad et al. (2004) found that
rhizobia strain and S affected grain yield of lentil, nodules per plant, S content in nodules, total N uptake, and nitrogenase activity. Additions of
micronutrients (Mo, Co, B) was shown to increase lentil nodule biomass,
plant biomass, N content, seed yield, seed size, and N and P content of seed
for lentil, chickpea and lupin (Yanni, 1992).
The concentration of soil N affects nodulation and nitrogen fixation of
all legumes. High levels of soil N reduce nodulation and fixation, thus minimizing the benefit of the symbiotic relationship. This occurs because nitrogen fixation requires considerable energy and nutrients, whereas the uptake
of soil N is metabolically less expensive for the plant. Also, removing N
from soils lessens competition from other species. Considerable research
has been conducted to select genotypes and strains of rhizobia that maximize nitrogen fixation in N-rich soils (Giller and Cadisch, 1995; Wani et al.,
1995; Herridge and Rose, 2000). However, it is unclear if this would actually
improve soil health and benefit cropping systems because, in the end, it
may increase the loss of N from soils through leaching and volatilization
(Giller and Cadisch, 1995). It doesnt seem to be a particularly sustainable
approach and is likely to degrade soil health. The maximum contribution
of lentils and other legumes to soil health occurs in N-poor soils. When soil
N levels are low, the growth of some legume species may benefit from a
small amount of added N until nodulation and nitrogen fixation begin (van
Kessel and Hartley, 2000). The greatest benefit of nitrogen fixation is realized in N-poor soils that have adequate amount of P and micronutrients,
and suitable strains of effective rhizobia.
235
Biotic soil factors

The contribution of the rhizobia-lentil symbiosis to soil health is affected by
intrinsic biotic factors, such as plant genotype, strain of R. leguminosarum
bv. viciae, and interactions between the two. Studies have shown significant
variation among lentil genotypes in nodule number, nodule biomass and
nitrogen fixation (Bhattacharyya and Sengupta, 1984; Hafeez et al., 2000),
root-hair density and micronutrient uptake (Gahoonia et al., 2006), and root
growth and weight (Sarker et al., 2005). Small-seeded genotypes of lentils
have greater nodule biomass than those of large seeds, and vary in total
fixed N and soil N uptake (Kurdali et al., 1997).
Several studies have shown considerable variation between strains of
R. leguminosarum bv. viciae in nodule number and biomass, nitrogen fixation, proportion of N derived from fixation, total plant N content, grain
yields and root growth (Bremer et al., 1990; Hafeez et al., 2000; MartinezRomero, 2003). May and Bohlool (1983) screened 31 strains of R. leguminosarum
for their ability to fix N in lentils. The rhizobia used in their experiment came
from a variety of locations and diverse sources, and included strains isolated
from lentil nodules. The isolates varied widely from ineffective (35% of isolates) to highly effective. Ventorino et al. (2007) assessed genetic diversity of
98 strains of R. leguminosarum bv. viciae nodulating Vicia faba plants, and
found that 53% of the isolates showed a high occurrence and persistence in
the soil.
Indigenous populations of rhizobia are often not adequate to support
nitrogen fixation in lentils. Soils that have not been conditioned to grow
lentils typically are devoid of effective rhizobial strains and must be inoculated. For example, in the Great Plains Region of the USA, most lentil production occurs on soils that were originally free of indigenous R.
leguminosarum (Bremer et al., 1990) and needed to be inoculated to increase
yield through nitrogen fixation. Slattery et al. (2004) conducted a survey of
the effectiveness of rhizobia communities at 50 sites in southern Australia.
They found that only 54% of the sites had sufficient resident populations of
R. leguminosarum bv. viciae for effective nodulation of lentils. Hafeez et al.
(2000) found that a specific strain of R. leguminosarum bv. viciae (i.e. Lc26)
fixed 243% more N than indigenous populations of rhizobia in N- and
rhizobia-poor soils in Pakistan. Although indigenous strains may not be
suitable for forming symbiotic relationships with specific legume species,
they often exhibit higher tolerance to adverse conditions found in their soils,
including tolerance to salt, higher temperatures and desiccation, and may
be excellent sources of genetic variation for improving the effectiveness of
rhizobia strains growing in marginal soils (Zahran, 2001).
Extrinsic biotic factors affecting nodulation and nitrogen fixation of lentils include the numerous other species that affect rhizobia in the soil, the
plant-rhizobia symbiosis, and the plant. Intra-strain competition among
rhizobia for nodule initiation sites is an important extrinsic biotic factor
because strains that are effective at fixing N must also be competitive. May
and Bohlool (1983) inoculated lentil with different strains of rhizobia and
236
M.A. Quinn
found that competitiveness depended on rhizobia strain, lentil genotype

and soil type. Moawad et al. (1998) found that in field-grown lentils, inoculant strains were not able to compete with indigenous rhizobia; the established rhizobia occupied 7688% of the total nodules formed on inoculated
plants. In contrast, Shah et al. (2002) reported that inoculated strains of
R. leguminosarum bv. viciae were highly competitive with indigenous rhizobia, being recovered from 7195% of lentil root nodules. The difference
between these studies reflects the previous crop history of the soil.
Because nodulation, nitrogen fixation and photosynthesis are linked,
factors that reduce photosynthate production and nodulation (e.g. insects
pests, plant pathogens, weeds) can adversely affect symbiosis and nitrogen
fixation in lentil. For example, Weigand et al. (1992) conducted a 3-year field
study on the effects of a nodule-feeding insect (Sitona crinitus), planting date
and fertilizer and insecticide applications on plant damage and yield of lentil in different locations in northern Syria. They found that the insect caused
significant nodule damage (up to 75%) and yield reductions. The overall
effect of the insect was related to precipitation and planting date.
Non-rhizobial microorganisms have been shown to enhance the symbiotic relationship that contributes to soil health. Chanway et al. (1989) examined the effect of plant growth-promoting rhizobacteria on grain legumes
and found that lentil inoculated with the bacteria had greater emergence,
growth, nodulation and root weight. In contrast, pea (Pisum sativum) was
not affected by the rhizobacteria. Zarei et al. (2006) conducted a greenhouse
study to evaluate the effect of arbuscular mycorrhizas (AM) and indigenous rhizobacteria strains on lentils in a calcareous soil with high pH and
low amounts of available P and N. They found that mycorrhizas, rhizobial
strain and addition of P had significant effects on nitrogen fixation in lentil,
and the effect of rhizobia and the AM was synergistic. Lentils inoculated
with phosphate-solubilizing bacteria can increase above- and below-ground
biomass, nodulation, and P-use efficiency (Singh et al., 2005).
Crop production factors
The contribution of nodulation and nitrogen fixation to soil health and sustainability depends on how the legumes and rhizobia are used by land
managers. Agricultural factors, such as the selection of plant genotype and
fertilizer regime have been discussed above. Additional factors that affect
the contribution of lentils to soil health are applications of inoculants, tillage
methods, their use in intercropping systems and crop rotation sequences,
and their use as green manures. These cropping system factors affect N
inputs into soils, as well as soil moisture and organic matter content.
Inoculants
Nodulation and nitrogen fixation are totally dependent on the presence of
effective strains of rhizobia in the soil, and there have been numerous cases
of crop failure because of the absence of the compatible and effective strains.
237
The adoption of rhizobia inoculants has allowed the successful introduction

of many legumes into new farming regions, and has enhanced crop productivity in regions where legumes have been grown previously. However,
widespread adoption of inoculants has not occurred in all regions and crops
(Graham and Vance, 2000). For example, Hall and Clark (1995) found that
only 30% of farmers in an established soybean-growing region in Thailand
were willing to adopt new inoculant technologies, perhaps because most
relied on N fertilizer to grow soybean.
Deaker et al. (2004) reviewed the current status of seed inoculation technology and concluded that there is a clear benefit from using commercial
inoculants, with several studies reporting yield increases of up to 25%. Singleton et al. (1992) conducted a review of 228 field experiments in more than
20 countries and 19 species of legumes. They found that growth responses
to inoculation were highly variable, but the majority of field studies showed
significant yield increases. Under standard field conditions, an average of
46% of studies showed significant positive responses to inoculation, ranging
from 10% with common bean to 70% with mung bean. Forty-eight percent
of 27 field experiments with lentils showed a positive response. Evans
(2005) attributed the failure of the standard inoculant strain, WSM1274, in
Western Australia, to survival problems in acidic soils. Cheminingwa and
Vessey (2006) estimated the abundance and effectiveness of resident populations of R. leguminosarum bv. viciae at 20 sites in Canada and compared
nodulation of uninoculated and inoculated pea. Nodulation was negligible
in pea at two sites with no history of growing grain legumes, and abundant
at sites with a previous history of inoculation. Commercial inoculation
resulted in a 19% increase in above-ground dry weight of pea plants and a
22% increase in number of nodules (but no effect on nodule biomass).
Bremer et al. (1990) compared 14 strains of R. leguminosarum bv. viciae with
two genotypes of lentils at three sites in Canada and found that inoculation
increased grain yields by up to 27166% over uninoculated controls. The
results depended on location, rhizobia strain and lentil genotype. These factors also affected the amount of fixed N. Shah et al. (2002) reported that
inoculation of lentil with R. leguminosarum bv. viciae strain Lc26 increased
yields by 393 kg/ha compared with uninoculated controls. Chandra and
Navneet (2003) tested the effect of inoculation of different lentil genotypes
on nodulation and yield over a 3-year period. They found that inoculation
of lentils increased nodule biomass from 29 to 64%, and mean grain yield
by 15 to 20%, compared with controls.
In a review of the benefits of inoculants, van Kessel and Hartley (2000)
concluded the following:
1. The response to inoculants is greatest when the legume is planted in new
areas where soils lack appropriate rhizobia.
2. The response to inoculants depends on the number of rhizobia already in
the eld.
3. Nitrogen xation via inoculated rhizobia is dependent on the availability
of soil N and the N demands of the crop.
238
M.A. Quinn
4. The response to inoculation is site specic and depends on numerous

other abiotic factors, such as soil pH, salinity, temperature and moisture.
5. There is signicant variation in the effectiveness of different rhizobia
strains and the nodulation and nitrogen xation of different plant genotypes.
Bullard et al. (2005) reviewed problems associated with the use of inoculants in Australian agricultural systems from 1953 to 2003. These problems
included loss of effectiveness and poor competitive ability of inoculant
strains, poor viability of rhizobia because of contaminants in peat carriers,
exposure to high temperatures or low temperatures in storage, ethylene
oxide residues left after sterilization, and high pH of material used to coat
seeds. As few as 100 rhizobia/seed may be sufficient in rhizobia-free, moist
soils. In contrast, greater than 106 rhizobia/seed may be required if soils are
not suitable for rhizobia growth and survival. Problems with quality control of inoculants have hampered their acceptance and use by farmers.
Gomez et al. (1997) evaluated 18 commercial soybean inoculants in Argentina, and found three contained no detectable rhizobia, five with fewer than
107 rhizobia/g of carrier, and 14 with more contaminants than rhizobia.
Currently, there are no international standards for quality and application rates for inoculants (Stephens and Rask, 2000). Many countries (e.g.
Australia, The Netherlands, Rwanda, Thailand, Russia, Canada, France) regulate minimum populations of rhizobia, contaminants per unit weight of
product, viable number of rhizobia per seed, and/or rate of application.
The USA and the UK leave product quality and rates to the discretion of the
manufacturer. As an example of quality standards, Australia has established
the following: (i) >109 rhizobia colony forming units (cfu)/g of moist peat;
(ii) contaminants not detectable at a dilution level of 106; and (iii) minimum
moisture contents of peat packets (Bullard et al., 2005). Standards were
established for peat and for pre-inoculated seeds.
There have been many improvements in inoculant quality, standards
for number of rhizobia per seed, and in storage and application methods. In
spite of quality controls and improved technology, the full potential of inoculation has not been realized because of the numerous site-specific abiotic
and biotic factors that affect the lentil-rhizobia symbiosis, and because the
survival of rhizobia on pre-inoculated seeds is still a significant problem
because of desiccation. Improvements are being made on strain selection
and the use of desiccant protectants and adhesives.
Inoculants are produced in three different forms: powder with a peat
moss carrier; liquid; and granular. Peat-based powdered inoculants are usually applied directly to seeds and are the most common form of delivery.
Sterile peat flour is generally used because it supports higher rhizobia populations and has a greater shelf life. Sticking agents are often used to
enhance coverage of seeds. A significant disadvantage of seed inoculants is
that seeds are limited in the number of rhizobia that coat the seed; smallseeded legumes may only carry a few thousand rhizobia. Another disadvantage is that seeds treated with fungicides or other chemicals may not be
suitable carriers of inoculant. Sedge peat carriers used in inoculants generally
239
support the growth of rhizobia because they have a high water-holding

capacity, a suitable pH, allow uniform dispersion of rhizobia and are nontoxic. Liquid inoculants can be applied to the seed or in the furrow during
planting and can have similar efficacy to peat-based inoculants. However,
liquid formulations may have a lower shelf life and need to be stored under
cool temperatures.
The direct application of inoculant to the soil near the seed (soil inoculation) is generally more effective than seed inoculation, particularly in
acidic, arid, cool or wet soils (Lupwayi and Kennedy, 2007). Granular inoculants, which are growing in usage, are applied at seeding in the furrow or
below the seed. They are most suitable with modern equipment that allows
the simultaneous placement of seed, fertilizer and/or inoculant. Granular
formulations allow much more control of application rates and placement
in the soil. It is currently recommended that granular inoculants be placed
directly below or to the side of the seed, instead of in the seedbed. An
advantage of granular inoculants is that they are more compatible with
other seed treatments because the rhizobia are not in direct contact with the
chemicals. Kyei-Boahen et al. (2002) evaluated the efficacy of different seed
and soil inoculation methods for chickpea at sites in Saskatchewan, Canada.
They found that nodulation after seed inoculation was restricted to the
crown region of the root system, whereas soil inoculation increased nodulation of lateral roots. Further, granular inoculant placed below the seed
increased seed yield by 1736% and 514% over the liquid and peat-based
inoculants, respectively, depending on chickpea cultivar. They concluded
that both peat and granular inoculants were equally effective in establishing successful symbiosis, but that placing granular inoculant 2.58.0 cm
below the seed may improve yield and quality.
The selection of the appropriate rhizobia strains are crucial to the successful use of the inoculant. As discussed above, the selected strains must be
competitive with indigenous strains, and must have high rates of nodulation
and nitrogen fixation. Ideally, the strains must be adapted to the particular
soils, climate and plant genotypes used in a geographic region.
Tillage
Soil health and sustainability are affected significantly by tillage. Conventional tillage reduces soil organic matter, lowers soil water-holding capacity, lowers soil fertility, increases wind and water erosion and increases
problem with nutrient runoff. Loss of organic matter after tillage is particularly severe in the tropics (Graham and Vance, 2000). Soil health can be
increased greatly by adopting conservation tillage methods of planting,
including minimum tillage and no-till. No-till involves the planting of a
crop into the stubble of a previous crop. It increases soil organic matter, soil
fertility and soil water-holding capacity, and decreases nutrient runoff, soil
erosion and fuel costs. Grain legumes are important components of no-till
systems because they provide an inexpensive source of soil N for subsequent
crops, use less soil water than non-legumes, depending on the specific legumes
240
M.A. Quinn
and its root system, and serve to break up the life cycle of crop pests, which
can be a problem in no-till and continuous cereal cropping systems. No-till
and minimum tillage methods lower mineralization and nitrification rates,
and increase immobilization of N (van Kessel and Hartley, 2000). This leads
to a decrease in available N, which stimulates nitrogen fixation of legumes
planted in no-till soil.
van Kessel and Hartley (2000) reviewed field studies involving conservation tillage systems, and found the following:
1. Under conservation tillage the percentage of N from xation in soybean
increased from < 75% under conventional tillage to 85%.
2. Soybean nodulation improved substantially under zero tillage and the
average amount of total N2 xed increased from 180 to 232 kg N/ha.
3. Nitrogen xation increased by 31% in pea and by 10% in lentil after 4
years in zero tillage in a semi-arid environment.
4. No-till had no effect on nitrogen xation of chickpea and soybean under
low rainfall conditions and high soil nitrate levels.
Other studies have shown that the benefit of legumes in no-till systems
occurs because of the extra soil moisture conserved from leaving standing
stubble over the winter, increasing snow trapping and moisture conservation, and the improved microclimate during the growing season (Miller et al.,
2002). Because of the low residue produced by lentils, they do not necessarily prevent erosion when used in no-till systems, at least not in comparison
with soil residues from no-till wheat. Thus, it is important to maximize
conservation of lentil residue when grown in highly erodible soils.
Intercropping
Intercropping and mixed cropping of legumes with other crops are common practices in Latin America, Africa and South Asia, which greatly contributes to soil health improvement. For example, more than half of lentil in
Bangladesh is grown mixed and intercropped with cereals and broadleaf
crops (Sarker et al., 2004). Several studies have shown increased yields, in
terms of land equivalent ratios, of non-legumes intercropped with grain
legumes, such as chickpea, peas and lentils (van Kessel and Hartley, 2000).
Gunes et al. (2007) studied nutrient uptake and yields of intercropped wheat
and chickpea and found that mean N content of wheat seed and chickpea
seed were 11.2 and 10.2% greater, respectively, when intercropped than when
grown alone. Phosphorus content of seed was also significantly greater for
both crops, and the land equivalent ratio for aboveground biomass and
grains was greater than 1.0, indicating significant benefits of intercropping.
Beck et al. (1991) conducted field experiments in France and Syria to examine the effect of intercropping lentils and other grain legumes with cereals
on nitrogen fixation, root biomass and the transfer of N from the annual
legumes to the cereals. They found no evidence of direct transfer of N from
pea to barley when intercropped. The percentage of N in pea derived from
nitrogen fixation was 39% higher when intercropped with barley, than
241
when grown alone. They concluded that on fertile soils, it is beneficial to

intercrop with a legume because of increased N production, greater nitrogen fixation efficiency, and/or greater N mineralization. Non-legume crops
grown with grain legumes reduce N pools in soils, thus increasing nitrogen
fixation of the companion legume crop. Differences in N demands also
reduce competition between legume and cereal intercrops.
Green manures
Lentils and other grain legumes can be used as both green manures and
cover crops. When used as a green manure, lentil is incorporated into the
soil before fixed N is transferred to grain to maximize the amount of N
added to the soil. Cover crops are typically used to prevent soil erosion
and loss of soil organic matter and nutrients. The main benefits of green
manures are the addition of organic matter, C, N and other nutrients to the
soil, and the protection they offer against erosion when used instead of fallow. Nitrogen inputs of legume green manure range from 45224 kg N/ha
and the amount of N made available to subsequent crops ranges from 40 to
60% of the legume grain N, depending on species (Sullivan, 2003). Bremer
and van Kessel (1992) found that about 40% of N in lentil green manure
potentially was available for subsequent wheat crops in Canada, but lentil
straw was not a significant source of N. Rochester et al. (1998) reported 169 kg
N/ha was added to the soil by lentils used as a green manure in Australia
(Table 15.1).
Biederbeck et al. (2005) showed that lentils and other grain legumes
used as green manure had significant effects on soil quality when compared
with traditional fallow-wheat cropping systems. In their study conducted
in Saskatchewan, Canada, lentils and the other legumes were incorporated
into the soil at full bloom and soils were analysed after growing wheat
in the sixth year of the experiment, 15 months after the last green fallow
period. Their results showed that green fallow returned 208% more N to
the soil than fallow-wheat, and increased soil organic matter and total soil
N by 11 and 9%, respectively. Lentil green fallow increased populations of
bacteria, filamentous fungi, yeasts, nitrifiers and denitrifiers by 101406%.
Activities of microbial degradative enzymes were similarly elevated. The
percentage of C and N as microbial biomass were 2.7 and 3.8, respectively,
much greater than the percentage sequestered in microbes in a fallow-wheat
system.
Zentner et al., (1996) conducted a study from 1988 to 1993 to compare
the effect of two rotations, green manurewheatwheat and fallowwheat
wheat, on N and water resources. They found that, in general, wheat yields
after green manure were lower than after fallow because of reduced availability of soil moisture. Grain protein was greater on green manure fallowed
mid-season compared to conventional fallow. Brandt (1996) reported on a
5-year study of summer-fallow alternatives. In his study, wheat after lentil
green manure produced comparable yields to wheat after fallow, wheat
after wheat, and wheat after grain lentil. Wheat after grain lentil was an
242
M.A. Quinn
excellent alternative to fallow because of profits from the crop, but lentil
green manure was also considered superior to fallow.
Crop rotations with lentils
Studies have shown increased yields and protein contents of cereals grown
in rotation with lentil and other grain legumes, compared to continuous
cereals (Wright, 1990; Stevenson and van Kessel, 1996; Gan et al., 2003;
Miller et al., 2003). This rotation effect is likely to be the result of several
causes including the addition of N to soils (Campbell et al., 1992; Badaruddin and Meyer, 1994; van Kessel and Hartley, 2000; Gan et al., 2003), more
soil moisture (Miller et al., 2002, 2003; Gan et al., 2003) and suppression of
diseases (Stevenson and van Kessel, 1996). Typically, the growth response
of cereals in rotation with grain legumes varies considerably among locations,
soil types, years and level of soil N.
Walley et al. (2007) provided an excellent review of the actual benefit of
grain legumes to the pool of soil N for crops grown in rotation with the
legumes. It is often assumed that nitrogen fixation will enhance soil N, but
the amount left in the soil depends on how much is removed in grain at
harvest. Legume seeds are generally high in protein and, thus, remove a
substantial amount of N from the soil. The N increment, Ninc, is the incremental change in soil N due to planting legumes and is highly correlated
with nitrogen fixation, but is also highly variable. If the amount of N fixed
is greater than the amount removed at harvest (i.e. Ninc >1), then there is a
net N benefit for subsequent crops. Walley et al. (2007) assessed a total of
230 published estimates on nitrogen fixation of different grain legumes,
including 79 estimates from lentil studies. As expected the estimates of the
percentage of N derived from fixation varied considerably because of differences in abiotic, biotic and crop production factors. However, residual
soil N seemed particularly important in determining the percentage of N
from fixation because of the general inhibitory affect of soil N on nitrogen
fixation. Their analysis indicated that lentils need to derive 47.8% of N from
fixation to achieve a positive net balance, which they typically exceed. They
concluded that lentil, field pea and faba bean are most likely to have a positive N balance, whereas, common bean and kabuli chickpea are likely to
have a negative balance.
Unless lentil is used as a green manure (see Table 15.1), the actual
amount of N returned to the soil is rather small. For example, Biederbeck
et al. (1996) estimated that lentil had a net N balance of 9 kg/ha in Canadian
prairie soils. Bremer and van Kessel (1992) estimated that about 7% of N in
lentil residue was mineralized in the following growing season, and concluded that lentil residue was not a significant source of N for subsequent
crops. Although lentils and other pulse crops may not contribute very much
N to subsequent crops, it still is greater than N released from cereal residue.
Overall, the contribution of lentils to soil health in rotation systems may be
cumulative, and may involve other non-N benefits such as improvement in
soil moisture and disease suppression.
243
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16
Mechanization
Jrgen Diekmann1 and Yahya Al-Saleh2

1International
Center for Agricultural Research in the Dry Areas (ICARDA),

Aleppo, Syria; 2University of Aleppo, Aleppo, Syria
16.1. Introduction
In comparison with most other field crops mechanization in lentil is an issue
because of the crops short stature and its tendency to lodge. Lentil mechanization is currently organized around an extremely wide range of techniques. Mediterranean areas are partially mechanized with a full range of
labour-intensive techniques still being practised in small-scale farming in
the majority of Middle Eastern countries, including hand broadcasting of
the seeds to establish the crop, often hand weeding and hand harvesting.
The reasons are cheap and available labour and limited availability of modern agronomic production schemes. Also, the crop often lodges and seedbeds have a preponderance of stones and are usually not level. Among the
harvest methods, hand harvesting is often preferred because of the market
value of the straw for feeding small ruminants and the fact that straw losses
are kept to a minimum. However, increased exports from Canada, Australia and the USA, where mechanization is universally applied, impacts world
lentil prices in a downward direction, whereas countries with high production costs because of lack of mechanization stand to lose market share.
At present the most important producers in the Middle East are Syria,
Turkey and Iran. A reduction of lentil production area in other countries of
the region and elsewhere may be related to high production costs where
mechanization was not achieved. India is presently one of the top producers with very little mechanization. At the same time India is planning for
increased use of zero-tillage techniques, particularly in connection with rice
production (Hobbs et al., 1997). This may be an opportunity for a greater
move towards mechanization.
Other parts of the world, particularly Australia and North America,
have low labour availability and high labour costs. In fact, their processing
power is expressed in ha/h/person, rather than man h/ha! While in traditional
248

Mechanization
249
production areas 2030 man days are required to harvest a single hectare,
14 ha/h/person can be harvested in a fully mechanized scheme.
Harvesting can be mechanized as a two-stage operation with cutting
and threshing being done separately, or as a single-stage (combine)
harvesting.
Preconditions for mechanized harvesting are:
1. The crop has to be upright and without pod dehiscence.
2. The minimum height of the lowest pod is 10 cm above the soil surface.
3. A level and smooth seedbed or soil surface rather than a ridged and uneven surface that is common with traditional production systems.
4. The combine must be adapted so that it has: low cutting heights by adjustment of the cutting table; small working widths (preferably 34 m) with
rigid headers so it is well adapted to cope with lateral soil-surface undulations (exible headers can operate with wider working widths); low drum
speeds and a concave (xed grid around part of the drum, through which the
thrashed grain has to pass) with smooth and rounded surfaces for low seed
breakage.
16.2. Field Selection

Selecting fields for lentil production considering mechanization involves a
check for physical obstacles to the passage of cutter bars/combines. The
ideal areas from this perspective are of course flat, level fields without
stones or other obstacles to low cutting heights. Cultivation on slopes is
manageable as long as the surfaces are levelled laterally and longitudinally
in a way to allow uniform cutting heights at about 38 cm. This aim can be
reached when all previous operations, particularly tillage and traffic on the
field (e.g. during previous harvest), have aimed to eliminate undulations
and improve on levelling.
Fields under zero tillage require special attention, as no levelling
through tillage is possible (Anon., 2008). The need for a flat surface calls for
rolling after planting in most cases. Additionally in zero-till areas stubble
from the previous crop may be a problem and so fields should be selected
where stubble has been cut short, chopped and evenly distributed.
16.3. Soil Tillage

Lentils are typically grown in rotation with cereals. In the Middle East cereal-crop residues are either removed for use in animal feeding, incorporated
by tillage or burned. In North America and Australia, straw is often kept at
the surface, or incorporated shallowly to reduce soil erosion in conservation
agriculture systems.
Three basic options exist for seedbed preparation: deep tillage, shallow
tillage or zero tillage.
250
Deep tillage
Deep tillage provides good incorporation of straw and stubble, mostly
through mouldboard ploughing. Deep tillage is best carried out when residues have been chopped to a maximum length of 10 cm, either by a chopper integrated into the combine or operated by a tractor following the
swathes of straw. The best timing for deep tillage is when soils are relatively dry to avoid compaction. In heavy clays mouldboard ploughs should
only be operated under dry conditions, because sticky soil hampers soil
inversion by sticking to the surface of the mouldboards.
Shallow tillage
Shallow tillage is an appropriate choice if only small quantities of residue
are to be incorporated and the soil is not compacted from previous management. Duckfoot tines or disc harrows can be operated more cheaply and
faster than mouldboard or disc ploughs and can leave a more levelled surface. Under conditions of residual moisture being left from the previous
season, two tillage operations may be required for efficient weed control.
Zero tillage
Direct seeding offers three possible advantages:
Reduced erosion risks.

Lower operation costs.
Reduced moisture evaporation losses.
This technique requires suitable planting equipment particularly under

stony conditions. Where residual moisture is available from the previous
season, chemical weed control (e.g. with herbicides such as glyphosate) is
required to control summer weeds.
For lentil harvesting there may be a problem with surface undulation,
as the degree of levelling by rolling is limited, particularly on heavy clays.
Zero-tillage techniques are in wide use in Australia and North America,
but are rarely adopted by farmers in the Middle East.
16.4. Seedbed Preparation

In the Middle East appropriate agricultural equipment for seedbed preparation is rarely available. The most frequently used equipment is a duckfoot
cultivator combined with a levelling bar. This equipment provides some
degree of levelling, but it leaves seedbeds far too loose and inadequately
levelled. Another factor producing uneven seedbeds is the widely used
hard tyres of the diagonal type which are usually of a size requiring about
Mechanization
251
a 1.6 bar inflation rate. It is recommended to use radial tyres with a size
large enough to require no more than 11.2 bar internal pressure.
Recommended equipment for seedbed preparation if used properly
will achieve a medium soil structure and sufficient firmness for good control of the planting depth and soil-to-seed contact, while preventing the formation of deep tyre tracks during planting and weed-control operations.
Combining seedbed preparation and planting in one operation improves
field levelling while reducing the number of equipment passes over the
field. This can be achieved by combining relatively short harrows with a
conventional seeder or through the use of other tillage-plus-planting units,
as often used in North America and Australia.
16.5. Seeding
Most seeders used in the eastern Mediterranean region are simple models
with actively digging tines that are able to complete the planting operation
in seedbeds that are minimally prepared. This is a robust and inexpensive
solution to seedbed preparation and can be improved for mechanized harvesting by rolling the fields after seeding; however, seed rates of two to
two-and-a-half times the normal rate are required to achieve an adequate
plant stand. Good seeding equipment can reduce seed rates by 50 kg/ha or
more and provide a saving of an estimated US$2040/ha. This is an amount
sufficient to pay for recommended solutions to the problem of seeding and
crop establishment. Recommended seeding machines are able to plant at
row spacings of 1020 cm and control the planting depth precisely enough
to place the majority of seeds at 35 or 46 cm depths or other preferred
depths. For further discussion of seeding rates see Ali et al. (Chapter 14, this
volume). A levelling device, either as a simple bar or tine system following
the coulters, is recommended for good seed cover. This is important to
promote high germination percentage rates as well as for safety with preemergence herbicides. A report (Anon., 2007a) from Australia indicates that
lentils are well suited to no-till, reduced tillage and stubble retention systems. If the crop is sown with the appropriate equipment direct seeding
into standing cereal stubble is often possible.
16.6. Rolling
Rolling after seeding generally smooths the soil surface and promotes rapid
germination and uniform emergence by improving seed-and-soil contact. In
addition, rolling smooths and levels the soil surface and presses stones into
the soil, which is a prerequisite for subsequent harvest by cutting plants
with cutter bars at a stubble height of less than 10 cm. Rolling should be
avoided when the seeds are germinating to avoid excessive damage to the
emerging plants. Suitable rollers are for example Cambridge or crosskill
rollers, but also rollers assembled with V-shaped rings are suitable. It is
252
important to leave a profiled surface, as otherwise flat, sealed surfaces

increase the risk of erosion with heavy rainfall.
Rolling is only recommended where surfaces are dry. Rollers should
have a weight of 400700 kg/m working width. Working widths of rollers
can range from just 3 m to 12 m, or more. Rolling after emergence is recommended until the 57 node stage is reached (Anon., 2007b). Also, a minimum
time span of 2 days between rolling and herbicide application is recommended. Rolling should not follow immediately after frost (Anon., 2000).
16.7. Weed Management

Lentil production in the Middle East includes manual weed control,
although mechanized inter-row cultivation and herbicides are available.
Weed management is also discussed by Yenish et al. (Chapter 20 this volume). Hand weeding does not have an impact on the choice of a harvesting
system.
Mechanized weed control, like inter-row cultivation, use of a weed
brush or a weeding harrow may have a negative effect on mechanized harvest, because surfaces may become undulating, and/or stones may be
brought up onto the surface and increase the necessary stubble length at
harvest time, which increases pod losses.
In most cases plants, at the time of inter-row cultivation, are too tall for
a subsequent rolling operation. From experience at the International Center
for Agricultural Research in the Dry Areas (ICARDA) it can be said that
only hand weeding or inter-row cultivation are economic and/or efficient.
Inter-row cultivation is the most efficient mechanized method of weed
control; weeding harrows that work across the surface have not worked
satisfactorily (Fig. 16.1). Weed brushes are not designed for cheap, economic
operations, but for (horticultural) high-value crops. While they do a good
and efficient job of weed control, they are too expensive and have insufficient capacity for large-scale agriculture.
Inter-row cultivators depend on the crop being planted by a seeder,
because the operation requires parallel rows with equal distances. The
working width of the seeder must match the working width of the inter-row
cultivator. Row spacings from 30 cm and wider are required (Diekmann et al.,
1992). For lentils, 3040 cm row spacings are ideal where inter-row cultivation is planned.
Soil problems can occur on heavy clay soils especially when the soil is
wet and sticky, while a preponderance of stones can obstruct the blades of
the inter-row cultivation equipment.
Working depths should be as shallow as possible and just deep enough
to cut off the weeds below the soil surface.
Where harvest is planned to be mechanized, either by cutter bars,
swathers or combines, inter-row cultivation will usually require longer
stubble heights and is likely to increase pod losses. Reasons for slow adoption of herbicides for weed control can be the costs and the risk of reduced
Mechanization
253
(a)
(b)
(c)
(d)
Fig. 16.1. Mechanical weed control: (a) hand-pushed weeding hoes; (b) inter-row cultivator
in double-row planting (most economic tool); (c) weeding brush for high-value crops; and
(d) weeding harrow (high capacity but low efficiency).
yields under dry and/or cold conditions. Lack of experience in the use of
herbicides may also be a factor preventing their more widespread use.
Complete mechanization is easiest to achieve when weed control is
entirely by chemical means. This is the standard approach in Australia and
North America (Anon., 2007a, b).
Parasitic weeds (Orobanche sp., Cuscuta sp.) are problematic for traditional and mechanized production. Thus far the only means of control is
through the use of systemic herbicides that reduce parasitism of host plants.
Manual removal of parasitic weeds is not proven to be efficient because of
the late visibility of the weeds and at the time of visibility of the parasite,
most of the damage to growth and yield has taken place.
16.8. Harvest
The harvesting method remains the most diverse part of lentil production,
with hand harvesting still being practised in the Middle East and Asian
countries (Fig. 16.2). Lentil harvest in North America and Australia is fully
254
(b)
(a)
(d)
(c)
(e)
Fig. 16.2. (a) Lentil harvest in Syria at present is often organized as hand pulling; (b) drying in
heaps for 510 days; (c) followed either by traditional threshing; (d) or threshing with a
combine; (e) which also chops and bags the straw.
Mechanization
255
mechanized, with either two-stage (swathing and threshing) or single-stage

(combining) methods.
Hand harvesting
Plants are pulled at the half-green/half-brown stage and left in small heaps
in the field or transported to the threshing area and left for drying before
being threshed. Threshing may be done by hand, by animal-drawn threshers or by stationary threshers.
It has been estimated that 20 man days are required to harvest a hectare
of lentils yielding 1200 kg of grain and 1800 kg of straw (Khayrallah, 1981).
Labour costs are a limiting factor for economical production using these traditional methods (Papazian, 1983).
Two-stage harvest
The two-stage harvest in the Middle East typically starts with use of a cutter bar, followed by manual collection of the dried material and transport to
the threshing place at the farm or village. During the 1980s ICARDA tested
several methods to mechanize the cutting as a first step (Erskine et al., 1991).
This included using existing cutter bars as well as other concepts.
Lentil pulling
As straw sales are part of the income from a lentil crop, several attempts
were made to simulate hand pulling by machine, without increasing the
loss of straw. The first development was made at Reading University in
1960, followed by Snobar (1979) in Jordan and Tauscher in Turkey and
ICARDA. The Tauscher system is functional and achieves lentil pulling at
the same stage as hand harvesting with acceptable losses (<10% for grain
and straw), but the 2 m wide machine was not cost effective. The capacity
of the trialed machine was about 0.5 ha/h and would leave the pulled crops
in a swathe (Friedrich, 1988).
Soybean blades
Another idea that was tested was to use soybean cutting blades to simulate
hand pulling, in order not to loose straw. The blades were designed to cut
the lentil crop at a shallow depth of 25 cm below the soil surface. The problem is that just too often moisture is available in that layer from recent rains.
This produces clogging of the blades, because of the sticky soil. The idea is
therefore impractical.
Cutter bars
Cutter bars are acceptable where straw losses of about 20% can be tolerated.
Preconditions for the use of cutter bars are reasonably levelled fields and no
256
(a)
(b)
(d)
(c)
Fig. 16.3. Knife arrangements for lentil harvest: (a) single knife and rigid fingers; (b) double
knife; (c) double knife with bottom knife fixed; and (d) flexible cutter bar.
large stones on the surface. The optimum height of cutting is about 5 cm

(Friedrich, 1988). For conventional cutter bars with rigid fingers, stones cause
the biggest problem. For this reason a double-knife system, without rigid fingers, is the most suitable for cutting close to the soil surface (Fig. 16.3).
Cutter bars of about 2 m working width are suitable for smaller areas and
can be a first step towards mechanization under Middle Eastern conditions.
The recommended cutter bar is designed to operate even if small stones
or other obstacles are at the cutting level. This is usually a problem for the
standard cutter bars, where damage to the knife blades is frequent with
very low cutting heights. The problem occurs when hard objects get stuck
between the rigid fingers, obstructing the passage of the knife.
The double-knife cutter bar has no rigid fingers, but two oscillating knife
assemblies (Fig. 16.4). Unfortunately this equipment is no longer widely
available, because the market for standard mowers has moved to disc-anddrum mowers, which are not suitable for lentil harvest. The discs and drums
rotate with high speed and cause pod losses through strong vibrations, even
at the half-green/half-brown maturity stage, which is the correct time for
this harvesting method. Double-knife cutter bars are still made and used in
Turkey.
The advantage of the use of cutter bars is that cutting is at about the
same maturity stage as for hand harvesting, which reduces the risk of pod
loss and pod dehiscence.
Mechanization
257
(a)
(b)
(c)
(d)
Fig. 16.4. Harvest machinery being tested: (a) self-mobile cutter bar; (b) double-knife cutter
bar; (c) lentil puller; and (d) soybean blades.
Swathers
In fully mechanized schemes, like in North America and Australia, higher
capacity equipment, like self-mobile swathers with 312 m working widths,
are used. These machines are also available with double-knife cutting systems. Machines of this type are used where separating cutting from threshing has an advantage. Reasons to carry out harvest in a two-stage manner
can be that the climate does not stop the crop growth. Lentils have an indeterminate growth habit and will not come to maturity without heat or
drought stress (Anon., 2007b). Swathing is recommended when one-third
of the bottom pods turn yellow to brown and seeds rattle when pods are
shaken. Also the machine capacity at cutting is higher than with singlestage techniques. The forward speed for swathers may be as high as about
15 km/h under ideal conditions.
The swathe formed by the machine is left to dry, which may take from
a few days to up to 2 weeks (Muehlbauer et al., 1998). A further advantage
of this technique is that all the vegetation, including weeds and volunteers,
is cut at an earlier stage, which may keep fields cleaner by avoiding renewed
seed setting. Threshing is then done with combine equipment with a pick-up
header, which is working at a width of 23 m.
A disadvantage of this technique is the need for two separate field
operations.
258
Single-stage harvest
Full mechanization can be achieved with standard combines.
Under Middle East conditions the limitation in width is required
because of the simple, rigid design of the older types of combines used, as
well as the relatively small field sizes that favour 3 m (maximum of 4 m)
wide tables which can follow contours easily. Similar to cutter bars and
swathers, combine headers may also be equipped with knife systems
adapted to very low cutting heights, such as replacing the rigid-finger plus
single-knife system with a double-knife system in which both knives are
active, or with a double-knife system in which one is fixed and the other
oscillates. The targeted cutting height is below 10 cm so that all pods can be
undercut. While the double-knife system with two oscillating knives is considered the best and most reliable, the demand was too low to permit manufacturers to offer it on these standard, narrow tables. Instead, various
combine manufacturers offer replacement of the fingers by additional triangular, fixed blades under the oscillating knife. The oscillating knife needs to
be held close to the fixed blades by additional plates every 2030 cm. This
system is much safer to operate and is almost undisturbed by stones. The
problem comes with the gaps between the fixed plates and the oscillating
blades. Any moisture in the plant stems or in the soil will often create a
layer of sticky coating. This must be removed to avoid gaps between the
knife and plate, making a clean cut impossible. A simple way to remove the
coating is by brushing at regular intervals with a broom dipped in water
with dish-washing liquid.
The simple replacement of the rigid fingers by triangular plates may
not be sufficient to achieve low losses. Many standard knives of combine
cutting tables are working with an inclination towards the soil surface. This
is a good working principle with stubble lengths over 10 cm, but where
short stubbles are required the knife should work horizontally. This can be
achieved by adjusting the angle between the steel holding the knife assembly and the front edge of the table.
The reel requires reel tine covers, which are metal boards with rubber
flaps, operating like paddles. Their function is to ensure the proper flow of
harvested material, particularly pods of lentils. The reel speed should match
the ground speed, with a maximum of 10% higher speed for the reel.
In North America and Australia the large areas to be harvested have
led to the development of headers which are mounted with flexible cutterbar assemblies. For very large working widths the left and right halves of
the headers can also be tilted. Both options together offer working widths
up to 12 m. Another factor that enables the use of such working widths is
that lentils are grown mostly on stone-free soils in these regions.
Beside losses at the header a second critical point with lentil threshing
is the risk of breaking seeds during the drum-concave passage. The drum
has to be operated at a relatively low speed, with a circumferential speed
below 13 m/s. This specification is required because the drum diameters of
various combine models and manufacturers range from 45 cm to 66 cm. A
Mechanization
259
drum with a 45 cm diameter performs well for lentils at a speed of 300 rpm,
about 7 m/s, provided the crop is dry. The concave must be as smooth as
possible. First of all it must be assured that the de-awning bars, used for
barley and sometimes durum, are removed. The grid of the concave must
not have any sharp edges. A wire concave is preferable to avoid breakage.
It performs better than a perforated steel sheet, because it has a higher percentage of clearance. Concaves with about 8 mm (and up to 12 mm) open
space can be used. Clearance of the concave at the inlet side is recommended
to be up to 17 mm, and at the outlet side up to 15 mm. This can be reduced
in case of unsatisfactory results. If reducing the concave clearance does not
work, the drum speed may be increased (Gerdes, 1987).
Timing of threshing is critical as the best performance is possible at
grain moisture contents of 1622%. This ensures low pod breakage, but
requires drying or aeration of the crop after threshing. If the crop is already
dryer than 16% moisture, it is best to work in the early morning with a low
drum speed and a wider concave opening. Below 12% moisture the risk of
cracking increases substantially (Anon., 2007b).
This requirement shortens the time window for any field down to a few
days, particularly under conditions in the Middle East. The sieve settings
are similar to cereal threshing. The top sieve (frogmouth) can be opened up
to 1.5 times of the lentil diameter and the sieve extension slightly wider.
The bottom sieve can be an adjustable frogmouth sieve with about 10 mm
clearance or a round-hole sieve with even larger holes. The use of flat sieves
reduces the amount of broken grains (Gerdes, 1987).
As lentils are relatively heavy, wind (draught) intensity has to be
relatively high. Cleaning needs to be adjusted so that returns are as low as
possible to avoid the risk of broken grains.
In the Middle East farmers value the straw almost as much as they do
that of the grain. It is therefore essential not only to maintain a low cutting
height, keeping short stubble, but also to enable collection of the non-grain
material. Farmers have therefore converted standard combines into units
that collect, chop and bag the straw. This is of interest as long as the market
pays high prices for the chopped and bagged product.
In other areas, such as in the USA, Canada and Australia, straw is not
harvested and collected but is incorporated into the soil. Also, the size of
combines in these areas is larger than in the Middle East and harvesting
may be done with cutting tables up to 10 m wide. For thinner, shorter or
lodged crops harvest direction may have to be one way (Anon., 2007a).
16.9. Cleaning and Storage

In West Asia and North Africa, after hand harvesting and two-stage harvest, it is now more common for threshing to be carried out by combines
being hand fed, rather than with the old village-type (stationary) threshers.
Basically the same settings apply as described under the heading
Single-stage harvest, from concave clearance to sieve opening and wind.
260
According to Muehlbauer et al. (1985) typical grading for marketable

products is specified as:
Red lentils: <3 mm diameter, with 50% being larger than 4.35 mm.
Yellow lentils: <5 mm, with 50% being larger than 7 mm.
Lentil seeds are more prone to mechanical damage than other pulses.
Handling and especially the use of augers should be kept to a minimum
(Anon., 2007a).
16.10. Losses
Al-Saleh (2000) compared losses from various harvesting systems. Hand harvesting had the lowest losses of about 8% of straw and 11% of grain. Systems
based on double-knife cutter bars created 15% straw losses and 16% grain
losses. Swathers with augers produced 22% straw losses and 23% grain losses.
The combine adapted at ICARDA with a 3 m header and a double-knife
cutter bar achieved 30% straw losses and 16% grain losses. These values are
not absolute, but show the range of losses and indicate that hand harvesting
and single-stage combining produce the least amount of grain losses.
16.11. Costs
In industrialized countries only fully mechanized techniques are feasible
because of relatively high labour costs. Also the value of straw is not a major
component of income to the producer. Therefore the only choices are to opt
for a two-stage harvest, if losses or forced ripening (drying) are an issue, or
otherwise choose the most economic way of direct combine harvesting. In
countries with lower labour costs all alternatives from hand pulling to full
mechanization are possible.
The costs of hand pulling are of interest for example in Syria or Turkey.
Hand pulling required 42% of the total production costs according to a survey conducted in Turkey (zcan, 1986). Logically, Turkey shows a considerable degree of mechanization mainly through the use of cutter bars. A
more recent survey in Syria and Turkey (Al-Saleh, 2000) revealed that 44%
of total production costs were required for hand pulling and 65% of the
total costs for pulling and threshing. A double-knife cutter bar reduced the
portion for harvesting costs to 33% and harvest-plus-threshing costs to 52%.
Single-stage (combine) harvesting reduced harvest-plus-threshing costs to
20% of the total.
References
Al-Saleh, Y. (2000) Suriye ve Trkiye de mercimek ve nohut hasadnda mekanizasyon
olanaklarnn belirmesi zerine bir arastrma. PhD thesis, ukurova Universitesi,
Adana, Trkiye.
Mechanization
261
Anon. (2000) Lentil production. In: Pulse Production Manual. Saskatchewan Pulse
Growers, Saskatoon, Saskatchewan, pp. 132. Available at: www.saskpulse.com/
media/pdfs/ppm-lentil.pdf (accessed 29 November 2007).
Anon. (2007a) Lentils in South Australia and Victoria. Pulse Australia, Bungwahl, New
South Wales, 20 pp. Available at: www.pulseaus.com.au/crops/lentils/476/lentils
%20fs%20/2007%20final.pdf (accessed 29 November 2007).
Anon. (2007b) Red Lentil Production. Government of Saskatchewan. Available at:
www.agriculture.gov.sk.ca/Default.aspx?DN=a88f57f0-242b-40f6-8755-1fc6df4
dfa14 (accessed 30 October 2007).
Anon. (2008) Lentil: Crop Establishment and Production. West Australian Department
of Agriculture. Available at: www.agric.wa.gov.au/content/fcp/lp/lent/cp/lentesaintr.
htm (accessed 22 May 2008).
Diekmann, J., Bansal, R.K. and Monroe, G.E. (1992) Developing and delivering
mechanization for cool season food legumes. In: Expanding the Production and
Use of Cool Season Food Legumes. Kluwer Academic Publishers, Dordrecht, The
Erskine, W., Diekmann, J., Jegatheeswaran, P., Salkini, A., Saxena, M.C., Ghanaim, A.
and El Ashkar, F. (1991) Evaluation of lentil harvest systems for different sowing
methods and cultivars in Syria. Journal of Agricultural Science, Cambridge 117,
333338.
Friedrich, T. (1988) Untersuchungen zur Mechanisierung der Linsenernte nach dem
Rupfprinzip im Vergleich zu anderen Linsenernteverfahren in Syrien. Dissertation
MEG-141, Universitt Gttingen, Gttingen, Germany.
Gerdes, J. (1987) Combine Instruction. Claas Company, Harsewinkel, Germany,
pp. 7578.
Hobbs, P.R., Giri and Grace, P. (1997) Reduced and zero tillage options for the
establishment of wheat after rice in South Asia. Consortium Paper Series 2. RiceWheat Consortium for the Indo-Gangetic Plaines, New Delhi, India.
Khayrallah, W.E. (1981) The mechanization of lentil harvesting. In: Webb, C. and
Hawtin, G. (eds) Lentils. International Center for Agricultural Research in the Dry
Muehlbauer, F.J., Cubero, J.I. and Summerfield, R.J. (1985) Lentil (Lens culinaris
Medic.). In: Summerfield, R.J. and Roberts, E.H. (eds) Grain Legume Crops.
Collins, London, UK, pp. 266311.
Welsh-Maddux, M.N. and Short, R.W. (1998) Principles and Practice of Lentil
Production. US Department of Agriculture, Agricultural Research Service. ARS
141. p. 26. Available at: www.ars.usda.gov/is/np/lentils/lentilsintro.htm?pf=1 (accessed
30 November 2007).
zcan, M.T. (1986) Mercimek Hasat ve Harman Yntemlerinin Is verimi, Kalitesi, Enerji
Tketimi ve Maliyet Ynnden Karslastrlmas ve Uygun Bir Hasat Makinesi
Gelistirilmesi zerinde Arastrmalar. Trkiye Zirai Donatm Kurumu Yaynlar,
Yayn No 46. Ankara, Trkiye.
Papazian, J. (1983) Lentil harvesting. Lens Newsletter 10(2), 16.
Snobar, B. (1979) A mechanical technique to harvest lentils. Research Journal of
University of Jordan VI(2), 714.
17
Diseases and their Management
Weidong Chen,1 Ashwani K. Basandrai,2 Daisy Basandrai,2 Sabine

Banniza,3 Bassam Bayaa,4 Lone Buchwaldt,5 Jenny Davidson,6
Richard Larsen,7 Diego Rubiales8 and Paul W.J. Taylor9
1USDA-ARS,
Washington State University, Pullman, Washington, USA; 2CSK Himachal

Pradesh Agricultural University, Dhaulakuan, India; 3University of Saskatchewan,
Saskatoon, Canada; 4International Center for Agricultural Research in the Dry Areas
(ICARDA), Aleppo-Syria; 5Agriculture and Agri-Food Canada, Saskatoon, Canada;
6South Australian Research and Development Institute, Adelaide, Australia;
7United States Department of Agriculture (USDA) Agriculture Research Service (ARS),
Prosser, Washington, USA; 8Institute for Sustainable Agriculture, CSIC, Crdoba,
Spain; 9The University of Melbourne, Victoria, Australia
17.1. Introduction
Lentil is an important crop in many parts of the world, cultivated under
various production systems. Lentil crops are affected by a number of diseases caused by bacteria, fungi, viruses and nematodes. Some diseases are
common in most lentil-growing regions worldwide, whereas others are limited to certain production areas. The economic importance of a particular
disease is not necessarily related to its geographic distribution. A disease
may have limited occurrence, but still it may cause significant economic
losses. Under conducive conditions, these diseases, singly or collectively
can cause significant reductions in both grain yield and quality. Hence,
proper management of these diseases is necessary to ensure sustainable
productivity and profitability of lentil.
This chapter reviews the important fungal and nematode diseases of
lentil and their management. Virus diseases and parasitic weeds of lentil
are subjects of Chapters 19 and 21, respectively, of this book. The important
foliar diseases include Ascochyta blight, rust, Botrytis grey mould, anthracnose, Stemphylium blight and powdery mildew, and important soil-borne
diseases are Fusarium wilt, Sclerotinia stem rot and nematode diseases.
Attempts have been made to include the most recent information on these
diseases, however, the readers are also referred to previous reviews by
Khare (1981), Agrawal and Prasad (1997), Morrall (1997), Bayaa and Erskine
(1998), Davidson et al. (2004) and Taylor et al. (2007).
262

263
17.2. Ascochyta Blight

Ascochyta blight is prevalent throughout the world where lentils are grown
and can cause yield losses of up to 70% (Gossen and Morrall, 1983). The
pathogen infects leaves, stems, petioles, pods and seeds (Taylor et al., 2007).
On leaves, young lesions consist of small necrotic spots that enlarge to 35
mm in diameter and change colour to tan or dark brown. Reddish-brown to
black pycnidia form in the middle of mature lesions. On stems, lesions are
brown and variable in size. In severe infections, the lesions often girdle the
stem or petiole, killing the epidermal and cortical cells, leading to stem
breakage. Pod infection may result in seed infection. Heavily infected seeds
are shrivelled and discoloured with a whitish mycelium and pycnidia.
There may be difficulties in distinguishing Ascochyta blight from
anthracnose based on leaf and stem symptoms alone when both diseases
occur together. Nevertheless, pycnidia in the lesions formed by Ascochyta
blight may be distinguished with a handheld lens from acervuli and/or
microsclerotia formed by anthracnose.
Causal organism
Ascochyta blight of lentil is caused by Ascochyta lentis Bond. & Vassil., which
produces conidia in immersed, depressed globose pycnidia (175300 m
in diameter). Conidia measure 1317 m in length and 46 m in width.
Ascochyta lentis is heterothallic, and requires two mating types to produce
its teleomorph state Didymella lentis W.J. Kaiser, B.C. Wang and J.D. Rogers.
Ascochyta lentis is morphologically indistinguishable from Ascochyta fabae.
However, host specificity and compatibility in mating separate the two species
(Kaiser et al., 1997).
Disease cycle and epidemiology
Ascochyta lentis relies on rain splash to spread conidia from infected debris
on the soil surface to newly planted lentil and from plant to plant. Long
distance dispersal occurs through seed-borne inoculum. Wetness periods of
2448 h and temperatures of 1015C are optimum for infection (Pedersen
and Morrall, 1994). Initial infection of leaves can take place within 1012 h
from germinated conidia. Macroscopic symptoms do not become evident
until 79 days after infection. Pseudothecia of the teleomorph stage D. lentis
may also develop on lentil debris and thus liberate ascospores under suitable environmental conditions at the beginning of the crop-growing season.
Management
Integrated disease management is partly effective in controlling Ascochyta
blight of lentil in developing countries where application of fungicides is not
264
W. Chen et al.
practical and economical. The impact of the disease can be reduced through
planting resistant varieties, crop rotation, early sowing by avoiding periods
of high leaf wetness and optimum temperature, use of disease-free seed and
burning of infected debris and stubble after harvest (Taylor et al., 2007).
Seed-borne inoculum can be reduced by sun drying, or hot water and
dry heat treatment of seeds, however, care must be taken that germination
is not adversely affected (Ahmed and Beniwal, 1991). Fungicides reported
to provide the best protection from seed-borne infection include benomyl,
carbendazim, carbathin, iprodion and thiabendazole (Bretag, 1989). The fungicides captafol, chlorothalonil, folpet and metiram, provide good protection against foliar infection.
In breeding for resistance to Ascochyta blight, knowledge of pathogenic
diversity is important when choosing appropriate isolates to screen for
resistance (Taylor and Ford, 2007). Since resistance to A. lentis has been
found to be controlled by specific resistance genes (Ford et al., 1999;
Nguyen et al., 2001), there is the likelihood that new pathotypes of A. lentis
may evolve, thus deployment of resistant varieties needs to be monitored
for potential resistance breakdown.
17.3. Rust
Rust is a widespread foliar disease of lentil and is an economically important disease in Algeria, Bangladesh, Ethiopia, India, Italy, Morocco, Pakistan and Nepal. Early infection accompanied by conducive environmental
conditions can result in complete crop failure, while yield losses in research
plots have been reported to vary from 30 to 60% depending on the cultivar
and disease severity (Sepulveda, 1985; Singh et al., 1986).
Lentil rust is characterized by lesions on the stems and leaves that result
in leaf drop and premature plant death. The pathogen can be seen as
yellowish-white pycnidia and aecial cups on leaflets and on pods, either
singly or in small circular groups (Plate 2A and 2B). Brown and oval to circular uredinial pustules, up to 1 mm in diameter, develop on either surface
of leaflets, branches, stems and pods. The dark brown to black and elongated telia are formed later in the season mainly on branches and stems.
Infected plants have a dark brown to blackish appearance, visible in affected
patches of the field or in the whole field. In severe infections plants dry up,
forming only small, shrivelled seeds.
Causal organism
Rust is caused by the autoecious fungus Uromyces viciae-fabae (Pers.) Schroet.
Spermagonia are subepidermal and globoid. Aecia and uredinia are initially
subepidermal, erumpent later. Aeciospores are spheroidal, 1826 m in
diameter, and yellowish brown in colour. Urediniospores are borne in
single pediceles, mostly echinulate, with three to four germination pores
265
and measure 2228 1922 m. Telia are subepidermal in origin, then

erumpent on leaves, but remain covered by the epidermis on stems for
an extended period. Teliospores are borne singly on pedicels, are globose
to subglobose, one-celled, measuring 2540 1826 m, with a single
germination spore.
Uromyces viciae-fabae was regarded as the causal organism of rust in faba
bean, vetch and lentil. However, some degree of host specialization is recognized, suggesting that U. viciae-fabae may be a species complex with different host ranges. Based on differences in host specialization, morphology
of spores, and substomal vesicle, a new classification into host-specialized
isolates (ex. Vicia faba L., Vicia sativa L. and Lens culinaris L.) has been suggested (Emeran et al., 2005). These groups were also clearly separated by
molecular markers (Emeran et al., 2008).

The disease first occurs during the flowering and early podding stages as
aecia which may further develop into secondary aecia or uredinia. The
resulting aeciospores and urediniospores are the source of further disease
spread during the cropping season. Uredinia usually appear a little later in
the season followed by the development of telia. The fungus survives
between crop seasons as teliospores and is also carried with seed as a contaminant in the form of small pieces of infected plant debris. High humidity
and cloudy weather with temperatures of 2022C favour disease development (Agarwal et al., 1993).
Management
Integrated management of rust includes control of volunteer plants and
removal of infected lentil debris. It is advisable to use clean seeds without
rust contamination, and to treat the seed with a suitable fungicide such as
diclobutrazole. Rust can also be effectively managed by a number of foliar
fungicides, although this needs to be adjusted in order to be economical.
Preventive fungicide sprays of mancozeb at the early disease development
stage have been recommended.
The use of host plant resistance is the best means of rust management.
Several rust-resistant cultivars have been released in different countries,
with resistance originally identified at the International Center for Agricultural Research in the Dry Areas (ICARDA), Syria and in India (Basandrai et al., 2000; Tikoo et al., 2005). There is no clear evidence for the existence
of races. The international testing of lentil genotypes differing in resistance
to rust has shown that the reaction to rust of individual lines is the same.
Although the rust resistance in lentils seems to be durable, it is likely to
break down eventually. Little is known on the inheritance of resistance to
rust. Both complete and partial resistance exist and monogenic resistance
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has been reported (Sinha and Yadav, 1989). Field resistance of lentil to rust
is governed either by a single dominant gene, by two independently inherited dominant genes, by two complementary genes or by three independently inherited genes (Basandrai et al., 2005).
17.4. Botrytis Grey Mould

Botrytis grey mould of lentil has been reported from Australia, Bangladesh, Canada, Colombia, Nepal, New Zealand and Pakistan. It is not
uncommon for yield losses to exceed 50% and in extreme cases total crop
loss occurs (Bayaa and Erskine, 1998; Davidson et al., 2004). Disease symptoms first develop on flowers and pods, and as dark green spots on the
lower foliage that later become pale tan coloured. Senescent or injured
plant tissues are also most prone to infection (Davidson et al., 2004).
Masses of conidia are formed within the canopy and are released into the
air when the canopy is disturbed. Infection of flowers and pods affects
seed formation. Infected seeds may be discoloured and shrivelled, but do
not always show clear symptoms. Infected seedlings from contaminated
seeds are yellow and stunted, with grey fungal growth on the stem at the
soil line and usually die after 1 or 2 weeks (Bayaa and Erksine, 1998;
Davidson et al., 2004).
Causal organisms
Botrytis grey mould is caused by Botrytis cinerea Pers. Ex Fr. and Botrytis
fabae Sard (Bayaa and Erskine, 1998; Davidson and Krysinska-Kaczmarek,
2007). Botrytis cinerea is ubiquitous and non-specific with the ability to infect
over 200 plant species including lucerne, beans, chickpea, field pea, safflower and sunflower (Ellis and Waller, 1974a), while B. fabae has a more
restricted host range occurring mostly on Leguminosae, particularly faba
bean (V. faba), common vetch (V. sativa) and lentil (Ellis and Waller, 1974b).
In culture, B. cinerea and B. fabae have similar white, cottony mycelial
growth that turns grey with age. Conidiophores are brown, erect and
septate, 1630 m thick, frequently 2 mm long and branched near the
cluster. Conidia are produced in clusters at the ends of branches and are
hyaline, single-celled, ovoid or spherical, with a thin wall. Conidia of B.
cinerea from lentil are generally smaller than those of B. fabae, but vary
depending on environmental conditions (Ellis and Waller, 1974a, b).
Sclerotia of B. cinerea are generally larger (24 13 mm) than those of
B. fabae (11.7 mm diameter), but the size and shape is highly variable
(Ellis and Waller, 1974b; Bayaa and Erskine, 1998). The teleomorphs of
B. cinerea, Botryotinia fuckeliana (de Bary) Whetzel and of B. fabae, Botryotinia fabae J.Y. Lu & T.H. Wu, have not been reported on lentil (Bayaa and
Erskine, 1998).
267

Botrytis spp. produce masses of wind-borne conidia on infected and
dead tissues under humid conditions at 1525C (Plate 2C), and epidemics can develop very quickly in comparison with other diseases
(Davidson et al., 2004; Davidson and Krysinksa-Kaczmarek, 2007). The
conidia are dry and dispersed by air currents in large numbers and may
also be carried by rain droplets (Ellis and Waller, 1974a). Botrytis spp.
can infect lentil at any growth stage, but severe epidemics generally
develop in humid conditions after the canopy has closed at flowering
and podding stages (Bayaa and Erskine, 1998; Davidson et al., 2004;
Davidson and Krysinksa-Kaczmarek, 2007). Sclerotia, which are highly
resistant to adverse conditions, are probably the main survival structure.
They may survive for long periods if they are not buried in the soil
where they decay more quickly. Mycelium can also survive saprophytically in plant debris for extended periods (Ellis and Waller, 1974a, b;
Bayaa and Erskine, 1998).
Management
Since Botrytis grey mould is more severe and frequent under humid conditions, avoiding dense canopies is an important component of integrated
disease management in lentil (Bayaa and Erksine, 1998). Delayed sowing
and reduced seeding rates as well as wide row spacings result in more open
canopies during crop maturation, and thus reduce the likelihood of epidemics. Weed control and optimum fertilizer use, particularly avoiding high
nitrogen levels, are also important strategies to prevent dense canopies and
reduce disease severity. It is important to plant seed with less than 5% grey
mould infection. Seed treatments with fungicides such as benomyl, carboxin, chlorothalonil, thiabendazole or thiram can reduce seed-borne inoculum and seedling blight (Bayaa and Erskine, 1998; Davidson et al., 2004).
Destruction of infected debris through grazing, burning or burying residues
reduces the carry-over of inoculum into the following season. A 3-year crop
rotation allows time for the infected residue to break down (Bayaa and
Erskine, 1998; Davidson et al., 2004).
The foliar fungicides benomyl, boscalid, carbendazim, chlorothalonil,
mancozeb, thiabendazole, tridemorph and vinclozolin have been found to
be effective against Botrytis grey mould, but can be uneconomic. Several
applications are required if conditions remain favourable for the disease
(Bayaa and Erskine, 1998; Davidson et al., 2004). However, precaution
should be taken to prevent development of fungicide resistance since Botrytis spp. are prone to evolve resistance to some synthetic, systemic fungicides
(Raposo et al., 1996). Resistant germplasm of lentil has been identified in
several breeding programmes and commercial cultivars have been released
with better resistance to Botrytis grey mould (Bayaa and Erskine, 1998;
Davidson et al., 2004).
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17.5. Anthracnose
Anthracnose is a serious disease of lentil in Canada where it was discovered
first in 1987. It has been subsequently reported in North Dakota, USA and
Bulgaria, and there is evidence of its occurrence in Bangladesh, Brazil, Ethiopia,
Morocco, Pakistan and Syria (Morrall, 1997). The first symptoms develop
when the lentil crop starts to flower. Necrotic lesions can be found on the
lower leaflets and many of them fall to the soil surface (Buchwaldt et al.,
1996). This premature leaf drop is characteristic for anthracnose. Lesions also
develop on the stems starting at the base and spreading to the top part of the
plant. They are tan coloured often with a defined reddish or black border,
and the deepest lesions create indentations along the stem (Plate 2E). Symptoms caused by anthracnose and Ascochyta blight are very similar, but the
presence of acervuli and microsclerotia produced by the anthracnose pathogen or pycnidia produced by the Ascochyta pathogen makes it possible to
distinguish the two diseases. Plants severely attacked by anthracnose disease
turn brown and areas of lodged and dying plants can be seen in the field.
Causal organism
Lentil anthracnose is caused by Colletotrichum truncatum (Schw.) Andrus &
Moore. The pathogen produces conidia and setae in acervuli on infected
plants and in culture. The conidia are single-celled, hyaline and slightly falcate, 1830 m long by 36 m wide. The black setae are 60120 m long by
3.58.0 m wide and protrude above the spore mass. The pathogen produces black pinhead-sized microsclerotia. These are particularly abundant
at the stem base, where they can be seen with the naked eye. Microsclerotia
are resting structures consisting of a couple of hundred heavily melanized
cells, able to survive in the soil in periods when host plants are absent
(Buchwaldt et al., 1996). A teleomorph stage of C. truncatum pathogenic on
lentil has not been found in nature, but perithecia with mature asci have
been induced in the laboratory by inoculating stem pieces with different
isolates (Armstrong-Cho and Banniza, 2006). Each ascus contains eight singlecelled ascospores measuring 1220 m long by 58 m wide. The sexual
stage is Glomerella truncata.
The pathogen survives in the field as microsclerotia on infected lentil debris
that is the primary source of inoculum. The initial infection occurs when
microsclerotia or conidia come in contact with lower leaves and stems of a
new lentil crop. Microsclerotia are also dispersed by wind either in dust
generated during harvest of infected crops or on plant debris from fields.
The disease is polycyclic and spreads rapidly in the field in periods of
rain. Conidia of C. truncatum are readily dispersed by rain droplets to the
269
surrounding plants. The optimum temperature range for infection and production of new conidia is 2024C, which results in a latent period of
approximately 13 days (Chongo and Bernier, 2000). Colletotrichum truncatum
is a hemibiotrophic pathogen. In the biotrophic phase, a conidium forms an
appressorium from which an infection peg penetrates the wall of a single
epidermal plant cell. Primary hyphae grow inside the host cell for 23 days
without killing the infected cell. In the necrotrophic phase secondary hyphae
colonize the surrounding tissue both inter- and intracellularly resulting in
appearance of necrotic symptoms 56 days after infection.
Management
Colletotrichum truncatum from lentil also attacks faba bean (V. fabae L.), other
Vicia species, and pea (Pisum sativum L.). A 3-year crop rotation is recommended to allow adequate reduction of the soil-borne inoculum. Tillage of
fields with anthracnose-infected stubble should be avoided in areas with
long cold winters, such as in Canada, to take advantage of the breakdown
of infectivity that occurs on the soil surface when the pathogen is exposed
to extreme temperatures and repeated wetting and drying. Undoubtedly,
the ability of the pathogen to survive in the soil differs under other climatic
conditions. The level of seed-borne inoculum is often less than 1%, and the
significance of seed-borne inoculum remains to be determined. Several foliar
fungicides, such as chlorothalonil and several strobilurin products, efficiently
control anthracnose when applied preventatively; some chemicals in the
latter group also have a slightly systemic effect. The optimum time of fungicide application in Canada is between the 8 and 12 node stage and approximately 1 week prior to flowering (Chongo et al., 1999). An above normal
and premature leaf drop at this time is the best indication that anthracnose
is likely to become a problem. It is important to apply a fungicide before the
canopy becomes too dense in order to obtain the best possible coverage of
the lower portion of the stems. A second application 1014 days later may
be necessary under high disease pressure and frequent rainfall.
Two morphologically identical races are present in Canada (Buchwaldt et al., 2004). Isolates of race Ct1 are unable to infect lines such as Indianhead, PI 320937, PI 345629 and PI 468901 (United States Department of
Agriculture (USDA) Agriculture Research Service (ARS), Pullman, Washington). Single major genes, either dominant or recessive, conferring resistance to race Ct1 have been utilized in Canada to develop cultivars with an
improved level of anthracnose resistance under field conditions (Buchwaldt et al., 2004). So far, sources of resistance to race Ct0 have not been found
in cultivated lentil, but are available in wild Lens species (Tullu et al., 2006).
17.6. Stemphylium Blight

Stemphylium blight of lentil is the most devastating disease in Bangladesh,
eastern Nepal and north-eastern India, where yield losses above 80% have
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been recorded (Bakr and Ahmed, 1992; Sinha and Singh, 1993). Reports on
the disease have also been published from Canada, Egypt, Hungary, Syria
and the USA (Bayaa and Erskine, 1998; Morrall et al., 2006). Initial symptoms often appear in the upper canopy in the form of small, light-beige
lesions that enlarge and coalesce (Plate 2D). Leaves, pedicels, flowers and
entire branches can become necrotic resulting in a characteristic blighted
appearance of lentil plants. Severe infection can significantly reduce plant
biomass, lower seed yield, decrease seed size and result in seed staining
and lower germination rates.
Causal organism
Stemphylium botryosum Wallr. produces light-brown to black conidia with a
length-to-width ratio of 1:1.5. Conidia (dictyospores) are partitioned by a
median transverse septum, and secondary trans- and longi-septa and range
in size from 24 14 m to 40 26 m. The fungus has been reported from a
wide range of host species including asparagus, canola, lucerne, cotton,
tomato, garlic, mango, pear, onion, clover and lentil. It sporulates profusely
on host tissue, but its pseudothecia have only been observed in culture.
The teleomorph stage is the ascomycete Pleospora tarda E.G. Simmons.
Septate ascospores of 40 17 m are formed in subcylindrical asci measuring approximately 200 40 m that develop in pseudothecia (Bayaa and
Erskine, 1998).
Airborne conidia of S. botryosum are commonly found in the environment
and are formed on conidiophores that produce successive generations of
spores. They germinate with several germ tubes that penetrate primarily
through stomata, and invade plant tissues including seed. The fungus produces the phytotoxin stemphol, which has been implicated in lesion formation on various hosts (Solfrizzo et al., 1994). Conidial germination and
infection of lentil can occur at temperatures from 5 to above 30C in the
presence of free water. Optimal conditions for Stemphylium blight epidemics are characterized by high relative humidity, cloudy days and moderate
to warm temperatures (Sinha and Singh, 1993). While Stemphylium blight
symptoms can be observed early in the growing season in Bangladesh and
India, the disease appears during the last third of the growing season in
Canada. Infected lentil debris and alternate hosts as well as saprophytic
growth enable S. botryosum to survive between crops.
Management
In Bangladesh and India, Stemphylium blight is managed through late
seeding of lentil, fungicide applications and the use of resistant cultivars.
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Three applications of iprodione, propineb, mancozeb or sulfur, sprayed at

7-day intervals once symptoms were observed, significantly reduced disease severity and increased yield by 2340% (Bakr and Ahmed, 1992). The
resistant cultivars Barimasur-4, Barimasur-5 and Barimasur-6 were developed in Bangladesh in cooperation between ICARDA and the Bangladesh
Agricultural Research Institute (BARI).
17.7. Powdery Mildew

Powdery mildew has been reported from many countries such as Cyprus,
Ethiopia, India, Siberia, Sudan, Syria, Tanzania and the former USSR. In
India and Sudan, the disease occurs in severe forms almost every year
on susceptible varieties. It becomes more severe in the Indian province of
Rajasthan, especially under dry weather conditions. In North America,
powdery mildew is often observed after lentil starts flowering in the field,
whereas it occurs at any growth stages under greenhouse conditions. The
disease poses a serious problem on breeding materials in plastic- or glasshouses in both India and Syria, and in India it is also recorded in off-season
nurseries in Trans Himalayan regions, such as Lahaul Spiti and Sangla, but
it is rarely seen in the field during the cropping season.
Symptoms are observed on the upper surface of older leaves (Beniwal et al., 1993). A fine powdery, white growth of conidia and mycelium
initiates as small spots and spreads rapidly to cover the entire surface
of leaves (Plate 2F), stems and pods. Later, the leaflets become dry and
curled, and are shed prematurely, causing considerable reduction in
yield and seed quality. The seeds from infected plants remain small and
shrivelled.
Causal organisms
Powdery mildew of lentil is caused by the ectoparasites Erysiphe pisi DC.
and Erysiphe polygoni DC. and the endoparasite Leveillula taurica (Lv.)
Arnaud. The anamorph stage of E. polygoni, Oidium sp., has been frequently
reported, and the anamorph stage of L. taurica, Oidiopsis taurica Salmon, has
been reported from the former USSR and Jordan. Recent evidence showed
that Erysiphe trifolii also infects lentil (Attanayake et al., 2008).

The anamorph stage is responsible for spread of the disease. The teleomorph stage has been reported to occur in India and Sudan (Chitale et al.,
1971). Moderately high temperatures and moderate relative humidity
favour disease development.
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Management
Many lentil genotypes are reported resistant to powdery mildew (Tikoo
et al., 2005), and should be planted whenever possible. Foliar sprays with
fungicides benomyl, tridemorph, aqueous sulfur, karathane (dinocap), calixin or sulfex (ferrous bisulfide) as well as certain insecticides (Quinalphos,
Tnazophos, Phoxim) applied at 1015 day intervals are effective in suppressing powdery mildew growth (Beniwal et al., 1993).
17.8. Fusarium Wilt

Fusarium wilt of lentil is an important soil-borne disease, and causes significant yield loss under dry and warm conditions. The disease occurs in
most lentil production regions, and has been reported to occur in at least 26
countries (Agrawal and Prasad, 1997; Bayaa and Erskine, 1998). Fusarium
wilt of lentil can be observed at the seedling and reproductive stages. It
usually occurs during the reproductive stages from flowering to pod filling,
causing yellowing, leaf curling and stunted growth. Infected plants show
reduced root development, sometimes with yellowish-brown discoloration
of vascular tissue and poorly developed nodules (Beniwal et al., 1993).
Causal organism
The fungal pathogen Fusarium oxysporum Schlecht. :Fr. f. sp. lentis Vasudeva
and Srinivasan causes Fusarium wilt disease of lentil. The fungus produces
three kinds of asexual spores: microconidia, macroconidia and chlamydospores. Microconidia are single celled, kidney shaped, ellipsoid to ovoid,
cylindrical, oblong or slightly curved, and hyaline, often agglutinated into
false heads. Macroconidia are 15 septate, nearly straight to fusiform, falcate, slender and thin-walled. Terminal or intercalary chlamydospores are
formed on mycelium. They are generally single celled, rarely two-celled,
spherical to pyriform, smooth and hyaline. The teleomorph state of the
fungus has not been observed.

Fusarium wilt is a monocyclic disease. The initial inoculum level is very important for determining the incidence and severity of the disease. The pathogen
survives as chlamydospores and dormant mycelium in infected plant debris.
The fungus can also be seed-borne either as systemic infection inside the
seeds or as a contaminant on the seeds. Seed-borne inoculum is an important source for the introduction of the disease into new production areas.
Warm soil temperature 2030C and dry soil conditions (25% water holding capacity) are the two most important environmental factors favouring
273
development of the disease (Khare, 1981). Other environmental conditions

such as soil type, aeration, pH and fertility, as well as soil microflora may also
affect the disease development. Soil-borne lentil-infecting nematodes, such as
Ditylenchus spp., Meloidogyne spp. and Pratylenchus spp., interact synergistically with F. oxysporum f. sp. lentis to increase disease severity (De et al., 2001).
Management
Proper diagnosis is the first step in managing Fusarium wilt of lentil. A
recent history of Fusarium wilt increases the likelihood of its reoccurrence.
Reduced root systems with discoloration, but without external fungal
growth on stunted plants suggest Fusarium wilt infestation. Definitive identification of the pathogen requires pathogenicity tests on susceptible lentil
plants. Planting of resistant cultivars is the most economical means for its
management. Resistance sources have been identified and incorporated into
lentil cultivars and resistant cultivars are available in several countries
(Bayaa and Erskine, 1998; Tikoo et al., 2005; Stoilova and Chavdarov, 2006).
Some high-yielding lentil lines have been reported to be resistant to multiple diseases including rust, Ascochyta blight and Fusarium wilt (Bayaa and
Erskine, 1998). However, resistance alone is not sufficient for satisfactory
management of the disease. So a number of cultural practices should be
used in conjunction in managing the disease.
Seeds for planting should be free of infection and contamination, particularly when expanding lentil production into new areas. If disease-free
and clean seeds are not available, fungicide treatments should be implemented. Seed treatment with carboxin and thiram, thiram, pentachloronitrobenzene, carbendazim, benomyl, or with boric acid plus KMnO4 can
reduce seed infection. Direct application of fungicides to the soil after planting for the control of Fusarium wilt is not economical because of the high
cost and technical difficulty during the growing season. Another management option is to select early maturing varieties or adjust planting time, so
that plants can escape hot weather conditions during the vulnerable growing stages. However, the most suitable planting date varies depending on
the climatic conditions of the production regions. The principle is to reduce
the overlap of susceptible growth stages with hot weather conditions.
Cultural practices like applying manure or composts that enhance populations of general microflora or supplementation with selected antagonistic
biocontrol agents may be feasible on subsistence or small farms.
17.9. Sclerotinia Stem Rot

Sclerotinia stem rot has a worldwide distribution and occurs on more than
400 plant species in 75 families (Boland and Hall, 1994). Economic loss can
occur in lush lentil canopies under wet conditions (Akem et al., 2006).
Infected plants exhibit bleached lesions on stems, leaves, pedicels and pods,
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W. Chen et al.
which are sometimes covered by a characteristic cottony white mycelium

occasionally harbouring dark sclerotia. Stem infection can cause wilting of
plants (Bolten et al., 2006).
Causal organism
The disease is caused by Sclerotinia sclerotiorum (Lib.) de Bary. The fungus
produces black, rounded to elongate sclerotia of up to 1 cm in length. Sclerotia can germinate directly by mycelium (myceliogenically) and can grow
rapidly over host tissue and ramify intra- and intercellularly. No spores are
formed, though globose spermatia of unknown function extruded through
phialides at the tips of microconidiophores which are often present in the
mycelium. Sclerotia can also germinate carpogenically to produce one to
several light- to dark-brown apothecia. These are generally 0.52 cm in
diameter, funnel shaped to discoid with a layer of asci (the hymenium) on
the upper surface. Eight elliptical to ovoid, hyaline, unicellular ascospores
develop within each ascus, and are forcibly ejected from asci (Clarkson et al.,
2003; Bolten et al., 2006).

The disease is promoted by high plant density, excessive vegetative growth
and high precipitation in the last third of the lentil-growing season (Akem
et al., 2006). Infection on lentil is initiated through ascospores released from
carpogenically germinating sclerotia, as well as through myceliogenically
germinated sclerotia that have overwintered on or near the soil surface.
Specific soil temperature and moisture levels are required to trigger carpogenic germination (Morrall, 1977). Senescent flower petals are the sites of
infection by ascospores providing them with nutrients required for successful invasion of the host tissue. Once abscised and in contact with healthy
tissue, the infected senescent petals are sources of inoculum for leaves and
stems. Evidence has also emerged on the ability of germinating ascospores
to directly invade healthy, green lentil tissue. Plants become more susceptible to infection as they mature. Contact with the soil by lodging plants and
between healthy and infected plant parts in dense canopies also result in
infection. Invasion of the tissue is facilitated by the production of cell-wall
degrading enzymes and oxalic acid, which cause yellowing and development
of bleached lesions. Sclerotia formed on infected lentil tissue are returned to
the soil upon harvest or threshed out with the seed (Bolten et al., 2006).
Management
Crop rotation has limited efficacy in controlling Sclerotinia stem rot because
of the longevity of sclerotia and the pathogens wide host range. High levels
275
of resistance to the disease have not been found in lentil, but some cultivars
consistently perform better. Whereas early fungicide applications can prevent or reduce flower petal infection, late season control through fungicides
is ineffective because the dense canopy prevents penetration of the fungicide to the lower plant canopy. Fungicide applications may not always control
the disease and may not be economical (Bolten et al., 2006).
17.10. Nematode Diseases

Nematode diseases of lentil are commonly observed worldwide and are
often important diseases in India, Syria, Turkey and in African countries.
The stem nematode is common in the temperate regions of the world
comprising Europe and the Mediterranean region, North and South
America, northern and southern Africa, Asia and Oceania (Agrawal and
Prasad, 1997).
Common above-ground symptoms of nematode diseases are similar to
those associated with plants with an impaired root system, because the
nematodes usually feed on and damage roots. Symptoms include stunting,
chlorosis, yellowing, wilting and plant death, which often appear in
patches in the field. However, these symptoms are not specific for nematode diseases. Therefore, definitive diagnoses require examination of root
systems and the attached soil. The characteristic feature of root-knot nematode is the presence of small galls on roots (Beniwal et al., 1993). Some lateral rootlets may develop out of the galls. The galls are similar to
nitrogen-fixing nodules in size and shape, but galls are easily differentiated from nodules by lack of colour and swelling of the root diameter. The
most characteristic features of infection by cyst nematodes are the presence
in roots of lemon-shaped cysts (i.e. leathery bodies of adult females)
(Agrawal and Prasad, 1997). The main symptoms and damage caused by
root-lesion nematodes are lesions or discoloration of roots and lack of
branching along the main roots (Beniwal et al., 1993). Stem nematodes
cause brownish necrotic lesions on the bases of stems, stunted and distorted leaves and stems. Lentil is very susceptible to stem nematodes at the
seedling stage.
Causal organisms
Many plant parasitic nematodes are associated with lentil. The economically important nematodes of lentil include: root-knot nematodes (Meloidogyne incognita (Kofoid & White) Chitwood and Meloidogyne javanica (Treub)
Chitwood); cyst nematode (Heterodera ciceri Vovlas, Greco & Di Vito); rootlesion nematodes (Pratylenchus thornei Sher & Allen, Pratylenchus penetrans
(Cobb) Chitwood and Oteifa and Pratylenchus mediterraneus Corbett); and
stem nematode (Ditylenchus dipsaci (Khn) Filipjev).
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W. Chen et al.

Plant parasitic nematodes have four moults of juvenile stages and then proceed to adult egg-laying females. Eggs hatch and produce second-stage
juveniles that seek and infect host tissue. Egg fertilization is not necessary
for many species. The egg and juvenile stages are resistant to adverse conditions and enable nematodes to survive from season to season. Some species take 1 year to complete a single life cycle (egg to egg), while some
others can complete several cycles during one growing season. Females
either deposit eggs into the soil around the infected roots or they are encased
in the female body (cyst). Cyst nematodes can survive for long periods of
time without a host. These are more prone than other plant parasitic nematodes to unintentional long-distance transport because their resistant cyst
stage can tolerate long periods of desiccation and adhere to soil particles on
vehicles and clothing. Stem nematodes at the fourth juvenile stage invade
seedlings and damage the growing tips of leaves and stems. After entering
the host plants, they mature into adults, feed and lay eggs. If the soil dries
up in late spring or summer, the fourth-stage juveniles can enter a state of
anhydrobiosis in either the plant or soil and can survive high temperature
and desiccation.
The host range of the cyst nematode is limited to Leguminosae, mainly
chickpea, lentil, pea and grass pea, while other lentil nematodes generally
have wider host ranges. Root-knot nematodes infect many vegetable, horticultural, cereal and legume crops. The host range of root-lesion nematodes
includes many cereal, legume and oilseed species. Stem nematode can attack
over 450 plant species including many weed species. Races are reported in
stem nematodes, and some races have limited host ranges. Nematodes are
notorious for interacting with other soil-borne fungal pathogens additively
or synergistically (Tiyagi et al., 1988; Anver et al., 1991).
Management
Crop rotation is a general practice in managing nematode diseases, and it
is particularly effective in controlling cyst nematodes because of their narrow host range. Even with the nematodes that have wide host ranges,
some crops are less favourable than others. In general, rotation with cereal
and Brassica species are effective in managing root-knot nematodes.
Another effective measure is to plant resistant or tolerant varieties whenever possible, which sometimes can significantly reduce the nematode
population. Where lentil is produced as a winter crop, solarization in the
summer for 68 weeks can also effectively reduce the nematode population (Linke et al., 1991). Since many weeds are alternate host of the nematodes, good weed control is another effective measure in reducing
nematode populations. Seed treatments with nematicides and soil amendments with organic matter can reduce disease severity as well (Gaur and
Mishra, 1990).
Table 17.1.
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List of bacterial and other fungal and nematode diseases of lentil.
Disease name
Pathogen
References
Alternaria blight
Alternaria alternata (Fries) Keissler
Aphanomyces root rot
Aphanomyces euteiches C.
Drechsler
Xanthomonas sp.
Pseudomonas radiciperda
Fusarium solani (Mart.) Sacc.
Gupta and Das (1964);

Kaiser (1992)
Lamari and Bernier (1985)
Bacterial leaf spot

Bacterial root rot
Black root rot
Black streak root rot
Cercospora leaf spot
Collar rot
Cylindrosporium leaf
spot and stem canker
Downy mildew
Dry root rot
Helminthosporium leaf
spot
Leaf yellowing
Phoma leaf spot
Pythium root and
seedling rot
Reniform nematode
Wet root rot
aPathogenicity
Thielaviopsis basicola (Berk. &

Broome) Ferraris
Cercospora lensii Sharma et al.
Sclerotium rolfsii Sacc.
Cylindrosporium sp.
Agrawal and Prasad (1997)

Agrawal and Prasad (1997)
Karahan and Katrcoglu
(1993)
Bowden et al. (1985)
Sharma et al. (1978)
Beniwal et al. (1993);
Khare (1981)
Bellar and Kebabeh (1983)
Peronospora lentis Gumann

Macrophomina phaseolina (Tassi)
Goidanich
Helminthosporium sp.a
Mittal (1997)
Kaiser (1992); Vishunavat
and Shukla (1979)
Karahan and Katrcoglu
(1993)
Cladosporium herbarum (Pers.)
Kaiser (1992); Karahan and
Link
Katrcoglu (1993)
Phoma medicaginis Malbr. & Roum. Kaiser (1992)
Pythium ultimum Trow, Pythium spp. Paulitz et al. (2004)
Rotylenchulus reniformis Linford &

Oliveira
Rhizoctonia solani Khn,
teleomorph: Thanatephorus
cucumeris (Frank) Donk
Anver et al. (1991); Gaur and

Mishra (1990)
Kaiser (1992); Karahan and
Katrcoglu (1993)
not conrmed.
17.11. Other Diseases

Due to limitation of space, a number of bacterial, fungal and nematode diseases of lentil not covered in details in this chapter are listed in Table 17.1,
some of which might be quite important locally. There are others for which
very little information is available. Reporting and more detailed studies on
those diseases are encouraged.
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18
Insect Pests and their Management

Ram Ujagir1 and Oonagh M. Byrne2
1G.B.
2The
Pant University of Agriculture and Technology, Pantnagar, India;

University of Western Australia, Crawley, Western Australia, Australia
18.1. Introduction
Lentil (Lens culinaris Medik. subsp. culinaris) is mainly cultivated in South
Asia, West Asia, North Africa, the Nile valley region, North America, South
America, Eastern Europe and Australia.
The lentil crop is attacked by a wide range of insect pest species, only
some of which are economically important, requiring control measures:
these include aphids, cutworms, pod borers, thrips, leaf and seed weevils.
Other pests infesting lentil, but not causing major economic damage, are
root aphids, leaf miner, lygus bug, lucerne weevil, semilooper and armyworms.
At the initial stage of crop growth, wire worms, lucerne flea and cutworms and Agrotis ipsilon cause the most damage to lentil. During the vegetative stage foliage feeders such as bud weevil, grasshopper, Sitona crinitus,
chrysomelids, cicadellids, Spodoptera exigua, Autographa gamma, Chromatomyia horticola, bugs, aphids and leaf miners impact most. Thrips and aphids
cause most damage at the reproductive stage of crop growth. At the podding stage grasshoppers and the pod borers, Helicoverpa armigera and Etiella,
cause most damage. Bruchus lentis and Bruchus ervi infest the lentil crop at
podding, causing seed losses in storage. Callosobruchus maculatus and Callosobruchus chinensis also attack lentil seed in storage. Viruses transmitted
by insect vectors such as aphids can impact on lentil production. Insect
damage can lead to subsequent fungal and bacterial diseases (Hariri, 1981).
18.2. Pest Complex Infesting Lentil Crop

The pest spectrum of lentil is wide with the crop being attacked by a large
number of insect pests. Bhatnagar and Sehgal (1989) reported about 20 insect
282

283
species infesting lentil in the northern Terai region of India. Among them
aphids, leaf miners and semiloopers attack the lentil crop at vegetative and
flowering stages, while pod-borer infestation occurs at late flowering and
post-reproductive stages. About 20 insect pests were found to be associated
with the crop in Ethiopia (Ali and Habtewold, 1993); most important among
these are the pea aphid, African bollworm and bean bruchid. The occurrence and status of different insect pests associated with lentil has been
reported to vary over geographical regions. A list of insects known to be
associated with lentil globally is given in Table 18.1.
One of the most important groups of insect pests of the crop are the
aphids, Aphis craccivora and Acrythosiphon pisum particularly in dry seasons in West Asia and North Africa (Thakur et al., 1984; Erskine et al.,
1994; Singh and Saxena, 1994). Sehgal and Ujagir (1980) reported the leaf
miner, C. horticola, as a major insect pest causing direct damage to the
lentil crop through foliage feeding. Hammad (1978), Singh and Dhooria
(1971), Erskine et al. (1994) and Singh and Sharma (2001) reported that
the pod borer Etiella zinckenella Treische, was a major pest of lentils in the
Indian Punjab and also in West Asia, and North and East Africa. Erskine
et al. (1994) observed that the species Apion ervi feeds on lentil. Grasshoppers, in particular Melanoplus bivittatus (Say), are reported as an economic
problem in lentil crops at flowering and early pod stages in Canada
(Olfert and Slinkard, 1999). Over the last decade in south-eastern Anatolia,
Turkey, lygus bugs, Exolygus (= Lygus) pratensis, have come to be considered the most important insect seed pest in red lentil because feeding
adults damage the seed, causing chalky spot syndrome (zberk et al.,
2006). In the Terai region of Nepal, Pandey et al. (2000) reported A. ipsilon,
A. pisum, H. armigera, C. chinensis and C. maculatus as major and Phyllotreta
sinuate, Athalia lugens proxima (Klug) and Adonia variegata as minor pests
of lentil.
Storage pests are important in all production areas and infestations can
result in complete loss of the crop (Singh and Saxena, 1994). The principal
storage weevils of lentil are C. maculatus and C. chinensis species. Bruchus
lentis is the most widespread and damaging of the seed-feeding bruchid
pests of lentil (Erskine et al., 1994).
18.3. Estimation of Losses Caused by the Insect Pest Complex

Perez-Andueza et al. (2004) reported aphids (A. craccivora and A. pisum), B.
lentis, thrips (Thrips tabaci and Thrips angusticeps) and leaf weevil (Sitona
lineatus) as key pests of lentil in Spain. The average damage caused by
insect pests to lentil during a 2-year study ranged from 24 to 59% (PerezAndueza et al., 2004). Leaf damage by Sitona weevil was assessed in former
Czechoslovakia under laboratory conditions by Sedivy (1972) who reported
175% leaf damage. Sitona weevil, S. crinitus Herbst, is the principal field
pest of lentil in West Asia and North Africa (Erskine et al., 1994; Singh and
Saxena, 1994).
284
Table 18.1.
Insect pests associated with the lentil crop.
Order and family
Scientic name
Orthoptera
Melanoplus bivittatus (Say)

Melanoplus sanguinipes (Fabr.)
Melanoplus packardii (Scudder)
Camnula pellucida (Scudder)
Bemisia tabaci (Gennadius)
Nature of damage/
plant parts attacked
Country/continent
References
Minor
Asia, North Africa,

Canada
Olfert and Slinkard (1999)
Suck the sap from

foliage and owers
Minor
Suck the sap from
tender parts of plant
Vector of viral diseases
Egypt
Hammad (1978)
Europe, Asia,
America, Africa
Worldwide
Suck the sap from

America, Europe,
Asia, Africa
Worldwide
Suck the sap from

Suck the sap from
North-west Pacic
region
India
Hariri (1981), Mokiyar (1985),

Bhatnagar and Sehgal (1989)
Makkouk and Kumari (2001),
Jones and Coutts (2008)
Tahhan and Hariri (1982),
Thakur et al. (1984),
Halfhill (1982)
Root damage
Syria
Miridae
Smynthurodes betae
(Westwood)
Lygus elisus (Vand)
Lygus hesperus (Knight)
Exolygus pratensis (Linnaeus)
Pod and seed damage

Pod and seed damage
Pod and seed damage
Pentatomidae
Nezara viridula (Linnaeus)
Suck the sap from

foliage
America
Europe
South-east
Anatolia, Turkey
Europe, Africa,
Asia, Australia
and America
Hemiptera
Aleyrodidae
Aphididae
Acrythosiphon pisum (Harris)
Aphis craccivora (Koch)
Macrosiphum creelii (Davis)

Myzus persicae (Sulzer)
Defoliators ower,
buds, foliage, early
pods
Worldwide
Sekhon et al. (1979),

Pandey et al. (2000)
Fye (1982)
Schotzko and OKeeffe (1986)
zberk et al. (2006)
Hariri (1981)
Status
Coreidae
Lygaeidae
Lepidoptera
Noctuidae
Phycitidae
Balclutha sp.
Empoasca sp.
Cletus signatus Walker
Graptostethus servus
(Fabricius)
Eusarcocoris ventralis
Westwood
Piezodorus rubsofasciatus
Fabricius
Agrotis ipsilon (Hufnagel)
Helicoverpa (Heliothis)
armigera (Hubner)
Laphygma exigua (Hubner)
Spodoptera praeca (Hubner)
Spodoptera exigua (Hubner)
Foliage
Foliage
Foliage
Foliage
Minor
Minor
Minor
Minor
India
India
India
India

Suck the sap from

foliage
Suck the sap from
vegetative parts
Cut the stem
Damage the pod and
seed
Defoliation
Defoliation
Defoliation
Minor
India
Minor
India
Syria, Egypt
Syria, India
Spodoptera littoralis (Boisd.)

Trichoplusia ni (Hubner)
Thysanoplusia oricholcea (Fab.)
Etiella behrii (Zeller)
Defoliation
Defoliation
Defoliation
Pod and seed
damage
Pod and seed
damage
Damage the pods
Zaazou et al. (1975)

Tahhan and Hariri (1982),
Paharia (1983)
Sacharov (1916)
Halfhill (1982)
Sacharov (1916), Brown and
Dewhurst (1975)
Brown and Dewhurst (1975)
Hariri (1981)
Etiella zinckenella (Tretsche)

Tortricidae
Diptera
Agromyzidae
Laspreyresia nigracana
(Fabricius)
Autographa gamma (Linnaeus)

Chromatomyia horticola
(Goureau)
Foliage feeder
Mines the leaves
Minor
Major
India
America, Syria
India, Africa, Near
East
Near East
Syria
Europe, India
Victoria, South
Australia
India, USA, Egypt
Cicadellidae
Singh and Dhooria (1971),

Hammad (1978)
Sedivy and Suchanek (1978)
North America,
former Czechoslovakia, Europe,
Mediterranean
Nepal, Syria
Europe, Africa, India Sehgal et al. (1980),
Mandal (1982), Bhatnagar
and Sehgal (1989)
285
(Continued)
286
Order and family
Cecidomyiidae
Coleoptera
Bruchidae
Nature of damage/
plant parts attacked
Ophiomyia phaseoli (Tryon)

Liriomyza spp.
Leaves and stem

Leaves
Contarinia spp.
Contarinia lentis (Aczel)
Damage leaves
Damage leaves
Bruchidius quinqueguttatus
(Oliv.)
Bruchus atomarius (L.)
Bruchus lentis (Froel.)
Damage pods and

seeds
Damage pods and
seeds
Damage pods and
seeds
Damage pods and
seeds
Pods and seeds
Bruchus ervi (Froel.)
Pods and seeds
Callosobruchus maculatus
(Fabricus)
Flowers, pods, seeds
Callosobruchus chinensis
(Linnaeus)
Flowers, pods, seeds
Apion ervi (Kirby)
Leaves, buds, owers
Apion trifolii (Linnaeus)
Leaves, buds, owers
Bruchus signaticornis (Gyll.)

Bruchus tristiculus (Fahrs.)
Apionidae
Status
Country/continent
References
Egypt
West Africa, North
Africa and South
America
Europe, Africa, India
France, former
Czechoslovakia
Mediterranean,
South-west Asia
Euro-siberian
Harakly and Assem (1980)

Stevenson et al. (2007)
Europe, North Africa,

America
Mediterranean, Near
East
Europe, Africa
and Asia
Europe, Nepal,
North Africa, Southwest Asia, India
Europe, Asia,
USA, Australia,
Mediterranean
Europe, Asia,
USA, Australia,
Mediterranean
Europe, South-west
Asia
Europe, USSR,
South-west Asia,
Mediterranean
Alkan (1966)
Kemkemian (1979)
Minssen and Pacqueteau (1969),
Kolesik and Kolesik (1989)
Hammad (1978)
Hammad (1978)
Hammad (1978)
Gibson and Raina (1973)
Staneva (1982), Pandey et al.

(1985)
Staneva (1982), Pandey et al.
(1985)
Sakamoto et al. (1983)
Melamed-Madjar (1968)
Scientic name
Apion pomonae (F.)

Apion punctigerum (Payk.)
Apion seniculus (Kirby)
Curculionidae
Sitona crinitus (Herbst)
Sitona macularius (Marsham)

Sitona limosus (Rossi)
Tychius aorominus
quinonepunctatus (Linnaeus)
Chrysomelidae Altica coerulea (Oliver)
Aulacophora foveicollis (Lucas)
Phyllotreata chotanica (Duvivier)
Hypera postica (Gyllenhal)
Coccinellidae
Thysanoptera
Thripidae
Brumoides suturalis Fab.

Epilachna spp.
Thrips anguisticeps (Uzel)
Kakothrips robustus (Uzel)
Frankliniella spp.
Thrips tabaci (Lind.)
Damage leaves and

seeds
Damage leaves and
seeds
Damage leaves and
seeds
Damage leaves and
seeds
Foliage and root
nodules
Major
Foliage and root

nodules
Damage seedlings and
nodules
Pod and seed
Foliage feeder
Foliage feeder
Foliage feeder
Foliage feeder
Minor
Minor
Minor
Minor
Minor
Damage foliage
Damage foliage
Minor
Suck the sap from
plant
Minor
Suck the sap from
plant
Suck the sap from
plant
Suck the sap from plant
Turkey and Syria
Zeran and Yabas (1984)
Europe, former
USSR, Turkey, Syria
Europe, former
USSR, Syria
Europe, former
USSR, Turkey
Europe, West Asia,
South-west Asia,
North Africa, former
USSR
Former USSR
Rivnay (1962)
Syria
Romania
Boguleanu et al. (1971)
India
India
India
West Asia, Europe,
USA
India
India
Syria

Nepal, Asia, Africa

Syria
Spain
Kilic et al. (1968)
Apion arrogans (Wenk.)
Hariri (1981)

Beniwal et al. (1993), Pandey
et al. (2000)
et al. (2000)
et al. (2000)
Perez-Andueza et al. (2004)
287
288
Several workers have reported C. horticola as an important pest of

legumes (Lefroy, 1909; Tahhan and Hariri, 1982). The damage to lentil
leaves has been reported to be as high as 5.5% from mid-April to mid-May
(Tahhan and Hariri, 1982). The bean aphid, A. craccivora, has been widely
reported from America, Europe, Africa, Asia and Australia. Tahhan and
Hariri (1982) observed that the feeding of bean aphids on young lentil
plants caused a reduction in the relative growth rate and efficiency of production of new tissues at an average plant infestation level of about 17.5%.
The pod borer, H. armigera, has been reported as the major pest attacking
the lentil crop. Field infestation of 26% by pod borer was observed during
a survey in Syria (Tahhan and Hariri, 1982). Paharia (1983) reported that
damage caused by the lentil pod borer, E. zinckenella, under field conditions was the main limiting factor in the production of lentil and gram.
Lentil pod borer can cause from 31 to 50% loss in yield (Memon and
Memon, 2005).
Singh and Dhooria (1971) reported that the pod borer E. zinckenella caused
10.6% damage to lentil grain and a loss in seed weight. Sandhu and Verma
(1968) assessed the losses caused by E. zinckenella to lentil pods as 1215%.
Experiments at the International Center for Agricultural Research in the
Dry Areas (ICARDA) showed that small lentil seed infested with B. ervi did
not germinate, while 20% of infested large seeds germinated (Hariri, 1981).
Infestation by C. chinensis also affects seed germination. Germination of
large and small lentil varieties was 84 and 25%, respectively, when the seeds
harboured one bruchid each; 32 and 5% when there were two bruchids per
seed; and 20 and 0% when each seed had three bruchids. In a survey in
Syria, Tahhan and Hariri (1982) observed that 93% of fields were infested
by bruchids with damage to 1.9% of plants.
Zeran and Yabas (1984) observed that bruchid feeding resulted in the
formation of swelling and spots on the buds and inhibited capsule production, leading to considerable crop losses. They also studied the morphology
and nature of damage caused by Apion sp. on the crop. Economic losses
resulting from damage by lygus bug, bud weevil and Sitona spp. are
reported for red lentil exports in Turkey (zberk et al., 2006). Summerfield
et al. (1982) reported that feeding by Lygus sp. on immature reproductive
structures caused formation of poor quality seeds (chalky spot) and reduced
economic yield.
Lentil weevil, B. lentis, and spotted pea weevil, Tychius quinquepunctatus, were reported to be the major pests of lentil in Italy. The seed damage
due to lentil weevil ranged from 6 to 29% under field conditions and 5.5 to
14.5% at harvest (Isidoro et al., 2001).
18.4. Insect-spread Viruses

The lentil crop is susceptible to a variety of insect-borne virus diseases
which can severely reduce the crop yield and are mostly spread by aphids
(see Kumari et al., Chapter 19, this volume, for details on viral diseases).
289
Lentil yellows disease

The disease is caused by Bean leaf roll virus (BLRV), Beet western yellows virus
(BWYV) or Subterranean clover red leaf virus (SCRLV). These viruses are
transmitted through the sap and by aphid in the non-persistent manner.
A number of aphids can transmit these viruses, but important ones include
A. pisum, Aphis fabae, A. craccivora and Myzus persicae.
Bean yellow mosaic

The disease is caused by Bean yellow mosaic virus (BYMV) and is transmitted
through aphids A. pisum, A. fabae, A. craccivora and M. persicae.
Broad bean stain

The disease is caused by Broad bean stain virus (BBSV). Vectors of this virus
are weevils Apion vorax and Sitona spp.
Pea seed-borne mosaic

The disease is caused by the Pea seed-borne mosaic virus (PSbMV). Transmission of the virus occurs by aphids A. pisum, A. craccivora and A. fabae.
Cucumber mosaic
The disease is caused by Cucumber mosaic virus (CMV) and is transmitted by
several species of aphids.
Other viruses
Other viruses which cause diseases in lentil include Bean leaf roll virus
(BLRV), Faba bean necrotic yellows virus (FBNYV), Soybean dwarf virus (SBDV)
and Lucerne mosaic virus. They are also transmitted by aphids (Makkouk
and Kumari, 2001; Jones and Coutts, 2008).
18.5. Pest Management

Lentil is grown in diverse environments under varying levels of pest pressure around the world. Very few insects have attained the status of economic pest of the crop. Low temperatures during the crop growth period,
290
short duration of crop growth, small leaf area and relatively small pod size
may be some of the reasons for low pest attack. However, certain situations
such as late planting increase the vulnerability to pest attack. Therefore,
successful cultivation of the crop should include effective management
through forecasting, monitoring and modelling of the economic insect
fauna.
Monitoring methods
Monitoring and forecasting systems for lentil are in their infancy. The use
of pheromone and sticky traps is becoming popular among growers.
The following examples will illustrate some of the different monitoring
programmes developed for a few important pests.
Aphids
Insecticide treatment for pea aphid control should be considered when an
economic threshold of 3040 aphids are collected per 180 sweep of a 38 cm
(15 inch) diameter insect net, when few natural enemies are present, and
when aphid numbers do not decline over a 2-day period (Homan et al., 1991).
An aphid-tracking network was recently developed for the Washington
State region of the USA.
Lygus bugs
Adult lygus-bug activity can be monitored during blooming and podding
by making 25 180 sweeps in at least five randomly selected places in a
field. Chemical control is applied when 710 adults are collected per 25
sweeps (OKeeffe et al., 1991).
Pod borer, Helicoverpa
Adult male populations can be monitored by pheromone-baited traps to
understand the onset of infestation and to study population dynamics. Lentils are very susceptible to native budworm damage from early podding to
pod fill peak moth flights coincide with the flowering to pod filling stage
of late-sown lentil. The crop should be monitored from late flowering and
sprayed when the economic threshold reaches >1 larva in 10 sweeps (one
larva/m2).
Etiella
Management is based on timing of sprays to target adult moths. Management is facilitated by downloadable Etiella DegreeDay models to identify
onset of flight activity within the crop. Monitoring should continue at 45
day intervals, using sweep nets, light or pheromone traps. Action is recommended for a threshold of 12 Etiella per 20 sweeps.
291
Cultural control
Cultural control includes some of the oldest pest control practices known,
such as manuring for vigorous growth, crop rotation, manipulation of
sowing date, mixed cropping and field sanitation, etc. Such practices may
be helpful in overall reduction of the population of harmful insects in the
crop. The sowing date may be manipulated in order to ensure that a susceptible stage does not coincide with peak pest numbers. However, different pests respond to time of sowing in different ways, and it may be
difficult to design a programme that offers protection against all potential pests. Early or timely sowing ensures that the crop escapes the damage by pod borers such as H. armigera in India. Thus lentil sown from
mid-October to mid-November is almost free from H. armigera whereas
late (December) sowing increases the risk of pod borer and leaf miner
infestations.
Certain weeds can act as hosts for lentil pests and such species should
be eliminated from the field. Since the suitability of the plant host depends
not only on the species, but also on its growth stage, generalization about
the suitability of the alternative host should be made with care. Vicia spp.,
Melilotus spp. and Chenopodium are among the hosts of H. armigera, leaf
miner and bean aphid. Intercropping is likely to find a place in reducing
pest problems. A few examples are available from India, where intercropping with mustard, coriander and linseed may lead to increase in spider
and braconid parasitoids.
Host plant resistance

Very little research has been conducted to identify resistant varieties
against various lentil pests. The use of resistant cultivars in an integrated pest management (IPM) programme depends on the efficacy of
other pest control options available. In a review of research achievements for plant resistance to insect pests of cool-season food legumes
by Clement et al. (1994), the consensus view was that insect pest resistance research and breeding was undervalued and underfunded. The
state of affairs has changed little since then, and although there has
been some notable progress in the identification of new sources of resistance, there has been less progress in resistant variety development. The
situation is probably in part influenced by economics and a lack of
resources, as well as the intricate nature of the plant-insect interrelationship.
Helicoverpa armigera
Six out of 18 cultivars (L 1282, LL 116, L 830, LL 78, LG 11 and HPL 5)
consistently showed low pod-borer attack compared to check variety Pant
L 639 and Pant L 406 in India (Ujagir, 1993).
292
Etiella zinckenella
Eleven genotypes out of 79 were categorized as least susceptible to the
infestation of E. zinckenella (Jaglan et al., 1993). Tolerance to E. zinckenella
was identified in lentil line LL 147 (Brar et al., 1989). However, there have
been no recorded attempts to incorporate these lines into breeding programmes (Erskine et al., 1994).
Bruchids
Singh and Sharma (2001) screened seven cultivars of lentil for their response
to the oviposition and development of the pulse beetle. Cultivar K-75 had
the minimum growth index and slowest developmental period of the insect.
In another study, lentil varieties 79-1 and Precoz were found resistant
against Callosobruchus analis (Shafique and Ahmad, 2002).
Ecological resistance to B. lentis under field conditions has been discussed by Clement et al. (1994). Genetic differences in L. culinaris genotypes
have been found against B. lentis (Chopra and Pajni, 1987), although no
attempt in resistance breeding has been undertaken (Erskine et al., 1994).
Sitona crinitus
Host plant resistance has recently become a potential option in the control
of S. crinitus. El-Bouhssini et al. (2008) identified resistance to S. crinitus in
wild species of lentil originating from Turkey, Syria and Croatia: Lens nigricans, L. culinaris ssp. odemensis, Lens ervoides and L. culinaris ssp. orientalis.
One of the resistant accessions, ILWL245 of L. culinaris ssp. orientalis is crossable with the cultigen, and the goal is now to transfer resistance to cultivated lentil, as well as to study the inheritance of resistance (El-Bouhssini
et al., 2008). Sedivy (1972) tested several lentil cultivars against Sitona for
feeding damage and found differences in the amount of damage. Weigand
and Pimbert (1993) recommend feeding damage as the scoring method of
choice in host-resistance screening for Sitona and described a laboratorybased screening protocol for this. The Yerli Kirmizi variety of lentil was
reported to be resistant against S. crinitus (Erman et al., 2005).
Aphids
Weigand and Pimbert (1993) reported that there were genotypic differences
between lentil genotypes in resistance screening for A. craccivora, using a
technique developed for screening in faba bean. Mechanisms of resistance
to A. pisum in lentil genotypes have been investigated by Andarge and
Westhuizen (2004). Genetic differences were observed in lentil germplasm
under natural infestation with A. craccivora and Acyrthosiphon kondoi in New
Zealand (Erskine et al., 1994). Muehlbauer and Kaiser (1994) reported that
resistance to aphids (and weevils) in lentil is incomplete and polygenic. To
date there are no reports of resistant varieties released to farmers from this
genetic material.
293
Emerging technologies
Over the past two decades the genomics revolution has enabled rapid
progress in functional genetics and proteomics of legume crops including
lentil. There has been some progress in our understanding of lentil-insect
interactions using these rapidly evolving technologies, which, in time,
should lead to practical crop protection applications. For a recent report
on the progress and applications of molecular research in lentil itself, see
Ford et al. (Chapter 11, this volume) and Muehlbauer et al. (2006).
Genomics and plant defence
The secondary plant compounds lectins, proteinase inhibitors (PIs) and
alpha-amylase inhibitors (AAIs) play a major role in plant defence and are
particularly well featured within the legume family (Ryan, 1990; Chrispeels
and Raikhel, 1991). Lentil PIs (trypsin and chymotrypsin inhibition sites)
have been used to investigate host-pathogen co-evolution (Sonnante et al.,
2005). There is a common functionality among many of these defence gene
families. Using functional and comparative genomics approaches, macrosyntenic relationships are being explored in model and crop legumes
including lentil (Zhu et al., 2002). For example, lectin genes on linkage group
VII in field pea have homologous regions in lentil. PIs and lectins from different plant origins have an inhibitory effect on the adult lucerne weevil,
Hypera postica (Elden, 2000). Comparative genomics provides an opportunity to fast-track lentil improvement by interpreting the information
obtained from equivalent genome regions in related legumes. This approach
is likely to target generalist insect pests; finding syntenic regions for specialist insect pests of lentil might be a greater challenge.
The following insect pests of Medicago truncatula may be candidates
for gene synteny comparisons with lentil: blue-green aphid (Acyrthosiphon
kondoi), pea aphid (A. pisum), cowpea aphid (A. craccivora), red-legged earth
mite (Halytodeus destructor), lucerne flea (Sminthurus viridus) and Sitona weevils (Sitona discoideus, Sitona humeralis).
Genes have been identified that control phytoalexins, chitinases, glucanases and lipoxigenases (the LOX genes). The jasmonic acid (JA) pathway plays a major role in inducing resistance to a broad spectrum of
insects. The omics revolution is providing us with increasing evidence
that many legume-defence mechanisms are conditioned by resistance gene
analogues (RGAs), damage response regions (DRR), flavonoids, plant
defence-related genes (e.g. str246N), gibberellin c-20 oxidase, maysin synthesis, glycosylflavone and NADH dehydrogenase, which may facilitate
new approaches to crop protection and improvement (Young et al., 2005;
Howe and Jander, 2008).
Genetic transformation
The Bacillus thuringiensis gene that controls insecticidal Delta endotoxin
production has been introduced into rhizobial strains a model that has
294
potential to control pea leaf weevil if introduced into its lentil host
(Erskine et al., 1994).
Mutation breeding
Mutation breeding may have potential for inducing monogenic resistance
to insects in lentil; however, it is likely to pose a major challenge for polygenic resistances underlying complex plant-insect interactions. Induced
mutations seem to be undergoing resurgence for a global perspective on
mutation-derived crop varieties see Ahloowalia et al. (2004).
In-vitro hybridization
Embryo rescue techniques have been successful in overcoming hybridization barriers to obtain hybrids between cultivated lentil and L. nigricans ssp.
ervoides and L. nigricans ssp. nigricans (Muehlbauer et al., 1994).
Biological control
The role of natural enemies has been studied for a number of insect pests,
and in several cases they have been shown to play a significant role, reducing the need for pesticide application or other control measures.
Aphids
Aphids have many natural enemies, including ladybird beetles, parasitic
wasps, lacewings and syrphid flies (Homan et al., 1991). Stark et al. (2004)
reported Coccinella septumpunctata (L.), predator and Diaertiella rapae (McIntosh), parasitoid of pea aphid Acrythosiphon pisum. The feeding potential of
spiders on A. craccivora was studied by Sebastian and Sudhikumar (2003) and
the spider species Lycosa poonaensis (Tikader and Malhotra) and Lycosa tista
(Tikader) consumed the highest number of prey. A predator complex including C. septumpunctata, Micraspis discolor, Menochilus sexmaculatus and Coccinella
transversalis were found feeding on A. craccivora by Sharma et al. (1991).
Sharma and Yadav (1993) reported the effectiveness of aphidophagous coccinellids in controlling the population of A. craccivora. They worked out the
possibilities of ecologically sound aphid management in lentil by exploiting
the insect-plant relationship against the pest in favour of its natural enemies.
Damage caused by the pea aphid, A. pisum, is a limiting factor in lentil
production in several countries. Andarge (2001) worked on the multifaceted
approach in order to keep A. pisum below economic threshold. Beauveria
bassiana was found to be significantly effective in reducing the population
of A. pisum compared to the control. The botanical product Neemolin also
proved to be highly efficient in reducing the bean aphid population.
Grasshopper
Dysart (1995) reported that egg pods of grasshopper species were parasitized by four species of scelionid wasps, including Scelio opacus, which was
295
the dominant parasitoid species, accounting for 92% of all parasitized pods
of lentil. Songa and Holliday (1997) reported the high feeding potential of
adult carabid beetles, Pterostichus corvus (Leconte) and Pterostichus femoralis
(Kirby) on the eggs of M. bivittatus.
Seed beetle, B. lentis
Isidoro et al. (2001) reported Uscana spp. as an egg parasitoid of B. lentis
with variable efficacy (121%). Pupal and larval parasitoids were also
observed.
Pod borer, E. zinckenella
Marwoto (2003) reported Trichogrammatoidea bactrae-bactrae as an effective
biocontrol agent for Etiella spp. Other important natural enemies include
Macrocentrus ancylovorus, Bracon piger, Apanteles beaussetensis, Bracon pectoralis, Phanerotoma planifrons and Cyrtotyx lichtensteini.
Cutworm, A. ipsilon
Among the wasps known to attack this cutworm are Apanteles marginiventris
(Cresson), Microplitis feltiae Muesebeck, Microplitis kewleyi Muesebeck, Meteorus autographae Muesebeck, Meterorus leviventris (Wesmael) (Hymenoptera:
Braconidae); Campoletis argentifrons (Cresson), Campoletis avicincta (Ashmead), Hyposoter annulipes (Cresson) and Ophion avidus Brulle (Hymenoptera:
Ichneumonidae). The cutworm larvae consume about 24% less foliage and
cut about 36% fewer seedlings when parasitized by M. leviventris. Thus considerable benefit is derived from parasitism in addition to the eventual death
of the host larvae. Other parasitoids known from black cutworm include flies
often associated with other ground-dwelling noctuids, including Archytas cirphis Curran, Bonnetia comta (Fallen), Carcelia formosa (Aldrich and Webber),
Chaetogaedia monticola (Bigot), Eucelatoria armigera (Coquillett), Euphorocera
claripennis (Macquart), Gonia longipulvilli Tothill, Gonia sequax Williston, Lespesia archippivora (Riley), Madremyia saundersii (Williston), Sisyropa eudryae
(Townsend) and Tachinomyia panaetius (Walker) (Diptera: Tachinidae) (University of Florida, 2008).
Leaf miners, Liriomyza trifolii and C. horticola
According to previous records, all natural enemies of agromyzids are
Hymenopterans of Chalcidoidea, Ichneumonidea and Cynipodea. Among
these, chalcid parasitoids are reported to constitute the most dominant
group (Murphy and LaSalle, 1999). Some of the other potential parasitoids
recorded are given in Table 18.2.
Pod borer, H. armigera
Over 78 species of arthropod parasitoids and 33 species of predators have
been reported to directly prey on a specific life stage of H. armigera within
296

Table 18.2.
Parasitoid species and their hosts (Source: Ltye, 2004).
Parasitoids
Pest
Diglyphus iseae
Diglyphus chabrias
Neochrysocharis arvensis
Neochrysocharis formosa
Pediobius acantha
Chrysocharis sp.
Hemiptarsenus sp.
L. trifolii and C. horticola

L. trifolii
Unknown species
Unknown species
L. trifolii
India (Manjunath et al., 1989). Similarly, a number of natural enemies have

been reported from Pakistan, South-east Asia, China, Australia, eastern and
southern Africa and Western Europe. Recently, Sithanantham et al. (2005)
prepared a list of about 134 parasitoids and 82 predators of eggs, larvae and
pupae reported from India and elsewhere. The most important orders are
Coleoptera, Hemiptera, Hymenoptera, Neuroptera, Orthoptera and Araneida. Among these, Trichogramma on eggs, Campoletis on larvae, and predatory birds are the most important biotic agents of H. armigera in northern
India.
Viruses, bacteria, protozoa, fungi and nematodes are important pathogens which cause diseases in H. armigera. The success achieved in chickpea
with commercial formulation of B. thuringiensis and nuclear polyhedrosis
viruses may be extended to lentil to reduce crop damage and increase grain
yield where H. armigera is a serious problem.
Chemical control
Due to ready availability, effectiveness and rapid control of pest populations, insecticides are popular among farmers. Sufficient numbers of selective pesticides are available to treat target pests without causing damage to
the predatory insects.
Sitona weevil
Effective control of Sitona by spraying with chlorinated pinene, trichlorphon
(Khlorofos), carbaryl (Sevin) and dimethoate (Bi-58) and dusting with methyl
parathion (Metafos) was reported by Petrukha (1970). Carbofuran and aldicarb were effective in reducing the damage caused by Sitona spp. (Solh et al.,
1986). Erman et al. (2005) tested different insecticides against S. crinitus on
the lentil crop and reported the application of oxydemeton methyl to be
most effective in decreasing the nodule damage caused by the weevil.
Weigand et al. (1994) reported that lentil seed treated with furathiocarb
(Promet) provided good control against Sitona spp. In a study in Syria, carbofuran effectively controlled Sitona spp. (Hariri and Tahhan, 1983).
297
Helicoverpa spp.
Memon and Memon (2005) worked on the comparative efficacy of four different insecticides against the pod borer (Helicoverpa spp.) on the lentil crop.
Spinosad was the most effective insecticide in reducing the pod borer population and decreasing pod damage, followed by chlorpyrifos and endosulfan. The maximum increase in seed yield per hectare was obtained with
spinosad. Barroga and Barrers (1969) reported that dimethoate, phosphamidon, methyl parathion, monocrotophos, promecarb, bromophos and naled
were effective in reducing the damage caused by Heliothis sp.
At Pantnagar (India), endosulfan, monocrotophos, quinalphos and triazophos, besides synthetic pyrethroids fenvalerate, decamethrin and cypermethrin gave effective control of pod-borer damage during the period
19791981 (Sehgal and Ujagir, 1980, 1982).
Tahhan and Hariri (1982) reported that the insecticides deltamethrin
and methidathion were effective in the control of H. armigera on lentil.
The effect of crude PI extracts from seeds of lentil and other pulse crops
on the insecticidal activity of B. thuringiensis var. kurstaki HD-1 against neonate larvae of H. armigera by the diet incorporation method was investigated
by Gujar (2004). A mixture of B. thuringiensis var. kurstaki and PIs of lentil
varieties ILL 8095 and L 4626 was found to act synergistically in controlling
H. armigera.
Aphids
Sharma et al. (1991) reported cypermethrin and endosulfan to be effective
against A. craccivora. Aphids transmitting viruses could be reduced by seed
treatment with imidacloprid (Makkouk and Kumari, 2001). Systemic insecticides also control aphids on lentil (Thakur et al., 1984).
Leaf miner
Several pesticides administered separately as well as in a combination of
sprays, dust and granular formulations have been found effective in controlling the damage by leaf miner (Atwal et al., 1969; Vyas and Saxena, 1982).
The application of phosphamidon, dimethoate, monocrotophos, endosulfan,
methyl demeton and dicrotophos effectively controlled this pest.
Bruchus lentis
Lambdacyhalothrin and endosulfan applied at the flowering stage of a lentil crop provide control of B. lentis.
Lygus bug
In Turkey, lygus bug has been indirectly controlled by aerial insecticide that
was applied to a cereal crop infested with sunn pest (Eurogaster integreceps L.)
(zberk et al., 2006). However, the government programme was suspended
leading to an increase in the population and damage cause by lygus bug.
298
Etiella
Application of deltamethrin and esfenvalerate to target adult moths prior to
egg lay were found to be effective in containing Etiella within the acceptable
industry threshold of <1% insect-damaged grain in 2004 and 2005 trials in
Australia.
Insecticide resistance
Cytochrome P450 DNA and mRNA expression has been investigated in different strains of lygus bugs that had different susceptibility to pyrethroid
insecticides (Zhu and Snodgrass, 2003). Annotated expressed sequence tags
(ESTs) and xenobiotic detoxifications have also been studied in the aphid
Myzus persicae (Sulzer) (Figueroa et al., 2007). There is scope for future
research into understanding gene regulation and biochemical pathways of
P450s and how they influence insecticide susceptibility, leading to protocols
that may reduce the risk of insecticide resistance developing.
18.6. Concluding Remarks

IPM includes effective use of various compatible pest control measures to
reduce pest populations below damaging levels and optimize yield with
minimum damage to the environment. This review clearly shows that considerable knowledge already exists about pest management systems in lentil to initiate formulation of IPM strategies. Modification to IPM approaches
can be made later on as our knowledge on host plant resistance, economic
threshold level (ETL) and economic injury level (EIL) for different pests in
the agroecosystem of the locality increases. Thus, based on the results and
experience gained so far in the lentil crop, the following agenda for IPM is
suggested.
Information on crop-loss assessment by major pests of lentil crop in different agroecological zones needs to be fully documented to prioritize our
research on pest management and host plant resistance studies on the key
pests in each area. For this, a survey needs to be planned, coupled with
methodology combining systematic sampling with statistics. Climate change
is likely to play a role in decision making in future IPM strategies, along
with greater adoption of novel technologies, over the coming years.
Manipulation of the date of sowing, intercropping and crop rotation
have given significant reduction in populations of pod borers, foliage feeders and soil pests. Promoting the right kind of cropping systems, by raising
companion and/or barrier crops such as coriander, mustard and linseed,
will certainly minimize the population of Helicoverpa and Etiella and other
pests.
Systematic studies on host plant resistance against key endemic pests
and the biochemical/biophysical basis for such resistance can make rapid
299
progress in the development of insect-resistant cultivars. Development of

multiple resistances or at least moderate resistance against key pests of the
area should be the research priorities. In recent years, there have been
reports of resistant germplasm to many key pests of lentil. For example,
Rembold and Schroth (1993) reviewed research achievements in the
improvement of resistance in lentil and other food legumes against a wide
range of insect pests, however, the factors responsible for this resistance are
yet to be identified. Sequencing of genomes in model legumes has led to
progress in our understanding of the underlying genetics and biochemistry
of lentil-insect interactions (Sato et al., 2007). Shared synteny of genes on
chromosomes with model legumes should lead to a better insight into
how lentil genes function in response to insect pests. Using molecular
technology, plant resistance (R) genes have been isolated and transferred to
susceptible varieties. Transferable molecular markers, such as microsatellites (simple sequence repeats; SSRs), have great potential in marker-assisted
selection for resistance, but adoption in lentil breeding programmes is
not yet available. It must be noted that so far, effective insect resistance has
not been obtained, and only a few durable R genes have been isolated.
These too have limitations for their applications across taxonomic barriers
(ICAR, 2007).
Certain biocontrol agents such as Chrysops spp., Campoletis spp., Trichogramma spp., Bacillus thuringiensis and nuclear polyhedrosis virus (NPV) are
suitable candidates for improving the biological control of some of these
pests. More potential strains of NPV, bacteria and other insect pathogens
need to be identified from the areas where they are virulent.
Research should be intensified into finding effective plant-derivative
pesticides that are within the reach of farmers in rural areas. This will
require identification of more such plants, evaluation of their extracts for
insecticidal properties and application methodology.
Studies on proper sampling methodology and ETLs and EILs of key
endemic pests of the lentil crop are needed to generate databases for judicious need-based application of ecofriendly chemical insecticides so as to
minimize risks of insecticide resistance and pest resurgence. Even with the
present level of knowledge, IPM can significantly reduce chemical pesticide
inputs. Governments, industry and funding bodies will need to invest in
research programmes that encompass these technical challenges, and
through education and training of key field and scientific personnel get
through this bottleneck.
References
Ahloowalia, B.S., Maluszynski, M. and Nichterlein, K. (2004) Global impact of
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19
Virus Diseases and their Control
Safaa G. Kumari,1 Richard Larsen,2 Khaled M. Makkouk1 and

Muhammad Bashir3
1International
Center for Agricultural Research in the Dry Areas (ICARDA), Aleppo,

Syria; 2United States Department of Agriculture (USDA) Agriculture Research Service
(ARS), Prosser, Washington, USA; 3Crop Diseases Research Institute, National
Agricultural Research Centre (NARC), Islamabad, Pakistan
19.1. Introduction
Biotic and abiotic factors limit yields and cause yield instability of lentil.
Many of the diseases that affect lentil, especially those induced by
viruses, can also infect other food and forage legumes. The relative
importance of virus diseases varies depending on geographical location
and agroecological conditions. Yield losses from virus infection vary
from little or none to complete crop failure depending on the time and
severity of infection.
Worldwide, lentils are susceptible to at least 30 different virus species
representing 16 genera from nine families with genomes comprised of
single-stranded RNA or DNA (Table 19.1) (Bos et al., 1988; Fonseca et al.,
1995; Makkouk et al., 2003a; Abraham et al., 2006; Taylor et al., 2007). Among
the most important viruses that infect lentil are Bean leafroll virus (BLRV),
Bean yellow mosaic virus (BYMV), Beet western yellows virus (BWYV), Cucumber mosaic virus (CMV), Faba bean necrotic yellows virus (FBNYV), Pea enation
mosaic virus-1 (PEMV-1), Pea seed-borne mosaic virus (PSbMV) and Pea streak
virus (PeSV).
In this chapter, we review transmission, ecology and epidemiology
of the economically most important viruses that infect lentil. We also
describe sensitive assays available for detection and appropriate measures for control.
19.2. Viruses Reported to Infect Lentil

Large-scale surveys carried out during the last two decades have shown
that the most widespread viruses in lentil include six that cause yellowing/
306

307
Table 19.1. Virus species reported to infect lentil by natural and/or articial inoculation. Virus
taxonomy adopted in this table is based on the most recent ICTV report (Source: Fauquet
et al., 2005).
Mode of
transmission
Virus species
Genomea
Abbreviation
Genus
Family
Vectorsb
Alfalfa mosaic virus

Artichoke latent virus
Bean common mosaic
virus
Bean leafroll virus
Bean pod mottle virus
Bean yellow mosaic
virus
Beet western
yellows virus
Broad bean mottle virus
Broad bean stain virus
Broad bean true
mosaic virus
Broad bean wilt virus-1
Chickpea chlorotic dwarf
virus
Chickpea chlorotic stunt
virus
Chickpea liform virus
Chino del tomate virus
Cucumber mosaic virus
Faba bean necrotic
yellows virus
Pea enation mosaic
virus-1
Pea seed-borne
mosaic virus
Pea streak virus
Peanut stunt virus
Quail pea mosaic virus
Red clover vein
mosaic virus
Soybean dwarf virus
Subterranean clover
stunt virus
Tobacco streak virus
Tomato black ring virus
Tomato spotted wilt virus
Tomato yellow leaf
curl virus
Turnip mosaic virus
ssRNA
ssRNA
ssRNA
AMV
ArLV
BCMV
Alfamovirus
Potyvirus
Potyvirus
Bromoviridae
Potyviridae
Potyviridae
Aphids NP
Aphids NP
Aphids NP
+
+
+
ssRNA
ssRNA
ssRNA
BLRV
BPMV
BYMV
Luteovirus
Comovirus
Potyvirus
Luteoviridae
Comoviridae
Potyviridae
Aphids P
Beetles
Aphids NP
+
+
ssRNA
BWYV
Polerovirus
Luteoviridae
Aphids P
ssRNA
ssRNA
ssRNA
BBMV
BBSV
BBTMV
Bromovirus
Comovirus
Comovirus
Bromoviridae
Comoviridae
Comoviridae
Beetles
Beetles
Beetles
+
+
+
ssRNA
ssDNA
BBWV-1
CpCDV
Fabavirus
Mastrevirus
Comoviridae
Geminiviridae
Aphids NP
Leafhoppers
ssRNA
CpCSV
Polerovirus
Luteoviridae
Aphids P
ssRNA
ssDNA
ssRNA
ssDNA
CpFV
CdTV
CMV
FBNYV
Potyvirus
Begomovirus
Cucumovirus
Nanovirus
Potyviridae
Geminiviridae
Bromoviridae
Nanoviridae
Aphids NP
Whiteies
Aphids NP
Aphids P
ssRNA
PEMV-1
Enamovirus
Luteoviridae
Aphids P
ssRNA
PSbMV
Potyvirus
Potyviridae
Aphids NP
ssRNA
ssRNA
ssRNA
ssRNA
PeSV
PSV
QPMV
RCVMV
Carlavirus
Cucumovirus
Comovirus
Carlavirus
Flexiviridae
Bromoviridae
Comoviridae
Flexiviridae
Aphids NP
Aphids NP
Beetles
Aphids
+
+
+
+
ssRNA
ssDNA
SbDV
SCSV
Luteovirus
Nanovirus
Luteoviridae
Nanoviridae
Aphids P
Aphids P
ssRNA
ssRNA
ssRNA
ssDNA
TSV
TBRV
TSWV
TYLCV
Ilarvirus
Nepovirus
Tospovirus
Begomovirus
Bromoviridae
Comoviridae
Bunyaviridae
Geminiviridae
Thrips
Nematodes
Thrips
Whiteies
+
+
+
ssRNA
TuMV
Potyvirus
Potyviridae
Aphids NP
a ss,
single stranded.
NP, aphids in non-persistent manner; Aphids P, aphids in persistent manner.
b Aphids
Sap
308
S.G. Kumari et al.
Table 19.2. Viruses of regional or global economic importance reported to naturally infect
lentil (Source: Bos et al., 1988; Makkouk et al., 2003a).
Geographical distribution
Virus species
AMV
BCMV
BLRV
BYMV
BWYV
BBWV-1
BBMV
BBSV
CpCDV
CpCSV
CMV
FBNYV
PEMV-1
PSbMV
PeSV
SbDV
TSWV
America
Europe
Africa
Asia
Australasia
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
stunting/necrosis and approximately ten viruses that cause mosaic or mottling symptoms (Table 19.2) (Makkouk et al., 2003a).
Viruses causing yellowing and stunting symptoms are the most important worldwide, and include BLRV, BWYV, FBNYV, Chickpea chlorotic dwarf
virus (CpCDV) and Soybean dwarf virus (SbDV) (= Subterranean clover red leaf
virus; SCRLV). These viruses induce symptoms of leaf rolling, reddening,
yellowing and stunting (Plate 3A, B, C) and plants infected at early growth
stages usually exhibit shortened internodes and pronounced proliferation
of axillary shoots. These viruses are phloem-limited and transmitted by
aphids except CpCDV which is always transmitted by leafhoppers. They are
not transmitted mechanically or by seed and can have a marked effect on
yield.
The most important viruses affecting lentil causing mosaic and mottling
(Plate 3D, E) are: PSbMV, CMV, PEMV-1, PeSV, BYMV, Alfalfa mosaic virus
(AMV) and Broad bean stain virus (BBSV). Symptoms caused by these viruses
are often confused with those resulting from nutrient deficiency, physiological disorders, herbicide damage or waterlogging.
Several viruses impair the quality of lentil seeds thereby rendering them
less attractive to consumers. For example, BBSV infection leads to undesirable staining of lentil seedcoats, which makes the grain unsuitable for marketing (Plate 3F).
309
Viruses causing yellowing and stunting symptoms

Bean leafroll virus (BLRV)
Lentil plants infected naturally with BLRV were first observed in Iran by Kaiser
et al. (1968), and it is known to be among the most important viruses that infect
lentil. The virus is reported to occur on lentil in Bangladesh, Ethiopia, Iran, Iraq,
Syria, Tunisia and the USA (Kaiser, 1973; Klein et al., 1991; Bakr, 1993; Tadesse
et al., 1999; El-Muadhidi et al., 2001; Kumari, 2002; Makkouk et al., 2003b).
A yield reduction of 91% has been reported when plants were infected
with BLRV at the pre-flowering stage, while at the flowering stage infection
may result in only 50% yield loss (Kaiser et al., 1968). In Washington, USA,
up to 80% disease incidence was noted in some fields during an epidemic
year (Klein et al., 1991).
Infection with BLRV is restricted to legume species including faba bean,
lentil, pea and chickpea in many parts of the world (Bos et al., 1988; Makkouk et al., 2003a). The main symptoms produced by the virus are interveinal
chlorosis, yellowing, stunting, leaf rolling, reddening and thickening of the
leaves and suppression of flowering and pod set. Entire leaflets can turn
red in cool temperatures.
BLRV is transmitted only by aphids in a persistent, non-propagative
manner. Acyrthosiphon pisum Harris, Aphis craccivora Koch., Aphis fabae (Scopoli) and Myzus persicae (Sulz.) are the most common aphid species reported
to transmit BLRV (Kaiser, 1973; Cockbain, 1983).
Beet western yellows virus (BWYV)
BWYV was first isolated by Duffus (1961) in California and is synonymous
with Malva yellows virus and turnip mild yellows virus. BWYV occurs on
lentil in Ethiopia, Iran, New Zealand, Pakistan, Syria and Turkey (Fletcher,
1993; Tadesse et al., 1999; Makkouk et al., 2001a, 2003b; Kumari, 2002).
BWYV is not restricted to leguminous species as in the case of other
legume luteoviruses (e.g. BLRV). It infects many crop and weed species,
which belong to the families Brassicaceae and Compositae in addition to
Fabaceae. Symptoms of BWYV on lentil include yellowing or bronzing with
leaflets becoming leathery and brittle. Usually, leaf margins become reddish, but the entire leaflets may turn red under cool conditions. Plants are
severely stunted or may die if infected early. Internodes become shortened
in the upper plant parts infected at a late growth stage. Pods in the infected
plants are reduced in size and poorly filled or may remain seedless.
BWYV is aphid transmitted in a persistent, non-propagative manner.
The main aphid vectors are M. persicae, A. craccivora, A. pisum and Aulacorthum solani (Kltb.) (Boswell and Gibbs, 1983; Cockbain, 1983).
Chickpea chlorotic dwarf virus (CpCDV)
CpCDV was first reported on chickpea in India (Horn et al., 1993), and later
in Egypt, Ethiopia, Iran, Iraq, Pakistan, Sudan, Syria and Yemen (Makkouk
310
S.G. Kumari et al.
et al., 2003a). It occurs on lentil in Pakistan and Iran (Makkouk et al., 2001a,
2003b). Crop loss is proportional to virus incidence in the field, as CpCDVinfected plants produce little or no grain.
The virus is reported to naturally infect chickpea, lentil, faba bean (Makkouk et al., 2003a), sugarbeet and bean in Iran (Farzadfar et al., 2002), and
Phaseolus bean and other wild species in Sudan (Ali et al., 2004). The main
symptoms of CpCDV are reddening or yellowing of leaflets with severe
stunting.
CpCDV is transmitted by leafhopper in a circulative non-propagative
manner, such as Orosius orientalis (Matsumura) in India (Horn et al., 1993)
and Orosius albicinctus Distant in Syria (Kumari et al., 2004).
Faba bean necrotic yellows virus (FBNYV)
FBNYV was first reported from faba bean near Lattakia, Syria (Katul et al.,
1993), and was later reported on lentil in Ethiopia, Iran, Iraq, Pakistan, Syria
and Turkey (Makkouk et al., 1992, 2001a, 2003b; Bayaa et al., 1998; Tadesse
et al., 1999; El-Muadhidi et al., 2001). In epidemic years the virus can be very
damaging, as observed in Middle Egypt during 1992, 1997 and 1998, and in
Tunisia in 2001. In Middle Egypt, losses due to FBNYV infection on faba
bean during 1992 reached 8090% (Makkouk et al., 1994). In West Asia and
North Africa, FBNYV is considered the most damaging virus on food
legume crops.
FBNYV is reported to naturally infect faba bean, pea, chickpea, lentil,
cowpea, Egyptian clover, French bean and many forage legume species
(Katul et al., 1993; Makkouk et al., 1998, 2003a). Leaves of infected plants
become thick and brittle and show interveinal chlorotic blotches starting
from the leaf margins. The uppermost young leaves remain very small and
cupped upwards, whereas the older leaves are rolled downwards. New
shoots, leaves and flowers develop poorly. Approximately 34 weeks after
infection is established, interveinal chlorosis becomes necrotic and plants
then die within 57 weeks. The host range of FBNYV is restricted to the
family Fabaceae.
Three aphid species, A. pisum, A. craccivora and A. fabae are the reported
vectors of FBNYV in a circulative non-propagative manner (Franz et al.,
1998; Makkouk et al., 1998).
Viruses causing mosaic and mottling

Bean yellow mosaic virus (BYMV)
BYMV was reported in lentil by Kaiser et al. (1968) in Iran, with high reductions in yield (Kaiser, 1973). The virus occurs on lentil in Bangladesh, Egypt,
Ethiopia, Iran, Iraq, Jordan, New Zealand, Syria and Turkey (Kaiser, 1973;
Russo et al., 1981; Makkouk et al., 1992, 1993, 2003b; Bakr, 1993; Fletcher,
1993; Bayaa et al., 1998; Tadesse et al., 1999; Al-Mabrouk and Mansour, 2000;
El-Muadhidi et al., 2001). BYMV is reported to occur on several other legumes
311
including pea, faba bean and chickpea in North America, Europe, Africa,
Asia and Australasia, as well as a few non-legume hosts (Bos et al., 1988;
Makkouk et al., 2003a).
The effect of BYMV on yield depends, to a great extent, on the time of
infection and virus strain. In a field experiment in Syria, sap inoculation of
lentil with BYMV at pre-flowering and flowering stages led to 96 and 34%
yield reductions, with seed transmission rates of 0.83 and 0.3%, respectively
(Kumari et al., 1994).
BYMV symptoms on lentil include chlorosis, mild mosaic or mottling,
and stunting. Leaves often become twisted or curled with necrosis along
the margins. Flowering and pod formation is reduced as a result of infection and consequently little seed is produced.
BYMV is transmitted mechanically and by aphids in a non-persistent
manner. Acyrthosiphon pisum, A. fabae, A. craccivora, M. persicae and Macrosiphum euphorbiae (Thomas) are among the most common aphid vectors
(Cockbain, 1983; Edwardson and Christie, 1986).
Broad bean stain virus (BBSV)
Lentil has been reported to be infected by BBSV under natural conditions
(Bos et al., 1988; Makkouk et al., 2003a). In addition to lentil, the virus occurs
naturally on pea, faba bean and several other legumes in Europe, Africa,
Asia and Australasia. It was reported also to occur on lentils in Ethiopia,
Iran, Jordan, Syria and Turkey (Makkouk et al., 1987, 1992, 2003b; Bayaa
et al., 1998; Tadesse et al., 1999; Al-Mabrouk and Mansour, 2000).
In lentils, very mild mottling occurs in response to BBSV infection
which is often not readily observable. Grain yield losses varied from 14% to
61% and the seed transmission rates of BBSV were found to range from
0.2% to 32.4% when 19 lentil genotypes were inoculated at the flowering
stage (Makkouk and Kumari, 1990). Infection of lentil plants at pre-flowering,
flowering and pod stages resulted in seed transmission rates of 20.6, 19.1
and 1.5%, respectively (Kumari et al., 1993). BBSV also has been detected in
growers seed in Turkey (Fidan and Yorganci, 1990).
BBSV is transmitted primarily by beetle species Apion aestivum Germ.,
Apion arrogans Wencher, Sitona crinita Herbst, Sitona limosa Rossi and Sitona
lineatus L. (Cockbain et al., 1975; Makkouk and Kumari, 1989, 1995).
Cucumber mosaic virus (CMV)
CMV was first recorded to infect lentil in Iran (Kaiser et al., 1968). The virus
naturally infects lentil in Australia, Ethiopia, India, Iran, Nepal, New Zealand, Pakistan and Syria (Kaiser, 1973; Rangaraju and Chenulu, 1981; Fletcher,
1993; Karki, 1993; Mouhanna et al., 1994; Jones and Coutts, 1996; Tadesse
et al., 1999; Makkouk et al., 2001a).
The virus is reported to cause economic losses in chickpea and lentil
crops in West Australia (Jones and Coutts, 1996), and has caused a significant reduction in lentil yields in Iran (Kaiser, 1973). In India, the incidence
of CMV on different lentil cultivars varied from 5 to 100% (Rangaraju and
312
S.G. Kumari et al.
Chenulu, 1981), and in New Zealand virus incidence was below 11%, and
three fields were found with 77, 87 and 93% CMV infection (Fletcher, 1993).
In 1995 experimental plots of lentil varieties Rajah and Titore inoculated
with CMV yielded 17% and 19% less, respectively, than the uninoculated
control (Fletcher et al., 1999). Kaiser (1973) reported yield reductions of 87%
and 75% when the lentil cultivar Ghazvin was inoculated with CMV at
pre-flowering and flowering stages, respectively.
CMV has a wide host range and infects more than 800 species of both
monocotyledonous and dicotyledonous plants from over 85 families. It is also
reported to naturally infect pea, faba bean and chickpea (Bos et al., 1988).
The symptoms produced in response to CMV infection include interveinal chlorosis, growth stunting, proliferation of axillary shoots and a
reduction in leaf size, with adverse effects on flower and pod formation.
Seeds produced are small, discoloured and shrivelled (Kaiser, 1973; Rangaraju and Chenulu, 1981).
CMV is transmitted mechanically and by over 60 aphid species in a
non-persistent manner. The main aphid vectors are A. pisum, A. fabae, A.
craccivora, M. persicae, M. euphorbiae, Rhopalosiphum padi L. and A. solani
(Edwardson and Christie, 1986). The virus is reported to be seed-transmitted in lentil (Jones and Coutts, 1996; Makkouk and Attar, 2003).
Pea enation mosaic virus-1 (PEMV-1)
What is commonly referred to as Pea enation mosaic virus is a symbiotic coinfection of two viruses, Pea enation mosaic virus-1 (PEMV-1, genus Enamovirus, family Luteoviridae) and Pea enation mosaic virus-2 (PEMV-2, genus
Umbravirus, not assigned to a family). The disease is often simply referred
to by the acronym PEMV. PEMV-1 is readily transmitted mechanically, a
property dependent on its multiplication in cells co-infected with PEMV-2.
PEMV-1 and PEMV-2 are transmitted by several aphid species, such as A.
pisum, A. craccivora, M. persicae, M. euphorbiae, R. padi and A. solani (Cockbain, 1983; Edwardson and Christie, 1986).
Natural occurrence of the virus on lentil was reported in Italy by Vovlas and Rana (1972), in Ethiopia (Tadesse et al., 1999) and Iran (Makkouk
et al., 2003b). It was found also to occur in the USA (Aydin et al., 1987), especially in the Palouse region (Kaiser, 1987). It is also found in Europe and
Asia (Bos et al., 1988; Kumari et al., 2001; Makkouk et al., 2003a). Surveys
conducted in Syria during the period 19972001 showed that PEMV-1 was
widely distributed in the major lentil-growing areas, with virus incidence
reaching more than 50% at some locations. When six lentil genotypes inoculated with PEMV-1 before flowering were evaluated, yield losses varied
from 16% (ILL 7706) to 50% in ILL 6031 (Kumari et al., 2001).
Symptoms caused by PEMV-1 and PEMV-2 in lentil include reduced
growth, leaf rolling, leaf mottling, tip wilting and plant necrosis. Infected
plants are severely stunted with twisted and malformed leaves that frequently exhibit vein clearing and translucent flecks. Pods from infected
plants also are misshapen, poorly filled and often show protrusions on the
313
surface. These symptoms were severe in lentil fields in Syria (Kumari et al.,
2001) and in chickpea and lentil fields in eastern Washington (Klein et al.,
1991). In addition to lentil, PEMV-1 is reported to infect pea, French bean,
soybean, faba bean, chickpea and several other legume hosts.
Pea seed-borne mosaic virus (PSbMV)
The natural occurrence of PSbMV in lentil fields was first reported from the
USA (Hampton and Muehlbauer, 1977). The virus is known to occur on lentils in Algeria, Egypt, Ethiopia, Iran, Iraq, Jordan, Morocco, New Zealand,
Pakistan, Syria, Tunisia, Turkey and the USA (Hampton and Muehlbauer,
1977; Hampton, 1982; Aftab et al., 1992; Makkouk et al., 1992, 1993, 2001a,
2003b; Fletcher, 1993; Kassim, 1997; Bayaa et al., 1998; Tadesse et al., 1999;
Al-Mabrouk and Mansour, 2000).
There are several strains of PSbMV but the three most common strains
include P-1 and P-4 isolated from Pisum sativum and the L-1 strain from lentil (Alconero et al., 1986). All three strains infect lentil and chickpea. The L-1
strain causes a more severe reaction on chickpea and lentil than P-1 or P-4.
In Pakistan Aftab et al. (1992) noted severe mosaic symptoms of PSbMV
on lentil cultivar Precoz showing chlorotic lesions on leaves, stunting of
plants with shortening of internodes and a reduction in flower and pod formation. The decrease in plant height, number of pods, number of seeds and
yield per plant as a result of infection was 51, 58, 66 and 73%, respectively.
Kumari et al. (1993) reported 28, 27 and 23% yield losses in the field due to
lentil PSbMV infection at pre-flowering, flowering and post-flowering
stages, respectively. Symptoms may vary in lentil due to genotype, virus
strain and environmental effects. Other symptoms of infection include narrowed leaves, downward leaf rolling, mottling or chlorosis of the leaves
with shoot tip necrosis and reduced seed size. Seed-filled pods may be
abnormal in shape and size. Necrotic line patterns that are common on
infected pea and faba bean seeds are rare on lentil seeds.
Yield losses from PSbMV infection in Syria varied from 3% in the cultivar Redchief to 61% in ILL 6245 when 20 lentil genotypes were inoculated
mechanically with PSbMV at flowering; however, four genotypes (ILL 6198,
Crimson, Palouse and Redchief) were found highly tolerant to infection
(Kumari and Makkouk, 1995).
PSbMV is transmitted mechanically and by several aphid species
including A. pisum, A. fabae, A. craccivora, M. persicae and R. padi in the nonpersistent manner (Makkouk et al., 1993). Seed transmission rates of PSbMV
in lentil vary widely (044%) depending on host and virus isolate (Hampton and Muehlbauer, 1977; Makkouk et al., 1993; Kumari and Makkouk,
1995; Abraham and Makkouk, 2002).
Pea streak virus (PeSV)
PeSV was first reported on lentils in the USA (Hagedorn and Walker, 1949),
Canada and Germany (Bos, 1973). The host range is relatively small and
includes pea, lentil, faba bean, chickpea (Kaiser et al., 1993) and the forage
314
S.G. Kumari et al.
legumes lucerne and clover. PeSV was found to infect lentil under natural
conditions in the Palouse region of the USA (Kaiser, 1987).
The most common symptoms associated with PeSV include stunting,
yellowing of leaflets and wilting of the terminal shoots. Vascular discoloration also may be observed, a condition that contributes to plant death,
especially at the seedling stage. Pods, when formed, are poorly filled and
seeds are misshapen. PeSV is transmitted by mechanical inoculation and by
the pea aphid (A. pisum) in a non-persistent manner. It is not transmitted by
seed or by pollen.
Other viruses
In addition to the viruses mentioned above, other viruses of lesser significance were reported to cause damage to lentil in specific countries and in
limited areas, such as AMV (Kaiser et al., 1968; Jones and Coutts, 1996; AlMabrouk and Mansoor, 2000; Makkouk et al., 2003b; Bekele et al., 2005),
broad bean mottle virus (BBMV) in Morocco (Fortass and Diallo, 1993),
Ethiopia (Bekele et al., 2005) and Syria (Mouhanna et al., 1994), Broad bean
wilt virus-1 (BBWV-1) in Syria (Makkouk et al., 1992), Tomato spotted wilt
virus (TSWV) in Brazil (Fonseca et al., 1995), Chickpea chlorotic stunt virus
(CpCSV) in Ethiopia (Abraham et al., 2006), Syria and Tunisia (Kumari et al.,
2007), and Soybean dwarf virus (SbDV) in Australia (Johnstone and Guy,
1986), Japan (Tamada, 1973), Ethiopia and Syria (Makkouk et al., 1997).
19.3. Epidemiology
All viruses of major importance in lentil crops are mostly transmitted by
insect vectors (Table 19.1). The viruses transmitted by aphids in a nonpersistent manner (e.g. BYMV, PeSV and PSbMV) are acquired by the vector within a few seconds, transmission can occur in an equally short period,
aphids remain viruliferous for only short periods of time, and they can
spread the virus over short distances. In contrast, the persistently transmitted viruses (e.g. BLRV, FBNYV and PEMV-1) can be retained and transmitted in most cases for the life of the vector although transmission efficiency
is reduced significantly in adults compared to the early nymphal stages.
The persistently transmitted viruses require an acquisition period of several
minutes to several hours. The latent period in the vector can range from a
few hours (PEMV-1) to more than 100 h (BLRV). The inoculation access
period can be for few minutes or as long as 1 h. The persistently transmitted
viruses can spread over long distances, with the ability of an individual
insect to transmit them to many plants. Table 19.3 summarizes the most
important vectors reported to transmit lentil viruses worldwide.
Weed hosts, and perennial crops such as lucerne, vetch and clover are the
main sources of inoculum for many of the viruses that infect lentil. Most nonpersistently transmitted viruses that infect lentil are also seed-transmissible.
315
Table 19.3. Major insect vectors reported to transmit lentil viruses.

Vector
Aphids:
Acyrthosiphon pisum Harris
Aphis fabae (Scopoli)
Aphis craccivora Koch.
Myzus persicae (Sulz.)
Macrosiphum euphorbiae
(Thomas)
Rhopalosiphum padi L.
Aulacorthum solani (Kltb.)
Beetles:
Acalymma trivittata
Mannerheim
Apion aestivum Germ.
Apion aethiops Hbst.
Apion arrogans Wencher
Apion radiolus Kirby
Apion vorax Hbst.
Viruses transmitted
References
AMV, BBWV, BLRV,

BWYV, BYMV, CMV,
FBNYV, PEMV, PeSV,
PSbMV, SbDV
AMV, BBWV, BLRV,
BYMV, CMV, FBNYV,
PSbMV
AMV, BBWV, BLRV,
BWYV, BYMV,
CMV, FBNYV,
PEMV, PSbMV
AMV, BBWV, BLRV,
BWYV, BYMV, CMV,
PEMV, PSbMV
AMV, BBWV, BLRV,
BYMV, CMV, PEMV
CMV, PEMV, PSbMV
AMV, BWYV, CMV,
PEMV, SbDV
Cockbain (1983), Edwardson and

Christie (1986), Makkouk et al.
(1990, 1993, 1997), Franz et al.
(1998), Kumari et al. (2001)
Kaiser (1973), Cockbain (1983),
Edwardson and Christie (1986),
Makkouk et al. (1990, 1993)
Boswell and Gibbs (1983),
(1990, 1993), Franz et al. (1998)
(1990, 1993), Kumari et al. (2001)
Christie (1986)
Edwardson and Christie (1986),
Makkouk et al. (1993)
Christie (1986)
BBMV
Walters and Surin (1973)
BBSV
BBSV, BBTMV
BBMV, BBSV
BBMV
BBMV, BBSV, BBTMV
Colaspis avida Say

Diabrotica undecimpunctata
Mannerheim
Hypera variabilis Herbst
Pachytychius strumarius Gyll
Sitina lineatus var.
viridifrons Motsch
Sitona crinita Herbst
Sitona limosa Rossi
Sitona lineatus L.
BBMV
BBMV
Cockbain et al. (1975)

Cockbain et al. (1975)
Makkouk and Kumari (1989, 1995)
Fortass and Diallo (1993)
Cockbain et al. (1975),
Cockbain (1983)
BBMV
BBMV
BBMV

Borges and Louro (1974)
BBSV
BBMV, BBSV
BBMV, BBSV, BBTMV
Smicronyx cyaneus Gyll

Spodoptera exigua Hbner
Leafhoppers:
Orosius orientalis (Matsumura)
BBMV
BBMV
Makkouk and Kumari (1995)

Makkouk and Kumari (1995)
Cockbain et al. (1975), Fortass
and Diallo (1993), Makkouk and
Kumari (1995)
Ahmed and Eisa (1999)
CpCDV
Horn et al. (1993)
316
S.G. Kumari et al.
Table 19.4. Viruses that can be transmitted through lentil seeds.

Virus
AMV
BYMV
BBSV
CMV
PSbMV
Seed transmission rate (%)

0.15.0
0.11.4
0.30.83
0.8
14.0
0.11.0
0.0537.0
0.19.5
5.044.0
6.0
References
Makkouk and Attar (2003)
Kumari et al. (1994)
Makkouk and Azzam (1986)
Fletcher et al. (1999)
Makkouk and Attar (2003)
Hampton and Muehlbauer (1977)
Makkouk et al. (1993)
Virus transmission via seed is of dual importance. The virus-infected seeds

act both as source of inoculum and as vehicle of virus dissemination. Of the
20 viruses reported to infect lentil, five are seed-transmitted (Table 19.4).
Viruses that infect the embryo may also be transmitted by pollen although
this occurs rarely in lentil. Infection of lentil after flowering rarely leads to
seed infection. Usually infected seeds appear normal except in some cases
where visible symptoms are observed on stained seeds, as in the case of
infection by BBSV.
19.4. Virus Detection

Early detection and accurate diagnosis of viral diseases is critical for application of appropriate control measures. It is also important to determine
the area of distribution. The last three decades have witnessed significant
developments in improving the sensitivity of the methods to detect lentil
viruses.
Immunological protein-based methods

The development of enzyme-linked immunosorbent assay (ELISA) for plant
viruses (Clark and Adams, 1977) was a major step forward that replaced
earlier serological methods such as gel diffusion, especially for large-scale
testing. This was enhanced further with the development of monoclonal antibody technology and its application to a large number of legume viruses.
Specific reagents (monoclonal and/or polyclonal antibodies) have been
developed for most viruses affecting lentil, and using them in a variety of
ELISA tests available has made lentil virus diagnosis simple and fast. At
present, antibodies for detecting most of the lentil viruses are available
(Kumari and Makkouk, 2007). A number of ELISA variants were developed
which further improved sensitivity of virus testing. In many laboratories in
317
developing countries, facilities for sophisticated tests are lacking. For this
reason tissue-blot immunoassay (TBIA) was developed to identify most
legume viruses (Makkouk and Kumari, 1996). TBIA is a reliable simple test
to detect all lentil viruses reported and it is especially recommended for
virus surveys (Makkouk et al., 2001a, 2003b) and for evaluating virus-host
interactions (Kumari and Makkouk, 2003). Plate 4 illustrates how lentil
viruses are detected in infected plants by using TBIA.
Molecular nucleic acid-based methods

Nucleic acid hybridization has been used successfully for the detection of
many viruses (Pesic and Hiruki, 1988; Martin and DArcy, 1990). Cloning of
plant viral nucleic acids and the development of non-radioactive detection
methods has increased the utility of nucleic acid hybridization for virus
detection. The development of polymerase chain reaction (PCR) has greatly
improved the sensitivity and utility of hybridization and other nucleic acidbased assays. Immunocapture PCR combines the advantages of serology
and PCR (or reverse-transcription PCR (RT-PCR) for RNA viruses) into a
very sensitive method of detection (Shamloul et al., 1999). Many primers for
detecting lentil viruses by PCR have been reported (Kumari and Makkouk,
2007). This technique has now been adapted for the detection of both DNA
and RNA viruses with either single- or double-stranded genomes. In addition to detection of viruses in infected plants, RT-PCR can also be used for
detection of viruses in their vectors (Shamloul et al., 1999).
19.5. Control of Virus Diseases

As there is no direct practical way of curing crops from virus infection, all
current control strategies emphasize measures that prevent or reduce infection. Control measures can be classified as those that: (i) control the virus;
(ii) are directed towards avoidance of vectors or reducing their incidence;
and (iii) integrated approaches, which combine all possible control components in a way to complement each other and be applied by farmers as a
single control package.
Methods directed to control the virus

Reducing sources of infection
Five viruses affecting lentil are seed-borne (Table 19.4). For these viruses, it
is always recommended to use virus-free seed for planting, especially when
the virus is also transmitted by active vectors. The use of genotypes or cultivars resistant to the virus or to seed transmission is an effective control
measure. Kumari and Makkouk (1995) reported that lentil cultivars Redchief
318
S.G. Kumari et al.
and Palouse had the lowest seed transmission rate of PSbMV with the
lowest yield loss among the 20 lentil genotypes evaluated. In another study,
they found that Redchief had the lowest seed transmission rate (0.2%) and
lowest yield loss (14%) among the 19 lentil genotypes evaluated (Makkouk
and Kumari, 1990). Seed transmission of BBSV was reduced to zero when
seeds were exposed to 70C for 28 days; however, this treatment caused an
unacceptable reduction (57%) in seed germination (Kumari and Makkouk,
1996). Such an approach could be useful to eliminate seed infection in germplasm accessions deposited in genebanks, but is not economical for
commercial production.
Roguing symptomatic plants is a widely used phytosanitary measure to
remove sources of virus infection from standing crops. However, when
virus incidence within a field is high, this control measure may not be practical. Over-wintering or over-summering crops, which could play the role
of sources of infection, should be avoided through spatial isolation. Such
methods are more effective with non-persistent viruses than with persistent
viruses. With non-persistent viruses, such as BYMV, a few hundred metres
isolation may adequately reduce virus spread. In contrast, persistently
transmitted viruses such as BLRV can be carried from lucerne fields over
long distances making it more difficult to avoid.
Breeding for virus resistance
Host resistance, when available, is the most acceptable component of virus
control because it is environment friendly, practical and economically
acceptable to farmers. Several workers have reported lentil lines resistant to
viruses. Four accessions (PI 212610, PI 251786, PI 297745 and PI 368648) are
immune to PSbMV. This resistance is controlled by the single recessive gene
sbv (Haddad et al., 1978). Accessions PI 212611, PI 212903, PI 343025, PI
370632 and PI 379368 were also found to be resistant, and PI 368648 was
immune to PSbMV (Hampton, 1982). Genotypes PI 472547 and PI 472609
were reported to be tolerant to PEMV (Aydin et al., 1987). The genotype ILL
7163 displayed a high level of resistance to BYMV and moderate resistance
to CMV (McKirdy et al., 2000; Latham et al., 2001), while genotype ILL 5480
was moderately resistant to AMV (Latham and Jones, 2001). Six lentil genotypes were identified with combined resistance to different viruses (Makkouk et al., 2001b): ILL 75 had resistance to BLRV, FBNYV and SbDV; and
ILL 74, ILL 85, ILL 213, ILL 214 and ILL 6816 were resistant to FBNYV and
BLRV. Two cultivars from the USA, Redchief and Palouse, were highly
tolerant to PSbMV infection. This tolerance was expressed as very low grain
yield loss due to infection and with a low seed transmission rate (Kumari
and Makkouk, 1995). Redchief was also reported to be tolerant to BBSV
earlier with low seed-transmission rates (Makkouk and Kumari, 1990). At
present, a number of legumes are being transformed with viral genes (coat
protein, replicase, etc.) to produce virus-resistant varieties (Chowrira et al.,
1998; Chu et al., 1999). No genetically transformed cultivars are available in
the market for immediate use by farmers, but genetic transformation could
319
be a useful alternative, especially for viruses against which no resistance

genes have been identified.
Methods directed towards avoidance of vectors or reducing their incidence

Cultural practices
Various practices such as planting date, high seeding rate and narrow row
spacing, use of early maturing cultivars, and using border plantings that
are a non-host of the virus have proved effective in reducing virus incidence
in lentil crops. Manipulation of planting date to avoid exposure of plants to
peak vector populations at the most vulnerable early growth stages is a
standard virus control measure widely recommended for use in legume
crops (Thresh, 2003).
Virus vector control (chemical control)
Application of insecticides helps to decrease the spread of some lentil
viruses vectored by insects. However, it is often ineffective because its success depends on the mechanism of virus transmission and the mode of
action of the insecticide selected. In general, success in reducing virus
spread by chemical control of vectors is considerably greater with persistently than with non-persistently transmitted viruses. This is mostly because
the incoming viruliferous vectors carrying non-persistently transmitted
viruses are not killed fast enough to prevent probing and subsequent virus
inoculation to the sprayed plants. Oil sprays can be used instead, but are
not always cost effective because of the repeated applications required. The
most effective insecticides to control non-persistently aphid-borne viruses
are the newer generation of synthetic pyrethroids (Loebenstein and Raccah,
1980), because of their rapid breakdown and greater anti-feedant activity.
Field experiments at ICARDA showed that seed treatment with the systemic insecticide imidacloprid (Gaucho) significantly improved yields of
moderately resistant and susceptible lentil genotypes, but had no effect on
the yield of resistant genotypes. Seed treatment was also effective in increasing seed yield of BLRV and FBNYV-inoculated plots, but had no effect in
SbDV-inoculated plots (Makkouk and Kumari, 2001).
Integrated approach
Each of the control measures described in this chapter provides only partial
control, but combining genetic resistance, cultural practices and chemical
sprays is expected to check virus spread. The use of host resistance, whether
obtained by classical breeding or genetic engineering, and one or two welltimed insecticide applications coupled with optimal planting date and early
roguing of virus-infected plants could offer reasonable and economic virus
disease control and stabilize lentil production. However, selecting the best
320
S.G. Kumari et al.
mix of measures for each virus-crop combination and production situation

requires knowledge of the epidemiology of the causal virus and the mode
of action and effectiveness of each individual control measure. Each strategy
must be affordable to the farmer and fulfill the requirements of being environmentally friendly and socially acceptable. It should also be compatible
with control measures already in use against other pests and pathogens.
19.6. Concluding Remarks

This chapter has described the major aspects of the economically important
viruses of lentil. Certainly, not all viruses have been discussed because of
space constraints. There are many other minor viruses which may be of
local importance in lentil that have not been discussed here. In addition,
new viruses or their strains are expected to be discovered or introduced that
will affect lentil. For example, CpCSV has recently been reported on lentil
and chickpea in Ethiopia and on other food legume crops in Syria (Abraham et al., 2006; Kumari et al., 2007). This new virus might become important for lentil with time. Work is currently in progress to study the etiology,
epidemiology and economic importance of this novel virus. Lentil is an
important crop for the resource-poor farmers in developing countries. Therefore disease control measures must be practicable, affordable and suitable
for situations in these countries. In addition, agricultural researchers, breeders, extension agents and growers must continually adjust disease management strategies to cope with changing conditions.
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20
Weed Management
Joseph P. Yenish,1 Jason Brand,2 Mustafa Pala3 and Atef Haddad3

1Washington
State University, Pullman, Washington, USA; 2Department of Primary

Industries, Horsham, Victoria, Australia; 3International Center for Agricultural
Research in the Dry Areas (ICARDA), Aleppo, Syria
20.1. Introduction
Lentil competes very poorly with weeds. This poor competitive ability is
due in part to the short stature and slow early season growth rates (Basler,
1981). Yield reductions of 2084% due to competition from weeds have
occurred (Saxena and Wassimi, 1980; Salkini and Nygaard, 1983; Knott and
Halila, 1986; Kumar and Kolar, 1989; Al Thahabi et al., 1994; Boerboom and
Young, 1995; Mohamed et al., 1997). Crop yield losses are primarily a result
of competition with weeds for nutrients, moisture and space. However,
additional losses are due to weeds harbouring insects and pathogens that
adversely affect lentils. Moreover, losses in harvest efficiency and reduced
crop quality may result from weed infestations. Actual yield, quality and
other losses will vary with the intensity of infestations and species makeup
of weed infestations (Bhan and Kukula, 1987).
Generally, weeds emerging before or at the time of crop emergence
have a greater competitive advantage than those emerging later (ODonovan
et al., 1985; Dieleman et al., 1995; Knezevic et al., 1995; Bosnic and Swanton,
1997). Slow emergence, short plant height and late canopy cover of coolseason legumes in general and lentils specifically, allow weeds to compete
effectively. A study by Moes and Domitruk (1995) showed average emergence for lentils, chickpeas, dry peas and wheat of 21, 23, 18 and 17 days
after planting, respectively, in a no-tillage system. Moreover, the same study
showed average canopy closure of 62, 69, 56 and 53 days after planting for
the same respective species. Slower crop emergence and canopy closure
provides opportunity for weeds to germinate and establish with little competition from the crop. The earlier weeds emerge relative to the crop, the
greater the yield loss if left uncontrolled (Kropff et al., 1992). Early establishing weeds, particularly those emerging before the crop, have a competitive
advantage over the later emerging crop and will severely reduce yields if
326

Weed Management
327
not controlled. Later emerging weeds will compete with the crop as it produces seed and can directly reduce seed size or quality. Additionally, several important weeds in lentil such as Lathyrus aphaca L., Vicia sativa L. and
Vicia hirsuta (L.) Gray, produce seed similar in shape and size to that of lentil and separation from the crop is difficult, resulting in lowered quality and
value of the harvested crop (Brand et al., 2007). Seed quality and yield may
also be reduced indirectly by weeds interfering with the harvest. Heavy,
wet or immature weed vegetation may interfere with mechanical or hand
harvest of lentil. Weeds containing sticky latex sap, such as prickly lettuce
(Lactuca serriola L.), may cause further problems as the latex disrupts the
normal operation of machinery and causes contaminants to adhere to the
seed (Yenish, 2007).
The critical weed-free period is defined as the period of crop growth
during which the crop must be kept weed free to prevent yield loss due to
weed interference (Weaver and Tan, 1987; Van Acker et al., 1993). Singh et al.
(1996) indicated that weeds could be allowed to grow in lentil from 34 or 41
days after emergence with crop yield losses not greater than 10% (Fig. 20.1).
(a)
(b)
100
Yield (% of weed-free control)
100
90
80
70
60
50
40
30
20
10
70
60
50
40
30
20
0
10 20 30 40 50 60 70 80 90 100 110
10 20 30 40 50 60 70 80 90 100 110
(d)
100
100
80
10
10 20 30 40 50 60 70 80 90 100 110
(c)
90
90
80
70
60
50
40
30
20
10
10 20 30 40 50 60 70 80 90 100 110
90
80
70
60
50
40
30
20
10
Time after lentil emergence (days)
Fig. 20.1. Lentil seed yield response to increasing length of weed-free period () and
duration of weed-infested period (---) in days after lentil emergence at: (a) Madaba, Jordan,
1988/89; (b) Mshager, Jordan, 1988/89; (c) Mshager, Jordan, 1989/90; and (d) Jubaiha, Jordan,
1989/90 (Source: Singh et al., 1996).
328
J.P. Yenish et al.
Seed yield
(% of weed-free control)
100
80
60
40
20
0
0
60
72
12
24
36
48
Time after lentil emergence (days)
Fig. 20.2. Lentil seed yield response to increasing length of weed-free period ()
and duration of weed-infested period (---) in days after lentil emergence at Rubatab,
Sudan, 19921994 (Source: Mohamed et al., 1997).
This research was conducted at four non-irrigated locations in Jordan and also
concluded that crop yield losses were not greater than 10% when weeds were
controlled from emergence up to 85 or 99 days after emergence. Figure 20.2
shows the critical weed-free period between 2 and 4 weeks after seeding for
a study conducted in Sudan under irrigation (Mohamed et al., 1997). A
study in Tunisia estimated the critical weed-free period for lentil at 16
weeks after emergence for a location with a low to medium severity of weed
infestation and 4 weeks for a separate location where the infestation was
described as severe (Knott and Halila, 1986).
An Australian study conducted by McDonald et al. (2007) indicated that
lentil loss due to weed density followed the yield loss model described by
Cousens (1985a, b). Additionally, the study showed that the actual yield
loss response varied with variety and environment. Maximum yield loss
due to weeds in the McDonald study varied from 50 to 60% between years
and locations (Fig. 20.3).
Tepe et al. (2005) showed yield losses due to weeds differed between
lentil cultivars at the same weed density in a study conducted in Turkey.
Comparisons of lentil yields in weedy and weed-free treatments resulted in
measured yield losses of 3567% depending on the cultivar. However, the
ranked order of yield loss for the cultivars was different between the 2 years
of the study. The study did not identify any particular lentil trait that may
have influenced its ability to compete with or tolerate competition with
weeds.
20.2. Problem Weeds

The species of weeds commonly infesting lentil fields vary with changing
production environments. Soil characteristics, moisture amount, precipitation
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329
2500
Minlaton 2002
Minlaton 2003
Horsham 2003
Lentil yield (kg/ha)
2000
1500
1000
500
0
0
10
20
30
40
50
60
Canola density (plant/m2)
Fig. 20.3. Relationship between weed (volunteer canola) density and lentil yield at
Minlaton, South Australia in 2002 and 2003 and Horsham, Victoria in 2003
(Source: McDonald et al., 2007).
pattern, crop rotation, temperature, latitude, altitude, fertility, weed control

technology and other factors interact to determine weed flora and intensity.
Generally, weeds causing significant losses in lentils are categorized as
annual monocots, annual dicots, perennial monocots, perennial dicots and
parasitic (Table 20.1). Production manuals for the various cropping regions
are the best local sources for identification, importance and management of
weed species (Wilding et al., 1998; Hnatowich, 2000; Singh et al., 2001; Moorthy et al., 2002; Holding and Bowcher, 2004; Day et al., 2006).
Typically the most problematic weed species in lentil have a similar
ecology and biology as the crop. Generally, cool-season broadleaf weeds are
the most difficult to control in lentils. Problem annual broadleaf or dicot
species in lentil include those of the Asteraceae, Brassicacae, Chenopodiaceae,
Fabaceae and Polygonaceae, among other families (Brand et al., 2007). While
many annual grass (Poaceae) or monocot species are selectively and effectively controlled with herbicides, Avena, Lolium, Phalaris, Poa, Setaria and
other species are commonly reported in lentils (Table 20.1). Generally, the
most troublesome grass weeds in lentil are classified as cool-season species
categorized as C3 plants based on their photosynthetic system. Perennial
weeds such as members of the Convolvulaceae, Asteraceae, Poaceae and other
families can also be problematic in lentil production.
Weeds that are specific problems in lentil are problems within a crop
rotation and not specific to the lentil crop alone. An exception to this is parasitic plants which are partially or completely obligate to lentil. Orobanche
and Cuscuta spp. have been reported to be parasitic weeds of lentil and
other legumes (see Rubiales et al., Chapter 21, this volume).
330
Table 20.1.
Common weed ora of lentil crop.
Category Family
Scientic name
Common name
Category
Liliaceae
Annual
monocots Poaceae
Asphodelus tenuifolius
Avena spp.
Wild onion
Wild oat
Bromus spp.
Brome grass
Fabaceae
Annual
dicots
(continued) Fumariaceae
Critesion murinum
Lolium spp.
Phalaris minor
Poa annua
Polypogon monspeliensis
Barley grass
Ryegrass
Little seed canarygrass
Annual bluegrass
Annual rabbitsfoot grass
Setaria viridis
Green foxtail
Vulpia bromoides
Amaranthus spp.
Bifora testiculata
Anthemis cotula
Gnaphalium indicum
Lactuca serriola
Launaea nudicaulis
Silver grass
Pigweeds
Bifora
Mayweed chamomile
Cudweed
Prickly lettuce
Skeletonweed
Annual
dicots
Amaranthaceae
Apiaceae
Asteraceae
Arrow weed
Sow thistle
Tarweed
Small bugloss
Sheep weed
Swine cress
Musk weed
Wild radish
Wild mustard
Corn spurry
Common lambsquarters
Kochia
Parasitic
Scientic name
Common name
Lathyrus aphaca
Medicago spp.
Medicago
denticulata
Melilotus alba
Melilotus indica
Vicia spp.
Vicia villosa
Fumaria spp.
Polygonum spp.
Wild pea
Medic
Bur clover
White sweet clover

Yellow sweet clover
Common vetch
Woolly pod vetch
Fumitory
Polygonaceae
Buckwheat/
Wireweed
Rumex dentatus
Wood sorrel
Rubaceae
Gallium spp.
Bedstraw
Solanaceae
Solanum spp.
Nightshade
Cyperaceae
Cyperus rotundus Nut grass
Poaceae
Cynodon dactylon Bermuda grass
Asteraceae
Canada thistle
Cirsium arvense
Convolvulaceae Convolvulus
Field bindweed
arvensis
Resedaceae
Reseda spp.
Mignonette
Convolvulaceae Cuscuta spp.
Dodder
Orobancheaceae Orobanche spp.
Broomrape
J.P. Yenish et al.
Pluchea lanceolata
Sonchus spp.
Boraginaceae
Amsinckia spp.
Anchusa arvensis
Buglosoides arvensis
Brassicaceae
Coronopus didimus
Myagrum perfoliatum
Raphanus raphanistrum
Sinapis arvensis and
Sisymbrium spp.
Caryophyllaceae Spergula arvensis
Chenopodiaceae Chenopodium album
Kochia scoparia
Perennial
monocots
Perennial
dicots
Family
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331
20.3. Methods of Weed Control

There are five recognized weed control techniques (Anderson, 1983). These
include preventive, cultural, mechanical, chemical and biological. Mixing
two or more of these weed management principles is the basis for integrated
weed management. Each of the five principal techniques is of equal value in
managing weeds in lentil with the exception of biological control. Few biological methods of weed control have proven effective in annual cropping
systems and individual weed species which are problematic in legumes are
largely a function of cropping systems or rotation rather than the individual
crop of lentil.
Preventive weed control
The most cost effective form of weed control is preventive. A quote attributed to the 13th-century English judge Henry de Bracton, an ounce of prevention is worth a pound of cure, is very true in the realm of weed
management. The definition of preventive weed management is to eliminate the introduction or spread of specified weed species in an area not currently infested with these plant species (Radosevich et al., 1997). The size of
the geographic area may vary from an individual field to an entire continent. While much of preventive weed control could be regulatory in nature,
there is still much that an individual farmer can do to prevent the introduction of weeds. In fact, individual farmer practices are more likely to be effective in preventing weeds than any regulatory practices.
Individual farmers concentrating on preventing weed introduction
often need only to plant weed-free seed, use manure generated from weedfree hay, feed or bedding, clean field equipment (particularly harvest equipment) and prevent weeds that are encroaching on their property from
spreading seed or other vegetative propagules onto their land (Young et al.,
2000). Typically, weed-free seed is easy to obtain, but normal cleaning or
screening methods may provide certain weeds a method of effective entry.
Specific weed pests such as wild pea or vetches produce seed similar in
shape and size to that of lentil and removing this seed through cleaning
may be difficult (Erskine et al., 1994; Brand et al., 2007). It is best to avoid
seed from fields or regions with known infestations of these and similar
weeds. Once seed cleaning is completed, growers need to adequately
destroy screenings through composting, hammer mill or other methods.
Livestock manure can also be a source of weed seed entering a farming
operation. Feed and hay should be inspected prior to feeding or mixing.
Even if feed, hay and bedding are thought to be weed free, manure should
be composted to further eliminate weed seed through the temperatures
generated. Temperatures as high as 83C may be required to adequately
reduce weed-seed viability, although some species may lose viability at
lower temperatures (Wiese et al., 1998). Acquired animals should also be
quarantined to allow the digestive system to completely empty of foreign
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J.P. Yenish et al.
feed. Additionally, hair, wool, hides and hooves should be inspected to make
sure no weed seed is trapped either directly or encased in mud or other foreign matter.
Equipment should be cleaned prior to removal from and entry into
fields to prevent weed dispersal (Anderson, 1983). Methods for cleaning
equipment may be as simple as removing seed, rhizomes, etc. by hand or as
high tech as radiation treatments. Special care needs to be paid to harvest
equipment since weeds typically mature at the same time as the crop and
modern harvest equipment is an effective means to disperse weed seed
within and between fields. In cases where residue is removed from crop
fields for the purpose of offsite separation of grain and stover, that residue
should be disposed of at the processing site rather than the farmer transporting it back to his own fields for disposal. When the stover is used for
the livestock feed, the manure should be composted or otherwise treated to
kill viable weed seed prior to spreading the manure back on to fields. Mixing of stover from multiple fields and then returning a portion to each field
will spread weeds quite effectively.
Irrigation water is also a common source of weed-seed influx. Kelley
and Bruns (1975) in a detailed study of the US Pacific Northwests Columbia River calculated as much as 15,000 seed/ha could be disseminated onto
fields by unscreened irrigation waters. Incoming irrigation water should be
screened or other apparatus used to remove weed seed from the water prior
to applying it to a field.
Controlling encroaching populations of weeds is also important for preventing new weed infestations. Most weed seed falls less than 1 m from the
mother plant (Radosevich et al., 1997). Thus, patches of weeds tend to creep
across a given area if allowed to naturally disperse. At the same time, longrange dispersal of weed seeds is important in the colonization of new weed
patches. Therefore, the nearest patches of weeds that are encroaching on a
particular area are the most threatening to that area, while weed seeds travelling long distances are a potential future problem after they colonize into
weed patches.
Preventive weed control could also be applied to weed species that are
already present in fields within the farming operation. This is particularly
important in regions where chemical weed control is a large component of
weed management programmes. Preventing external influxes of weed seed
from outside sources is important in keeping herbicide-resistant weed biotypes out of the farming operation. Moreover, preventive weed management could isolate a weed infestation to one field or a portion of one field
within a grower operation. In areas with high herbicide use, a weed seed in
screenings or as a seed contaminant probably survived a herbicide application to the field in which it was produced. Therefore there is a greater
potential for that seed to be resistant to the herbicide than the wild type. Of
course, the mother plant may have emerged after the herbicide application,
or it may not have been sprayed because of poor herbicide application or
equipment problems, in which case the plant is unlikely to be herbicide
resistant. However, the potential exists to bring herbicide resistance into the
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333
farming operation by using screenings or contaminated seed or to move

resistant weeds within and between fields.
Cultural weed control

Cultural methods of weed management utilize practices that are common
to good crop management (Smith and Martin, 1995). The goal is to manage
a crop or cropping system to maximize crop competition with weeds. In
other words, make the crop as healthy and competitive as possible to lessen
or better tolerate weed competition.
Tools for cultural weed control include use of competitive cultivars,
timeliness of planting, cover crops, and competitive rotational crops or rotational crops with different life cycles. Planting crops with differing life
cycles in rotation prevents population increases of any single species of
weed (Radosevich et al., 1997). Additionally, practices such as optimal fertilizer placement and timing, optimal irrigation timing, and other input timing or placement can benefit the crop at the expense of weeds (Di Tomaso,
1995). Cultural practices are generally applied within, between, and
throughout the crop rotation. Unfortunately, lentil is usually the weakest
competitor in the rotations in which they are grown. Typically, cultural
weed control in rotational crops keep weed populations low and benefits
are realized in the yield and quality of lentils. Lentil provides little to no
opportunity for cultural weed control within the rotation. Ultimately, the
value of lentil as a cultural weed control tool within a crop rotation is
directly related to differences in the ecology and biology between the lentil
and other crops in the rotation.
A primary rotational weed control benefit to lentil is the management
of parasitic plants. Unfortunately, with the dormant seed life of Orobanche and
Cuscuta spp. being 14 (Lopez-Granados and Garcia-Torres, 1999) and 60 years
(British Columbia Ministry of Agriculture, Food and Fisheries, 2002), respectively, the rotational interval between susceptible crops may be impractical.
There are several cultural practices commonly involved with lentil production that makes them more prone to losses from weed competition. Very
early spring or winter planting of lentil have shown greater yield potential
than later spring seeding (Sarker and Erskine, 2007). However, effective
weed control is critical to fully realizing this increased yield potential.
Delayed sowing reduces early vigour, crop competitiveness, and resulting
yield potential (Mishra et al., 1996; Holding and Bowcher, 2004). However,
delayed seeding provides greater opportunity for weed control by mechanical or chemical methods prior to sowing (Brenzil et al., 2006; Day et al., 2006).
In North America, autumn-seeded lentils have a substantial yield advantage over spring-seeded lentils. However, weed competition and limited
post-emergence control options are major obstacles to fully realizing the
greater yield of autumn-seeded lentils. Moreover, weed competition may
be equal or greater from winter annual weeds and cold-weather-tolerant
weeds than with spring annual weeds present in a spring-seeded crop.
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J.P. Yenish et al.
Lentil cultivars vary in growth habit and morphology (Erskine and

Goodrich, 1991) and differences in competitive ability could be expected.
However, research has been inconclusive with Tepe et al. (2005) reporting
minor differences in competitiveness between cultivars, but indicating that
none of the cultivars were really effective as a stand-alone component for
weed control in lentils. McDonald et al. (2007) also showed that differences
in early vigour between genotypes were insufficient to affect the competitive ability of lentil.
Seeding depth, rate and row spacing influence lentil establishment,
competitiveness and yield. Seeding at 35 cm depth increases plant emergence, height, plant dry weight and grain yield compared to seeding at 810
cm. However, greater crop injury has been observed due to pre-emergence
herbicide applications with shallow compared to deeper seed placement
(van Rees, 1997; Brand et al., 2001). Greater seeding rates also reduce weed
populations due to greater crop competition (Boerboom and Young, 1995;
Ball et al., 1997; Paolini et al., 2003; McDonald et al., 2007). Narrow row spacing theoretically improves crop competition against weeds as the lentil crop
reaches full canopy closure earlier than with wider spacing. However,
Chaudhary and Singh (1987) found no effect of row spacing on final weed
populations. Although the crop is likely to be less competitive with weeds
with wider row spacing, wider spacing can allow more efficient weed
removal by tillage or hand weeding.
Rotational crops in lentil production systems vary greatly within and
between production areas. In some areas of the Indian subcontinent, lentil
may be grown in rotation with millet, sorghum, maize, cotton, guar, sesame
or rice (Ali et al., 1993). In other areas, lentil may be grown in rotation with
wheat, barley or forage crops (Yenish, 2007). While the most common practice of growing lentil is as a sole crop, there are instances where lentil is grown
as part of a crop mixture. In other situations, the crop may be grown in relay
with other crops to allow two harvested crops a year. Regardless of crop rotation, lentil is an attractive crop because of the disease, fertility and monetary
benefits it brings to the rotation. Lentil is not grown to provide a weed control benefit. Moreover, lentil could become a liability with regards to weed
control because of the increased weed-seed production due to poor competition or limitations of the use of soil-persistent herbicides in rotational crops.
Cultural control of weeds is an important part of cropping systems.
Managing weeds by cultural methods means reduced expenditure on pesticides, fuel and labour. While cultural management of weeds is an important
part of crop rotations that include lentil, lentils are usually the beneficiary
of the positive aspects of cultural weed control rather than a benefactor.
Mechanical weed control

Mechanical methods are the primary approach to weed control in lentil.
Mechanical practices range from tractor-powered tillage to weed
removal by hand hoeing or pulling. Pulling weeds by hand or removal
Weed Management
335
by human-powered equipment such as a wheel hoe is common in less

industrialized countries (Knott and Halila, 1986; Bhan and Kukula, 1987).
However, even in these countries labour costs are becoming prohibitive
(Solh and Pala, 1990). In more industrialized production, mechanical control of weeds is limited by the narrow row spacing commonly used. Row
spacing is too narrow to allow between-row cultivation without damaging
the crop. Mechanical weed control is limited to aggressive and multiple tillage operations prior to planting, with ploughs, cultivators or discs and postplanting to early post-emergence use of a harrow, cultipacker or rotary hoe.
In order to be effective, mechanical weed control must be repeated
throughout the critical weed-free period. This often means that the initial
weeding may occur before lentil emergence. Light tillage with a cultipacker,
harrow or rotary hoe can be effective to some degree in removing small
weed seedlings without excessive damage to lentil seedlings. Generally,
more aggressive tillage will damage lentil and may result in reduced yields
compared to non-weeded fields. Thus, mechanical control of weeds in lentils after crop emergence is largely limited to hand weeding only.
The frequency of weed removal by mechanical methods varies with
production systems and environment. Experimentally, up to five mechanical weed removals have been necessary to ensure weed-free conditions in
lentils or other legumes (Knott and Halila, 1986). However, competition
from weeds may occur as weeds are allowed to develop into a stage that
allows for proper identification and grasping for removal. Weeds are in
competition with the crop during this time and often there is physical damage to the crop as weeds are removed by hand or machine.
Traditionally, due to its short stature lentil has been grown in systems
where the soil has been cultivated to create a flat seedbed and stubble
removed to allow good establishment and ease of harvest. These conditions
are also generally regarded as optimal for pre-emergence herbicide applications. Cultivation of soil can incorporate weed seed into top soil layers
allowing for maximum germination when soil is moist and the potential for
a knockdown herbicide or follow-up cultivation to control emerging weeds
prior to sowing in some situations (Roberts, 1981). Deep tillage can also
bury weed seed below their optimum emergence depth inducing dormancy
and inhibiting germination by reducing the oxygen concentration (Preston,
2007). Future disturbance may reintroduce the seed to an appropriate depth
for germination. Tillage can also stimulate the germination of some weed
species by damaging the seedcoat, which breaks seed dormancy. Stubble
from previous crops can be removed by burning or physical incorporation
into the soil. Burning can reduce weed numbers of some species. Walsh and
Newman (2006) found that all seed of annual ryegrass (Lolium rigidum Gaudin) and wild radish (Raphanus raphanistrum L.) were destroyed by burning,
provided burning temperatures were above 400 and 500C, respectively.
Removing stubble and allowing the soil to be solarized can have a significant impact on weed populations. Linke (1994) showed that soil solarization
reduced the population in most weed species present, with only a minority
increasing in population.
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J.P. Yenish et al.
Currently, farmers in the more developed regions of the world are

adopting no-tillage systems with minimal soil disturbance and crop residue
conservation. No-tillage systems greatly reduce the opportunity for preplant mechanical weed control. However, reduced tillage systems may have
an advantage in managing the soil weed seedbank. Minimum soil disturbance ensures that most weed seeds are left on the soil surface. Seed survival of weed species such as green foxtail and wild oat (Avena fatua L.) are
reduced fivefold by leaving seed on the surface in comparison with burying
by tillage (Banting et al., 1973; Saga and Mortimer, 1976; Thomas et al., 1986).
Modelling and field observations have confirmed trends that preventing
seed entry into the surface seedbank will reduce weed populations in the
long term (Mohler, 1993; Anderson, 2004; Preston, 2007). Additionally, standing stubble can provide trellising for the crop, resulting in higher canopy and
improved machine harvest.
Chemical weed control

Chemical weed control is synonymous with using herbicides to control
weeds. Herbicides are regulated by various government agencies and laws.
Products available vary greatly between regulatory entities. Further complicating matters are the limited number of herbicide manufacturers and relatively limited markets for potential lentil herbicides. Moreover, herbicides
that are effective for controlling the weed spectrum in one particular lentil
production system in one particular geographic area may have limited
activity against weeds in another production system due to poor efficacy or
soil persistence. Thus, discussing specific herbicides is somewhat pointless
as recommendations for one country may be ineffective or illegal in another
country or even different regions of the same nation. That being said, discussion in this section will be general in nature and specific only to make a
point. Readers need to be aware to follow all local laws and regulations,
seeking out specific recommendations from local agronomists, extension
personnel or other qualified individuals or entities.
Non-selective herbicides may be used alone or as an aid to tillage in
controlling weeds prior to planting or emergence of grain legumes (McKay
et al., 2002). Herbicides such as glyphosate or paraquat do not have activity
once they come into contact with the soil (Vencill, 2002). These herbicides
are commonly used in no-tillage or reduced tillage production systems to
ensure control of weeds with no or minimal tillage prior to lentil planting
(Baker et al., 1996). More typically, control of weeds prior to planting is done
using tillage operations which are applied specifically for weed control or
provide effect weed control while preparing a suitable seedbed (Knott and
Halila, 1986).
Often residual herbicides are applied prior to planting lentil (Bhan and
Kukula, 1987). These residual herbicides will provide control against weeds
that emerge over the following few days to several weeks following herbicide
application. The exact duration of weed control will vary with herbicide
Weed Management
337
characteristics, rate, environmental conditions or other factors. Ideally, residual herbicides will provide effective weed control from time of application
until the lentil crop progresses beyond the critical weed-free period or the
full-bloom stage as noted earlier. Herbicides that have provided effective
broadleaf-weed control with no or acceptable crop injury include several
dinitroanalines (trifluralin, pendemethalin and others), triazines (metribuzin),
acetanalides (metolachlor) (Bhan and Kukula, 1987; Solh and Pala, 1990)
and imidazolinones (imazethapyr) (Lyon and Wilson, 2005). These herbicides may be mechanically incorporated with sweep cultivators, discs, harrow or other tillage tools prior to planting or applied following planting,
but prior to crop emergence. In some instances, residual herbicides may be
tank-mixed with glyphosate or other non-selective herbicide. Often, it is
necessary to combine or tank-mix one or more herbicides in order to
broaden the spectrum of weeds controlled (Bruff and Shaw, 1992; Hydrick
and Shaw, 1994; Zhang et al., 1995).
Post-emergence herbicides available for lentils are very limited relative
to the number of products available for more widely grown legume crops
such as soybean (Zollinger, 2006). Grass weeds can be effectively controlled
by aryloxyphenoxy propionate or cyclohexanedione herbicides with excellent crop safety (McKay et al., 2002). However, resistant grass weed populations have developed through heavy use of these herbicides in legumes and
rotational crops such as cereals (Delye, 2005). Herbicides for post-emergence
broadleaf control are extremely limited for lentil and include only a few
herbicides, the legal use of which is changing with regulating agencies
between countries. Crop safety is often limiting with post-emergence broadleaf herbicides in lentils.
Controlling weeds or desiccating a crop to aid in harvest can be done
by mechanical or chemical options (McKay et al., 2002). While mowing or
swathing the crop is possible, it is often not the best method of control since
lentils often do not cure well in the swathe (Hnatowich, 2000). Similarly, a
crop with uneven maturity will have great variability in seed size and quality when swathed or mowed prior to harvest. Chemical desiccation can be
achieved using products such as glyphosate or paraquat (McKay et al.,
2002). Given that weed control options are limited (particularly for broadleaf weeds), chemical desiccation is often necessary to aid in the mechanical
aspects of harvest or the timeliness to ensure successful dry-down of weeds
and crops prior to harvest.
20.4. Integrated Weed Management

Maredia (2003) defines integrated pest management as:
A pest management system that, in the context of the management associated environment and the population dynamics of the pest species, utilizes
all suitable techniques and methods in as compatible a manner as possible
to maintain the pest populations at levels below those causing economically
338
J.P. Yenish et al.

unacceptable damage or loss. Consideration is given to social acceptability,
ecological stability, environmental safety and human resource development.
More simply, integrated weed management uses all weed-control strategies to effectively control weeds in a safe, cost-effective and environmentally sound manner. Currently, all or nearly all lentils are grown under
some form of integrated pest management. Integrated weed management
in lentils is a necessity due to its poor competitiveness, lack of effective herbicides, the need to rotate crops as part of disease or other pest management, and other reasons. Generally, lentils are able to be successfully grown
due to integrated pest management throughout the rotation. Lentils, by
themselves, have limited benefit in a truly integrated cropping system.
20.5. Summary
Lentil is a very poor competitor against weeds. Most previously published
compendia on lentil production note lack of effective herbicides and the
need to expand available herbicides. Most herbicides labelled for use in lentils were initially developed for the soybean market and happened to have
safety in lentils. However, the introduction and phenomenal success of glyphosate-resistant soybeans have largely lessened the development of herbicides
for use in soybeans. Moreover, while the development of herbicide-resistant
technology was mentioned by some as a potential method to provide outstanding weed control in minor crops, actual development of herbicide-resistant minor crops has been non-existent (Devine, 2005). Given the reduction
in new herbicide molecules being discovered and labelled for use in any
crops, it is unlikely that new herbicides for use in lentils will be developed
without the development of herbicide-resistant lentils.
Even if herbicide-resistant technology develops for use in lentils, growers must learn to effectively control weeds using integrated weed management. Herbicide-resistant lentils might serve to limit cultivar development
of lentils and limit the germplasm available with resistance to certain diseases and pests (Devine, 2005). Heavy use of a single herbicide would tend
to be increased through the use of herbicide-resistant crop systems and it
would be likely that herbicide-resistant weeds would develop or there
would be a weed species shift (Ball, 1992). Thus, the development of additional herbicides for lentils through transgenic means or otherwise, are
unlikely to produce a sustainable system of lentil production by itself. Integrating existing practices is the most likely answer to sustained lentil production within a cropping system.
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21
Parasitic Weeds
D. Rubiales,1 M. Fernndez-Aparicio1 and A. Haddad2

1Institute
for Sustainable Agriculture, CSIC, Crdoba, Spain; 2International Center for

Agricultural Research in the Dry Areas (ICARDA), Aleppo, Syria
21.1. Introduction
Over 4000 species of angiosperms are able to directly invade and parasitize
other plants. A few of them are weedy and parasitize cultivated plants.
They belong to various plant families, and attach to host roots, shoots or
branches. Aerial parts of lentil plants can be infected by dodders (Cuscuta
spp.), whereas roots can be infected by broomrapes (Orobanche spp.) causing significant damage.
21.2. Dodders
Dodders have a broad host range, being widely distributed. The most
important species is Cuscuta campestris Yuncker that is a problem in some
areas of the Middle East, and a threat to lentil in certain locations. It has
fine, yellow or orange, thread-like branches which grow and entwine
around the stems and other aboveground parts of the host (Plate 5AC).
The stem of the parasite is tough, curling and leafless and bears only minute scales in place of leaves. Cuscuta hyalina Roth. has also been observed on
lentil in India.
Grey to reddish-brown seeds are produced in abundance and mature a
few weeks after bloom. Seeds may germinate immediately after they fall to
the ground, or may remain dormant until the following season. Dodderinfested areas appear as patches in the field and continue to enlarge during
the growing season (Plate 5A). In late spring and early summer, dodder
produces a massed cluster of white flowers. Infected host plants become
weakened, decline in vigour and produce poor yields. Several patches
might coalesce to form large areas that can be easily recognized by the yellowish colour of the parasite strands.
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D. Rubiales et al.
21.3. Broomrapes
Broomrapes are holoparasitic plants, completely dependent on the host due
to the lack of chlorophyll. Once attached on the host roots, broomrapes are
able to survive and develop, sucking carbohydrates from the phloem and
water and minerals from the xylem of the host by means of a bridge to the
host vascular tissues.
Lentil can be parasitized mainly by two different species of broomrapes,
namely crenate broomrape (Orobanche crenata Forsk.) (Plate 5D, E) and
Egyptian broomrape (Orobanche aegyptiaca Pers., syn. Phelipanche aegyptiaca
(Pers.) Pomel) (Plate 5F).
Orobanche crenata has been known to threaten legume crops since antiquity. This parasitic weed is mainly restricted to the Mediterranean basin,
southern Europe and the Middle East, and is an important pest in grain and
forage legumes, as well as in some apiaceous crops such as carrot and celery. Sauerborn (1991) estimated that over 1 million ha of faba bean in the
Mediterranean region and western Asia are infested or at risk from O. crenata. Dispersion of seeds is easy by machinery, wind, water or mixed with
crop seeds, but populations of O. crenata will only establish in areas with a
distinct precipitation seasonality, with a warm and dry period being followed by warm and wet conditions and when host crops are grown under
this warm and wet period. Climatically suitable regions include all Mediterranean climate areas and part of the monsoon, savannah and winter-dry
climate regions of Central America, Africa, Australia and South Asia (Grenz
and Sauerborn, 2007). There is evidence that O. crenata is expanding to new
areas (Rubiales et al., 2008). Losses up to 95% have been reported for lentil
(Sauerborn, 1991) depending on the infestation level and the planting date.
Orobanche crenata is characterized by large erect plants, branching only from
their underground tubercle. The size of spikes is influenced by host plant
vigour. Orobanche crenata plants infecting lentils are 1040 cm tall, bearing
many flowers of diverse pigmentation, from yellow, through white to pink
and violet. The calyx is 1318 mm, with segments free and bidentate. The
corolla is 1828 mm, glandular pubescent, the lips large and divergent, not
ciliate but often with lilac veins. The anthers are brown, glabrous or subglabrous. The filaments insert 23 mm above the base of the corolla, and are
hairy at the base with glandular hair at the apex (Pujadas-Salv, 1999).
Orobanche aegyptiaca parasitizes lentil and other legumes, but also many
other crops such as tomato, potato, cabbage, sunflower, parsley or watermelon (Joel et al., 2007). It is widely distributed in eastern parts of the Mediterranean, in the Middle East and in parts of Asia. It is very similar to
Orobanche ramosa L. (syn. Phelipanche ramosa (L.) Pomel) that is more common as a weed in Europe, parts of Africa and Asia, with some overlap with
O. aegyptiaca in the northern parts of the Middle East. However, O. aegyptiaca is more aggressive and adapted to more arid regions. Both species have
branched stems with flowers that may significantly differ in colour, from
white to dark blue. A simple feature that allows field distinction is the hairiness of the stamens at the connective, which occurs only in O. aegyptiaca.
Parasitic Weeds
345
The upper leaves and the corolla are hairy glandular and the lobes of the
lower lip of corolla (2035 mm) are rounded.
Orobanche foetida is another broomrape species that can infect legume
crops in Tunisia. Fortunately, it is not yet reported infecting lentil, and lentil
germplasm is highly resistant (Fernndez-Aparicio et al., 2008b).
Broomrapes propagate via seeds. Broomrape seed germination occurs
only in response to a chemical signal from the host root. Before germination, broomrape seeds must undergo conditioning under suitable temperature and moisture conditions. Optimum temperatures for conditioning and
germination are different among broomrape species, being about 18C for
O. crenata and about 23C for O. aegyptiaca, with substantial differences
among populations within the species.
Seeds are very small, approximately 0.200.35 mm long. During germination, only a radicle emerges out of the tiny seed (Plate 5G). The small
storage reserves, mainly composed of lipids, are capable of supporting a
few days of autonomous growth of the emerging radicle that can grow only
to a limited extent (few millimetres). When a root host is reached the radicle
tip develops into an attachment organ, and a haustorium is formed. Subsequently the parasite develops a tubercle (Plate 5H, I) that grows underground on the host root surface for several weeks or months. At maturity,
the parasite develops flowering shoots (a single non-branching shoot in the
case of O. crenata, branched in O. aegyptiaca) that emerge above the soil near
host plants and set seeds (Plate 5E, J). In contrast to other broomrapes that
are self-pollinating, both O. crenata and O. aegyptiaca are believed to be at
least partially outcrossing being pollinated by insects, although they will
self-pollinate if not visited by insects. The number of seeds produced by a
healthy O. crenata or O. aegyptiaca plant can exceed 200,000. Large quantities
of long-lived seeds assure the parasite genetic adaptability to changes in
host resistance and cultural practices. The minute seeds may easily be transferred from one field to another by cultivation, and also by crop contaminated seeds, water, wind, animals, and especially by vehicles and farming
machines. Broomrape seeds may remain viable in soil for decades. Harvesting parasite shoots in lentil straw and forage crops may also assist in spreading the seeds if used to feed animals, because their manure may then be
contaminated with parasite seeds, which remain viable after passing
through the alimentary system of animals.
21.4. Management
Several strategies have been developed for the control of parasitic plants,
from cultural practices to chemical control (Parker and Riches, 1993; Joel
et al., 2007), but none with unequivocal success. Proposed methods were
not feasible, uneconomical or resulted in incomplete protection. So far, the
effectiveness of conventional control methods is limited because of the
numerous factors involved, in particular the complex nature of the parasites, which reproduce by tiny and long-living seeds, and that, in the case of
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D. Rubiales et al.
broomrapes, are difficult to diagnose until they irreversibly damage the

crop. The intimate connection between host and parasite also hinders efficient control by herbicides.
The efficient management of dodders starts with the use of dodder-free
seed and the spraying of the first affected patches with a contact herbicide
to prevent spread. Infested equipment should be cleaned before and after
use, and the movement of domestic animals from infested to dodder-free
areas should be limited. The application of pendimethalin at 0.71.0 kg
active ingredient (a.i.)/ha pre-emergence or early post-emergence is effective against Cuscuta in lentil.
The only way to cope with the broomrapes is through an integrated
approach, employing a variety of measures in a concerted manner, starting
with containment and sanitation, direct and indirect measures to prevent
the damage caused by the parasites, and finally eradicating the parasite
seedbank in soil. So far, a combination of sowing Orobanche-free lentil seeds
and delaying the sowing date has been the only available strategy for controlling broomrape infection. Autumn-sown cultivars have a greater yield
potential in dry areas than spring-sown cultivars, but early sowing is known
to increase the incidence of O. crenata (Silim et al., 1991). An alternative
strategy could be the use of very early maturing cultivars that could escape
O. crenata damage due to early pod filling, such as in the lentil cultivars
Chaouia and Abda.
Seedbank demise can efficiently be achieved only for the cultivated soil
layer by fumigation or solarization, but once soil is disturbed by deep ploughing, deeply buried broomrape seeds will affect the roots when brought into
the lentil root area. However, this is uneconomic in a low-value crop like lentil. Some biological control agents have shown promise, but the technology is
not ready yet to provide acceptable control.
Chemical control of broomrape is complicated as herbicides have been
effective only as a prophylactic treatment, since in most cases the actual
infestation level in the field is usually unknown. Also, if the herbicide is to
be applied to the parasite through the host via its conductive tissues, then
the host should show some tolerance to the herbicides phytotoxicity. Herbicide treatments have to be adapted to the broomrape life cycle that is very
much affected by sowing dates and climatic factors. Application must be
repeated in a time interval of 24 weeks, because Orobanche seeds may germinate throughout the season, and may therefore re-establish on newly
developed host roots. This would also minimize lentil phytotoxicity by
using only a few tolerated applications.
It has been shown that good broomrape control can be achieved in faba
bean by glyphosate at low rates. However, insufficient selectivity is found
in lentil, although post-emergence treatments of 3040 g a.i./ha can be tolerated by some cultivars but the herbicide still reduces yield. Lentil tolerates pre-emergence treatments of imazapyr (25 g a.i./ha) and imazethapyr
(75 g a.i./ha) and post-emergence treatments of imazaquin (7.5 ml a.i./ha)
(Arjona-Berral et al., 1988; Jurado-Expsito et al., 1997) and imazapic (3 g
a.i./ha) (Bayaa et al., 2000). Two post-emergence applications of imazapic
Parasitic Weeds
347
are recommended, with the first application recommended when lentil has
between five and seven true leaves, which is when broomrape usually starts
to develop attachments on lentil roots, followed by a second application 23
weeks later. Lentil may need a third application of 2 g a.i./ha when late
rain comes and extends the growth period, or when lentil is supplementary
irrigated (Bayaa et al., 2000). Phytotoxicity might be higher in the presence
of water, low temperature and heat stresses and varies with the cultivar of
lentil used (Hanson and Hill, 2001). An additional problem of these imidazolinone herbicides for broomrape control is that they are not registered in
every country, and that doses and timing for application need to be adjusted
case by case. Also, traditional imidazolinones are being replaced in some
countries by imazamox that is less residual in the soil, so doses and timing
of application need re-adjustment.
An alternative to cultivars resistant to parasitic weeds is the development of cultivars resistant to herbicides. This can be achieved by genetic
engineering or simply by induced mutation. This second option has been
successful and a number of cultivars are being introduced to the market
under the trademark CLEARFIELD lentils (BASF Agsolutions, 2008) that
are not genetically modified and that tolerate higher doses of imidazolinone
herbicides.
Crop rotation is of little value because of the persistence of the seeds for
extended periods and the broad host range. However, there is promise in a
number of other strategies. For example suicidal germination by application of germination stimulants has shown some success under laboratory
conditions, although so far there is no evidence of success in the field.
Another strategy is the use of trap or catch crops, which have yet to be
proven successful and economical. An alternative might be intercropping,
which is widely recommended for use for Striga control on maize and sorghum in Africa, and has recently been shown to reduce O. crenata infection
in legumes (Fernndez-Aparicio et al., 2007).
Resistance against parasitic weeds is difficult to access, scarce, of complex nature and of low heritability, making breeding for resistance a difficult task (Rubiales et al., 2006). In spite of these difficulties, significant
success has been achieved in some other crops, such as sunflower against
Orobanche cumana, or faba bean and vetch against O. crenata (Rubiales et al.,
2006; Joel et al., 2007). However, no resistant lentil cultivars are available.
Only very recently, sources of resistance against O. crenata have been
reported in lentil germplasm (Fernndez-Aparicio et al., 2008) and in wild
Lens accessions (Fernndez-Aparicio et al., 2009). Accessions have been
identified that escape infection because of their lower root density (Sauerborn et al., 1987), that induce lower seed germination, or that hamper establishment of broomrape radicles in contact with host roots thus limiting
development of established tubercles. In addition, necrosis of tubercles was
observed in some accessions. This can be exploited in lentil breeding. Combining different resistance mechanisms into a single cultivar should provide
a more durable outcome. This can be facilitated by the adoption of markerassisted selection techniques, together with the use of in vitro screening
348
D. Rubiales et al.
methods that allow dissecting parasitic weed resistance into highly heritable
components.
References
Arjona-Berral, A., Mesa-Garca, J. and Garca-Torres, L. (1988) Herbicide control of
broomrapes in peas and lentils. Food and Agriculture Organization (FAO) Plant
Protection Bulletin 36, 17578.
BASF Agsolutions (2008) Available at: http://www.agsolutions.ca/basf/agprocan/
agsolutions/WebASClearfield.nsf/mainLentils.htm (accessed 1 June 2008).
Bayaa, B., El-Hossein, N. and Erskine, W. (2000) Attractive but deadly. ICARDA
Caravan 12. Available at: http://www.icarda.cgiar.org/publications1/caravan/
caravan12/Car128.Html (accessed 1 June 2008).
Fernndez-Aparicio, M., Sillero, J.C. and Rubiales, D. (2007) Intercropping with cereals
reduces infection by Orobanche crenata in legumes. Crop Protection 26, 1166
1172.
Fernndez-Aparicio, M., Sillero, J.C., Prez-de-Luque, A. and Rubiales, D. (2008)
Identification of sources of resistance to crenate broomrape (Orobanche crenata)
in Spanish lentil (Lens culinaris) germplasm. Weed Research 48, 8594.
Fernndez-Aparicio, M., Sillero, J.C. and Rubiales, D. (2009) Resistance to broomrape
in wild lentils (Lens spp.). Plant Breeding (in press).
Grenz, J.H. and Sauerborn, J. (2007) Mechanisms limiting the geographical range of
the parasitic weed Orobanche crenata. Agriculture, Ecosystems and Environment
122, 275281.
Hanson, B.D. and Hill, D.C. (2001) Effects of imazethapyr and pendimethalin on lentil
(Lens culinaris), pea (Pisum sativum), and a subsequent winter wheat (Triticum
aestivum) crop. Weed Technology 15, 190194.
Joel, D.M., Hershenhorn, Y., Eizenberg, H., Aly, R., Ejeta, G., Rich, P.J., Ransom, J.K.,
Sauerborn, J. and Rubiales, D. (2007) Biology and management of weedy root
parasites. In: Janick, J. (ed.) Horticultural Reviews. Vol. 33. John Wiley and Sons,
Hoboken, New Jersey, USA, pp. 267350.
Jurado-Expsito, M., Garca-Torres, L. and Castejn-Muoz, M. (1997) Broad bean
and lentil seed treatments with imidazolinones for the control of broomrape
(Orobanche crenata). Journal of Agricultural Science 129, 307314.
Parker, C. and Riches, C.R. (1993) Parasitic Weeds of the World; Biology and Control.
CAB International, Wallingford, Oxon, UK.
Pujadas-Salv, A.J. (1999) Species of the family Orobanchaceae parasitic of cultivated
plants and its relatives growing on wild plants, in the South of the Iberian
Peninsula. In: Cubero, J.I., Moreno, M.T., Rubiales, D. and Sillero, J.C. (eds)
Resistance to Orobanche: the State of the Art. Publisher Junta de Andaluca,
Sevilla, Spain, pp. 187193.
Rubiales, D., Prez-de-Luque, A., Fernndez-Aparicio, M., Sillero, J.C., Romn, B., Kharrat,
M., Khalil, S., Joel, D.M. and Riches, C. (2006) Screening techniques and sources
of resistance against parasitic weeds in grain legumes. Euphytica 147, 187199.
Rubiales, D., Fernndez-Aparicio, M. and Rodrguez, M.J. (2008) First report of crenate
broomrape (Orobanche crenata) on lentil (Lens culinaris) and common vetch
(Vicia sativa) in Salamanca province, Spain. Plant Disease 92, 1368.
Sauerborn, J. (1991) Parasitic Flowering Plants: Ecology and Management. Verlag Josef
Margraf, Weikersheim, Germany.
Parasitic Weeds
349
Sauerborn, J., Masri, H., Saxena, M.C. and Erskine, W. (1987) A rapid test to screen
lentil under laboratory conditions for susceptibility to Orobanche. LENS Newsletter
14, 1516.
Silim, S.N., Saxena M.C. and Erskine, W. (1991) Effect of sowing date on the growth
and yield of lentil in a rainfed Mediterranean environment. Experimental Agriculture
27, 145154.
22
Seed Quality and Alternative Seed

Delivery Systems
Zewdie Bishaw,1 Mohamed Makkawi2 and Abdoul Aziz Niane1

1International
Center for Agricultural Research in the Dry Areas (ICARDA),

Aleppo, Syria; 2Dhaid Research Station, Ministry of Environment and Water, Dhaid,
Sharjah, United Arab Emirates
22.1. Introduction
Lentil has been traditionally cultivated in South and West Asia, North and
East Africa since its domestication. It spread from the Near East to the Mediterranean and then to Africa, Asia, Europe, America and lately to Australia
owing to its plasticity in adaptation. Lentil has been introduced to developed countries as part of crop diversification, where traditionally production is dominated by cereals. Although Australia and North America
(Canada and the USA) are becoming major global players in production
and export of lentil, Asia remains the major producer, consumer and
importer of lentil, with India being the single most important country in
terms of production and consumption.
Modern varieties and seed technology have a significant role in the transformation of agriculture. The availability, access and use of quality seed of
adaptable crop varieties are critical to realize the impacts of investments in
agricultural research. Despite a well-functioning lentil seed sector in developed
countries the situation falls far short of the desired level in many developing
countries. Lack of access to seeds of improved varieties, mechanization problems, high production costs, etc. are some of the major constraints hindering
the development of the lentil seed industry. In a nutshell, both the public and
the private sectors do not pay sufficient attention to lentil seed provisions.
22.2. What is Seed Quality?

Seed is a primary input in crop production and is a means for delivering
crop-based innovations to farmers to realize the impacts of investments in
agricultural research. Since seed is one of the main factors limiting crop
production potential it should reach farmers in a good quality state.
350

Seed Quality and Alternative Seed Delivery Systems
351
Seed quality is complex and determined by many factors (Fougereux,

2000), but four key attributes may be explicitly identified (Bishaw et al., 2007):
1. Genetic quality the inherent genetic potential of the variety for higher
yield, better grain quality and greater tolerance to biotic or abiotic stresses.
2. Physiological quality the potential germination and vigour leading to
subsequent seedling emergence and crop establishment in the eld.
3. Physical quality free from contamination with other crop seeds, common, noxious or parasitic weed seeds, mechanical damage, discoloration,
and with uniform seed size and seed weight.
4. Health quality absence of infection/infestation with seed-borne pests
(fungi, bacteria, viruses, nematodes, insects, etc.).
Genetic purity
McNeil et al. (2007) reported that in the language of trade and commerce in
the Western countries, two types of lentils can be identified based on the
cotyledon and seedcoat colour: red lentil (orange cotyledon and pale grey
to dark seedcoat) and green (yellow cotyledon with green to brown seedcoat). This terminology is not followed in the major lentil-growing region of
Africa and Asia. Moreover, genotypes with pure green cotyledons (Emami
and Sharma, 1996a, b; Sharma and Emami, 2002; Sharma, Chapter 7, this
volume) and a range of seedcoat colours may become available for consumption and trade. The existing grouping into red and green lentils will
then become totally redundant. Genetic or varietal purity is crucial for grain
quality and uniformity of grain traded in bulk because of differences in consumer preferences. A mixture of lentil grain with different cotyledon colours
will certainly lower grain quality and hence its market value. Since the crop
stand in lentil is usually dense, plant canopy is short and cotyledons are
concealed, the best practical procedure to maintain varietal purity is the use
of seed with high genetic purity for sowing.
Physical purity
Use of physically pure seed is always desirable for lentil production. Contaminated seed is a major vehicle for introducing and spreading parasitic
weed such as Orobanche and Cuscuta into new areas and fields (see Rubiales
et al., Chapter 21, this volume). These two weeds have attained great economic significance in the Middle East although they are not a major issue in
the countries of the Orient. For this reason, Orobanche and Cuscuta are considered as quarantine pests and a reason for rejecting seed lots in most certification schemes. However, seed cleaning using dodder mill and magnetic
separator machines can eliminate Cuscuta from infested seeds, but these
procedures are less practicable for large-scale application in lentils.
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Z. Bishaw et al.
Germination
Like most field crops, physically sound lentil seed germinates well under
laboratory conditions. Moreover, lentil is among those crops in which laboratory germination is reliable for predicting field emergence (Makkawi
et al., 1999). This indicates that if proper care is taken, germination may not
be a major constraint in lentil production.
Seed vigour
Seed vigour is defined as the capacity of a seed lot to germinate and produce normal seedlings under various field conditions (i.e. the rate and uniformity of seedling emergence and stand establishment in the field). It is a
quantitative character, controlled by several factors including mechanical
injury to embryo or seedcoat, environment and nutrition of the mother plant,
stage of maturity at harvest, seed size, senescence, and attack by pathogens.
Seed health
Seed health has a significant effect on germination and seedling establishment leading to low germination, reduced crop establishment, severe yield
loss or total crop failures. Lentil Ascochyta blight, which is also seed-borne,
anthracnose and Botrytis can cause seed death and result in seedlings with
necrosis or seedling blight. The infected seedlings die soon after emergence
and reduce the plant stand, with significant consequences on the crop yield.
For example, Ascochyta lentis has been associated with low seedling emergence in lentil (Morrall and Sheppard, 1981). Lentil plants from Ascochytainfected seeds had fewer branches, smaller roots and shoots, reduced vigour
and lower seed yields (Kaiser and Hannan, 1987).
22.3. Factors Affecting Seed Quality

Lentil seed quality can be extensively affected by several factors such as the
production environment, agronomic practices and handling operations during production. Apart from genetic factors, environmental conditions during crop growth, seed development and maturity (soil conditions, nutrient
deficiency, water stress, extreme temperatures, biotic stresses), harvesting
(time, maturity, moisture), mechanical damage (during harvesting, conveying, cleaning, treatment, packaging) and improper storage (insects, moisture content, temperature) have their effect on seed quality.
Genetic factors
Seed quality can be affected by genetic factors as differences exist among
species and varieties. Lakhani et al. (1986) found genetic variability in seed
353
vigour and hard seed among lentil genotypes. There is also evidence that
seedcoat thickness has an effect on germination and emergence in lentil
(Sarker et al., 1995). Differences were also detected among lentil varieties in
their response to ageing treatments (Makkawi and van Gastel, 2006). The
results showed that high-vigour varieties are capable of withstanding ageing treatments for longer duration compared to the low-vigour varieties.
Therefore a vigour test could be used in breeding programmes to select
high-vigour genotypes with greater potential for storability.
The physical integrity of seed also determines seed quality. Hence, testa
quality is one of the principal components of legume seed quality and is
influenced by harvesting. Mechanical damage influences the conditions of
testa. The degree of cracking from a given handling treatment depends on
seed moisture content. Mechanical injury exposes the seed to insect infestation and increases water absorption during storage, resulting in poor germination, reduced vigour and poor seedling establishment. Physiological studies
showed that lentil seeds are very sensitive to injury during imbibition,
especially at high temperatures and high humidity (Makkawi et al., 2008).
Production environments
Climatic conditions during the pre-harvest period have greater influence on
seed quality. Lentil produces good quality seed in semi-arid or drier areas.
High humidity and high rainfall during the season encourage excessive
vegetative growth and reduce yield and seed quality. Excessive drought
and/or high temperatures during flowering and the pod-fill period cause
flower drop and reduce yields. Lentil is sensitive to heat and drought
stresses and exposure to these conditions during crop growth causes a considerable reduction in yield (Johansen et al., 1994). It was reported that high
temperatures during seed development in the field can dramatically reduce
seed quality (Hampton and Coolbear, 1990).
Harvesting
The time and method of harvesting also influence seed quality. Delaying
harvest subjects the seed to more deteriorative field stress and causes
greater loss of quality (increases physiological age). Moreover, delaying the
harvest leaves the mature seeds under weathering conditions, which may
reduce seed germination significantly. It was reported that delay in harvest
by 1 week reduces germination by 20% in lentils (Ellis et al., 1987).
Mechanical damage
Mechanical damage is one of the primary causes for significant loss in seed
quality. Mechanical damage can be caused during harvesting, cleaning and
handling. Lentil seeds, due to their convex surface on both sides, are more
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Z. Bishaw et al.
vulnerable to mechanical damage than seeds with a rounded shape such as

faba bean, pea or chickpea (Muehlbauer et al., 1985). The net result is broken or cracked embryos, split or chipped seed, and ultimately abnormal
seedlings. The damage is greater when the moisture content at the time of
mechanical constraint is low. Further investigation on the effect of moisture
content showed that the breakage percentage increases with a decrease in
moisture content below 14% and also when the moisture content is higher
than 20% (Fougereux, 2000).
22.4. Lentil Seed Treatment for Better Field Performance

Lentil is grown under harsh rainfed conditions such as drought, heavy clay
soil, extreme temperature and variable soil moisture, hence field establishment is a key determinant for crop production. Rapid and uniform germination with subsequent seedling development and crop establishment are
considered important factors influencing yield potential, since the plant has
limited ability to compensate for low populations. Therefore, knowledge of
seed vigour, how it is acquired during seed development and subsequently
lost through deterioration, is relevant to plant breeding, seed production,
quality control and marketing. Several seed treatment options are associated with a productive crop as described below.
Seed priming
Seed priming, an ancient practice of soaking the seed with water for a specific period of time prior to planting, has been shown to improve crop establishment and crop yields in dry areas. Seed priming, controlled hydration
and dehydration of seed, pelleting, etc., are used extensively to increase the
rate and uniformity of seedling establishment of commercial crops (Ali
et al., 2005). Harris (2006) gave an excellent review of on-farm seed priming
including improved nutrition from Asia and Africa for various crops such
as cereals (wheat, barley, rice, millet, sorghum, maize), legumes (chickpea,
lentil, mungbean), bambara groundnut and cotton. Neupane (2002) reported
that soaking seeds for 12 h followed by drying in the shade for 2 h was the
best combination for lentil. During 2-year (season) trials priming reduced
the average emergence time to 50% (from 9.4 to 7.8 days) and increased
average grain yield by 34%. Gupta and Bhowmick (2005) also reported that
priming lentil seed increased plant stand and yield significantly. From Bangladesh, where lentil is almost totally cultivated as a rainfed crop, Ali et al.
(2005) reported after 2 years of experimentation that seed priming (water
soaking) for 810 h increased the crop stand (from 74 to 85 plants/m length),
plant height (from 34.4 to 40 cm), branches per plant (from 3.47 to 4.91),
pods per plant (from 61 to 75.5), 1000-grain weight (from 17.19 to 17.59 g)
and yield per ha (from 1528 to 2098 kg/ha, i.e. by 37.3%) in the variety Barimasur 2. This variety gave a better response to seed priming than Barimasur
4. This suggests that response to priming is genetically controlled.
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Rhizobial seed treatment

Lentil seed must be inoculated at planting time with specific rhizobium
(Gan et al., 2005), particularly if they are planted in new lentil-growing
areas. In India, under a maize-lentil cropping sequence, seed inoculation
with rhizobium increased yield by 20.3% over uninoculated seed (Sardana
et al., 2006). Huang and Erickson (2007) found that seed treatment with Rhizobium leguminosarum bv. viceae is effective in controlling Pythium damping-off
disease.
Chemical seed treatment

Lentil is affected by several seed-borne pests. Chemical seed treatment
becomes a standard procedure for disease control in many crops. Lentil diseases and control measures have been reviewed by Taylor et al. (2007) and
Kaiser et al. (2000). Seed treatment with benomyl, carbendazim, carabthiin,
ipodion or thiabendazole showed varying levels of efficacy against various
diseases. In Spain, seed treatments with imazapyr did not affect germination or crop growth but controlled 8595% of broomrape (Orobanche crenata
Forsk.) and increased crop biomass and seed yield (Expsito et al., 1997).
22.5. Requirements for Quality Seed Production

In seed production, maintaining quality is essential if the variety is to meet
the expectation of farmers and consumers. Special attention should be given
to ensure the genetic, physical, physiological and health quality. Following
a combination of key regulatory, administrative and technical control measures and procedures can ensure seed quality.
The regulatory measures establish a framework for variety release, set
up field and seed standards, and enforce the regulations and standards
through an impartial quality assurance authority. The agency establishes
administrative guidelines and technical procedures to implement robust
quality assurance systems. Bishaw et al. (2007) summarized the key technical components for producing quality seed. They also gave a detailed
account of lentil seed production from variety maintenance through to field
production, cleaning, treatment, storage and quality assurance.
22.6. Status of the Lentil Seed Industry

Yadav et al. (2007) reported that 20 countries accounted for almost 99% of
world lentil production from 2001 to 2005. The top ten countries include
Australia, Canada and the USA from developed countries and Bangladesh,
China, India, Iran, Nepal, Syria and Turkey from developing countries. The
second tier countries comprise Ethiopia (12th), Morocco (13th), Pakistan
356
Z. Bishaw et al.
(14th) and Yemen (20th) from the dry areas of Central and West Asia and
North Africa (CWANA) region. The top lentil producers remain India followed by Turkey from the developing countries and Canada followed by
Australia from the developed countries.
Several authors have provided detailed accounts of diverse lentil production systems (Materne and Reddy, 2007; Yadav et al., 2007). At least two
contrasting lentil production systems exist in the developed and developing
countries. Some of the key distinguishing features include the scale, degree
and level of farm operation, mechanization, processing (value addition),
commercialization and integration in the lentil seed industry. In developed
countries (e.g. Australia, Canada and the USA) lentil is grown as a crop for
export, characterized with high adoption of improved varieties and production technologies. In the traditional lentil-producing developing countries
(e.g. India, Iran, Syria and Turkey) lentil is grown as a subsistence crop primarily for local consumption with limited use of improved varieties and
technologies. The status of the lentil seed industry in the developed and
developing countries is presented below.
Developed countries
Australia, Canada and the USA are the major global players in production
and export of lentil, overtaking traditional producers (and exporters) from
developing countries such as India, Iran, Syria and Turkey. In the developed
countries, lentil has been introduced as part of an agricultural diversification
programme in a cereal-dominated production and export market. Bishaw et al.
(2009) summarized key features that distinguish the legume seed industry in
the developed and developing economies, which is equally applicable to lentils. First, in developed countries, lentils are produced on a large scale (>200 ha
by individual farmers) and considered as an export commodity to diversify
the product portfolio replacing cereal crops. Legume production is market oriented where farmers are willing to adopt technology and adjust production
factors accordingly compared to subsistence production in the developing
countries. Second, there is strong public-private partnership where both the
government and the private sector are funding agricultural research and variety development (Gareau et al., 2000). Accordingly, farmers in Australia and
Canada pay a levy on total production or the amount of grain marketed which
is matched or supplemented by the government or the industry to support
research and commercialization. A similar approach was initiated in Turkey
(Ksmenoglu, 2003). Third, there is strong integration of agricultural
research, seed production and seed use where the private seed companies
and/or farmers play a key role in commercialization of released varieties.
Fourth, although legumes are inherently low-profit crops, the availability of
good infrastructure and socio-economic conditions ensure a reasonable profit,
thus providing sufficient incentive to attract private sector investment.
The presence of market-oriented and/or export-led commercial agriculture remains the major driving force for the development of a vertically
357
organized and sustainable lentil seed industry. The net result is that agricultural research, variety development, seed production and marketing,
and grain processing and marketing are well integrated across a wide spectrum of the legume industry (i.e. the government, farmer growers, industrial
processors and exporters).
Developing countries
Among the developing economies India, Iran, Nepal, Syria and Turkey are
leading lentil producers. Lentil production is largely subsistence and produced by small-scale farmers (holding size <10 ha) with the primary objective of household consumption and a little surplus for market. Lentil
production is characterized by a low level of mechanization, minimal use of
technology-related inputs (e.g. seed, fertilizer, irrigation, rhizobial treatments and agrochemicals such as herbicides, etc.) and is mostly in dry rainfed environments.
In most developing countries, in view of national food security the formal seed sector focuses on major food crops at the expense of low-priority
legume crops (e.g. lentil). For example, in Ethiopia and Syria the seed industry for all food crops is handled by the public sector, but from the total
amount of seed supplied, legumes, including lentil, account for less than
5% compared to a single major cereal crop like wheat which accounts for
almost 70% of seed production (Bishaw, 2004). Farmers, particularly in dry
areas, not only lack access to modern varieties and good quality seed, but
may also not be able to afford to pay a higher price for seed, and therefore
use their own seed or any seed, even commercial grain, purchased from
local markets or traders for planting. In India, the leading global lentil producer, more than 95% of lentil seed comes from the informal sector (Materne
and Reddy, 2007). Similarly in Turkey, a large proportion of seed for planting comes from the informal sector and a very small fraction from a formal
sector with no or limited private sector involvement (Ksmenoglu, 2003).
There is a missing link between public plant breeding and public seed
delivery. As lentils are considered to be a low-profit crop by the private sector, organized seed production did not show a better record even in countries where privatization of the seed sector has made some progress (e.g.
India, Morocco, Turkey). There are many policy, regulatory, institutional,
organizational, technical and socio-economic constraints that affect the seed
industry of legumes in general and lentil in particular in the developing
countries.
Constraints to the lentil seed industry

Bishaw et al. (2009) summarized the constraints of the legume seed industry
which also apply to lentils in the developing countries. Apart from policy
and regulatory constraints there are technical constraints which hinder the
358
Z. Bishaw et al.
lentil seed sector including the availability, access and use of varieties and
seeds, agricultural mechanization, high disease pressure (diseases, pests,
parasitic weeds), high seed production costs, etc.
Availability of quality seed

Table 22.1 shows seed production and distribution from a few selected
CWANA countries. For example Ethiopia, Iran, Morocco, Pakistan, Syria
and Turkey are major lentil producers in their respective regions (see
Erskine, Chapter 2, this volume). However, the amount of seed distributed
by the formal sector remains insignificant compared to cereals. In almost all
countries, lentil seed production accounts for less than 5% of the formal sector seed supply. Moreover, the amount of seed supplied compared to
national demand is pitiable. In Ethiopia the amount of seed distributed by
the formal sector met only 11.2% of demand for lentil seed in 2005, and this
was because of specific government intervention which was otherwise nonexistent earlier (e.g. 0.7% in 2004). Similarly, in Morocco the formal sector
on average provides 7% of seed for food legumes (1% each for faba bean,
chickpea and lentil and 34% for peas). Lentil seed supply is not only low,
but also fluctuates between years. For example in Syria lentil seed supply
from the formal sector was 1.2 and 11.2% in 2003 and 2004, respectively.
Low commercial seed supply was also reported in Morocco, Syria,
Tunisia and Turkey (Oram and de Haan, 1995; Ksmenoglu and Kugbei,
Table 22.1.
region.
Seed production and/or distribution (t) in selected countries of the CWANA
Lentil (%)
Countrya
Totalb
Cereals
Legumes
CSFLc
Lentil
Total
Legumes
Ethiopia*
Morocco*
Pakistan*
Syria*
Turkey**
Yemen**
Total
13,656
67,620
191,043
104,137
263,105
1,643
641,204
11,929
64,900
188,582
102,364
260,688
1,643
630,106
1,628
2,460
1,011
1,768
66
6,933
1,380
2,460
572
1,768
729
60
818
5
1,612
5.3
0.1
0.8
0.002
0.3
44.8
2.4
46.3
7.6
23.3
aFigures
6,180
are from different years (*for 2005, **for 2004) and sources (from unpublished internal reports,
documents, presentations and personal communications with seed sector institutions in the respective
countries including: Ethiopian Seed Enterprise (ESE); Service du Controle des Semences et des Plantes
(DPVCTRF), Morocco; Federal Seed Certication and Registration Department (FSCRD), Pakistan; General Organization for Seed Multiplication (GOSM), Syria; and General Corporation for Seed Multiplication
(GCSM), Yemen).
bThe total includes cereal, legume and oilseed crops, whereas CSFL comprise faba bean, pea, chickpea
and lentil seed only.
cCSFL, cool season food legumes.
359
2000). Materne and Reddy (2007) also cited over 95% seed supply from the
informal sector as a major constraint for lentil production in India. There is
no reliable information on lentil seed supply in some other major lentilproducing countries like Iran, Pakistan and Yemen.
Mechanization problems
Lentil production has been mechanized in the developed countries.
Despite the availability of mechanical operations elsewhere lentil is still
under a traditional production system with little mechanization in the
developing countries (Sarker and Erskine, 2002). Such technical constraints,
for example, limited contractual lentil seed production with state farms in
Ethiopia. Traditional planting and harvesting methods still continue to
persist among small-scale farmers who lack the resources to adopt new
technologies.
High production costs

Legume production in general, and lentil in particular, requires regular
sprays for diseases and parasitic weeds, need for special equipment for
planting and harvesting, etc., which add to the cost of production of a crop
which is already at a low level of profitability. Among production costs,
harvesting alone contributes more than half of the total cost in some countries. It is reported that hand harvesting has led to a reduction in area under
lentil due its high cost (Erskine et al., 1991). Oram and de Haan (1995) cited
high seed cost as a disincentive against adoption of improved varieties. All
these technical problems combined with price uncertainty exacerbate the
problem of the lentil seed industry.
22.7. Alternatives for Lentil Seed Delivery

Substantial increase in lentil production and productivity can be achieved
only with the use of quality seed of improved varieties. A concerted effort
is required to develop a strategy to achieve sustainable lentil production
through conventional and participatory approaches in plant breeding and
formal and farmer-based seed provisions. The demand for lentil seed is
diverse because of diverging interests, production environments and differences between socio-economic conditions of commercial and subsistence
farmers. Moreover, lentil is a high-volume, less profitable crop and neither
the public sector nor the private sector is willing to invest in seed production and marketing, particularly in the developing countries. An innovative
alternative approach needs to be designed to ensure sustainable lentil seed
supply to farmers.
360
Z. Bishaw et al.
Formal seed supply

In the developed countries, the success of the cereal and legume seed industry has often resulted from integration of agricultural research, production
technology, input supply, market support and extension information.
According to Byerlee and White (2000), the success and rapid expansion in
soybean production is attributed to investments in research, effective extension programmes, price support, encouragement to the processing industry,
and facilitating export markets. They suggested that similar efforts are
needed for legumes in developing countries.
Farmer-based seed supply

In this chapter farmer-based seed production is used loosely and broadly to
describe seed production and supply with or by farmers even when they
differ greatly in scope and ownership among themselves. Several approaches
are used by different stakeholders to involve farmers in local seed production, including genetic resource conservation, crop improvement, variety
popularization and seed supply. In the context of seed delivery, farmerbased seed production and marketing implies farmers ownership and
responsibility for operating independently an enterprise with commercial
intent to ensure profitability and sustainability. It could be a small-scale
enterprise owned and managed by one or a few persons who are engaged
not only in production but in marketing of seed as well. At the community
level, these may be individual farmers, a group of farmers, traders or merchants, cooperatives, or farmers associations (Kugbei and Bishaw, 2002).
Non-commercial seed supply

Finding alternative strategies to enhance rapid dissemination of new crop
varieties and seeds among farming communities are crucial to realize the
impact of crop improvement research. Non-market intervention is one of
the alternative strategies for seed distribution owing to the failure of public
and private sectors in providing seed to small-scale farmers.
Grisley (1993) suggested a non-commercial approach for crops not handled by the formal sector, where seed packets are distributed in small quantities to farmers to encourage informal varietal diffusion and adoption
rather than regular seed production and supply for beans in sub-Saharan
Africa. A follow-up study showed farmers continue growing the variety
and exchange with other farmers after the initial supply of seeds. Jones et al.
(2001) also reported successful dissemination of a modern pigeon pea variety from injection of seed in a single demonstration trial in Kenya. It was
reported from Ethiopia that 211 farmers in ten districts produced 88 t of faba
bean, pea, chickpea and lentil (4 t) seeds for informal diffusion (ICARDA,
2006) in the absence of seed supply from the formal sector.
361
Village-based seed enterprises

David (2004), drawing on experience from Uganda, reported on the role of
farmer seed enterprises as a tool to ensure sustainable seed production and
marketing of new crop varieties as well as a regular source of clean seed.
Therefore, it is important to institutionalize local seed production and marketing by establishing a technically sound and economically viable seed business by mobilizing and ensuring the participation of farming communities.
ICARDA is currently exploring alternative approaches for seed delivery
for crops where neither the public nor the private sector is able to serve smallscale farmers. The objective is to establish village-based seed enterprises (VBSE)
to produce and market quality seed in selected communities. The VBSEs are
farmer-based seed production and marketing schemes operating at local level
to ensure availability and access to varieties and seeds by farmers in less
favourable environments and remote areas. The system works primarily by
organizing volunteer farmers into local seed production and marketing enterprises. There are several advantages for organizing local seed production and
marketing units as summarized below (Bishaw and van Gastel, 2008).
Key elements for the success of village-based seed enterprises

There are a few prerequisites for establishing and successful operation of
VBSEs:
Regular seed demand from farmers within community, neighbouring

villages or districts.
Reasonable seed price seed should be affordable by farmers and protable for producers.
Appropriate seed quality seed should be consistently of higher quality
than farm-saved or locally exchanged seed.
Enterprise ownership farmers should take responsibility for managing
and operating the enterprises.
Business plans there should be appropriate training in developing
tailor-made business plans based on demand analysis.
Establishing village-based seed enterprises

The initiatives involving farmers is often top-down, based on presumptions
of development agencies rather than on critical appraisal of situations on
the ground. A number of steps to be followed for successful establishment
of VBSEs are given below (Fig. 22.1).
Stakeholders consultation convene multi-institutional multi-stakeholders consultation meeting to build consensus and solicit interest in
and support for VBSEs, and determine their roles and responsibilities in
supporting operations and implementations.
362
Z. Bishaw et al.
Profitability
Seed marketing
Performance analysis for profitability and

sustainability
Pricing, market information, marketing

linkages created
Quality control
Guidelines and procedures for quality control

adopted
Seed production
Informal seed-production scheme developed

and adopted
Enterprise management
Operational mechanism established and

business plan developed
Establish farmer groups
VBSE formally constituted and trained

(technical, financial, managerial)
Identify seed demand
Farmer participation in problem diagnosis

and making decisions
Fig. 22.1. Steps in establishing village-based seed enterprises (VBSEs) (Source: Bishaw and
van Gastel, 2009).
Seed system analysis conduct seed system analysis prior to establishing VBSEs to assess whether there is real seed demand in partnership
with stakeholders identied during the consultative process.
Identifying target areas and groups VBSEs should target: (i) farmers
lacking access to improved crop varieties and seeds from the formal sector; (ii) resource-poor small farmers with limited opportunities; (iii) remote and isolated areas with poor infrastructure; and (iv) less favourable
areas with limited commercial crop production.
Selecting farmers the participating farmers must be interested in setting up a seed business as an alternative to grain production; and they
should be selected based on criteria of high reputation in the community,
experience in farming/seed production, relatively bigger/better land
holdings, entrepreneurial skills and nancial resources.
Forming seed-producer groups farmer participation and empowerment are key elements of the VBSE programme. Farmers should take full
responsibility and leadership and elect their own leaders, whereas partners facilitate, provide guidance and advice.
Selecting seed-production sites the land selected must be suitable for
quality seed production: fertile soils, reliable rainfall (or irrigation), low
363
incidence of diseases, pests and weeds, proximity and accessibility to

main roads and other facilities.
Preparing a business plan the plan should consider all factors affecting
the enterprise: strengths, weaknesses, risks, products, markets, costs,
prots, etc. It also includes ownership, management, legal structure,
staff, equipment and budget.
Producing and marketing seed all seed-production and marketing operations are carried out by the VBSE members. Promotional efforts and
marketing are required to ensure success.
Conducting protability analysis farmers commitment in ownership
and protability of the business are the driving forces for long-term sustainability. This will help members in making informed decisions in
planning and diversication.
Linking and supporting village-based seed enterprises

The strategy of involving stakeholders and encouraging them to work towards
an annual business plan based on demand-led production is critical to
develop financially profitable and sustainable seed production and marketing enterprises. Key aspects of partner support are described below:
Sourcing seed and other inputs partners help the VBSEs to source early
generation seed of varieties most adapted to their areas from National
Agricultural Research Systems (NARS) or the formal sector. Similarly,
the partners assist the VBSEs to source inputs (fertilizers, pesticides) required for quality seed production.
Producing seed partners provide training, guidance and assistance, to
ensure that VBSE members have the skills and knowledge necessary to
produce seed that meets quality standards.
Processing and storing seed the VBSEs ensure that they are able to acquire simple low-cost mobile cleaner and treater prototypes which can
be easily maintained and modied locally. The partners may also help
building appropriate seed storage facilities.
Controlling seed quality partners train VBSE members to carry out
eld inspections and simple seed quality tests or provide services through
the formal sector.
Marketing seed promotional activities should be conducted through
on-farm demonstrations of varieties, organizing eld days for neighbouring farmers and providing market information through ministries,
extensions services or non-governmental organizations (NGOs).
Accessing credit the VBSEs need access to credit for purchasing farm
machinery, inputs (e.g. source seed, fertilizers and pesticides) and seed
handling equipment (for cleaning, treatment and packaging).
Building capacity training will be implemented to build, step-by-step,
farmers technical (planting, harvesting, cleaning, treatment, testing, storage), nancial and management skills (daily operation of enterprises,
record keeping, developing business plans).
364
Z. Bishaw et al.
Evaluating and monitoring establish procedures for regular reporting

and identify key indicators against which the progress and performance
of VBSEs are measured.
Establishing a network of VBSEs assist to establish a network of VBSEs
to link with input providers and stakeholders to facilitate market information and sharing of experience.
Linking with local agro-industries create linkages between farmers and
local agroprocessing enterprises to create a market for produces which
will then stimulate use of better technology, creating a demand for seed
among farming communities.
A detailed work plan and timetable should be developed for the implementation of the VBSEs. The commitment of all partners to the work plan and
timetable ensures timely and successful execution of the planned activities.
22.8. Conclusion
Although global lentil production has increased during the last few decades,
its availability per capita has declined with increasing population, particularly in South Asia (McNeil et al., 2007). There is an increasing demand for
lentil in the developing countries where it is a prominent part of the human
diet. Developing improved varieties with a high and stable yield that is
tolerant to biotic and abiotic stresses is a prerequisite for increasing lentil
production. Apart from improved varieties, developing and transferring
appropriate production technologies, reducing production risks and costs,
ensuring markets and a well-defined pricing policy are the basic requirements for sustainable legume production (Byerlee and White, 2000).
In the developing countries, lentil seed supply is primarily dominated
by the informal sector, as is the case with India (Materne and Reddy, 2007).
However, addressing policy, regulatory, technical, institutional, organizational and socio-economic constraints will encourage development of a
robust seed industry involving both formal and informal sectors to serve
the diverse seed users. The integration of research, seed supply, processing
and marketing is the only viable option in this endeavour. The private sector initiative from Turkey could serve as a model for promoting the adoption and diffusion of new lentil varieties elsewhere in the face of public
sector failures in delivery of improved varieties and seeds. While encouraging the formal private sector to increase the availability of seed, at present
the establishment of technically feasible and economically viable farmer
seed enterprises (David, 2004) is the best alternative option for lentils.
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23
Nutritional and Health-beneficial

Quality
Michael A. Grusak
USDA/ARS Childrens Nutrition Research Center, Baylor College of Medicine,

Houston, Texas, USA
23.1. Introduction
Lentil (Lens culinaris subsp. culinaris Medik.) is an ancient, domesticated
legume that has been nourishing humans for millennia (Erskine, 1997;
Sandhu and Singh, 2007). As with most pulse seeds, lentil can provide several dietary nutrients; these include amino acids (in the form of protein),
energy (primarily in the form of starch), most essential minerals and several vitamins. They also contain various compounds that are believed beneficial to the promotion of good health, as well as several anti-nutrients. In
this chapter, we will review existing data on the types of nutrients and
health-promoting phytochemicals reported for lentils, and will provide
information on the general range of values measured in lentil seeds. We
will discuss to what extent these levels might contribute to human dietary
needs when lentils are consumed in normal serving sizes, taking into
account possible anti-nutrients also contained in the seeds. Finally, we will
discuss the influence of different processing methods on the nutritional
value of lentil products. Additional information on some of these topics
can be found in recent and earlier reviews (Abu-Shakra and Tannous, 1981;
Hulse, 1994; Urbano et al., 2007).
23.2. Nutritional Composition

The nutritional value of any given lentil variety will be dictated by its genetics, in combination with its ability to make use of various environmental
resources (e.g. soil parameters, agronomic practices, temperature, rainfall,
etc.) and/or its ability to cope with diverse environmental constraints (biotic
and abiotic stress). In other words, the compositional phenotype of harvested
lentil seeds is as dependent on genotype-by-environment interactions as are
368

Nutritional and Health-benecial Quality
369
Table 23.1. Proximate composition of whole, dry lentil seeds (per 100 g dry matter)
(Source: see table footnotes).
Component
Energy (kJ)
Protein (g)
Carbohydrates (g)
Fat (g)
Total bre (g)
Ash (g)
Range of published valuesa

14182010
15.931.4
43.474.9
0.33.5
5.126.6
2.26.4
Lentil (mean values)b

1638
28.3
67.1
2.5
12.2
2.2
a Values derived from the following references: Kylen and McCready (1975); El-Nahry et al. (1980); Souci
et al. (1989); Jood et al. (1998); Solanki et al. (1999a, b); Sotomayor et al. (1999); Ereifej and Haddad
(2000); Cai et al. (2002); Porres et al. (2002); El-Adawy et al. (2003); Wang and Daun (2004, 2006); de
Almeida Costa et al. (2006); Iqbal et al. (2006); Amir et al. (2007); Ryan et al. (2007); USDA/ARS (2007).
b Values derived from USDA/ARS (2007), Nutrient Database No. 16144, lentils, pink, raw.
other phenotypic traits, such as yield, disease resistance, time to flowering,

etc. None the less, there are general tendencies that have been reported for
the components of lentil seeds; these will be discussed in detail in the following sections. The gross proximate composition for lentil (range of values)
is presented in Table 23.1, along with mean values for one lentil type.
Energy
The energy value of a food provides a gross assessment of the potential
energy stored within that foods chemical bonds. This energy is critical for
humans, as it is needed to sustain all the bodys functions, including protein synthesis, respiration, circulation, general metabolism and physical
activity. Energy values for lentil range from 1418 to 2010 kJ/100 g dry matter (Table 23.1), which is equivalent to caloric values of 339480 kcal/100 g
dry matter. Most of the energy comes from starch, the primary component
of lentil seeds, although non-starch carbohydrates, protein and lipids also
contribute to the energy value. The energy range for lentil is similar to that
of chickpea (Cicer arietinum L.) (Wood and Grusak, 2007), but is much lower
than that of legumes such as soybean (Glycine max L.) or groundnut (Arachis
hypogea L.), with their higher oil content (USDA/ARS, 2007).
Protein and amino acids

Legume seeds are known for their high protein content, especially when
compared to other plant foods such as cereal grains, most root crops, fruit
and vegetables. Reported protein values for lentil range from 15.9 to 31.4
g/100 g dry matter (Table 23.1). Most of the seed protein is stored in the
cotyledons, with the majority of the protein consisting of salt-soluble globulins (storage proteins) that are stored in protein bodies. The remainder
370
M.A. Grusak
belongs to the albumin fraction, which includes many housekeeping proteins, lectins and lipoxygenases (Bhatty and Christison, 1984; Bhatty, 1986;
Wang et al., 2003).
The nutritional value of lentil protein is determined by its amino acids,
which are needed by humans to build and/or repair structural proteins,
enzymes, peptides, antibodies, neurotransmitters and other important components (Reeds and Beckett, 1996). Lentil has a generally good amino acid
pattern, in that it contains all the human essential amino acids, and with
most of them occurring in proportions recommended for humans by the
World Health Organization (WHO) (Table 23.2). The exceptions are the sulfurcontaining amino acids, methionine and cysteine, which (as with most
legume seeds) are the most limiting of the amino acids. On the other hand,
lentil has adequate to high levels of lysine. This is why lentil and other
legumes are suggested as ideal complementary foods to cereals (rice, wheat,
maize), which are low in lysine and generally better sources of the sulfur
amino acids (Shewry and Halford, 2002).
Table 23.2. Amino acid composition (g/16 g N) for whole, dry lentil seeds and Food and
Agriculture Organization (FAO)/World Health Organization (WHO)/United Nations University
(UNU) recommended proportional pattern for essential amino acids (g/16 g N) in children
(Source: see table footnotes).
Amino acid
Histidine (His)
Isoleucine (Ile)
Leucine (Leu)
Lysine (Lys)
Threonine (Thr)
Tryptophan (Trp)
Valine (Val)
Tyrosine (Tyr)
Phenylalanine (Phe)
Methionine (Met)
Cysteine (Cys)
Alanine (Ala)e
Arginine (Arg)e
Aspartate (Asp)e
Glutamate (Glu)e
Glycine (Gly)e
Proline (Pro)e
Serine (Ser)e
Published range for

lentila
1.33.4
2.65.5
5.78.7
4.08.2
2.54.9
0.62.6
3.36.1
1.13.6
3.65.8
0.81.3
0.71.5
2.45.0
3.911.1
9.315.9
12.818.5
3.35.6
1.27.2
2.96.4
Pattern for 1-year-old

childb
Pattern for pre-school

child (25 years old)b
2.6
4.6
9.3
6.6
4.3
1.7
5.5
7.2c
1.9
2.8
6.6
5.8
3.4
1.1
3.5
6.3c
4.2d
2.5d
a Values derived from the following references: Shekib et al. (1986); Combe et al. (1991); Kavas and Nehir
(1992); Urbano et al. (1995); Carbonaro et al. (1997); Porres et al. (2002); Wang and Daun (2004, 2006).
b Values from FAO/WHO/UNU (1985).
c Combined recommended pattern for tyrosine plus phenylalanine.
d Combined recommended pattern for methionine plus cysteine.
e Non-essential amino acids.
371
A more direct way to view the potential amino acid value of lentil is to
compare its essential amino acid composition in an average serving size
(100 g cooked, half a cup in the USA, for a young child; 200 g cooked, one
cup in the USA, for a boy or adult) with the daily US Recommended Dietary
Allowance (RDA) for these nutrients (Table 23.3). Again it can be seen that
methionine and cysteine are the most limiting amino acids, providing only
1920% of the RDA for a boy or an adult and 29% for a young child. These
single servings provide roughly 3060% of the RDA for most of the other
essential amino acids, and effectively the full RDA for tryptophan. Note
that tryptophan levels are low in lentil (Tables 23.2, 23.3), relative to most of
the other amino acids; however, human dietary requirements are also lower
for tryptophan.
Carbohydrates
The principal nutritional component of lentil seeds is the carbohydrate
fraction, which can range from 43.4 to 74.9 g/100 g dry matter (Table 23.1).
Carbohydrates can be divided into four major classes: monosaccharides,
disaccharides, oligosaccharides and polysaccharides.
Monosaccharides and disaccharides
Monosaccharides are single sugar moieties, such as glucose and fructose,
while disaccharides contain two monosaccharides connected via a glycosidic bond (e.g. sucrose, made from glucose plus fructose). Free mono- and
disaccharides are readily absorbed and/or metabolized in the human digestive tract, and thus are a quick source of energy. However, mature lentil
seeds contain low levels of free soluble sugars (2.38.9 g/100 g dry matter),
with fructose (0.010.30% dry matter) reported as the main monosaccharide
and sucrose (1.13.0% dry matter) as the predominant disaccharide (Table
23.4). Free glucose has not been found or has been reported in only trace
amounts in mature lentil seeds (0.04% of dry matter), whereas maltose (a
disaccharide formed from two glucose subunits joined by an (14) linkage) has been detected in higher amounts (up to 0.33% dry matter (Nutrient
Database No. 16069), USDA/ARS, 2007).
Oligosaccharides
Oligosaccharides are short polymers that generally are composed of three
to ten monosaccharides. Most oligosaccharides are not digested in the small
intestine and thus move on to the large bowel where they are metabolized
(fermented) by colonic bacteria. This activity releases gases (e.g. hydrogen,
carbon dioxide, methane) that can promote flatulence. Lentils contain several raffinose-family oligosaccharides, including stachyose (up to 3.1% dry
matter), raffinose and verbascose (Table 23.4). Lentils also contain the oligosaccharide ciceritol (up to 2% dry matter), whose name comes from chickpea (Cicer arietinum) where it was first identified. Ciceritol does not belong
372
Table 23.3.
Contribution of lentil to human essential amino acid needs at different life stages (Source: see table footnotes).
Amino acid
g/16 g Na
mg/200 g
cooked
(one cup
serving)b
His
Ile
Leu
Lys
Thr
Trp
Val
Tyr + Phe
Met + Cys
2.4
3.3
6.4
5.7
4.1
2.5
4.0
6.9
1.8
379
521
1010
900
646
394
632
1088
284
Child RDAb
(mg) for
48 year
old @
22 kg
352
484
1078
1012
528
132
616
902
484
RDA (%)
in 100 g
cooked
54
54
47
44
61
149
51
60
29
Child RDAc
(mg) for
913-yearold boy @
37 kg
RDA (%)
in 200 g
cooked
629
814
1813
1702
888
222
1036
1517
814
60
64
56
53
72
177
61
72
35
Adult RDAc
(mg) for
19+ year RDA (%)
in 200 g
old @
cooked
75 kg
1050
1425
3150
2850
1500
375
1800
2475
1425
36
36
32
32
42
105
35
44
20
Pregnant
woman
RDAc (mg)
for all ages
@ 60 kg
RDA (%)
in 200 g
cooked
1080
1500
3360
3060
1560
420
1860
2640
1500
35
35
30
29
41
94
34
41
19
a Mean
values from Wang and Daun (2004) for Canadian green lentils.
calculated from previous column, using an average % protein = 26.3 (equivalent to % N = 4.21) (Wang and Daun, 2004), and assuming 70% water in
the cooked lentil seeds.
c Calculated from US Recommended Dietary Allowance (RDA) Tables for amino acids (Institute of Medicine, 2006).
b Values
M.A. Grusak
373
Table 23.4. Carbohydrate composition of whole, dry lentil seeds

(g/100 g dry matter) (Source: see table footnote).
Carbohydrate
Starch
Total soluble sugars
Sucrose
Fructose
Glucose
Total -galactosides
Rafnose
Stachyose
Verbascose
Ciceritol
Maltose
Reported rangea
34.765.0
2.38.9
1.13.0
0.010.30
00.04
1.86.8
0.161.49
1.13.1
01.35
0.241.99
0.050.33
a Values derived from the following references: Reddy et al. (1984); Souci et al.
(1989); Vidal-Valverde et al. (1993a, b); Fras et al. (1994, 1995, 1996);
Jood et al. (1998); Sotomayor et al. (1999); Fasina et al. (2001); Cai et al.
(2002); Porres et al. (2002); El-Adawy et al. (2003); Wang and Daun (2004,
2006); Martn-Cabrejas et al. (2006); Amir et al. (2007); USDA/ARS (2007).
to the raffinose series of oligosaccharides, nor is it believed to contribute

significantly to flatulence (Quemener and Brillouet, 1983). Additionally,
lentil contains -galactosides (galactose polymers joined by (16) linkages) in amounts ranging from 1.8 to 6.8% dry matter, which can also lead
to flatulence (Fras et al., 2003).
Polysaccharides
Starch is the predominant carbohydrate in lentil seeds, ranging from
34.7 to 65.0% of dry matter. It serves as the principal energy source from
this food. Starch is composed of two types of polymer: amylose and amylopectin. Both types are built from glucose subunits, although the linkages
vary. Amylose is an essentially linear polymer of glucose, built up from
repeated (14) linkages. Amylopectin has the same linkages, but also contains (16) linkages every 2430 glucose units; this gives amylopectin a
highly branched structure. These two polymers occur together in starch
granules, and the relative proportions of amylose and amylopectin give each
starch granule its unique structure and crystallinity, as well as confer specific
physical properties to the combined starch (e.g. swelling factor, gelatinization temperature) (Hoover and Ratnayake, 2002). Lentil starch can contain
from 20 to 45.5% amylose (Urbano et al., 2007), which is similar to that found
in chickpea (Wood and Grusak, 2007).
Digestion of starch, via pancreatic -amylase and the intestinal enzymes
sucrase-isomaltase and maltase-glucoamylase (Nichols et al., 2003), yields
glucose units that can be absorbed in the small intestine and utilized for
energy. However, not all of the starch is digested, presumably because of
STARCH
374
M.A. Grusak
inhibitory components within the grain. This leads to the so-called resistant
starch fraction (Sajilata et al., 2006), which escapes the small intestine and
passes to the large bowel (also leading to flatulence). De Almeida Costa et al.
(2006) reported a resistant starch value of 3.7 g/100 g dry matter in lentil
cultivar Silvina.
Resistant starch and all other undigested carbohydrates, including non-starch polysaccharides and oligosaccharides, are classified as dietary
fibre. Non-starch polysaccharides are components that are not starch, but
which are polymerized carbohydrates. These include cell wall components
such as cellulose, hemicelluloses, pectic substances and b-glucans (mixedlinkage (13), (14)-b-D-glucan). In total, these can range from 5.126.6%
dry matter (Table 23.1). Specifically, cellulose comprises 4.15.7% and hemicelluloses 615.7% of dry matter (Reddy et al., 1984; Amir et al., 2007;
Urbano et al., 2007), and b-glucans in lentil germplasm have been reported at
0.41.1% dry matter (Demirbas, 2005).
DIETARY FIBRE
Lipid and fatty acids

Lentil has a low concentration of lipids (fats), ranging from 0.33.5 g/100 g
dry matter (Table 23.1), which places it similar to other cool-season legumes
(chickpea, pea (Pisum sativum L.)) and most cereals (rice, wheat) (USDA/
ARS, 2007). Although lipids are energy-dense compounds, the low concentration in lentil means that little of the energy content of lentil seeds
comes from fat. Compositionally, lentil contains both of the human essential fatty acids: linoleic and linolenic acids (Table 23.5). Linoleic is by far
the predominant fatty acid, comprising from 41 to 57% of oil in several
cultivars, whereas linolenic acid ranges from 0.3 to 16% of oil. The monounsaturated oleic acid (C18:1) and the saturated palmitic acid (C16:0)
account for most of the remainder of the fatty acid profile. Myristic, palmitoleic, stearic, arachidic, gadoleic, eicosadienoic, behenic, erucic and lignoceric acids have also been reported in lentil seeds, with most occurring at
<1% of oil (Table 23.5).
Minerals
Lentil seeds contain several mineral elements that are required by humans
(i.e. essential), although not all are necessarily required by lentil itself. Plants
are known to take up many mineral ions that they encounter in the soil
environment (via root absorption), even if these elements are not needed
for plant growth and development; many of these elements are subsequently partitioned to seeds (Grusak and DellaPenna, 1999; Grusak, 2002).
In total, these elements constitute the ash fraction that is frequently reported
in proximate analyses. The reported ash range for lentil seed is 2.26.4 g/100 g
dry matter (Table 23.1).
Fatty acid composition of lentil (% fatty acid in oil) (Source: see table footnotes).
Fatty acid
Myristic (C14:0)
Palmitic (C16:0)
Palmitoleic (C16:1)
Stearic (C18:0)
Oleic (C18:1)
Linoleic (C18:2)
Linolenic (C18:3)
Arachidic (C20:0)
Gadoleic (C20:1)
Eicosadienoic (C20:2)
Behenic (C22:0)
Erucic (C22:1)
Lignoceric (C24:0)
var. LB Meta
NAc
18.3
0.3
2.5
22.1
55.7
0.3
0.6
NA
NA
NA
NA
NA
var. 81sa
NA
24.5
0.6
2.0
21.6
47.8
1.9
1.2
NA
NA
NA
NA
NA
Canadian greenb
Canadian redb
Australianb
0.000.93
10.7915.36
0.000.61
1.271.82
17.0425.63
40.9746.14
11.9316.23
0.771.11
1.211.58
0.000.22
0.811.13
0.360.74
0.471.99
0.420.73
13.2515.77
0.000.33
1.341.65
17.0522.17
42.9145.23
11.6814.66
0.800.92
1.121.24
0.001.86
0.810.91
0.800.92
0.560.70
0.600.60
12.7013.70
NA
1.802.10
22.7028.00
41.9057.14
11.6012.70
NA
1.201.30
NA
NA
NA
NA
Table 23.5.
a Data
from Amir et al. (2007).

from Wang and Daun (2004).
c NA = not available.
b Data
375
376
M.A. Grusak
With respect to human mineral needs, reported values are available for
lentil seeds for 12 of the 17 human essential elements (Table 23.6). Data are
not available for chloride, fluoride or iodine. Two other human essential
elements not listed here are nitrogen and sulfur, because these are viewed
as being acquired in the form of amino acids (Table 23.3). Additional elements, such as boron, nickel and cobalt, are reported here, due to availability of data in the literature; these elements are not deemed essential in
humans, but have been suggested to provide various health benefits (Nielsen,
1996). In general, it is seen that mineral concentrations are quite variable,
presumably because of both genotype and environmental influences. On a
weight basis, potassium and phosphorus constitute the bulk of the mineral
profile, although even these minerals can be relatively low in some cases
(Table 23.6).
The mineral profile for lentil can also be viewed in terms of its percentage contribution to human dietary needs. Table 23.7 provides values for the
current maximum adult US Recommended Dietary Allowance (RDA) or
Adequate Intake (AI) for several essential elements, and the amount of these
elements in a single serving of cooked, whole lentil (200 g fresh weight;
equivalent to one cup in the USA). Lentil is clearly a very good source of
copper and phosphorus, providing about half of the RDA for these elements. A single serving can also provide roughly one-quarter to one-third
of the RDA or AI for iron, zinc and manganese. Lentil is a poor source of
Table 23.6. Mineral concentrations of whole, dry lentil seeds (mg/100 g

dry matter)a (Source: see table footnotes).
Mineral
Concentration (mg/100 g dry matter)

(Ca)b
Calcium
Magnesium (Mg)b
Phosphorus (P)b
Potassium (K)b
Iron (Fe)b
Zinc (Zn)b
Manganese (Mn)b
Copper (Cu)b
Sodium (Na)b
Boron (B)
Chromium (Cr)b
Nickel (Ni)
Cobalt (Co)
Selenium (Se)b
Molybdenum (Mo)b
a Range
42165
13167
2401287
381360
3.113.3
2.310.2
0.61.0
0.49.9
0.479
0.61.0
0.03
0.120.35
0.04
0.0090.012
0.080.22
of values derived from the following references: Kylen and McCready

(1975); Souci et al. (1989); Sika et al. (1995); Sharma et al. (1996); Solanki et al.
(1999a); Ereifej and Haddad (2000); Koplk et al. (2002); El-Adawy et al. (2003);
Wang and Daun (2004, 2006); Iqbal et al. (2006); USDA/ARS (2007).
b Elements conrmed as essential for humans (Grusak and DellaPenna, 1999).
377
Table 23.7. Contribution of a 200 g serving of cooked lentils to the US RDA or Adequate
Intake (AI) of various minerals for adults (Source: see table footnotes).
Maximum adult US
RDA or AI (mg)a
Potassium
Calcium
Phosphorus
Magnesium
Iron
Zinc
Manganese
Copper
Selenium
4700
1200
700
420
18
11
2.3
0.9
0.055
Amount (mg) in 200 g

of cooked lentilb
738
38
360
72
6.6
2.6
0.98
0.50
0.006
Contribution to adult
RDA or AI (%)
16
3
51
17
37
24
43
56
11
a RDAs are the daily levels of intake of essential nutrients judged to be adequate to meet the known
nutrient needs of practically all healthy persons. Values presented are the highest RDA either for male or
female adults, excluding pregnant or lactating women. The values for potassium, calcium and manganese
are the AI value, which is that amount believed to cover the needs of all individuals in a group, but lack of
data prevent being able to specify with condence the percentage of individuals covered by this intake.
All values are from Institute of Medicine (2006).
b Values are from USDA/ARS (2007) for mature, cooked, boiled without salt lentil seeds (Nutrient
Database No. 16070), which contain 70% water. Note that these are only a subset of the minerals that
can be found in lentil seeds (see Table 23.6).
calcium, providing only 3% of the AI, and it is questionable how much of

this is even available, because most of this calcium would probably be present as insoluble calcium oxalate crystals (Zindler-Frank, 1987; Franceschi
and Nakata, 2005).
Vitamins
Lentils contain many of the water-soluble and fat-soluble vitamins required
by humans (Table 23.8), although they are devoid of vitamins B12 and D,
which are not synthesized by plants. As with other seed components, the
vitamin concentrations vary across cultivars, with ranges generally much
larger for the water-soluble than fat-soluble vitamins. In fact, vitamin C and
biotin were totally undetected in some sample surveys (Table 23.8). When
the vitamin compositional quality of lentil is assessed in a single, cooked
serving, relative to maximum US RDA or AI values for adults (not including pregnant or lactating women), it is clear that lentil is an excellent source
of folate. In various regions of the world, processed foods are fortified with
folate in order to ensure adequate intakes, especially for women of childbearing age (Buttriss, 2005). Folate deficiency can lead to neural tube defects
in infants and may be linked to a higher incidence of certain types of cancer
and heart disease (Selhub and Rosenburg, 1996). Other vitamins for which
lentil provides good levels are pantothenic acid, pyridoxine and thiamin,
Vitamin
Water-soluble:
Vitamin C
Niacin (vitamin B3)
Pantothenic acid
Pyridoxine (vitamin B6)
Riboavin (vitamin B2)
Thiamin (vitamin B1)
Folate
Biotin
Vitamin B12
Fat-soluble:
Vitamin E (-tocopherol)
Vitamin E (-tocopherol)
Choline
Vitamin A
Vitamin K
Vitamin D
Units
Reported range (per

100 g dry matter)a
Maximum adult
RDA or AIb
mg
mg NEd
mg
mg
mg
mg
g
g
g
0.07.7
0.63.6
0.42.4
0.160.60
0.110.46
0.130.90
40535
0132
Not found in plants
90
16
5
1.7
1.3
1.2
400
30
2.4
mg -TEf
mg -TE
mg
g REh
g
g
0.361.60
0.310.64g
109
2.23.4
5.6
Not found in plants
15
550
900
120
15
Amount in 200 g
cooked lentil seedc
378
Table 23.8. Vitamin composition of whole, dry lentil seeds, of cooked lentil seeds, and percentage contribution of a 200 g serving of cooked
lentils to the US RDA or AI of several vitamins for adults (Source: see table footnotes).
Contribution to adult RDA
or AI (%)
3.0
2.12
1.28
0.36
0.15
0.34
362
106e
0
3
13
26
21
11
28
91
353
0
0.22
65.4
16
3.40
0
1
12
2
3
0
M.A. Grusak
Range of values derived from the following references: Kylen and McCready (1975); Savage (1988); Souci et al. (1989); Wang and Daun (2004, 2006); Ryan
et al. (2007); USDA/ARS (2007).
b See Table 23.7 for explanation of RDA and AI. All values in this column are from Institute of Medicine (2006). All values reect the RDA, except for vitamins D,
K, pantothenic acid, biotin and choline for which the AI is listed.
c Values are from USDA/ARS (2007) for mature, cooked, boiled without salt lentil seeds (Nutrient Database No. 16070), which contain 70% water.
d NE, niacin equivalent; 1 mg NE is equal to 1 mg of niacin or 60 mg of dietary tryptophan.
e Value calculated from biotin concentration reported in Savage (1988).
f -TE, -tocopherol equivalent; 1 mg a-TE is equal to 1 mg (R,R,R)-a-tocopherol.
g Range calculated using a 10% conversion factor for g-tocopherol (i.e. 10 mg g-tocopherol = 1 mg -TE).
h RE, retinol equivalents. Preformed vitamin A is not found in plant foods. However, plants contain a number of provitamin A carotenoids (e.g. b-carotene)
which can be metabolized to vitamin A. Vitamin A activity is expressed in RE; 1 g RE is equal to 1 g all-trans retinol, 6 g all-trans -carotene, or 12 g of
other provitamin A carotenoids.
379
supplying 2128% of the RDA or AI in a single 200 g serving. Lentil is not a

particularly good source of fat-soluble vitamins, potentially providing only
12% of the AI for choline, and less than 3% of the RDA or AI for vitamins A,
E or K. The low concentration of fat-soluble vitamins is probably a reflection,
in part, of the low oil content of lentil (Table 23.1).
23.3. Non-nutrient Components

Plant foods contain a myriad of secondary metabolites, peptides, enzymes
and other components that have gained attention for their potential to
reduce the incidence of disease and to promote good health in humans.
These components are non-nutritive in nature, although some of them can
be metabolized to release energy, or to generate nutritional compounds.
They can serve as antioxidants, as prebiotics, as regulators of gene expression and modifiers of cell division, to name a few (Lila and Raskin, 2005;
Weaver et al., 2008). Although much has still to be determined regarding
the bioavailability of various phytochemicals, and thus the effective dose
that one can gain from a given food (Holst and Williamson, 2008), researchers have none the less been quantifying the non-nutritive components of
different seeds, fruit and vegetables, including lentil. In addition to the
health-beneficial components, plants also contain anti-nutritional factors
that can reduce the availability and/or utilization of various dietary nutrients. Several compounds have been identified in lentil that can interfere
with the utilization of minerals, protein or carbohydrates. These are discussed
below.
Fibre
Several non-nutritive carbohydrates were mentioned in an earlier section,
including raffinose-family oligosaccharides, -galactosides, -glucans and
other polysaccharides, which along with resistant starch constitute the fibre
fraction of lentil. Dietary fibre is important for general functioning of the
gastrointestinal tract, as it assists faecal motility and helps maintain proper
faecal water balance (Gallaher and Schneeman, 1996). Much of the fibre
(especially the oligosaccharides) can be fermented in the large bowel,
thereby providing energy and metabolites to the gut microflora. The population of good microbes in the gut, referred to as probiotic bacteria, are
proving important for their role in the suppression of unwanted pathogenic
bacteria (Baumgart and Carding, 2007), stimulation of the immune system
(Holzapfel and Schillinger, 2002) and the enhancement of calcium absorption and bone mineralization (Abrams et al., 2005). Thus, a diet containing
lentils can serve as a prebiotic source that provides needed dietary fibre for
the maintenance of a balanced gut ecosystem (Guillon and Champ, 2002;
Blaut and Clavel, 2007).
380
M.A. Grusak
While fibre has many benefits, it also has some anti-nutritional aspects.
Fibre has a high cation exchange capacity and thus can bind several minerals
including calcium, iron and zinc. This can lead to reductions in the solubility and bioavailability of these minerals, from the fibre-containing food, or
from other sources in the diet (Kelsay et al., 1988; Weber et al., 1993). It is
interesting that fibre has both a positive (prebiotic) and a negative (binding
capacity) effect on mineral absorption.
Phenolics
Phenolics are a huge class of secondary compounds that play diverse roles
in the physiology of the plant (e.g. growth, reproduction, pigmentation,
pathogen resistance), but which also are proving important as dietary constituents due to their antioxidant properties and potential promotion of
good health. Phenolics range from simple molecules, such as phenolic acids,
to highly branched and polymerized compounds, such as tannins (Bravo,
1998). In legume seeds, the principal phenolics are phenolic acids, flavonoids and lignans. In lentil, most of the phenolics are localized to the
seedcoat, which has a broader diversity of phenolic compounds as well as
higher concentrations overall (Table 23.9). In general, although the cotyledons make up most of the seed mass, they provide very little in the way of
total dietary phenolics.
The health-beneficial significance of these compounds is derived from
their antioxidant properties, which allows them to scavenge free radicals
(Seifried et al., 2003). This activity presumably explains their role in the
reduction of chronic diseases (Williamson and Manach, 2005). For instance,
catechins and procyanidins (oligomeric catechins) have been shown in human
Table 23.9. Phenolic composition of lentil seedcoat and cotyledons (mg/100 g
dry matter) (Source: see table footnotea).
Total catechins
Hydroxybenzoic acid
Free hydroxycinnamic acid
Combined hydroxycinnamic acid
Dimer procyanidins
Trimer procyanidins
Tetramer procyanidins
Galloylated procyanidins
Prodelphinidins (dimers plus trimers)
Flavone glycosides
Flavonols
trans-Resveratrol glycoside
a Range
b ND,
Seedcoat
Cotyledons
91.9163.3
2.84.5
1.23.0
ND
61.9112.2
44.149.8
1.96.0
6.912.3
36.972.5
3.318.6
1.024.1
0.60.9
0.020.03
0.180.22
0.30.6
0.11.4
NDb
ND
ND
ND
ND
ND
ND
ND
of values derived from: Dueas et al. (2002, 2003) and Dueas (2003).
not detected.
381
intervention studies to increase plasma antioxidant activity (Nakagawa et al.,

1999). In an in vitro comparison of antioxidant capacity of several food
legumes (lentil, pea, chickpea, common bean (Phaseolus vulgaris L.) and soybean), lentil cultivars possessed the highest antioxidant activities (Xu et al.,
2007). On the other hand, several phenolic compounds (especially the catechins) have the ability to chelate metals (Khokhar and Owusu Apenten,
2003; Andjelkovic et al., 2006). Foods high in phenolics have been shown to
inhibit iron absorption in human subjects (Tuntawiroon et al., 1991), and
may inhibit the absorption of other micronutrient metals (Le Nest et al.,
2004; Viadel et al., 2006).
Another group of polyphenolics is the condensed tannins, also known
as proanthocyanidins, which are polymers of two to 50 (sometimes more)
flavonoid units. Tannins have been identified in lentil, usually associated
with the seedcoat fraction, and are found at concentrations of 0.021.01
g/100 g dry matter (Solanki et al., 1999a; Wang and Daun, 2006). As with the
phenolic acids, tannins have the ability to chelate iron and other metals, thus
they can reduce the bioavailability of metals in plant foods (Viadel et al.,
2006). Tannins also interact with protein, starch and other carbohydrates,
forming insoluble complexes that reduce the digestibility of these nutrients
(Carbonaro et al., 1996; Muzquiz and Wood, 2007).
Phyto-oestrogens
Plants contain several oestrogen-like compounds (referred to as phytooestrogens) that can induce oestrus in immature animals or interfere with
normal reproductive processes (Kurzer and Xu, 1997). Phyto-oestrogens
include the isoflavonoids: genistein, daidzein, biochanin A and formomonetin; the coumestan: coumestrol; and the lignans: secoisolariciresinol and
matairesinol (Mazur et al., 1998). Although many plant groups contain phytooestrogens, members of the legume family exhibit the highest levels of phyto-oestrogens. For example, soybeans are rich in the isoflavonoids genistein
and daidzein (Kurzer and Xu, 1997), and populations that consume high
levels of soy products appear to have lower incidences of disease, including
reductions in certain types of cancers and type 2 diabetes (Dixon and Ferreira, 2002; Villegas et al., 2008). In lentil, phyto-oestrogenic compounds
have been identified, but their concentrations are quite low, relative to several common crop legumes. Daidzein (3.310.4 g/100 g dry matter) and
genistein (7.118.8 g/100 g dry matter) are basically present in trace
amounts, relative to soybean (values range from 10,500 to 56,000 g/100 g
dry matter for daidzein and 26,80084,100 g/100 g dry matter for genistein)
(Mazur et al., 1998). Lentil also contains only minute amounts of the lignans,
anhydrosecoisolariciresinol and secoisolariciresinol (combined range of 8.9
12.3 g/100 g dry matter) and trace to non-detectable amounts of matairesinol (Mazur et al., 1998). However, only a few lentil cultivars have been
characterized and it would be interesting to quantify these compounds in
additional germplasm, or in seeds of lentil plants subjected to various stress
382
M.A. Grusak
conditions during seed development (which may stimulate isoflavone synthesis).
Phytate
Plants synthesize various myo-inositol phosphates, denoted IP1 through to
IP6, which contain from one to six phosphate groups. A significant antinutrient from this group is myo-inositol(1,2,3,4,5,6)hexakisphosphate, (IP6,
phytic acid, or phytate), which can complex with calcium, iron and zinc
ions in the food matrix (Raboy, 2001). This complexation reduces the solubility of these minerals and inhibits their absorption in humans (BrinchPedersen et al., 2007; Hotz and Gibson, 2007). However, it should be noted
that the metal scavenging ability of phytate (and some of the other inositol
phosphates) may also reduce the free radical activity of bound transition
metals, thereby simultaneously conferring health benefits and disease prevention attributes to phytate (Zhou and Erdman, 1995). Lentil seeds contain from 0.41.6 g phytate/100 g dry matter (Sharma et al., 1996; Wang
and Daun, 2006). The inositol phosphates IP3, IP4 and IP5 have also been
identified in lentil seeds, although at less than 10% the amount of IP6
(Kozlowska et al., 1996; IP1 and IP2 could not be resolved in this study).
Saponins
Saponins are bioactive compounds that are composed of a steroidal or triterpene aglycone linked to one, two or three saccharide chains of variable size
and complexity (Fenwick and Oakenfull, 1983). Saponins have traditionally
been classified as anti-nutritional factors, due to their ability to form stable,
soap-like foams that can interact with components of the digesta (Muzquiz
and Wood, 2007); however, specific saponins are also being recognized for
health benefits, such as hypocholesteraemic and anticancer properties, or
stimulation of the immune system (Champ, 2002; Mathers, 2002). The saponin
content of lentil seeds includes both soyasaponins I and VI, with concentrations ranging from 0.065 to 0.126 g/100 g dry matter (Ruiz et al., 1997).
Enzyme inhibitors
Legume seeds contain two main classes of enzyme inhibitors: those that
inhibit proteases (trypsin and chymotrypsin) and those that inhibit -amylases
(Muzquiz and Wood, 2007). The former disrupt protein digestion, thereby
influencing the utilization of protein and amino acids from the seeds. The
-amylase inhibitors interfere with starch breakdown, and thus limit the
available energy found within seeds. Lentil demonstrates trypsin inhibitor
activities (TIA) of 2155 TIA/mg protein, which places it in line with the
activities found in pea, broad bean (Vicia faba L.) and mung bean (Phaseolus
383
aureus L.), but much lower than that measured in chickpea, common bean
or soybean (Guillamn et al., 2008).
Toxins
Legume seeds contain lectins or haemagglutinins, which are glycoproteins
that possess at least one non-catalytic domain and can bind to specific
monosaccharides, oligosaccharides or other glycoproteins (Peumans and
Van Damme, 1995). When bound to these components on the surface of
cells lining the intestine, they can interfere with digestive processes and
absorption. Lentil seeds do contain lectins, but their concentration is low
and they provide only minimal reactivity in the erythrocyte agglutination
test; thus, lentils have been deemed to have low lectin toxicity when compared
to other highly reactive legumes such as common bean, Phaseolus coccineus
L. or Phaseolus acutifolius L. (Grant et al., 1983).
Allergens are another form of toxin in plant seeds, with some individuals reacting quite severely to specific proteins or peptide fragments after the
ingestion of a food (Cordle, 2004). Lentil is frequently associated with IgEmediated hypersensitivity reactions, particularly in pediatric populations in
Mediterranean countries where lentil is a common food (Kalogeromitros
et al., 1996). A major allergen from the vicilin protein family, designated Len
c 1.01, has been identified in lentil (Lpez-Torrejn et al., 2003).
23.4. Processing Types and Effects on Nutritional Quality

When lentils are prepared for human consumption, they are usually processed in some manner to enhance their palatability and to improve their
overall nutritional value. Different processing methods can be employed,
including decortication (seedcoat removal), soaking, germination, fermentation, sprouting, cooking, microwaving, enzyme treatment or irradiation
(see Urbano et al. (2007) for a detailed discussion of these processes). In
general, these treatments lead to a significant reduction or total elimination
of anti-nutritional components, although losses of certain nutrients can
also occur. For instance, decortication reduces the levels of phenolics and
tannins (Dueas et al., 2002), which might otherwise interfere with mineral
or protein utilization; cooking inactivates enzyme inhibitors (Urbano et al.,
1995), but may also cause thermal degradation of vitamins (FernandezOrozco et al., 2003). Soaking, germination, sprouting or fermentation processes lead to metabolic changes of the lentil seed components, which can
result in enhanced availability of some compounds (e.g. protein, starch,
some minerals and vitamins) (Urbano et al., 1995; Kuo et al., 2004; Ghavidel
and Prakash, 2007), as well as reductions in the concentration of others
(e.g. phytate, lectins, some phenolics) (Kozlowska et al., 1996; LpezAmors et al., 2006).
384
M.A. Grusak
23.5. Acknowledgements
This work was funded in part by funds from USDA/ARS under Agreement
No. 58-6250-6-001 and from the Harvest Plus Project under Agreement No.
58-6250-4-F029. The contents of this publication do not necessarily reflect
the views or policies of the US Department of Agriculture, nor does mention of trade names, commercial products or organizations imply endorsement by the US Government.
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24
Postharvest Processing and Value

Addition
Albert Vandenberg
University of Saskatchewan, Saskatoon, Canada
24.1. Introduction
Lentils are usually consumed as a natural food product in the form of whole
seeds or decorticated seeds. In most countries they are used in traditional
food preparations such as soups and stews which are served with other staple foods based on rice, wheat or other major cereal grains. As is the case
with most pulse crops, consumer preferences for lentils are very specific in
terms of seed size and shape, seedcoat appearance and colour, cotyledon
colour and uniformity of appearance. Processing consists of a series of
mechanical separations and milling operations that prepare lentils for direct
consumption.
The lentil processing industry has two major components. Primary processing consists of cleaning and delivery of whole seeds to consumers. The
basic principle of primary processing is to use gravity to separate lentil
seeds into desired quality classes defined by diameter, thickness, density and
colour. Various screens and air-flow mechanisms are used to remove
unwanted organic and inorganic materials and to sort the retained lentils. In
regions where lentils have been grown as a traditional crop, simple seed separation systems based on sieves are used for small-scale primary processing.
In regions where lentils are accumulated, stored and then processed for delivery to urban consumers, the primary processing may be done on a feefor-service basis, or may be done in-house by a lentil marketing company.
The secondary processing component mostly involves decortication,
splitting, sorting and polishing of the lentils. In some countries, secondary
processing of whole lentils involves thermal processing into cans or jars.
The tertiary level of processing, which involves using whole or decorticated
lentils in further milling, grinding or fractionation for use in processed food
products, is the least developed segment of the processing industry.
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A. Vandenberg
Value addition through processing is essentially the economic concept

of preparing lentils and lentil-based products in such a way as to convince
consumers to pay more for the improved convenience or the perceived nutritional or culinary benefits. The simplest form of value addition is the cleaning
of whole lentil seeds to acceptable culinary standards. Removal of the seedcoat reduces cooking time and is therefore the primary form of value addition for lentils. Value addition through processing also may be based on
improving the value of the by-products of processing, for example, the
seedcoats and small particles and broken bits of lentils that are cleaned out
from the cleaning, decortication and splitting processes.
24.2. Primary Processing

The principles of primary processing used for cleaning lentils are essentially
the same as for cereal grains. Specific modifications used for processing lentils include changes to elevators, conveyors and storage systems to reduce
damage to the seeds and the seedcoats and to maintain high visual quality.
The components used in primary processing systems are very specific in
terms of what type of equipment is economical and necessary in a given
lentil production region For example, specific requirements for weed seed
removal may determine what type of separation equipment is essential.
Figure 24.1 presents a generalized scheme of the steps in primary processing beginning with storage and ending with bagging of cleaned lentils. The
actual configuration in a processing facility is highly variable and flexible
depending on the economics of meeting consumer demand according to a
generally agreed level of quality based on providing lentils with specific
diameter, thickness, colour, damage and admixture specifications that are
agreed to by buyers and sellers.
Storage considerations
After harvest, lentils are either stored in bins or other containers on farms or
sold and delivered to a primary processing facility for storage and eventual
cleaning. In regions where on-farm grain storage is not common, lentil growers
deliver lentils to marketers immediately after harvest. In this case, the purchaser may have storage facilities where lentils are kept in bulk, in bags or in
steel bins. Where on-farm grain storage is common, for example in Canada,
lentils are normally stored on-farm in steel bins and delivered to primary processing facilities when required by sales contracts. The initial condition of the
grain and the storage environment may have an impact on processing quality.
In some cases, the lentils undergo rough cleaning before storage using stationary screens built into the conveying systems on combine harvesters or grain
conveyer systems to remove large and small particles from the grain.
Storage conditions prior to processing can affect the quality of the lentils and cause processors to accommodate the flow of product through the
Postharvest Processing and Value Addition

Operation
393
Comments
Stored lentils
Specific conditions are location specific, may differ depending on end use
Pre-cleaning
Removal of coarse and fine materials based on stationary

screens
Air screening
Removal of material based on particle density, shape and size

using gravity and air flow
Indent cleaning
Physical separation of lentils based on diameter
Gravity
separation
Further separation of lentils based on density
Destoning
Additional separation of stones based on density
Sizing
Colour sorting
Packaging
Optional separation into diameter or thickness based on

customer requirements
Optional separation of lentils based on colour
Final product delivery into bags, totes, containers, bulk

containers or rail cars for delivery
Fig. 24.1. General outline of the steps involved in primary processing of lentils.
cleaning process. For example, seedcoats of green lentils darken during

storage, slowly changing from green to an oxidized brown colour. The rate
of colour change is increased with increases in temperature, light or humidity. Cenkowski et al. (1989) and later Tang et al. (1994) studied the equilibrium moisture profiles of green lentils during storage and their effects on
quality. The rate of darkening of green lentil seedcoats increased over time
based on biochemical changes in the seedcoat. Desiccation of green lentils
with diquat may also cause seedcoat colour to darken more rapidly in the
field during the final stages of seed maturation, and possibly during storage (Davey, 2007). Similar research on storage conditions of red lentils with
brown or grey seedcoats is currently underway in Canada (K. Agblor, 2008,
Saskatoon, personal communication) to determine storage effects on subsequent dehulling and recovery percentage.
In production regions where lentil is grown as a subtropical dry cool
season (South Asia, for example) or as a winter crop in Mediterranean climatic zones (south-eastern Anatolia, Australia) the moisture content of the
harvested lentils is generally below 12% and is often much lower because
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A. Vandenberg
the crop is harvested during seasons with increasing temperature and

decreasing humidity. Problems with seed damage during handling may
occur if the lentils are too dry. Combine harvesting, augering and elevation
of very dry lentils into storage may cause problems with seed breakage and
with chipping of the seedcoat at the edge of the seed. This is more likely to
be a problem with large-diameter lentils (usually large green types), particularly for varieties that produce seeds with a thin instead of a thick edge.
In general, for lentil types that are sold as whole seeds to the consumers, storage moisture conditions must be managed so that the seedcoat
remains intact. In temperate regions with summer production, like the prairie regions of North America, harvest progresses into cooler and more
humid conditions. In some seasons it is necessary to reduce the moisture
content of the lentils immediately after harvest using aeration fans in the
bins for lentils that have been harvested at moisture content above 15%. The
safe storage moisture level for lentils is considered to be below 15%, but
storage moisture below 12% may lead to problems with handling damage.
One particular problem in the North American prairies is damage caused
by handling dry lentils at extremely cold temperatures during winter
months. Some environmental condition experienced during the growing
season may lead to development of brittle or wrinkled seedcoats in lentils,
which in turn contributes to damage during storage and postharvest handling. Alternating wet and dry periods during late stages of maturity can
cause wrinkling, colour loss and brittle seedcoats. Chemical desiccation of
the crop prior to harvest may also be a factor in causing brittle seedcoats
that chip and split more easily. For lentils that are destined for markets that
prefer whole lentils with intact seedcoats, primary processing facilities are
designed to minimize damage by using bristle-flighted augers and belts in
the place of standard conveying equipment used for cereal grains.
In most parts of the world, red lentils are produced for dehulling.
Recent research from Canada indicated that the critical moisture content for
maximizing the recovery of decorticated red lentils during milling is 12.5%
(Wang, 2005). In some countries, delivery contracts specifically state the
maximum allowable moisture content for delivered lentils. In Canada, recommended storage moisture for green lentil is 14% to help minimize seedcoat breakage, while the recommended storage moisture for red lentil is
13% to help maximize dehulling recovery.
Air screening
The primary method of cleaning lentils is the air-screen separator. It uses a
combination of air, gravity and screens to separate seed based on size, shape
and density. Commercial machines can be equipped with two to eight
vibrating screens. The lentils are fed into a hopper where they are evenly
distributed by a feed roller and transferred through a controlled gate on the
top sieve. The lentils are subjected to primary aspiration by the use of an air
trunk which drains off chaff, straw, dust or deceased grains. Then the lentils
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are passed through the sieve layer for separation according to their width
and thickness. After the separation, the graded material is subjected to an
air sifter and aspiration chamber where remaining light particles are sucked
off by a strong upward draught of air. The graded material and the impurities are automatically discharged in separate chutes.
Indent cleaning
Indent cleaning machines are generally used for separating lentils by diameter into different sizes. After air screening, the lentils are fed into the indent
cleaner through a conveyor into the centre of a rotating cylinder. The lentils
of smaller diameter are lodged in the indents and are lifted by the cylinder
until they fall out under gravitational force into the internal trough. The large
particles remain in the trough for further separation. Due to the rotating
speed and inclination of cylinder, the large lentils are separated and are discharged out while the small lentils and other crop seeds are removed separately with the help of an auger or a vibrating device fitted with a trough.
Gravity separation
This type of grain cleaner is used for separating lentils that have the same
size but differ in specific weight. A gravity separator can be used effectively
to remove partially eaten (for example by storage insects), immature or broken seeds to ensure maximum quality of the final product. Gravity separators have a rectangular deck so that the product travels a longer distance,
which results in effective separation of light and heavy particles. The product flows over the rectangular vibrating deck as pressurized air is forced
through from underneath, causing the material to stratify according to its
specific weight. The heavier particles travel to the higher level and the
lighter particles travel to the lower level of the deck. In order to obtain efficient separation by specific weight, the pressurized air supply needs to be
accurately adjusted to control the volume of air distribution at different
areas of the vibrating deck. The table inclination, the speed of eccentric
motion of vibration and the feed rate can be easily and precisely adjusted to
maximize separation efficiency.
Destoning
Destoning machines are used to separate heavier materials like stones, glass
and metal from lentils. The operating principle is the stratification of the
product into heavier and lighter fractions by introducing air through the
bed of material. Pressure fans are located in the body of the machine, below
the vibrating deck. The vibrating deck pushes the heavier material which
is in contact with the deck upwards towards the stone discharge spout.
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A. Vandenberg
The lighter material flows with air assistance down the inclined, vibrating
deck and exits through the clean product spout. It is possible to individually
adjust the inclination of the vibrating deck, speed of eccentric motion, feed
rate and the air flow in order to achieve the optimum degree of separation.
Sizing
Cleaned lentils may be sized after cleaning based on consumer demand for
uniformity of specific diameter or thickness of seeds. Sizing adds cost to the
final product and demand for sizing can be very specific. For example, premiums may exist for extra-large green lentils in Spain. In some parts of
South Asia, for example in Bangladesh, premiums are paid for extra-small
red lentils that are dehulled but not split. In most cases, the separations
used prior to the destoning step will provide sufficient sizing to meet the
general consumer demand.
Colour sorting
Various types of electronic colour-sorting machines are available. They are
used opportunistically to add value to lower grade lentils, or to ensure
maximum quality for premium products. All colour-sorting machines used
in the lentil industry were adapted from other industries, for example from
the rice or coffee industries. The basic principle is to use computerized digital imaging to compare the colour of lentils passing by sensors that are
adjusted to an acceptable predetermined colour range. Lentils with colours
that lie outside the selected colour range are rejected by removing them
from the product stream using air jets that change the trajectory of rejected
lentils. Some sorting machines convey the lentils on horizontal belts with
downward-facing air jets. Most colour sorters are designed to have downward flow of lentils combined with horizontal air jets.
Colour sorters are used to add value to lentils by improving uniformity
of colour for specific market classes, but only when it is economically feasible to do so. Typical situations where colour sorting of whole lentils may be
used are situations where lentils that are discoloured as a result of weathering of seedcoats during wet harvests or discoloration due to frosts (both can
occur in Canadian lentil production). Other situations where optional colour
sorting may be economical could be removal of seeds that are stained by
fungal disease like Ascochyta blight or admixtures of lentils with contrasting seedcoat colour. The decision as to whether or not colour sorting is economically feasible is determined by the differential in price between the
sorted (value-added) lentils and the unsorted material, the operation and
investment cost of the colour-sorting system, and the proportion of lentils
required to be removed from the unsorted product. A requirement to
remove too many or too few lentils reduces the economic efficiency of the
sorting process and the potential added value.
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Packaging
After whole lentils pass through the various types of cleaning and separation, they are packaged and shipped as demanded by local or export specifications. Various types of bagging, containerization and shipping methods
can be used. Cleaned lentils may be shipped in bulk-hopper cars for delivery to bulk shipping vessels for export to bulk unloading facilities in port
facilities for bagging. In Canada, some exporters may temporarily position
a completely mobile grain-cleaning system beside the railway to fill hopper
cars for bulk delivery to the port. For some customers, primary processors
fill large polypropylene tote bags that are loaded into standard 20 or 40 t
shipping containers. In some cases, polyethylene-lined shipping containers
are filled with bulk-cleaned lentils. Bulk shipments are often received at
facilities for re-cleaning and packaging in smaller consumer-ready packages
under individual brand names. Typical consumer packages are polyethylene
bags in 500 or 100 g portions. In regions with high lentil consumption, packages may be up to 15 kg, particularly for restaurant and institutional use.
One of the most common forms of shipping-cleaned lentils is preprinted polypropylene bags or jute bags that are loaded into shipping containers or trucks, depending on the export origin and the destination. Bag
size, bag quality and labelling information are predetermined by the purchaser. Typical size ranges from 25 to 100 kg. The bags may be loaded manually or mechanically into shipping containers, or bags may be piled and
then shrink-wrapped on pallets, depending on customer preference. Based
on changing consumer requirements and market trends, bagging systems
are flexible. The packaging system for lentils is also highly influenced by
packaging trends in the other crops such as rice.
Screenings or dockage from the various cleaning processes are generally accumulated and then used to feed livestock. Each stream in the cleaning process may be kept separate, for example split lentils, damaged lentils
and weed seeds may be stored together as they accumulate through the
cleaning system. Products such as hulls, pods and chaff that are removed
by air separation may accumulate separately. This material is generally sold
to farmers with livestock or sold to feed-compounding companies. Screenings are passed through hammer mills, and then blended with other feed
grain products and dietary supplements. Water is added and the mixture is
then heated and pelleted for use as animal feed.
24.3. Secondary Processing

The main form of secondary processing of lentils is decortication or dehulling, which is the removal of the seedcoat from the cotyledon. The dehulling
process is usually followed by polishing or splitting or both before the lentils are consumed in soups or stews. The basic economic concepts that relate
to milling of lentils are milling efficiency, dehulling efficiency and football
recovery. The term milling efficiency in pulses may be defined in a variety
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A. Vandenberg
of ways. Ehiwe and Reichert (1987) described dehulling efficiency in terms

of the percentage of hull removal from the cotyledon and the yield of the
dehulled grain obtained from this process. Wang (2005) defined milling
efficiency as the sum of the percentage of whole dehulled seeds and split
dehulled seeds recovered after decortication. Football recovery is defined as
the proportion of dehulled unsplit seeds recovered after decortication. Football lentils are often sold at a premium to splits. Regardless of terminology,
milling quality of the lentils influences economics because the by-products
of milling (seedcoat, endosperm, broken pieces and flour) are of comparatively low value. A more precise definition of dehulling efficiency was provided recently (J. Bruce, 2008, unpublished results) as the percentage of the
total split and unsplit cotyledons whose outer surface has more than 98% of
the hull removed during the decortication process. Maximizing milling efficiency, dehulling efficiency and football recovery define the profitability of
secondary processing of lentils.
This form of secondary processing is known as dhal milling in South
Asia, where it has been practised for millennia. An up-to-date review of the
current status of dhal milling of the wide spectrum of pulses produced in
India is provided by Ilyas and Goyal (2005). Removal of the seedcoat is an
ancient process, originally practised at the household level with mortar and
pestle or hand-driven rotating horizontal millstones. Removal of the seedcoat reduces cooking time for pulses, and generally increases the concentration of nutrients in the decorticated seeds because the seedcoat is mostly
cellulosic material. Seedcoat thickness varies depending on the crop, from
as low as 5% in some varieties of cowpea (Vigna unguiculata) to 30% in some
lupine species (Kurien, 1984).
The seedcoat of lentil tends to be thinner than that of most other
legumes (Hughes and Swanson, 1986). It normally ranges between 6 and
7% of the seed weight (Singh et al., 1968; Erskine et al., 1991). Wang (2005)
reported the seedcoat seed weight for Canadian red lentil as 7.3%. Largerseeded lentils tend to have lower percentage loss during decortication
because the proportion of hull to seed mass is lower. Erskine et al. (1991)
found that lentil seeds with a mean diameter of 4 mm lost about 8.2% of
their weight during decortication, whereas losses from seeds sized 3 mm
averaged 9.8%.
The two countries with the greatest amount of lentil milling capacity
are Turkey and India (Williams et al., 1993). Virtually all lentil milling is
focused on red lentils, but there are some countries, for example the Aegean
coast of Turkey, where dehulled yellow cotyledon lentils are consumed.
Yellow cotyledon lentils are also milled to a very limited extent in Canada.
Since India produces the largest volume of pulses and also of red lentils in
the world, it also has the most dhal mills. Ilyas and Goyal (2005) stated that
India has more than 7000 dhal mills, and that no standard process is followed because of the wide variety of pulse crops, varieties, environments
and treatments. Turkey has a large red lentil milling industry for both
domestic consumption and export. Most mills were originally located in the
Gaziantep region of south-eastern Anatolia. More recently, larger mills were
399
built or relocated primarily to the Mediterranean port city of Mersin. In the

past 10 years, lentil milling capacity has expanded in both Sri Lanka and
Egypt, two of the worlds largest importers of red lentils. Significant red
lentil milling capacity also exists in Syria and the United Arab Emirates.
With the expansion of red lentil production in Australia and Canada that
started in the mid-1990s, milling capacity has also begun to expand. Rising
costs for energy ultimately raised crop production and transportation costs.
It is expected that milling capacity will continue to expand in exporting
countries in order to reduce shipping costs.
The decortication process for pulse seeds is influenced by the structure
of the seedcoat and the extent to which it is bound to the cotyledon. Usually
a gum (such as galactomannan) or lignin layer binds the cotyledons to the
hull (Siegel and Fawcett, 1976). Muller (1967) demonstrated that among
pulse species, variability in the depth and tackiness of this layer results in
different binding ability between the hull and the cotyledons. He considered lentils to be in the category of pulses that are relatively easy to dehull
compared to pigeon pea (Cajanus cajan), green gram (Vigna radiata) and
black gram (Vigna mungo).
The general process for abrasive dehulling of lentils is outlined in
Fig. 24.2. The process begins with sized-cleaned lentils and proceeds
through various necessary and optional steps. Similar to the scenarios of
primary processing, the actual commercial setup of modern dehulling
plants is highly variable depending on environment, production system,
primary processing capability, economics, availability of red lentils, and the
cost of purchase, maintenance and operation of machinery. No two dehulling plants will be set up the same way. Much of the equipment used in the
process is borrowed from other industries that process other grain products, and in some cases the equipment is modified or manufactured exclusively for red lentil dehulling.
The pitting, water addition, heating and cooling steps are designed to
reduce the natural binding of the seedcoat to the cotyledon. The dehulling
step produces both whole dehulled footballs, plus dehulled splits which
can be separated by optional sieving or left mixed. If customers specify split
lentil products, it is also possible to use a second set of horizontal millstones
that can be adjusted to produce 100% split products from the dehulled lentils. Individual lots and varieties may dehull more or less easily, depending
on environmental factors and interactions that affect final seedcoat and
seedcoat development. One of the key factors in maintaining high milling
efficiency is deciding how to handle specific lots of lentils to maximize
return.
Specific information on methods and techniques are usually held in
strict commercial confidence. Through in-house design, modification and
innovation, some steps in the general procedure outlined in Fig. 24.2 may
be eliminated, modified to suit local conditions, or added as a means of
reducing cost or improving quality of the final milled lentil products.
Dehulling also has a seasonal influence. Many millers report that successful
milling requires that the lentils be allowed to fully equilibrate and mature
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A. Vandenberg
Operation
Comments
Cleaned and sized lentils

Pitting
Abrasion allows moisture transfer
Re-cleaning
Removes fines (very small particles of

broken grains) and brokens
Water mixing
Adjusts moisture content
Steeping
Equilibration
Heating
Separation of hull and cotyledon tissue
Cooling
Dehulling
Hulls
Feed and
waste
Removes seedcoat
Sieving
Fines Wholes
Separates splits and footballs

Splits
Aspiration
Colour sorting
Polishing
Removes discoloured material

Water or oil
Packaging
Fig. 24.2. General outline of steps in the manufacturing of whole and split dehulled
lentils (Source: adapted from Matanhelia, 1980; cited in Ilyas and Goyal, 2005).
in storage for at least 6 weeks after harvest before milling can begin. The
cold winter temperatures of Canada also pose specific problems for milling
and handling of lentils. One of the key factors in successful dehulling is
management of the moisture content differential between the seedcoat and
the cotyledon. Modern mills have the ability to pit seedcoats and manage
the moisture content before dehulling through a tempering process that
involves water addition, heating and cooling.
Approximately 80% of the worlds total lentil production is of the red
cotyledon type. Most of these red lentils are consumed after secondary processing that removes the seedcoat by abrasive milling. When the seedcoat is
removed, the cotyledons may remain intact (known as the football type)
and the product is sold to consumers specifically in the unsplit form. In
many countries, the final product from splitting mills is the red split lentils.
In general, there are very few published scientific investigations related
specifically to the decortication and splitting process in lentil. Partly in
response to the recent expansion of red lentil production and secondary
processing facilities in Canada, Wang (2005) published a useful baseline
401
laboratory method for assessing milling efficiency in lentil, based on conducting evaluations at 12.5% moisture using a Satake dehuller. This procedure
can be easily used to assess the effects of various agronomic, environmental
or genotypic effects on lentil milling efficiency. This protocol was recently
used to assess the effect of the agronomic practice of chemical desiccation
by diquat on milling efficiency (Bruce, 2008). Desiccation treatments were
applied in four environments at early, recommended and late pod maturity
stages for eight red lentil varieties. These were compared to similarly timed
swathing treatments. Under ideal harvest conditions, differences due to
treatments were minor. In environments that experienced harvest delay
caused by wet weather conditions during the harvesting period, all three
milling parameters (milling efficiency, dehulling efficiency and football
recovery) were significantly reduced for all varieties by the early desiccation treatment compared to all other treatments.
Split or football lentils are usually subjected to a polishing process to
remove fine flour dust as a means of improving the visual quality of the
product. Polishing with water is a common practice that is accomplished in
some facilities by adding a small amount of water to the product stream as
it passes in a horizontally mounted screw conveyor prior to bagging. Adding a small amount of vegetable oil (usually locally available products like
sunflower or canola, for example) to the product stream is also used for red
lentils destined for certain markets, especially in the Middle East. The oil
penetrates the tissues of the dehulled seeds and deepens the colour of the
red cotyledons. In some cases the lentils are double-oiled for specific markets. Oiling is also a traditional method of preparation of dehulled pulses in
South Asia. The practice is commonly used in Africa and India at the household level to control grain storage insects such as Bruchus ervi Froel.
(Coleoptera: Bruchidae occurring in Europe, North Africa and South-west
Asia) and Bruchus lentis Froel. (which occurs in the USA, Europe, North
Africa and South-west Asia) and India (Periera, 1983).
In some regions, lygus bugs such as Exolygus pratensis L., are a serious
insect pest of lentil crops because feeding adults damage the seed, causing
chalky spot syndrome (zberk et al., 2006). Seeds with chalky spot have pitted, crater-like depressions in the seedcoat with or without a discoloured
chalky appearance that reduces quality and in turn causes economic losses
of 510 % to the producer. In Turkey, the value of dehulled lentils is maintained somewhat by ensuring a high recovery rate of lentils in the football
form, specifically in red lentil where split cotyledons may display chalky
spot on the inner surface of the cotyledon. By avoiding or minimizing the
splitting process, the chalky spot problem is not as obvious and the economic losses are minimized.
The colour-sorting process (Fig. 24.2) after dehulling or splitting can
add considerable cost to the final product depending on the frequency and
nature of the defects that are obvious after milling. Lentils with adhering
hulls can be detected by electronic scanning devices; this is achieved based
on colour comparisons of particles passing the scanning device. The machines
can be adjusted to reject particles of a specified colour or colour contrast.
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A. Vandenberg
Removal from the product stream occurs by using finely tuned compressed
air jets to change the trajectory of the rejected lentils. The reject stream can
be redirected to the dehulling process if the reason for rejection is failure to
remove the seedcoat. Reduction in final product quality can be caused by
variation in the cotyledons, for example, admixture of yellow cotyledons in
red lentils, or vice versa. The specific problem of dark green immature cotyledons from normal-sized seeds can occur in some seasons or regions, for
example in Mediterranean climates that experience the rapid onset of high
temperatures prior to harvest. Extreme high temperature may cause rapid
dehydration of the cotyledons, preventing normal colour development. The
efficiency and cost of colour sorting of dehulled and split lentils is influenced by the colour contrast of the defect, the frequency of the defect and
the initial and maintenance cost of the colour-sorting devices which are
available from several manufacturers.
Thermal processing in glass jars or cans is probably the second most
common form of secondary processing of whole lentils, especially in regions
where consumers purchase these products instead of dehulled lentils. European and North American consumers are generally more familiar with
pulses in this form of consumer product. Thermal processing may alter the
nutritional profile of the lentils, depending to some extent on the temperature used in the process and the chemical composition of the brine solution
used in the canning procedure. Whole lentils with intact seedcoats may be
added to other ingredients to make stews or soups in ready-to-eat form. In
some countries, for example Spain, whole lentils in jars or cans, usually
large green or Pardina types, are readily available for use as a quick side
dish. Variations on the thermal process include packaging of prepared lentil dishes in foil or plastic envelopes or bags.
Another form of secondary processing of lentils and other pulses is
infrared drying (Cenkowski and Sosulski, 1997). Infrared treatments are
applied to rehydrated whole lentils that are rehydrated to a moisture content
of 1939%. Using near infrared wavelengths of 10001500 nm, this process
can quickly reduce moisture content and cause changes to cooking properties after dehydration. The process reduces cooking time and changes starch
gelatinization and solubilization due to changes in porosity and water diffusivity (Scanlon et al., 2005). This process has been commercialized for
whole lentils and other grain products (Infraready Products Ltd, 2008) to
produce quick-cooking whole grains. For lentil, the cooking time is reduced
to 1520 min, similar to the cooking time for dehulled lentils.
Lentils in dehulled or whole form may also be ground into flours for
speciality applications in food processing. For example, dehulled green
lentils may be used as an ingredient to make fortified pasta products
(Barilla, 2008). Addition of lentils increases both the protein and the fibre
content of the pasta. Whole or dehulled lentils may also be used as ingredients in extruded food products such as spiced snack mixes, which are
especially common in South Asia. Other speciality uses of lentils include
deep frying of soaked lentils for use in snack mixtures. In some parts of
the world lentils are used as sprouts, much like mung bean, to increase
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the nutritional value. In North America and Europe, lentils in cooked form
are used in salads with other vegetables. For most of these uses, lentils
represent one of many similar ingredients. Most of the value addition for
lentil is done at the level of street vendors, small restaurants or boutique
food manufacturers.
Over the past decade (19982008), food scientists have increased their
efforts into studying the potential uses of lentil in tertiary food processing
applications such as substitutes or enhancements for existing ingredients
derived from other foods. In order to understand the potential for using
lentils in food processing, it is necessary to understand the fundamental
technical processing characteristics of the fibre and starch in lentils (see
Grusak, Chapter 23, this volume). For example, processing properties of
fibre include composition (amounts and types of cellulose, pectin, neutral
polysaccharides, cellulose, lignin, oligosaccharides) and processing properties such as solubility, viscosity, gelling, swelling, fat binding, water binding and protein compatibility. This matrix of properties affects the potential
application of using lentils in processes such as cooking, extrusion, air classification, germination, fermentation, and fractionization and separation.
Recent scientific publications and patents focus on the possibility of using
lentil for further food processing and inclusion of lentils in meat products,
bakery products, dairy products, pasta products, speciality food products
in dietetic applications (for example, gluten-free products for celiac diets)
and snack foods (K. Ablor, 2008, Saskatoon, personal communication). Similarly, the specific properties of starches from lentils can be described in
terms of concentration of amylose, amylopection, gelatinization temperature,
enthalpy, retrogradation temperature, resistant starch content, starch digestibility, pasting temperature, viscosity and starch granule structure.
Many gaps exist in global knowledge about food processing applications for lentils. Attributing processing properties to specific genotypes
and environmental conditions is even more challenging, since many of the
processing characteristics have not been fully described for even one genotype. Fundamentally, the question is one of economics. Most food characterization and processing techniques were developed for other major
commodity grain crops such as maize, rice, wheat and among the food
legumes, soybean and groundnut. Lentil is a relatively minor crop on a
world scale. About 3.5 million t are produced on an annual basis, spread
out over 48 countries (see Erskine, Chapter 2, this volume; FAO, 2008). In
the 19972007 period, global lentil production increased from 2.7 to 3.5
million t. In comparison, during the same period, global soybean production increased from 123. 7 to 196.4 million t, an average annual increase
of 7.3 million t equal to more than double the entire annual total world
lentil production. The major crops are also the major focus of the food
processing industry because of their relatively high availability and relatively low price in comparison to lentils. The major crops have higher
returns on investment in value-added processing because they are often
subsidized in the production systems because of their fundamental importance to national food supply.
404
A. Vandenberg
24.4. Future Trends in Processing and Value-added Activity

Export-oriented pulse-crop industry associations and pulse-crop producer
organizations in many parts of the world have begun to engage in research
and development activity related to improving the consumer acceptance of
pulses, including lentils. A major theme of these efforts is to raise the profile
of pulse crops as an excellent source of nutrition in the human diet. One of
the current trends in North America is a focus on health and wellness issues
related to obesity, type 2 diabetes and other metabolic syndrome conditions.
Increasing pulse consumption combined with significant changes in lifestyle (exercise) and diet could be one way to provide increased health and
wellness benefits. However, this must be counterbalanced as a marketing
strategy by the fact that every other plant food producers are pursuing a
similar strategy.
An accompanying trend in North America is a focus on using pulses as
a food-processing ingredient. For reasons described above, the relatively
high cost of lentils reduces their prospects as a source of ingredients, except
for the lowest grade lentils that have reduced value as whole food. If consumers do respond to the use of pulses as ingredients, it is likely that the
pulse-crop ingredient of choice will be field pea because of its higher yield
which makes it cheaper. The underlying assumption is that consumer trends
for increased convenience and improved nutrition will lead to expanded
use of lentils and other pulses in value-added processing of new specific
consumer products. Based on the above economic comparisons with other
competing food grains, this may be a major challenge.
A third trend in food marketing, is that of re-introduction or expansion
of pulses in the diet as a traditional cultural food item as part of the return
to the concept of eating wholefoods that are not highly processed. In North
America and in Europe, lentils and other pulses can be re-positioned in the
marketplace as part of a healthier diet with cultural origins in the Mediterranean, the same region where lentil was domesticated. In contrast, in most
parts of Asia and the Middle East, lentils have never lost their traditional
status as a healthy wholefood.
The potential for value addition for lentils in the food industry anywhere in the world is ultimately dependent on the perceived nutritional
and culinary value that lentils can provide. Similar to the macronutrient
profile of many other pulses, lentils are approximately 25% protein, with
the remainder providing carbohydrates and fibre, along with small amounts
of micronutrients and vitamins. The carbohydrate profile is relatively high
in resistant starches which digest more slowly, providing fibre-like nutritional benefits. If pulses are processed to add value on the basis of the economics of yield and unit cost of macronutrients, lentil will not be competitive
even with higher yielding pulse crops such as pea. In comparison to the
major cereal grains and food legumes, many of which are much higher
yielding, the unit cost of macronutrients in lentils is simply not competitive.
The concept of value-added processing is based on the premise that the
ingredients are inexpensive. For food products that are already relatively
405
high in value because of lower yield and lower availability, like lentils, the
value-added potential is restricted to boutique food products. The availability of a wide variety of seedcoat colours and sizes and red, yellow and green
cotyledons may help add value as lentil products are more closely tailored
to specific consumer preferences.
Alternatives to value-added processing include value-added marketing,
a scenario in which lentils are marketed as a balanced and healthy wholefood alternative to highly processed foods which will increasingly be made
from a much smaller group of global crops with high productivity and volume. One distinct advantage and highly competitive advantage of lentils,
particularly in dehulled form, is the fact that they cook in about the same
time as milled rice, providing both convenience and energy savings.
Another avenue of current scientific investigation that could influence
future value-added marketing concepts for lentils, is the role that lentils
may play in providing sustainable metabolic energy for athletic performance (Little, 2007). This may lead to innovative marketing strategies that
promote the perception of improved nutritional value for lentils. A third
value-added marketing strategy is one that emphasizes some of the inherent nutritional benefits of lentils, more specifically their micronutrient and
vitamin content. They are known to be excellent sources of Fe, Zn, other
micronutrients and folate and may provide high levels of low-cost bioavailable nutrition. This is a wide-open area for future research that may help
prevent the relative global decline in lentil consumption.
Based on recent economic development trends in the Middle East and
other regions where lentil milling was historically established, it is likely
that many of the small dhal mills in South Asia will be replaced by less
labour-intensive and larger modern mills in the major port regions of consuming countries. This has already occurred in Turkey, where many small
lentil mills were replaced by more modern facilities at the port in Mersin,
where exporting and importing could be accommodated more easily. A
similar situation may develop in South Asia as labour rates and demand for
higher milling efficiencies increase. Concurrent with this trend could be
increased milling capacity in large and growing production regions like
Canada and Australia. Rising energy costs that cause transportation costs to
increase will probably make it more economical to ship dehulled lentils. As
dehulling mills become larger it may be feasible to explore the potential for
value addition from waste products like hulls that are now used only for animal feeding. The economic feasibility of phytochemical extraction for specific
compounds such as antioxidants requires evaluation in future.
At this point in time, the future of the global lentil industry may depend
entirely on developments in the South Asian countries, where most lentil
consumption occurs. If South Asian consumers, in particular in India, develop
food habits that reduce lentil consumption in favour of other pulses and soybean, lentil production and consumption will decline globally. Although
recent trends suggest that per capita consumption of all pulses is declining
in South Asia, there is debate as to whether the decline is due to higher
prices or changing consumer preferences. Some evidence (Kyi et al., 1997;
406
A. Vandenberg
Kumar, 1998) suggests that as incomes rise in South Asia, pulse consumption increases. If this is true, the future of value-added processing in lentil
may be closely linked to the degree that lentils produced in other countries
can successfully supplement the role that summer pulses such as pigeon
pea, black gram and green gram play in traditional South Asian cuisine. A
similar situation has occurred over the past 15 years with respect to the substitution of yellow pea dhal and flour for desi chickpea dhal and flour. Consumption of imported yellow pea in India has risen from almost nil to more
than 1 million t in the past 15 years.
The future of lentil consumption globally may also be related to the
extent to which South Asian cuisine is popularized in the rest of the world.
Food trends now move easily around the globe, and if consumers in developed countries develop greater acceptance of food products based on South
Asian dishes, lentil consumption will also increase.
Value-added processing will increasingly come under the influence of
the cost of energy inputs into total food systems, from the production end
right through the chain of value to the end consumer. This will affect all
aspects of lentil production, including the scale of primary, secondary and
tertiary processing.
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barillaus.com/Home/Pages/Barilla_Plus__Protein.aspx (accessed 8 July 2008).
Bruce, J.L. (2008) The effects of preharvest treatments on the milling efficiency of red
lentil. MSc thesis, University of Saskatchewan, Saskatoon, Canada.
Cenkowski, S. and Sosulski, F.W. (1997) Physical and cooking properties of micronized
lentils. Journal of Food Process Engineering 20, 249.
Cenkowski, S., Sokhansanj, S. and Sosulski, F.W. (1989) Equilibrium moisture content
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Davey, B.F. (2007) Green seed coat colour retention in lentil (Lens culinaris). MSc
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Erskine, W., Williams, P.C. and Nakkoul, N. (1991) Splitting and dehulling lentil:
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Kumar, P. (1998) Food Demand and Supply Projections for India. Agricultural Economics
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20(1), 313.
25
Food Preparation and Use

Rita S. Raghuvanshi and D.P. Singh
G.B. Pant University of Agriculture and Technology, Pantnagar, India
25.1. Introduction
Cereal grains and grain legumes are the major source of calories and proteins for a large proportion of the worlds population. Legumes are a rich
source of proteins and the essential amino acid, lysine, but are usually deficient in the sulfur-containing amino acids, methionine and cystine. On the
other hand, cereal grains contain lower concentrations of proteins that are
deficient in lysine but have adequate amounts of sulfur-rich amino acids
(Eggum and Beame, 1983). It is therefore emphasized that legumes are the
natural supplement to cereals in ensuring essential amino acid balance.
Also, it is being increasingly recognized that legumes are good source of
vitamins and minerals.
In countries where non-vegetarian food is expensive and its consumption restricted for cultural reasons, the protein-rich lentil provides a cheap
substitute. Lentils are lower than most pulses in antinutritional factors such
as haemagglutinin, oligosaccharides and flavogens. They contain tannins in
the seedcoat but not in cotyledons (Vaillancourt et al., 1986). In small-seeded
lentil (microsperma) with red cotyledons even this is not important as the
testa is frequently removed before use in culinary preparations. The macrosperma (large-seeded) lentils also contain tannins, which can cause digestive
disorders. Lentils are a major component of food in the Mediterranean
region and the Middle East. In the Indian subcontinent lentil is mostly eaten
as porridge or thick soup called dal.
25.2. Physio-chemical Properties and Cooking Quality of Lentil

The chemical composition of lentil seeds indicates that lentils can be the
perfect diet as they contain little fat (~12%) and large concentrations of
408

409
proteins with an energy value of 2432% and minerals (iron, cobalt and
iodine) (Kowieska and Petkov, 2003). Savage (1988) reported that lentils by
virtue of their high protein content could be processed to produce many
products similar to those produced from soybeans. Leucine was the major
essential amino acid in lentil flour. Lentil is rich in lysine and arginine, and
can fulfill the essential amino acid requirement except for the sulfur-containing amino acids and tryptophan (El-Zoghbi, 1998).
Pasantes and Flares (1991) reported taurine content of lentil as 25
nmol/g, 50% of which is leached out during soaking, while the cooking
procedures in the absence of presoaking did not affect the same. Sanchez
et al. (1994) reported that lentil contained higher concentrations of nonstarch polysaccharides than cereals and a greater portion was in the form of
soluble non-starch polysaccharide. Paliwal et al. (1987) reported that lentil
chuni (groats), a by-product from the lentil processing industry produced
during dal manufacturing, contains 23.3% crude protein, 1.6% ether extract,
21.6% crude fibre, 48.2% nitrogen-free extract and 5.2% total ash. Candella
et al. (1997) reported that cooking decreases the mineral content of lentils
whereas warm holding practices resulted in decreased isoleucine, valine
and leucine and significant increase in lysine, phenylalanine and tyrosine.
The analysis of seven cultivars of lentil for cooking time revealed that the
least amount of time required for cooking was 40 min and was positively
correlated to total ash (r = 0.24), fibre content (r = 0.18) and seed size (r =
0.12), and negatively correlated with protein content (r = 0.19) (Raghuvanshi
et al., 1994).
Even though lentils are considered to be highly nutritious, they contain
antinutritional factors such as trypsin inhibitors, haemagglutinins and oligosaccharides that cause flatulence. These problems can be greatly reduced
by heating and germination. Estevez et al. (1991) found that heat treatment
reduced activity of trypsin inhibitors and significantly increased digestibility of seed proteins. Germination of lentil for 6 days at 20C followed by
cooking resulted in breakdown of disaccharides and destruction of phytic
acid thus improving digestibility and availability of calcium, magnesium,
iron and zinc (Garcia, 1991). In another study, Camacho et al. (1992) found
that germination of overnight-soaked lentil at 25C and 85% relative humidity (RH) for 3 days in a solution of sodium hypochlorite resulted in an
increase in crude protein content with an increase in almost all amino acids;
however, the concentrations of ash, ether extract and phytates decreased.
25.3. Food Uses

Lentil is the most desired legume in many lentil-producing regions because
of its high average protein content and fast cooking characteristics (Fig. 25.1).
It can be used as a starter, main dish, side dish or in salads. Seeds can be
fried and seasoned for consumption as snacks. Flour is used to make soups,
stews and pures, and mixed with cereals to make bread and cakes, and as
a food for infants. Young, green, fresh lentil pods are eaten raw or steamed
410
Food uses
Green pods
Whole grain
Vegetable
Curry
Salad
Salad
Soup
Dal
Namkeen (salty and
spicy snacks)
Snack sprouts
Lentil fritters
Lentil sprout spread
Lentil burgers
Canned food mix
Decorticated/split
Flour
Medicinal uses
Fodder uses
Soup
Dal
Khichri
Kebab
Dosa
Soup
Papad
Vermicelli
Infant food
Lentil loaf
Therapeutic diet
Paste for surface application
Lentil water for external washes
Broken stem
Branches
Pod walls
Leaflets
Fig. 25.1. Uses of lentil.
like green peas, while lentil sprouts are added to salads, soups, breads and
savoury dishes.
25.4. Methods Used in Lentil Preparations

There are several methods used in lentil preparation, which provide different texture and nutritional quality to the final products. The methods are
detailed with their effects on nutritional quality.
411
Soaking
Washing/cleaning of lentil is done before soaking, which helps to remove
chemicals and other unwanted matter. However, water-soluble nutrients
are lost during the process. This can be reduced by quick thorough washing. Soaking is done in fresh or saline water with sodium chloride or sodium
bicarbonate. It hastens the process of cooking. Prodanov et al. (2004) found
that soaking lentil in different solutions (citric acid solution, pH 4.96 0.02;
distilled water, pH 7.00 0.02; and sodium bicarbonate solution, pH 7.85
0.02) caused losses of 510% thiamin and 2642% available niacin while
riboflavin increased up to 98%. The losses were greater with soaking in alkaline solution. An appreciable increase in hemicelluloses and neutral detergent fibre was seen when lentils were soaked in 0.1% citric acid solution
and 0.07% sodium bicarbonate solution (Vidal et al., 1992).
Boiling
Boiling is probably the most frequently used method for cooking lentils. It
may be open-pan boiling or pressure cooking. In open-pan boiling, lentils are
cooked by immersing them in boiling water until tender. While using openpan boiling for cooking, the heat is turned down when the contents start boiling as violent boiling throughout may result in breakage and disintegration
of grains. Continued vigorous boiling only results in excessive evaporation of
water, as a result the lentils may be burnt at the bottom. If excess water is
used for cooking and the water is discarded, 3070% of the water-soluble
nutrients may be lost. Over-boiling of lentil may make it mushy. Pressure
cooking raises the temperature above 100C and reduces cooking time.
The losses of fats and carbohydrates in lentil are more or less similar for
open-pan boiling and pressure cooking, but more proteins are retained in
pressure cooking (Pratibha et al., 1999). The in vitro protein digestibility of
proteins (81.0%) was greater in pressure-cooked lentil as compared to openpan boiling (Naveeda and Jamuna, 2006). Pressure cooking caused 23.5%
loss of tryptophan against 32.9% in open-vessel cooking (Parihar et al., 1996).
Although boiling results in the loss of nutrients, it simultaneously reduces
concentrations of antinutritional factors and improves the nutritional value
of lentil. Heating lentils at 120C at 1 atmospheric pressure for 30 min
decreased trypsin inhibitor activity, phytates and tannin content by 76%,
8% and 12%, respectively (Porres et al., 2003). Warm holding is a widely
used technique in catering services. Candella et al. (1997) studied the effect
of warm holding on the total composition and amino acid content of kidney
bean, chickpea and lentil, of which lentil was found to be most affected.
Sprouting/germination
Lentil seeds are easy to sprout. To have good home-made sprouted lentils,
seeds are soaked in water overnight and the free water is drained in the
412
morning. The grains are washed two to three times without rubbing. The
soaked seeds are covered and kept in a warm dark place. One more washing is done at night and all the water is drained and the seeds are kept
again as before. Sprouts of about 2.0 cm length are ready by the next morning. Porous containers like an earthen pot or humidity chamber should be
used for best results.
The presence or absence of light during germination had no significant
effect on the thiamin and riboflavin content in germinated lentils while niacin
content was higher when germination took place under light. The increased
number of rinses during the germination process had no significant effect on
the thiamin content while riboflavin and available niacin increased significantly. However, no effect was seen on riboflavin when seeds were germinated for 6 days and on niacin when germination was carried out for 3 days in
the dark. A longer germination time led to recovery of thiamin content and a
significant increase in riboflavin and niacin content (Prodanov et al., 1997).
Hulled seeds should not be used for sprouting, as they may split and the
embryos be injured. Soaking lentil seeds for 12 h and then sprouting for 34
days has been found to completely remove all haemagglutinin and amylase
inhibitor. A marked increase in moisture content and reduction in protein, fat,
fibre and ash content was observed in sprouted lentil; however, the protein content of sprouts increased on a dry weight basis (Vanderstoep, 1981). Germination resulted in total elimination of a-galactosides and reduction in total starch
content and trypsin inhibitor activity (Urbano et al., 1995). The vitamin content,
that is thiamin and niacin (Urbano et al., 1995), riboflavin and biotin (Hozova,
1994) and ascorbic acid (Hsu et al., 1980), also increased in sprouted lentils.
Fermentation
Fermentation is the process of breaking down complex compounds into
simpler ones with the help of enzymes and bacteria. Fermented foods are
more nutritious than their unfermented counterparts because of improved
protein quality, increased vitamin B synthesis and reduced antinutritional
factors. Fermentation of lentil for 4 days resulted in a slight increase in
crude protein content and significant increase in in vitro protein digestibility (Shekib, 1994), whereas fermentation for 2 days reduced total galactosides and trypsin inhibitor activity by 26% and 50%, respectively (El-Rahman
et al., 1998). Lentils can also be processed by extrusion cooking. However,
application of lactic acid fermentation as a pretreatment was suggested by
El-Rahman et al. (1998) to increase the consistency coefficient, yield stress
and thixotropy values of lentil slurries during the first 24 h of fermentation.
Frying
The various products prepared from lentil using the frying technique are
papad, cutlets, dosa, pancakes, etc. Frying may involve deep-oil frying,
413
shallow frying and sauting. Cooking is rapidly completed in deep-oil

frying as the temperature is 180220C. Deep frying of food increases
the calorific value due to absorption of oil. In shallow frying, the food is
cooked in a larger volume of oil than sauting but not enough to cover
it. Heat is transferred to the food partially by conduction (i.e. by contact
with the heated pan) and partially by the convection current of the foods,
for example cutlets and patties. Sauting involves cooking in just enough
oil to cover the base of the pan (greasing the pan), for example dosa and
pancakes. The food is tossed or turned over occasionally with a spatula
to enable all the parts to come in contact with the oil and get cooked
evenly.
Dry-heat methods
When food is cooked uncovered in a metallic frying pan, the method is
known as pan broiling or roasting. The advantage of the method is that it
reduces the moisture content of the food and thus improves its keeping
quality; however, losses of nutrients such as amino acids can occur if the
food becomes brown. Urbano et al. (1995) found that roasting resulted in
reduced starch content and trypsin inhibitor activity in lentils.
Microwave cooking does not require any medium of transfer of heat in
the cooking process as heat is generated inside the food, whereas in conventional cooking methods the heat is transferred to the exterior of the food
by conduction, convection or radiation. Both methods are equally effective
in destroying the antinutritional factor in lentil (i.e. trypsin inhibitor) without any adverse effect on the protein efficiency ratio (PER) of raw seeds
(Hernandez et al., 1998). The nutrient composition of microwave-cooked
lentil was also more or less similar to that in pressure-cooked lentil except
for thiamin, which was lost to a significant extent in microwave cooking.
The in vitro digestibility of proteins in microwave-cooked samples was
75.2% (Naveeda and Jamuna, 2006).
Baking is a dry-heat method of cooking but the action of dry heat is
combined with that of steam, which is generated while the food is being
cooked. The baking properties of lentil flour were evaluated by incorporating 5, 10 and 15% flour of germinated or dry lentils with wheat flour before
baking and it was found that mixing of up to 15% lentil flour with wheat
flour had no adverse effect on loaf volume (Hsu et al., 1980).
25.5. Preparations and Recipes

Usually, lentils are boiled to a stew-like consistency with or without vegetables and then seasoned with a mixture of spices to make many side dishes
such as sambhar, rasam and dal, which are usually served over rice and sometimes with roti (unleavened bread).
414
Salad
A salad is a dish that usually (but not always) takes off from a green leafy
base, usually piquant in flavour, mostly served cold with a cold dressing
made of oil, vinegar and special flavourings. The word salad comes from a
Latin word meaning salted. A simple recipe to prepare lentil salad at home
requires about 225 g whole lentils, salt (two tablespoons; tbsp), plain flour
(one tbsp), freshly ground black pepper, olive oil (four tbsp) and one small
finely chopped onion. The method involves soaking the lentils overnight in
lukewarm water with cooking salt and flour. The following day the lentils
are removed from the water and the water is brought to a boil and then
cooled. The rinsed lentils are then added to this water and brought to the boil
and then simmered gently for 30 min. They are then strained, the water discarded and the lentils put into a large saucepan with plenty of fresh, lightly
salted, boiling water, covered and cooked until tender and then well drained.
While the lentils are still warm, they are seasoned to taste with salt and pepper and olive oil and finely sliced onion is stirred into them (Russell, 1974).
Soup
Lentils are used to prepare an inexpensive and nutritious soup all over
Europe and North and South America, sometimes combined with some form
of pork or chicken meat. Soup can be prepared from mature whole seeds,
dehulled lentil flour or splits. To prepare lentil soup, onion and celery are
fried lightly for 10 min in 25 g melted butter in a large saucepan, and 150 g
lentils and 1.1 l of stock with spices (quarter of a teaspoon of ground cloves
and quarter of a teaspoon of ground allspice) are added to the saucepan and
brought up to the boil. The heat is turned down when it starts boiling and
the lentils are allowed to simmer gently for 3035 min or until they are soft.
Then it is transferred to a liquidizer or food processor and processed until
smooth. The soup is reheated gently and seasoned to taste with salt and
black pepper (Good Housekeeping Institute, 1966). Some of the typical ingredients added into a lentil-vegetable soup may include green beans, lentils,
carrots, celery, tomatoes, spinach, potatoes, garlic and various seasonings.
The soup is made from a water base and as it cooks, a thick sauce-like consistency develops that enhances the flavours of all other ingredients.
In Morocco, a similar process is used to prepare lentil with vegetables
and the recipe is known as lentil tagine. The method of preparation involves
boiling 450 g lentil, 1.1 l of fresh vegetable stock and four cloves of garlic in
a saucepan and then simmering for 20 min. Sweet potatoes (675 g), tomatoes (550 g), capsicum (325 g), onions (325 g), ginger (one tbsp), cumin (one
teaspoon), cayenne pepper (one teaspoon) and salt (to taste) are added to
this and cooked for another 30 min or until the sweet potato is soft. The
soup is garnished with chopped coriander while serving.
Use of lentil in dehydrated form for preparation of lentil soup can offer
a unique opportunity for its consumption, providing value addition apart
415
Lentil seeds
Cleaning impurities
Dehulling
Frying onions (15 g) with olive oil (20 g) for 2 min
Adding lentil splits (200 g), NaCl (8 g) and deionized water (500 ml)
Cooking for 45 min

Moulding into aluminum frames (1 1 0.05 cm)
Drying at 75C using air-forced oven
Removal from frames
Dehydrated traditional lentil cubes
Fig. 25.2. Process flow chart for producing dehydrated lentil soup cubes (Source:
Ereifej, 1995).
from protecting it from insect damage (lentil seeds are prone to insect damage but dehydrated lentil soup cubes are packaged and not exposed to environmental factors leading to infestation). A process for the preparation of
acceptable and nutritious traditional lentil soup cubes has been developed in
Jordan by Ereifej (1995). Whole and sound seeds of Balady landrace and three
newly developed lentil cultivars (Jor-1, Jor-2 and Jor-3) were used. The
process consisted of cleaning, dehulling, frying cut onion with olive oil, addition of salt, lentil splits and water, cooking, moulding and drying (Fig. 25.2).
The soups prepared from all lentil cultivars had similar appearance and flavour and produced acceptable soup cubes, however, the commercial soup
scored higher for appearance and flavour. The cultivar Jor-1 was superior to
commercial soup with respect to protein content and energy value.
Dal
Lentil is predominantly eaten in South Asia as boiled or fried dal from
whole seed (from macrosperma) or split (microsperma) as lentil (masur) dal.
For preparing dal, whole grains are soaked in water overnight, drained and
then boiled in fresh water for 3040 min. It is consumed after salt and spices
416
are added. The split grains with or without husk are cooked by boiling in
water for 3040 min. Salt and spices are added to taste and the cooked mashed
dal is consumed along with a cereal preparation. The preparation appears as
a thick soup and is eaten with unleavened bread roti (chapatis) made from
cereals such as wheat or maize, etc. Boiled rice is also eaten with lentil dal. To
remove the seedcoats, the seeds are moistened with oil and water, dried in
shade and passed through a mill two or three times. To give an attractive
appearance, the dal is polished with magnetite powder and gritty powder.
To prepare lentil dal, approximately five cups of water are used for one cup
of lentil. Water is added to the saucepan and lentil is added when the water
begins to boil. It is boiled for 23 min and then the heat is reduced to simmering and dal is cooked until tender. Flavourings such as garlic and onion can be
added if desired; however salt should be added at the end otherwise lentil
tends to be tough. Different cultivars require different cooking times. Green
and brown lentils are cooked for approximately 45 min and red (dehusked)
lentils for 25 min. People in Hyderabad (South India) make a preparation called
khatti dal which is seasoned with tamarind juice and ginger garlic paste during
cooking. Mildly sweet, pungent and tart, the khatti dal dazzles the taste buds
and tastes great on its own or with rice and chapatis (Indira Singari, 2008).
In Assam (north-east India), lentil dal is prepared with several variations
such as cooking with bamboo shoots, Colocasia leaves (Colocasia anti-quorum)
or ferns. Lentil dal with bamboo shoots is prepared by pressure cooking 50 g
lentil dal in 600 ml water with salt (10 g), green chilli (5 g) and turmeric (5 g),
and then seasoned with grated bamboo shoot (30 g) and fenugreek seeds (5 g).
The nutrients provided by this amount of dal are 21.1 g protein, 12.9 g fat, 5.0 g
minerals, 20.5 g fibre, 4.8 g carbohydrate, 427.5 kcal energy, 97.8 mg calcium
and 7.2 mg iron. To prepare lentil dal with Colocasia leaves, 50 g lentil is
cooked with chopped Colocasia leaves, 10 g salt and 10 g sugar in 800 ml
water and then seasoned with 15 g garlic and 10 g black pepper powder using
15 g mustard oil. It has a higher amount of calcium (415.4 mg) and iron (12.3
mg). Lentil dal with fern is prepared by pressure cooking 100 g lentil with 40 g
tomatoes, 50 g ferns, 5 g green chilli, 5 g turmeric, 10 g salt and 10 g sugar in
650 ml water. The seasoning is done with 5 g mustard seeds in 15 ml mustard
oil with 15 ml lemon juice added while serving. It gives about two servings
and the nutrients available are 20.9 g protein, 10.8 g fat, 4.2 g minerals, 1.1 g
fibre, 62.6 g, carbohydrate, 432 kcal energy, 250 mg calcium and 8 mg iron.
The conventional method of cooking resulted in about a 2% increase in lentil protein. Diets containing cooked lentil and supplemented with different types
of meat (10% poultry, 15% mutton and 20% beef) significantly improved PER
(from 1.45 to 1.65), true digestibility (from 76 to 87%) and net protein utilization
(NPU) (from 43 to 51%) over non-supplemented diets (Bhatty et al., 2000).
Khichri
A combination of cereals and legumes cooked together is called khichri. It is
made from a mixture of dry masur dal and cooked wheat mixed in the ratio of
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3:7 or a mixture of split/dehulled lentil with rice. The mixture is cooked to a

soft consistency, salted and served as a staple (Nezamuddin, 1970). In West
Asia and North Africa the macrosperma lentils are used in mjeddarah made of
whole lentil and immature wheat seed. Mjeddarah, a Lebanese staple is also a
khichri-like product but the difference is that here lentil is combined with
brown rice and it is seasoned with onion, pepper and a few more spices with
vegetables added as optional ingredients. To add variety to mjeddarah, Parmesan cheese is sprinkled as a garnish (Wikimedia Foundation Inc., 2008a, b).
Another similar dish of mjeddarah is a basic component of Palestinian cuisine
with lentil and wheat. Sometimes rice is used in place of wheat with the same
procedure and the recipe is gluten free (Wikimedia Foundation Inc., 2008a, b).
Lentils are frequently combined with rice, which has a similar cooking
time and is known by the name of khichri, pulav or koshary. Because of time
constraints, various kinds of ready-to-eat, heat-eat and instant mixes are
available in the market. Shekib et al. (1986a) prepared an instant koshary (a
lentil-rice blend) using decorticated lentil and rice which has a shelf life of 21
days at 37C (moist atmosphere) and 60 days at 25C with a high nutritive
value as compared to rice and lentil alone. The methionine content was 0.94,
2.62 and 2.01, cystine 0.96, 1.92 and 1.54 and lysine 7.09, 3.41 and 4.70 g/16 g
nitrogen in lentil, rice, and the lentil-rice mixture in 1:2 ratio in koshary,
respectively. Their respective in vitro protein digestibility values were 63.9,
73.5 and 75.6% compared to 98.9% of casein (Shekib et al., 1986b). Khichri,
mjeddarah, lentil pulav or khoshary are all lentil-rice blends in different proportions. These preparations make the main dish of a meal. In Gujarat (western India), khichri is very popular, particularly for dinner. The process involves
soaking one teacup each of rice and lentils in plain water for 1015 min and
then pressure cooking with 56 cups of water with salt and other spices
added. Heat is reduced after the first whistle and the khichri is cooked for an
additional 45 min. Hot khichri is served with ghee (clarified butter) or curd.
Sweet pongal is another form of khichri made with one part rice (100 g), one
part dehusked split lentil (100 g), two parts jaggery (200 g), grated coconut
(20 g), raisins and cashew nuts (5.0 g), refined oil (15 g) and ghee (30 g). The rice
and lentil are slightly roasted in oil and then cooked in boiling water. When the
rice is about 75% cooked, jaggery, coconut and raisins are added and cooked till
soft but grainy in texture. It is served with ghee to give the distinct flavour. The
dish is part of the main meal; however, it is served towards the end in South
Indian cuisine. Sweet pongal is a protein-energy rich product, 100 g having
around 4.5 g protein, 4.4 g fat, 31 g carbohydrates and 185 kcal energy.
Vegetable
The hulled green seeds of macrosperma lentils are used as green vegetable.
The green pods of lentil are shelled and washed. One cup of shelled lentil is
boiled in water with salt. In a saucepan 20 g butter is melted. Cut garlic
(one teaspoon) and cubed ginger (one teaspoon) are put in hot butter. When
slightly pinkish, one cup of carrot cubes, half a cup of capsicum and half a
418
cup of boiled maize are added to the saucepan. It is kept on a medium heat
for 5 min then the boiled lentil is added to the mixture. It is sauted for a
while. Salt and black pepper are added to the mixture. It is again cooked for
a few minutes. The vegetable is garnished with chopped tomatoes, green
chilli and coriander leaves. It is usually a side dish with the meal. Having
green lentil as the main ingredient, the preparation has good quality protein along with vitamins and minerals.
Masur curry is prepared using 100 g lentil, 80 g tomatoes, 40 g spring
onion, 30 g oil, 60 g onion, two cloves of garlic, half a teaspoon of cumin
seed, a 2.5 cm length of ginger (diced), half a teaspoon of chilli powder, two
teaspoons of coriander powder, half a teaspoon of turmeric and salt to taste.
The lentil is cleaned, washed and soaked overnight. It is pressure cooked
with 400 ml water, turmeric and salt for 1012 min. In a separate vessel, the
seasoning is prepared by heating the oil and tempering it with cumin seeds.
Thereafter, garlic, ginger and onion are added. When the onion becomes
pinkish to brown, the chilli powder, coriander powder and tomatoes are
added and cooked for 23 min. The seasoning is put in the cooked dal. The
preparation will be around 300 ml of semi-solid consistency. One serving of
this curry (approximately 150 ml) has 8 g protein, 5 g fat, 12 g crude fibre,
78 kcal energy, 37 mg calcium and 2.89 mg iron.
25.6. Lentil Snacks

A fried and crispy salty snack prepared from legumes is called namkeen. In
the Indian subcontinent whole lentil grains are soaked, dried and deep
fried. The salt and spices are added and the finished product is eaten as a
snack. The fried lentil namkeen is made up of 16.6% protein, 20.5% fat, 0.59%
ash, 1.95% crude fibre, and provides 465 kcal energy, 45.9 mg/100 g calcium and 7.5 mg/100 g iron (Verma and Raghuvanshi, 2002).
To prepare salty snack sprouts, a half cup of lentils are soaked in water
for 812 h, drained and sprouted for 35 days until the length of shoots
equal the seed length, with two or three rinses daily. To these sprouted
grains, onion powder (two tbsp), garlic (one teaspoon), tamarind (two tbsp)
and pepper (a pinch) are added and baked at 120C until brittle and crispy,
which takes approximately 1 h and the tasty snack sprouts are ready.
These snacks can be stored in an airtight container which will keep them
dry and crisp.
Lentil fritters are prepared by blending one cup of sprouted lentils in a
processor and combining with grated cheese (one cup), half a teaspoon of
soy sauce, salt, pepper, half a cup of finely chopped onion, half a cup of
beef stock and one cup of fresh breadcrumbs. The mixture is then made into
balls and put into an oiled frying pan and shallow fried.
Kebabs are small balls (2.5 cm in diameter) made with dal paste that has
been dried over a low heat and these can be used dry or in soup like dumplings. To prepare lentil kebabs, one cup of red lentils (washed and soaked
overnight) are ground in a mortar and the mixture is whipped into a paste.
419
One tablespoon of oil is heated in a heavy pan and the above paste is cooked
over a low heat until the mixture thickens and changes colour. Salt and a
mixture of black pepper, coriander, cloves, cumin seeds, cinnamon sticks and
nutmeg are added and the paste is allowed to cool. This dough is made into
balls of 2.5 cm in diameter and shallow fried until golden brown.
Dosa is a thin, crisp, fried pancake-like staple food of South India, considered mainly a breakfast food but has become popular as a snack food
throughout India. Black gram with rice is generally used as the starting substrate for production of a variety of fermented products like dosa, idli, adai,
etc. Different cookbooks use the ratio of 1:3 to 3:1 of pulse and rice. Black
gram is at times replaced with lentil to make an easily digestible and less
flatulent product. It is prepared by soaking three parts of rice and one part
of black gram or lentil dal in water for 46 h at ambient temperature, grinding to a fine paste by adding 2.02.5 parts (w/w) water and mixing the two
together to make a free running batter. The batter is mixed with 1% salt and
allowed to ferment for 1224 h by natural microflora or by inoculating with
the fermented batter of the previous batch. On a hot and greased griddle
about 6080 ml of the batter is thinly spread for a few minutes where it
assumes a thin, crisp pancake. It is then soaked and served with coconut
chutney (pungent sauce) and sambhar (thin pigeon pea dal cooked with a
heavy dose of spices and vegetables, tamarind and curry leaves).
As per recorded history, idli have been used in India since 1100 ad. It is
a small, white, acid-leavened steamed cake made from fermented thick batter of carefully washed, soaked rice and dal (black gram or lentil). These are
soft and spongy with a mildly sour flavour and eaten with sambhar, coconut
chutney, pickles or dry chutney powder. Normally the proportion of dal
with rice used is 1:3. The amount of water required to prepare the batter to
be of a desirable and uniform consistency varies around 1:5 times on a dry
weight basis of the ingredients. In this case, rice is coarsely ground in comparison to the dosa batter. The coarsely ground batter with 1% salt is fermented for 1830 h with natural microflora. The fermented batter is then
poured with a ladle on to greased round-shaped perforated moulds and
steam cooked for 1020 min to give a fine-textured idli ready to eat. The
soaking of rice and dal brings about the rapid disappearance of common
aerobic contaminants. Leuconostoc mosenteoids and Streptococcus faecalis
develop concomitantly during soaking and continue to multiply after grinding and play a significant role in fermentation. More than a dozen bacteria
and yeast have been identified in the fermented idli and dosa batter. Proteins
are partially converted to amino acids in the fermented products; dosa usually
has 1520% protein, 1525% fat and around 400 kcal energy. Water-soluble
group B vitamins, namely thiamin, riboflavin, niacin and cyanocobalamin,
increase significantly as the batter fermentation progresses (Soni and Arora,
2000).
Sprouted lentil can be used to prepare a variety of food items such as
spread (lentil sprout spread), cutlets (lentil fritters), dal and other snacks. Lentil sprout spread can be prepared by sprouting half a cup each of lentil (for
34 days) and sesame seeds (for 12 days). Then they are put in a processor
420
to which ripe avocado, ground spices and fresh herbs are added and blended
until smooth. This spread is used on bread or savoury biscuits.
Easy to prepare no-meat burgers are useful on their own or in buns.
They require half a cup of sliced almonds, one teaspoon of salt, half a cup of
carrots, one onion, five green chillies, finely chopped cilantro (or other herbs
of preference such as celery or thyme) and two teaspoons of lemon juice. To
prepare the burgers, the lentils are pressure cooked with a little water or
cooked in water until tender; the water is then drained. Meanwhile, one
teaspoon of oil is heated on a griddle and carrot, onions, chillies, cilantro
and almonds are added. They are sauted until the almonds are light brown
and then cooled. The mixture is then transferred to a food processor, and
the cooked lentils and salt added. The mixture is stirred several times,
scraping the sides, until the mixture is coarsely ground. The mixture is then
put in a bowl and lemon juice is added and mixed thoroughly. The lentil
mixture is then formed into round patties. One teaspoon of oil or butter is
then heated on a cast-iron griddle and the patties are added, cooked for 34
min on a medium low heat until light brown. The patties are delicate, so a
large flat spatula is required to turn them. They are served hot between
sliced buns with shredded cabbage, onion, tomato slices along with tomato
ketchup and shredded cheese.
25.7. Processed Lentil

Modern processing technology has allowed lentil to be used in a range of
foods including bakery products, snacks, ready-mixes and ready-to-eat
foods. Lentil being rich in protein is important to vegetarians and those
allergic to dairy products (or on diets specifying reduction of dairy products) as an alternative source of protein. Papad, also known as appalam, is a
popular snack item of India. It is consumed either as such after frying or
roasting or as adjunct along with vegetable soups and curries. Papads are
made from a blend of cereal flour, edible starch and pulse flour with common salt, spices, edible oil, alkaline and mucilaginous additives. The main
varieties of papads are those made from spiced/un-spiced black gram, green
gram, sago and potato. In northern India, papads are mainly prepared from
black gram dal, blends of green gram and black gram dals and starch. Saxena et al. (1989) explored the possibility of preparing papads from blends of
different dal (dehusked split pulses) flours with or without the addition of
black gram dal. Blends of dal flours comprising black gram and chickpea
(70:30), black gram + mung bean + lentil (60:25:15) and black gram with
lentil (80:20) yielded papads with acceptable physio-chemical and organoleptic attributes. The papads prepared from the blend of black grammung
beanlentil (60:25:15) showed better colour, aroma, taste and texture.
Vermicelli is prepared from refined wheat flours, lentil and pea flour
made into a tough dough which is pressed through a vermicelli mould and
dried. Singh et al. (2003) prepared vermicelli fortified with 20% malted lentil
and 25% pea flour. The protein content increased to 1416.7% as compared
421
to the traditional vermicelli (11.5% protein). The author reported that incorporation of 25% malted lentil showed higher nutritional value with respect
to vitamins and minerals.
Lentil was used to develop low-cost food supplements for infant feeding along with other ingredients such as soybean, rice, wheat, chickpea and
semolina and the formulation was found to be a very rich source of protein
with high amounts of essential amino acids (Hegazy et al., 1989).
A canned vegetable-mushroom mix was developed. The main ingredients of the product were grass pea (Lathyrus sativus) and lentil seeds, which
were mixed with white button mushrooms (Agaricus bisporus) and oyster
mushrooms (Pleurotus ostreatus). High temperature ensured the required
seed softness and degradation of the non-nutritional substances. The canned
food thus manufactured exhibited highly positive sensory properties, and
was rated as 4.14.8 points on a five-point scale. The dry matter content of
the canned food ranged between 20.6 and 25.5%, and the average protein,
total carbohydrates, nutrient cellulose and ash contents were 5.8, 2.6, 1.8
and 1.3%, respectively (Kalbarczyk, 2003).
Lentil loaf features a crunchy breadcrumb topping and is served with a
savoury vegetarian gravy, mashed potatoes and peas. To prepare lentil loaf,
two cups of lentils are cooked in boiling water until tender, which may take
about 40 min. This cooked lentil is then combined with six small pieces of
sliced white bread, two eggs, one cup of vegetable broth, two tbsp tomato
paste, half a teaspoon of dried basil, quarter of a teaspoon of garlic powder,
half a teaspoon of ground black pepper, one teaspoon of dried parsley, one
tbsp of olive oil and a packet of dry vegetable soup mix. The mixture is then
spread into a greased and preheated 25 15 cm loaf pan and baked at 205C
for 40 min. The dry breadcrumbs are then sprinkled over the loaf and baking is continued for another 10 min. This lentil bread has high contents of
protein (15.4 g) and dietary fibre (12.2 g) as compared to wholewheat flour
or refined flour bread.
25.8. Medicinal Uses of Lentil

Lentil is also known to have medicinal value with complex carbohydrates
that regulate the bodys metabolic system, help stop diarrhoea and are useful in treatment of constipation and other intestinal afflictions. The seeds
are mucilaginous, rich in dietary fibre and laxative. They are easy to digest
and help in formation of stools. Made into paste, lentils are useful as a
cleansing application in foul and indolent ulcers. In India, lentil paste is
poulticed on to the ulcers that follow smallpox, measles, chicken pox,
rashes, boils and other slow-healing sores.
In the Ayurveda system of medicine, the human system is divided into
three components, that is humour of phlegm (khaff), bile (pitta) and wind
(vayu), and it is believed that a balance of the three makes the person healthy.
Lentil is believed to retain stools, is light for the digestive system and causes
the formation of wind humour (flatulence) in the body. It eliminates the
422
excessive humour of phlegm and bile from the body. It also cures fever.
Roasted lentil seeds are ground to make flour and dissolved in fresh pomegranate juice. This solution stops vomiting caused by the above three
humours in the body.
Lentil is being used in beauty-care products. The water in which lentils
are washed is rich in minerals and it is used as a face wash or hair rinse. Lentil paste is applied to pimples for an early cure and to remove pimple marks.
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26
The Impact of Improvement Research:

the Case of Bangladesh and Ethiopia
Aden A. Aw-Hassan,1 Senait Regassa,2 Q.M. Shafiqul Islam3 and

Ashutosh Sarker1
1International
Center for Agricultural Research in the Dry Areas (ICARDA), Aleppo,

Syria; 2Oxfam America-Horn of Africa Regional Office, Addis Ababa, Ethiopia;
3Bangladesh Agricultural Research Institute, Joydebpur, Bangladesh
26.1. Introduction
Global studies on the impact of lentil research have revealed the advances
of lentil improvement research in developing countries (Aw-Hassan and
Shideed, 2003a, b) and the spillover effect in developed countries such as
Australia (Brennan et al., 2002). The first study covered seven developing
countries (Bangladesh, China, Egypt, Iraq, Jordan, Pakistan and Syria) and
showed that the diffusion of improved lentil cultivars was at its early stage,
but steadily increasing. The authors estimated combined annual gross
research benefits of US$7.7 million for 1997, with Bangladesh, Syria and
Pakistan getting the highest gains in that order because of their large cultivated areas. The estimated overall adoption rate was 17%. The slow down
in improved varietal diffusion to larger areas was attributed to external factors. These factors include: (i) the difficult nature of the production environment; (ii) high climate and soil variability, which limits the likelihood of a
single cultivar to dominate large areas; and (iii) the inability of seed systems
to get high quality seed to small farmers in marginal environments. The
authors suggested that decentralized breeding with the strategy of adapting
germplasm to the specific agroecological conditions of different subregions
as one way to achieve higher varietal diffusion. Increased diffusion of
improved cultivars of lentil also requires innovative ways to involve extension and seed systems in the technology development and transfer process.
The second study examined the global impact of lentil research and
specifically identified anticipated spillover benefits of lentil improvement
research at the International Center for Agricultural Research in the Dry
Areas (ICARDA) in terms of cost reduction for lentil growers in Australia.
Such cost reductions are expected to result from yield increases attributable
to germplasm development at ICARDA or acquired by ICARDAs germplasm
improvement programme and incorporated into genotypes that will be grown
425
426
A.A. Aw-Hassan et al.
in Australia, or from particular resistances in the germplasm (Brennan et al.,

2002). Overall, Australia was estimated to receive an average of A$4.9 (US$2.5)
million annually from ICARDAs research over 20 years. The same study estimated global annual average discounted benefits of A$17.5 (US$9.1) million
from ICARDAs germplasm improvement over 20 years, 20012021.
But these studies mainly relied on secondary data and projected adoption
levels, and lacked field-based adoption data. As a result the analyses were
mostly ex ante providing estimates of potential impacts. These global studies
also do not explain why adoption levels of new cultivars are low in less developed countries. In this chapter we present field-based evidence of adoption
of improved lentil cultivars from two cases: Bangladesh and Ethiopia.
Bangladesh and Ethiopia are low-income countries with high population density and incidence of rural poverty (41% of the population in Bangladesh and 23% in Ethiopia are living below US$1/day), the majority of
whom depend on agriculture for their livelihoods. New crop cultivars and
good management practices are required to increase productivity and
income, and to improve the welfare of many rural households in these countries. Lentil is an important crop for the small-scale farmers of both countries. Bangladesh grows about 148,000 ha of lentils annually and Ethiopia
approximately 82,000 ha. The people of both countries have strong traditions of food legume consumption, which is the main source of dietary protein. But yields are low with averages of 0.81 t/ha in Bangladesh and 0.66
t/ha in Ethiopia over the 20012006 period (FAO, 2008a), hence the potential for increasing productivity and farm incomes is substantial.
The aim of this chapter is to determine the state of adoption of improved
lentil cultivars in these two countries using survey data, analyse the factors
constraining the diffusion of new lentil cultivars and estimate ex post economic impacts based on the adoption that has already taken place. We also
aim to highlight the costs of the time lag in new lentil cultivar adoption.
The chapter is organized in eight sections. The first section provides the
background to lentil research and development, the second section introduces the study areas and the third section describes the household surveys. The fourth section provides estimates of the profitability of new lentil
technologies and the fifth section documents the adoption of new lentil cultivars, which is followed by an analysis of the factors influencing adoption
of new lentil varieties in the sixth section. The economic impacts of lentil
improvement research as well as the methods used in estimation are discussed in the seventh section. Finally a discussion of results, their implications or lessons for research and extension are presented in the eighth section.
26.2. Research and Development

Bangladesh
The Bangladesh Agricultural Research Institute (BARI) has been engaged in
lentil improvement since the 1980s. The development of high-yielding lentil
The Impact of Improvement Research: Case Studies
427
cultivars with a short production cycle was adopted as a breeding strategy

through the introduction of new germplasm from ICARDA. The early
maturing germplasm of ICARDA was supplied through the International
Nursery Network. These included segregating populations specifically created for Bangladesh using its elite landraces and parents of diverse origin
(Sarker et al., 2004). This was a result of changes in ICARDAs international
lentil breeding programme to better serve the needs of the national programmes (Erskine et al., 1998). The change was towards a decentralized
breeding approach where ICARDA develops segregating populations using
parents from national programmes and incorporates improved agronomic
traits. The targeted segregating populations were sent to Bangladesh for
selections to be made under local agroclimatic conditions. The result of that
effort was the release of high-yielding cultivars with high resistance to rust
(Table 26.1).
The research programme simultaneously included capacity building in
the national lentil improvement programme through short- and long-term
training and visits to international research centres. This research and capacity building programme was supported by the International Development
Research Center (IDRC) and the Canadian International Development
Agency (CIDA) of Canada and the Australian Centre for International Agricultural Research (ACIAR). The Food and Agriculture Organization (FAO)
also provided technical support to BARI. The research programme covered
a wide range of agronomic practices including on-farm seed priming, relay
cropping, mixed and intercropping and disease management (Sarker et al.,
2004).
The lentil improvement programme in Bangladesh owes its success to
its strong outreach strategy. The demonstration and dissemination of lentil
technology including new cultivars, agronomic practices and disease management were conducted by extension staff (Afzal et al., 1999). This programme was implemented under a technology transfer programme called
the Lentil, Black gram and Mungbean Development Pilot Project (LBMDP)
launched in the 1996/97 cropping season and funded by the Government of
Bangladesh from its own resources (Sarker et al., 2004). The programme
coordinated a wide range of technology transfer activities including foundation seed production of new lentil cultivars, large-scale demonstration
plots, assessment of varietal performance with farmer participation, free
Table 26.1. The origins, characteristics and yield of new lentil cultivars in Bangladesh
(Source: Sarker et al., 2004).
Cultivar
Uthfala (Barimasur-1)
Barimasur-2
Barimasur-3
Barimasur-4
Year of
release
1991
1993
1996
1996
Origin
Local selection
ICARDA
Local crosses
ICARDA
Maturity
(days)
100-Seed
weight (g)
Yield (t/ha)
110
110
115
116
1.6
1.5
2.5
1.7
1.31.5
1.8
2.0
2.3
428
distribution of seeds, fertilizers and other inputs, farmer training, organization

of mobile workshops for researchers, extension staff and non-governmental
organization (NGO) staff, field days, awards to the best farmers and extension workers, procurement of seed from farmers for next seasons distribution and popularization of technologies through national media, technical
bulletins, leaflets, booklets and posters. This programme was carried out in
collaboration with five major institutions led by BARI. These were the Bangladesh Institute of Nuclear Agriculture (BINA), the Directorate of Agricultural Extension (DAE), Bangladesh Shaik Mujibur Rahman Agricultural
University (BSMRAU), Bangladesh Agricultural Development Cooperation
(BADC) and local NGOs (World Vision, Ghoroni). As a result, from 1997 to
2003 the programme trained about 5740 farmers, over 2000 research and
extension staff and conducted 49 field days with over 5000 farmers participating. It also conducted over 1100 field demonstrations covering 1548 ha
in 12,400 farmers fields, and distributed about 4 t of certified seeds of the
four new Barimasur cultivars (Sarker et al., 2004).
Ethiopia
The Debre Zeit Agricultural Research Center initiated a lentil improvement
programme in the early 1970s, with the focus on increasing yields, development of disease resistance and improvement of management practices. In
recent years, quality aspects, particularly export-quality lentils, were added.
In 1976, the Center took the responsibility of coordinating lentil research at
the national level. Currently, research on lentil is being undertaken at Debre
Zeit, Adet, Holetta, Sinana, Debre Berihan and Sirinka Agricultural Research
Centers (Regassa et al., 2006). Since the 1970s the lentil research programme
has collaborated with ICARDAs Lentil Improvement Program. The national
programme has benefited from the collaboration in human resources development, information exchange and acquisition of new germplasm and
advanced materials for the breeding programme. This collaboration has so
far led to the release of one regional and nine nationwide cultivars with
their full management packages (Table 26.2). Among these cultivars are
Chekol, Chalew, Gudo, Adaa, Alemaya, Alemtena and Teshale.
EL-142 was released through selection of germplasm from Ethiopia.
In addition, the Ethiopian lentil improvement programme progressively
improved non-yield attributes of lentil cultivars. For example, the seed size
has increased from 1.52 g/100 seeds of the local cultivars to 45 g/100
seeds of the new cultivars. Productivity at the research farm has increased
three- to fourfold (i.e. from 0.50.6 to 33.5 t/ha). Besides higher yields and
rust resistance, some of these new cultivars also have other important attributes such as high efficiency of zinc accumulation.
Since the late 1990s, the Debre Zeit Agricultural Research Centers outreach and on-farm technology popularization programme has conducted
on-farm trials and demonstrations of improved lentil production packages
(improved cultivars and management practices) in lentil producing areas
Origin, characteristics and yield of new lentil cultivars and their recommended agroecological areas in Ethiopia.
Adaptation zone
Days to
maturity
Variety
Source
Year of
release
EL-142
R-186
NEL158 (Chalew)
NEL2704 (Chekol)
FLIP 84-78L (Gudo)
Ethiopia
ALAD
ICARDA
ICARDA
ICARDA
1980
1980
1985
1994
1995
85103
122143
111128
8491
86151
FLIP 86-41L (Adaa)

FLIP 89-63L (Alemaya)
ICARDA
ICARDA
1995
1997
86157
81136
FLIP 96-49L (Alemtena) ICARDA

FLIP 96-46L (Teshale)
ICARDA
2003
2003
81115
97129
Seed
colour
Brown
Green
Yellow
Brown
Reddish
brown
Grey
Reddish
yellow
Grey
Grey
Altitude (m above
sea level)
Rainfall
(mm)
Seed yield (kg/ha)

Farmers
elds
Research
trials
Change
(%)
16502000
18002400
18502450
16002200
18502450
400600
5001200
5001200
400600
5001900
912
1416
1418
1416
1418
1420
1725
1926
1522
1825
5667
2156
3644
738
2939
18502450
16502600
5001100
4501200
1624
1824
1926
2030
198
1125
16002400
18002400
400600
400800
1418
1626
1723
1831
2128
1319
Table 26.2.
ALAD, Arid Land Agricultural Development.
429
430
particularly in Ada-Liban, Akaki, Gimbichu and Alam-Gena districts in east

and south-west Shewa zones to promote technology adoption. The technologies were also promoted through the establishment of Farmer Research
Groups (FRGs) in the Gimbichu, Ada-Liban and Alam-Gena districts
who have taken part in evaluation and dissemination of seeds of released
lentil cultivars. Seeds of improved cultivars have been distributed to
FRG members so that these farmers help in further diffusion of seeds to
their neighbours. This mechanism increased farmers access to information
and evaluation of new cultivars. As a result, some farmers visited research
centres and requested new seeds. In addition, the technologies have been
demonstrated to farmers through extension activities of the regional agricultural bureaus.
The on-farm research programme facilitated farmer-based seed multiplication and farmer-to-farmer exchange of new cultivars on a commercial
basis. In this process researchers and the Ethiopian Seed Enterprise (ESE)
experts provided technical support to ensure quality seed is produced by
farmers, which led to the production of a significant amount of improved
seeds. Gimbichu district is now considered as a major source of improved
lentil seeds for growers in other parts of the country. After realizing the
potential of new cultivars and the potential for expanding areas, pioneer
farmers started producing seeds for sale to other farmers, because the formal seed system has no major involvement in lentil seed production. In
2004, one farmer produced 10 t of the cultivar Alemaya and sold it to the
ESE at Birr 4900/t1 and over the 3 years (20032006) more than 100 t of lentil seed was produced and sold either to ESE, agricultural offices or directly
to farmers (Regassa et al., 2006).
Data from field demonstrations in Ada-Liban, Lume and Akaki districts
have shown that the adoption of the improved lentil package (cultivar and
agronomic practices) increased lentil yields by about 60% (DZARC, 1995).
The lentil demonstrations conducted in Ada-Liban, Akaki, Gimbichu,
Enewary, Minjar and Tulu Bolo areas at different farmers fields showed
that the improved lentil package performed better than the traditional practice in most of the cases by about 68% (Fasil and Kiflu, 2003).
26.3. Study Areas

In Bangladesh, lentil is traditionally grown in the north, north-western and
south-western parts of the country. The study was conducted in the lentil
growing areas in the Gangetic floodplain. The districts of Jhinaidah, Chawadanga, Kustia, Meherpur, Magura, Jessore and Norail were included in
the north-western part. In the south-western part, the districts of Rajbari,
Faridpur and Madaripur were covered. In the northern region, which
includes Pabna and Natore, lentil is intensively grown in multiple cropping
systems. These are the traditional areas of lentil cultivation. The Comilla
district, which is a non-traditional area for lentil, was also brought under
lentil as a new crop in the production system.
431
Lentil in Bangladesh is grown as a rainfed crop, but it is occasionally

subject to waterlogging in low-lying areas if there is heavy winter rain. Lentil is usually grown as a sole crop, but also as an intercrop and in mixed
cropping with mustard, linseed, sugarcane and wheat.
In Ethiopia the study was conducted in the four districts of Akaki,
Alam-Gena, Ada-Liban and Gimbichu in the central part of the country,
which constitute the main lentil producing areas (Regassa et al., 2006). Lentil is the third most important crop among pulses after faba bean and chickpea in the area under study. The Ada-Liban district (woreda) is located in
mid-highland with the mean elevation of 1900 m, annual precipitation over
800 mm, and has mainly vertisol (60%) and clay-loam (24%) soils. Lentil is
in the fourth place in cultivated area after faba beans, field peas and chickpea. The Akaki district in the south-east of Addis Ababa is a highly urbanized district situated at an average altitude of about 2000 m and is dominated
by vertisol soils (90%). Lentil is considered as a cash crop because growers
have better access to markets. The Alam-Gena district at about 2100 m altitude is mainly covered with vertisol and alluvial soils. Lentil, grown on the
residual moisture and in the Awash River floodplain when the water
recedes at the end of rainy seasons, is the most important pulse crop. The
Gimbichu district is located at an elevation of 2400 m. More than 50% of
this area is classified as highlands with average annual rainfall of about 900
mm. Most soils (75%) are vertisols. Lentil is one of the important pulse crops
grown in the district.
26.4. Household Surveys

In Bangladesh a random sample of 125 lentil growers who participated in
the demonstration blocks and 125 non-participants from the main lentil
growing region of the country covering 13 pulse-growing districts (Madaripur, Rajbari, Faridpur, Magura, Jhinaidah, Chawadanga, Norail, Jessore,
Kustia, Pabna, Meherpur, Natore and Comilla) was analysed in the 2003/04
cropping season to determine the adoption of new lentil cultivars. They
were typical smallholder producers with average landholding of 1.41.5 ha,
with 2630% illiteracy rate (Table 26.3). The data collected included basic
household asset data (land holding, education, etc.), adoption of new lentil
cultivars, the area planted with new and traditional cultivars, yields, costs
and their perceptions of the performance of the new cultivars. Farmers also
indicated how they spent the additional income, if any, from adopting the
new cultivar.
In Ethiopia, a sample of 289 lentil growers was drawn by multi-stage
random sampling from four main lentil-growing districts: Ada-Liban,
Gimbichu district from East Shewa zone, Alam-Gena district from southwest Shewa, and Akaki district from Addis Ababa administration in the
2002/03 cropping season. Secondary data were also collected from Ministry
of Agriculture, district offices of agriculture, the cooperative union and seed
suppliers.
432

Table 26.3.
Sample farmers land holding and education levels in Bangladesh.
Item
Land holding (ha)
Education (%)
Own land
Rented in land
Rented out land
Mortgaged in land
Mortgaged out land
Total
Illiterate or signature
Primary
Secondary
SSC and above
Adopters
Non-adopters
1.40
0.11
0.04
0.11
0.03
1.68
26
23
25
26
1.34
0.16
0.09
0.05
0.03
1.67
31
16
29
24
SSC, Secondary School Certicate
Table 26.4.
Landing and crop mix of sampled farmers in selected districts, Ethiopia.

Crops (% area)
Study area
Ada-Liban
Akaki
Gimbichu
Alam-Gena
All districts
Mean
Sample holding
size
(ha)
50
47
99
93
289
1.5
3.01
1.7
3.04
2.31
SD
Barley
Tef
Wheat
Faba
bean
0.50
1.22
1.05
1.04
1.23
0
1
1
1
1
28
46
18
50
41
42
22
49
13
26
5
0
4
1
2
Chickpea Lentil
13
17
11
17
15
9
5
14
10
10
Grass
pea
Field
peas
3
8
4
7
6
1
0
1
1
6
The surveyed farmers practise a mixed crop-livestock system with most

households having at least one or more livestock species. The interdependence of crop production and animal husbandry is critical for livelihood in
this mixed farming system. The average family size was about eight persons. Overall 38% of household heads were illiterate. This was higher in
Ada-Liban (52%) and Akaki (43%) districts than in the other two districts,
Alam-Gena (36%) and Gimbichu (19%). The study areas are typical of smallholder farming with overall average holding size of 2.31 ha, with AlamGena and Akaki farmers holding almost double the holding area of the
other two districts (Table 26.4).
As shown in Table 26.4, the main staple traditional food crop (tef) and
wheat were given the highest priority in land allocation (over 40% and 26%,
respectively) ensuring household-level food security. Chickpea was the second most important crop in the study area and lentil the third most important crop in terms of cultivated areas with 10% of the cropped area.
The livestock in the mixed farming system are cattle, sheep, goats, donkeys, horses, mules and poultry. On the average, each family owned three
or more oxen, which are an important power source for tillage. Livestock
ownership was estimated at 7.36 tropical livestock units (TLU) in Ada-Liban,
433
11.4 in Akaki, 8.1 in Gimbichu and 9.6 in Alam-Gena. The differences were
significant.
26.5. Profitability of New Lentil Technology

In Bangladesh, crop enterprise budget analysis shows that the surveyed
growers adopting new lentil cultivars had increased their gross margins of
return on land, labour and management by 120%, receiving Taka 15,576/
ha, compared to Taka 7096/ha by the non-adopters.2 The high financial
benefits are a result of low additional costs typical of new crop cultivar
adoption. These lentil growers reported that they mostly spent the extra
income on clothing (68% farmers), food (62%), cultivation costs of other
crops (62%), education (58%) and medicare (58%) (Table 26.5). They also
spent extra income on rebuilding houses, and purchases of land and farm
equipment.
In the Ethiopian case, the gross margin per hectare for the cultivar Alemaya was estimated at Birr 8238/ha,3 while the gross margin for the local
cultivar was estimated at Birr 2881/ha, increasing lentil profitability by
186%. These estimates are based on data from demonstration farms which
are monitored by extension staff and have applied recommended agronomic practices. Normally increase in farm profitability by new technologies would be lower than these figures as farmers generally do not apply
optimal management practices, thus reducing their yields or increasing
their costs of production, hence reducing their profits.
Although national-level economic impacts are important, these farmlevel financial impacts of new crop cultivars have direct and immediate
impact on the welfare of rural households that grow lentils. These immediate impacts on the livelihoods of rural households and the long-term effect
on human development such as education and health and capital asset
building, and the self-esteem gained and satisfaction from the sense of
Table 26.5. Utilization of additional income accrued from new

lentil cultivars, Bangladesh.
Items of expenditure
Clothes
Food
Cultivation costs
Education
Medical
Loan servicing
Land, cattle or thresher
House improvement
Marriage of daughter
Small business
Frequency
Farmers (%)
86
78
77
73
73
9
14
6
1
1
68.8
62.4
61.6
58.4
58.4
7.2
11.2
4.8
0.8
0.8
434
achievement are, perhaps, the most important impacts of agricultural

research that builds the foundation for sustainable livelihoods in the long
term. The long-term outcomes of these intangible impacts are harder to
assess.
26.6. Adoption of New Cultivars

Bangladesh
The total lentil area and areas of new lentil cultivars and number of growers participating in the demonstration programme and non-participants are
shown in Table 26.6. The number of non-participating farmers growing new
lentil cultivars increased from 0 to 47 during the period 19992002, giving
an adoption rate of 38%. Clearly, these farmers adopted new cultivars after
becoming aware of the new technology from other farmers who had participated in the demonstration programmes. Informal farmer-to-farmer exchange
is the main source of new seeds and information in developing countries,
and is key to the diffusion of crop cultivars (Aw-Hassan et al., 2008). The
area of new cultivars increased from 191 ha in the 1998/99 cropping season
to 10,908 ha in 2002. This is an impressive 24-fold increase.
The adoption of Barimasur cultivars was accelerated by the technology
transfer programme described above. The adoption rate (given as percentage of area cultivated) of modern Barimasur cultivars and local cultivars for
1998/992001/02 is presented in Table 26.7.
The survey data show that the proportion of the area cultivated with
new cultivars grew faster among farmers who participated in the demonstration blocks, increasing from 6% in 1998/99 to 63% in 2001/02 cropping
season, compared to a rise from 0 to 24% for non-participants during the
same period. This result shows the impact of the technology dissemination
Table 26.6. Total area cultivated in Bangladesh with modern lentil cultivars (Barimasur) by
farmers participating in demonstrations and non-participants.
Cultivar
category
Modern
cultivars
Local
cultivars
Demonstration
farmers
Year
1999
2000
2001
2002
1999
2000
2001
2002
Number
4
42
118
106
37
92
68
61
Area (ha)
191
2,484
7,516
8,344
3,132
8,506
5,964
4,846
Non-demonstration
farmers
Number
4
16
47
38
103
109
100
All farmers
Area (ha)
Number
183
855
2,564
3,359
9,055
9,783
8,074
4
46
134
153
75
195
177
161
Area (ha)
191
2,667
8,371
10,908
6,491
17,561
15,747
12,920
Cropping
seasons
1998/1999
1999/2000
2000/2001
2001/2002
1998/1999
1999/2000
2000/2001
2001/2002
1998/1999
1999/2000
2000/2001
2001/2002
a MV,
Lentil cultivars
Farmer group
Demonstration participants
Non-participant farmers
All farmers
Barimasur-4 Barimasur-3 Barimasur-2 Barimasur-1

2
16
36
40
0
2
7
18
1
9
22
29
4
6
15
16
0
0
1
6
2
3
8
11
0
1
2
3
0
0
0
0
0
0.5
1
2
0
0
4
6
0
0
0
0
0
0
2
3
All MVa
6
23
56
63
0
2
8
24
3
13
32
44
Local cultivars
94
77
44
37
100
98
92
76
97
88
68
57
Table 26.7. Area cultivated (%) with lentil cultivars among farmers participating in demonstration blocks and non-participants, in selected
Bangladesh sites, 1998/992001/02.
Modern varieties.
435
436
and extension programme. Clearly, Barimasur-4 which accounted for 67%

of that adoption rate was the most favourite cultivar, followed by Barimasur-3 which claimed a quarter of the adoption rate. By 2002, all the surveyed
farmers cultivated 44% of their lentil area with improved lentil cultivars.
This rapid diffusion rate is a result of the success of the ICARDABARI
collaboration in lentil breeding and developing new cultivars with clear
productivity advantages, disease tolerance and other agronomic traits, and
the effective multi-institutional technology transfer programme coordinated
by BARI. Farmers reasons for preferring the Barimasur cultivars over the
local cultivar are given in Table 26.8. The farmers who experienced these
new cultivars are convinced of their advantages in terms of yield, seed
colour, disease tolerance, grain size and straw yield, in that order. Farmers
also used other indicators given in the table. Other studies have shown that
farmers take several plantings to assess new germplasm in comparison with
their own germplasm before they are convinced of the superiority of the
new material (Aw-Hassan et al., 2008).
Ethiopia
The overall adoption rate of new lentil cultivars among the surveyed farmers was 19%, with higher adoption rates in Ada-Liban and Gimbichu districts (Table 26.9). The most adopted was the cultivar Alemaya. After this
survey was completed, a rapid increase in the area under Alemaya was
reported, particularly in Gimbichu district from 156 ha in 20032004 to 2460
ha in 2005 because of the high profitability of this technology (Regassa et al.,
2006). However, most of this adoption is localized to a small part of the lentil growing areas. In fact, it is estimated that cultivar Alemaya is the dominant cultivar in Gimbichu district covering 24% of the lentil area in 2004.
The success of the Gimbichu district, none the less, was not repeated in
other lentil growing areas and as a result, these estimates corresponded to
only a 2.6% adoption rate nationwide in 2005.
Table 26.8. Bangladeshi farmers reasons for

preferring Barimasur cultivars of lentil.
Traits
High grain yield
Preferred seed colour
Disease resistant
Large grain size
High straw yield
Drought tolerant
Good taste
High price
Short duration
Frequency
Percentage
121
114
114
112
73
32
18
9
6
99
93
93
92
60
26
15
7
5
437
Table 26.9. Adoption rate of improved new lentil cultivars in different districts of Ethiopia, 2003.
District
Ada-Liban
Number of farmers
Adopters (%)
Non-adopters (%)
Total (%)a
aThe
50
30
70
100
Akaki
47
11
89
100
Gimbichu
99
35
65
100
Alam-Gena
93
0
100
100
Total
289
19
83
100
small differences between Total (%) and percentage adoption is due to rounding error.
These low adoption rates are not surprising given that farmers have
been largely unaware of the cultivars and extension services do not reach
the majority of farmers. The extension services were weak as only 43% of
the farmers reported they had contact with extension services at least twice
in every 3 months, about 25% had no contact with extension services at all
and the remaining had contact but over longer intervals. The farmers
involved in demonstrations, on-farm trials and farmer research groups and
training comprise only 6% of the sample. The inadequate extension services
were reflected in the level of farmers knowledge about new lentil cultivars.
Farmers who knew at least one of the improved lentil cultivars were estimated at 59% in Gimbichu, 49% in Ada-Liban, 36% in Akaki and 7% in
Alam Gena. Almost all participants of extension activities were fully aware
of the new cultivars, but the majority of the sample (94%) comprised nonparticipants and only about one-third (33%) of those were aware of the new
cultivars. The most popular cultivars across all districts were Alemaya
(11.4%) and Ada (4.8%). Other cultivars were not known to farmers. Farmers knowledge and evaluation of new cultivars through formal extension
systems or through farmer-to-farmer exchange of seeds and information is
the first critical step in the technology diffusion process. The outreach of
extension services is essential to improve the level of farmer awareness of
new cultivars.
26.7. Factors Influencing Adoption of New Lentil Cultivars

Adoption of new crop cultivars mainly depends on their performance, profitability and feasibility within the farming system. Adoption also depends
on both internal household factors and external factors. Internal factors
include age, gender, education level, exposure to extension and research
activities, and the level of entrepreneurship. Further, adoption of new crop
cultivars is influenced by household-level resources (human, natural and
physical assets) and social assets. These personal characteristics make an
individual more or less conservative to new ideas, shape the persons willingness to take risk and seek information making the person more or less
knowledgeable of new farming technologies. External factors influencing
438
technology adoption are institutions, services and markets that facilitate the
flow of information, seeds and capital to farmers. Effective extension services and markets greatly influence the uptake of new crop cultivars and
impact the well-being of producers and consumers. Analysis of how these
factors influence adoption provides useful information that can strengthen
knowledge transfer programmes and increase the likelihood of technology
adoption, ultimately leading to improved food security, reduced poverty
and improved rural livelihoods. This also increases the economic and social
returns to public investment in research and extension.
In Ethiopia the adoption analysis using the logit model showed that the
gender of the head of the household, access to seeds of improved cultivars,
participation in extension activities such as demonstration, verification trials, training and membership of a farmers research group significantly
affected adoption of improved lentil cultivars (Table 26.10). Livestock ownership was also significant. Besides livestock ownership, all other variables
had the expected direction of relationship with the dependent variable. The
analysis indicated that male farmers had a higher probability of adopting
improved lentil cultivars than female farmers. This is because men have
better access to resources and information. Although education was hypothesized to positively affect adoption, no significant relationship was found
between education and the adoption of improved cultivars. The other variables, which had a significant positive relationship with cultivar adoption
were access to extension services and improved seeds. The households participating in verification and demonstration trials and training courses were
members of the Farmers Research Group (FRG) and they had a higher
probability of adoption. This indicates the crucial role that agricultural
extension plays in promotion and popularization of technologies. The percentage of correct prediction by the model was 85%.
Wider coverage of the extension service, mass popularization of the
new cultivars and seed distribution are needed if the adoption rates are to
be increased from these low levels. The major reasons these farmers do not
use newly introduced cultivars are lack of information about the cultivars
(42%) and source of seed (38%). Lack of information about the cultivars,
seed sources and shortage of seeds were mentioned as causes of non-adoption
by 86% of non-adopters.
Table 26.10. Logit model coefcients of factors affecting the adoption of
improved lentil cultivars in selected sites in Ethiopia.
Factor
Coefcient
Standard error
t-ratioa
Gender of head of household

Education
Access to improved seed
Access to extension services
Livestock ownership
1.56429
0.00077
0.00190
2.37721
0.00100
0.18870
0.00156
0.00063
0.56547
0.00058
8.299**
0.4942
3.0162**
4.2039**
1.7183*
aPercentage
of correct prediction = 85%,; *P<0.1; **P<0.05.
439
26.8. Economic Impacts

The economic impact of the adoption of new crop cultivars can be estimated
with the economic surplus (ES) model (Alston et al., 1995). The model allows
computing annual flows of research benefits and costs. The procedure
requires data on yield advantage of new cultivars, adoption rates, change in
cost of production due to the adoption of new cultivars, producer prices,
and the demand and supply elasticities (price in response to supply and
demand changes). The small open economy formulation of the ES model
can be represented as:
ES = PtQtKt(1 + 0.5Kte); Kt = [g/e - c/(1 + g)].At
where Pt = prices in year t, Qt = production in year t, Kt = the supply shift
down in year t as a proportion of the initial price, e = supply price elasticity,
g = percentage yield increment of the new variety over traditional variety,
c = proportion of production cost increase due to technology adoption, and
At = adoption rate in proportion of areas under improved cultivars. Both
Bangladesh and Ethiopia are considered as small economies since they are
not globally dominant producers of lentils. The small open economy representation implies that international prices will not be affected by increased
supply and domestic prices directly in relation to the international prices. In
that formulation demand elasticities are not needed. The estimates and
assumptions of the parameters in the above model are described next.
Production (Qt) and prices (Pt)

In this study, we used the FAO production data for 19992006 (FAO, 2008a)
and average production of the preceding 5 years for the period 20062013
(working on the basis that future production would be a similar average).
Similarly, the cost, insurance, freight (c.i.f.) prices were taken from the FAO
trade data FAOSTAT (FAO, 2008a) for the period 19992005 and the average prices of the preceding 5 years were used for the period 20052013.
Yields gains (g)

In the Bangladesh case, average yields of new lentil cultivars and local cultivars in the fields of the farmers participating in demonstrations and the nonparticipants show a yield increase of 51% (non-participants) to 92%
(participants) (Table 26.11). Under controlled conditions researchers reported
a yield advantage of 20% for Barimasur-2 and 53% for Barimasur-4. A
higher yield advantage of 77% of Barimasur-4, at 2.3 t/ha, over the improved
local cultivar Uthfala (1.3 t/ha) was also reported (Sarker et al., 1999). An
average yield gain of 61% was estimated from the 5-years yield data
(1998/992001/02) from the survey of farmers participating in extension
demonstrations and non-participants (Table 26.11). In this analysis we used a
440

Table 26.11. Average yields (kg/ha) of modern and local lentil cultivars for
demonstration plots and non-demonstration farmers elds in Bangladesh,
19992002.
Variables
Modern cultivars
Local cultivars
Difference (%)
Participant
farmers
Non-participant
farmers
All farmers
1097
572
92
967
641
51
1050
651
61
conservative estimate of half that yield gain (30%) of new Barimasur cultivars
over the local lentil cultivar. The lower yield increase is assumed to account
for average farmer practices that may not employ all the necessary agronomic
practices and hence may not achieve the expected technical efficiency.
In Ethiopia, average yields of 1.62.4 t/ha were reported for the Alemaya cultivar compared to 0.6 t/ha for the local cultivar (Dadi and Bekele,
2003; also see Table 26.2). This very high yield advantage of over 100% is
because the high rust resistance of Alemaya allows it to be sown early in
the rainy season taking full advantage of the rainfall, while the local cultivar
has to be planted later in the season to escape rust infection but faces moisture stress, reducing yield significantly (Geletu Bejiga, personal communication). However, a yield advantage of 37% was estimated from the 2003/04
survey. We therefore used this conservative yield advantage in order to
account for the non-optimal agronomic practices under farmers conditions.
Production cost changes (c)

Any technological change involves change in production costs. Generally,
improved agricultural technologies reduce production costs, but in some
instances cost of production may increase due to additional inputs or operations needed to achieve full technology potential. In Bangladesh, the total
variable cost per hectare increased by only 6% but average cost per tonne of
output decreased by 13%. So no cost change was assumed for the Bangladesh case. This is a conservative approach and will in effect underestimate
the research benefits. Similarly, in the Ethiopian case, the cost per hectare
increased by 42% which was due to additional weeding cost, as mentioned
earlier. But this amounted only to 6% increase in the cost of production per
unit, which was used in the Ethiopian case.
Adoption rate (A)

National level estimation of the research and extension impacts requires
national level adoption data over time. Such data are seldom available. In
Bangladesh, cultivar adoption rates were estimated from the 2003 farm
441
survey. Since those estimates do not represent national level adoption rates
we used four adoption scenarios (Table 26.12). In the first scenario, we used
national level adoption estimates of 9% for the year 2003/04 and increased
it to 10% annually until 2013. The 9% nationwide adoption rate is based
on the estimates of agricultural offices for selected lentil growing areas. In
the second scenario, we used the adoption rate of 25% which is close to the
actual estimates of the 2003 survey for the non-participant farmers of the
extension demonstration programme (24%) and projected forward until it
reaches a 30% adoption ceiling. This scenario assumes a reasonably wellfunctioning extension programme facilitating information and material flow
to farmers. This estimate is lower than the 60,000 ha (38%) estimated to be
under new lentil cultivars in 2004 (Sarker et al., 2004). The third scenario is
the same as the second except that a lag of 5 years in reaching the 25% adoption rate was introduced. In the fourth scenario we used the adoption rate
of 45% which is close to the estimates from the 2003 survey for the whole
sample (pooled average of the participants and non-participants 44%) and
projected up to an adoption ceiling of 60%. This scenario assumes excellent
extension and seed systems similar to the one implemented by BARI and
partners, and explained previously in this chapter, which provide effective
services to small-scale farmers. The adoption ceilings of less than 100%
implies that the current cultivars may not fit in all the conditions of the lentil growing regions, or factors such as lack of seed and access to markets
may limit the potential adoption of these cultivars. Again, this is a conservative approach. The four scenarios represent different intensity and effectiveness of technology transfer which results in different diffusion rates over
time, and time lag in actual and potential technology adoption (ceiling)
Table 26.12. Gross annual research benets for different scenarios of farmers access to
seeds and extension services (average over 19992013), Bangladesh.
Scenario description
1. National level adoption
estimates based on secondary
data
2. Assumes modest nationwide
extension programme similar
to one BARI and partners
implemented
3. Same as scenario 2 except the
25% adoption has a 5-year lag
4. Assumes effective nationwide
extension programme
Adoption Adoption
Year
(%)
ceiling
ceiling
2003/04
(%)
reached
Annual gross research

benets (million US$)
under three supply
elasticity scenarios
e = 0.5 e = 1.0 e = 1.5
21
2013
5.6
2.8
1.9
25
30
2007
11.5
5.7
3.8
30
2010
7.3
3.7
2.4
45
60
2008
21.9
10.9
7.3
442
because of the manner in which the extension and seed systems reach out to
small-scale farmers.
In Ethiopia, as mentioned earlier, nationwide adoption estimates are
not available. The adoption data available are localized to small parts of the
country, making it difficult to extrapolate over the whole lentil growing
area. Therefore, we used two scenarios; a low adoption rate scenario which
is 3% of the lentil areas, and the second one which is higher with a modest
rate of 25% to compute the forgone economic impacts or losses due to slow
technology diffusion (Table 26.13). We limited the Ethiopian case to these
two scenarios because of the limited adoption data. These two scenarios
also represent different levels of extension activities and farmers access to
information and seeds.
Supply elasticities
Price supply elasticities for most agricultural commodities are expected to
be greater in the medium- and long-term (Alston et al., 1995). To see effects
of elasticity assumptions on the estimates of gross annual research benefits
we considered three supply elasticity levels: 0.5, 1.0 and 1.5. These supply
elasticities are within the range of generally reported commodity supply
elasticities.
Estimates of economic impacts

In the absence of accurate estimates of research costs we computed only
gross research benefits. Zero increase in cost of production was assumed
throughout the analysis because the new lentil cultivars do not require
additional inputs to achieve the yield advantage used in the analysis. A 5%
discount rate was used to compute the present value of research benefits
and the estimates were deflated to maintain 2008 constant US dollars for all
periods using the FAO price index (FAO, 2008b). In the Bangladeshi case,
with low price supply elasticity scenario (e = 0.5), the gross annual economic
impact of new (Barimasur) lentil cultivars was estimated at around US$5.6
million. This amounts to about US$84 million for the 15-year period of
19992013. In the second scenario assuming a relatively well-functioning
nationwide extension programme with higher adoption rates of 25% (2004),
the economic impact is estimated at about US$11.5 million annually. In the
third scenario where the adoption was lagged for 5 years the impact of
research benefits were estimated at US$7.3 million annually; and in the
fourth scenario with a more optimistic nationwide adoption rate of 45%,
which represents a situation with a highly effective nationwide extension
programme, the economic impact of lentil technology is estimated at
US$21.9 million annually. These results show the economic impacts of lentil
technology and how those impacts can be affected by the effectiveness of
technology transfer programmes.
443
In the Ethiopian case, under the low price supply elasticities (e = 0.5),
the gross annual research benefits for 2005 was estimated at US$0.33 million when the low adoption rate of 3% was used and these annual benefits
increased to US$2.29 million annually with the higher adoption of 25%. This
is a potential annual flow of research benefits (gross annual research benefits) since adoption still remained lower at national level. The results show
an unrealized economic benefit of approximately US$2.0 million annually
from research that has already generated profitable technologies which has
been verified by extension work with farmers, but is not spreading rapidly
enough.
As shown in Tables 26.12 and 26.13, the estimates of the research benefits were sensitive to the price supply elasticities. These estimates, however,
provide useful information for research administrators and policy makers
to consider in allocating resources for research and extension.
26.9. Discussion and Lessons

We have presented two cases (Bangladesh and Ethiopia) where poor smallscale farmers in developing countries have benefited from improved lentil
cultivars using farm survey data. We selected these two cases among the
poorest countries to highlight the importance of international and national
public research for small-scale farmers in the less developed countries. Bangladesh and Ethiopia are both developing countries with a high concentration of small farmers and rural poverty. In both cases the new lentil cultivars,
originating from ICARDA-developed germplasm, have proved to be superior to the local germplasm under farmers conditions in yield, disease resistance and other traits such as plant structure, seed size and colour. In both
cases the technology has significantly increased the profitability of lentil
production. The gross margins have consistently increased over 100% when
the new package (modern cultivars and agronomic practices) was used.
These results are based on demonstration plots and surveys of a sample of
adopters and non-adopters of technology.
Table 26.13. Gross research benets from lentil improvement in year 2005 for two adoption
scenarios, Ethiopia.
Scenario
Adoption estimates from 2003/04 survey and
secondary sources
Higher projected adoption assuming greater
farmers access to information and seeds
through more effective extension
Adoption
area (%)
Gross annual research benet

for year 2005 for three supply
elasticity scenarios
e = 0.5
e = 1.0
e = 1.5
0.33
0.15
0.09
25
2.29
1.07
0.66
444
Similarly, in both cases the research programmes had clear outreach

activities that initiate popularization of new lentil technology, albeit at different intensities and with varying performance. The Bangladeshi case had
a more systematic outreach programme, which involved a number of institutions that participated in technology transfer, farmer workshops, demonstrations, farmer training, and seed production and distribution. There was
much greater national commitment in technology promotion through the
formal programme of the LBMDP launched in 1996/97. The programme
was successful in increasing farmers awareness of technology and increased
the area under new lentil cultivars by participants, non-participants and all
farmers to 66%, 24% and 44%, respectively, in 4 years. The Ethiopian case
had also an outreach programme, which was carried out by the Debre Zeit
Agricultural Research Centers Outreach and On-Farm Technology Popularization Program, involved FRGs and the ESE in several districts. The
result was a significant increase in farmers awareness and adoption of the
new technologies among farmers who were able to access information about
the technology and seed through this programme. In this case, farmers who
realized the benefits of the technology actively sought the new seed, and a
local seed market of the cultivar Alemaya started developing. However,
the adoption rate of new lentil cultivars among the surveyed farmers
remained low (19%) in 2004. None the less, a high adoption rate (24% of the
lentil area) was reported but remained localized in small areas such as Gimbichu district.
A number of important lessons can be learnt from these two cases. First,
the outreach programmes were effective in increasing farmers awareness
and adoption rates of technology. Without such efforts technology adoption
will not occur. The importance of extension work was supported by the
analysis of the constraints in technology adoption discussed previously in
this chapter. It was demonstrated, using logit model analysis, that both
access to information about the technology and seeds had significant effects
on technology adoption. They were more important than farmers personal
characteristics such as education, age and gender in influencing technology
adoption.
Second, the coverage of these farmers by participatory extension and
research programmes was limited and, as a result, although adoption was
high at the local level, it was still low at the national level. In both cases
modest adoption rates of 24% (in area coverage) were estimated for the
areas where the farmer outreach programmes were implemented, excluding the farmers who participated in the demonstrations. The coverage was
much wider in Bangladesh where the programme had better resources. The
strength of the Bangladeshi farmer participatory extension programme had
almost doubled the adoption of the new cultivars (44%) taking the participants and non-participants together compared to the localized adoption
rate in the Ethiopian example.
Third, the economic impacts generated from these technologies both at
the farm level and at the national level can be substantial, given the high
acceptance by farmers of these technologies in both countries to the tune of
445
about US$11.5 million for Bangladesh and nearly US$2.3 million for Ethiopia annually using modest adoption rates of 25% in the both countries.
However, there is a long time lag in lentil technology adoption. The three
most popular cultivars, Barimasur-4 and Barimasur-3 in Bangladesh
(1996) and Alemaya in Ethiopia (1997), were released over 10 years ago.
This time lag in adoption is largely due to the lack of an effective farmer
participatory extension programme that can out-scale the local success of
technology adoption. National institutions already have the capacity to
implement such programmes as demonstrated in the two cases presented
here. The policy commitment to support and implement such programmes
as part of agricultural research and development is lacking. These policy
and institutional failures have significant costs. The cost of the lag in lentil
technology adoption could reach up to US$2.0 million annually in Ethiopia
even using modest adoption targets of 25%, and in Bangladesh the adoption lag of 5 years at the same modest adoption rate could cost just over
US$4.0 million annually (at the lower supply elasticity). Policy makers
should consider these economic costs while supporting farmer-oriented
extension programmes.
Finally, the results presented above show that the ICARDA lentil
improvement programme had yielded superior germplasm for Bangladesh
and Ethiopia; the National Agricultural Research Systems (NARS) of the
two countries have tested and adapted these germplasm to local conditions,
leading to the release of new lentil cultivars in both countries. The full
impact of this technology, however, requires vigorous farmer participatory
extension programmes that can out-scale the adoption of such technologies
without delay. Moreover, adequate monitoring of technology adoption is
required both for evaluating the overall impacts of agricultural research
and extension more accurately and for timely assessment of the effectiveness of extension programmes. The current global food price crisis is a clear
warning that without adequate investment in such programmes that are so
vital to modernize small-scale farming and to bridge the persistent yield
gaps, the growth in food supply may not keep pace with demand, which
could have serious social, economic and humanitarian consequences.
Notes
1. US$1 = 8.5 Birr in 2004, 9.09 Birr in 2008.
2. US$1 = 68 Taka (May 2008).
3. US$1 = 9.09 Ethiopian Birr.
References
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Gazipur 1701, Bangladesh, 64 pp.
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impact of international and national investment in barley germplasm improvement
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Association of Agricultural Research Institutions, 38 pp.
Index
Page numbers in bold type refer to figures and tables.

Acidity, soil 201202, 232234
Acyrthosiphon pisum (pea aphid) 284,
290, 294, 314
Adaptations, for crop environments
4445, 4849, 5859
Adoption rates, cultivar 105, 425,
434438, 440442
Agrotis ipsilon (cutworm) 282, 284,
295
Air screening 394395
Alemaya 430, 433, 436, 440
Alkalinity 54, 201, 203204
Allergens 383
Allozyme 91
Alternaria alternate (Alternaria bight)
277
Altica coerulea 287
Amino acids
composition, in lentil protein 370,
370
essential, human 370371, 372, 408,
409
Alfalfa mosaic virus (AMV) 307, 308, 314,
315, 316, 318
Anatomy, lentil plant 4344
Anthracnose disease 91, 268269
Anti-nutrients 379, 380, 381, 382383,
409
Antioxidants 380381, 405
Aphanomyces euteiches (Aphanomyces

root rot) 277
Aphids
infestation 288, 290
insecticides 294, 297, 298, 319
natural enemies 294
pest species 283, 284
plant resistance to 292, 293
virus vectors 288289, 314, 315
Aphis craccivora (cowpea aphid) 284,
288, 292, 294, 297
Apion sp. 283, 286, 288, 289
Archaeology, lentil distribution data 25,
27, 28, 2930
Artichoke latent virus (ArLV) 307
Autographa gamma 282, 285
Ascochyta lentis (blight pathogen) 157,
263264, 352
Aulacophora foveicollis 287
Backcross breeding 146

Bacterial diseases of lentil 277
Bagging, crop 259, 397
Balclutha sp. 284
Bangladesh
cropping systems 430431
cultivars 427, 434, 434436, 435, 436
farmers 431, 432, 433
447
448
Index
Bangladesh Agricultural Research
Institute (BARI) 124, 271,
426428, 436
Barimasur cultivars 354, 427, 434436,
436, 439440
Bean common mosaic virus (BCMV) 307,
308
Bean leafroll virus (BLRV) 307, 308, 309,
314, 315
Bean pod mottle virus (BPMV) 307
Bean yellow mosaic virus (BYMV) 289,
307, 308, 310311
Beet western yellows virus (BWYV) 306,
307, 308, 309, 315
Beetles (Coleoptera) 286287, 311, 315
see also Bruchids; Sitona crinitus
(pest weevil)
Bemisia sp. 284
Biological control
pests 294296, 296, 299
weeds 331
Blight diseases
Ascochyta 9091, 263264, 352
Stemphylium 91, 269271
Boiling 411
Boron 53, 110111, 198, 204
Botrytis spp. (grey mould pathogen) 56,
266267
Branching pattern (of stems) 3940, 40,
4445
Breeding programmes
aims 104, 107, 137138
disease resistance 8991, 111, 125,
148, 318319
environmental tolerances 5859,
104, 109111, 149
food value enhancement 112, 113,
150151
high seed vigour 353
mechanization, adapting to
111112, 148
parasitic weed resistance 347348
pest resistance 291294, 298299
short season crops 125126
future prospects 116117, 131132,
147
genebank use 7072, 104, 115116,
139
methods 106, 108, 126128, 131,
142147
national 112, 114, 124, 151, 427
see also Improvement programmes,

lentil
Broad bean mottle virus (BBMV) 307, 308,
314, 315
Broad bean stain virus (BBSV) 289, 307,
308, 311, 315 316, 318
Broad bean true mosaic virus (BBTMV)
307, 315
Broad bean wilt virus-1 (BBWV-1) 307,
308, 314, 315
Broomrapes (Orobanche spp.) 344345,
346348, 355
Bruchids 282, 283, 286, 288, 292, 401
Bruchus lentis (seed weevil) 282,
283, 286, 295, 297
Bruchus ervi (seed weevil) 283, 286
Callosobruchus spp. 282, 283, 286
Brumoides suturalis 287
Bulk population breeding 106, 108, 128,
143145
Calcium oxalate 44, 377

Camnula sp. 284
Canning 402, 421
Carbohydrates 371, 373, 373374
Cercospora lensii (Cercospora leaf spot)
277
Chalky spot syndrome 283, 288, 401
Chickpea chlorotic dwarf virus (CpCDV)
307, 309310, 315
Chickpea chlorotic stunt virus (CpCSV)
307, 308, 314, 320
Chickpea filiform virus (CpFV) 307
Chino del tomate virus (CdTV) 307
Chromatomyia horticola 282, 285
Chromosomes, structure 23, 9293,
166
Cladosporium herbarum (leaf yellowing)
277
Cletus signatus 284
Climate
effect on seed quality 353
for successful lentil growth 4748,
216217, 219220
variability 49, 50, 182
Collections, lentil 6468, 124125, 139,
140
composite 6768, 70, 159
Colletotrichum truncatum (anthracnose
pathogen) 91, 268269
Index
449
Colours
flower 81
foliage 4041, 7778
seed 9, 43, 8284, 8687, 393
sorting machinery 396, 401402
Combine harvesting 249, 258259, 392
Conservation reserves 68
Consumption of lentils 1012, 11
current trends 149150, 404406
traditional preferences 121, 139,
391, 396
see also Marketing
Contarinia spp. 286
Cooking time 392, 402, 409, 416
Core collections see Collections, lentil
Cotyledons 43, 44, 83, 8687
Cover crops 241
Cropping systems 213216, 430431
double cropping 122, 215
integrated management strategies
176180, 207, 298
intercropping 45, 122, 215216,
240241, 291
mixed cropping 215
monocropping 214
relay cropping 122, 216, 218, 220,
221
see also Rotation, crop; Summer
crops; Winter crops
Cross breeding
as source of cultivars 36, 106, 115,
115116
techniques 141142
see also Recombination breeding
Cucumber mosaic virus (CMV) 289, 307,
308, 311312
Cultivars
adoption rate 105, 425, 434438,
440442
disease-resistant (named) 8991,
269, 271, 317318
herbicide-resistant 57, 146, 158159,
338, 347
micronutrient-dense 112, 113,
150151
sources 4849, 106, 115, 115116,
128
types 138139
variation in yield 217, 439440, 440
see also Alemaya; Barimasur
cultivars; Precoz
Cuscuta spp. (dodder) 343, 346, 351

Cutter bars 255257, 256, 257
Cutworms 282, 295
Cylindrosporium sp. (Cylindrosporium
leaf spot) 277
Cyst nematodes 275276
Dal 2, 102, 415416, 419

Debre Zeit Agricultural Research Centre
(DZARC) 428430, 444
Decortication see Dehulling
Deficiencies, soil mineral 53, 5455, 198,
201204, 234
Dehiscence, pod 35, 82
Dehulling 150, 383, 397401, 405
Desiccation, chemical 57, 337, 393, 401
Destoning machines 395396
Dhal mills 398399, 405
Dietary allowances, recommended
(RDAs)
amino acids 371, 372
minerals 376, 377
vitamins 377379, 378
Dietary fibre see Fibre, dietary
Digestibility 381, 409, 411, 412
Diseases of lentil
control methods 55, 56, 148,
317320
effect on N-fixation 236
foliar 263272, 277, 306314
locally important 277, 314, 320
resistance 8991, 111, 157, 166167
seed-borne 263264, 272273, 316,
316
soil-borne 272276
Distribution, geographical
in archaeological remains 25, 27,
2930
of L. culinaris varieties 26, 2831
lentil pests 283, 284287
of wild Lens spp. 22, 2829, 68
Ditylenchus dipsaci (stem nematode)
275276
Diversity, genetic 36, 67, 68, 159
Dodders (Cuscuta spp.) 343, 346, 351
Domestication of lentils 27, 2831, 86,
103
Dormancy, seed 43, 86, 333, 335
Dosas 413, 419
Double cropping 122, 215
450
Index
Drought tolerance 88, 110, 180186
dehydration resistance 186
escape, by rapid growth 5051,
181182
morphological mechanisms 36,
182183
physiological mechanisms 183186
Dry-heat cooking 413
Dry matter accumulation 39, 172173,
174175
Drying, of harvested seed 224, 259, 394
Economic surplus (ES) model 439442

Elasticities, supply 442
ELISA (enzyme-linked immunosorbent
assay) 316317
Emergence, crop 251, 352, 354
weed competition 326328, 327,
328, 335
Empoasca sp. 284
Energy value 369, 374
Enzyme inhibitors 293, 382383
Epilachna sp. 287
Erysiphe spp. (powdery mildew
pathogens) 271272
Ethiopia
cropping conditions 431
cultivars 429, 436438, 437, 438,
444445
farmers 432, 432433
lentil improvement
programme 428430
Ethiopian Seed Enterprise (ESE) 430,
444
Etiella zinckenella (pod-borer) 285, 290,
292, 295, 298
Eusarcocoris ventralis 284
Evapotranspiration (ET) 176177, 214
Evolution 16, 3031
Exports of lentil 9, 9, 138139, 150
delivery systems 397
production systems for 356357
Expressed sequence tag (EST) markers
157, 160, 162
Faba bean necrotic yellows virus (FBNYV)

307, 308, 310
FAO (Food and Agriculture
Organization) 4, 427
Farmers
access to seed services 438, 438, 441
income 426, 433434
land holdings 356, 357, 431432,
432
Research Groups (FRGs) 430, 438,
444
Fasciation 80
Fatty acids 374, 375
Fermentation 412, 419
Fertilizers
application methods 205
cost/benefits analysis 55, 206207,
221
and nitrogen fixation 195196, 229,
234
recommended types 196, 197, 200
Fibre, dietary 374, 379380, 403, 421
Fields, preparation 217218, 249251
Flatulence 371, 373, 374, 409
Flours, lentil 402, 420421
Flowers 4142, 8182, 141, 173
flowering time 4950, 123124,
147148, 181
genetic control 8081, 108109
Football lentils 150, 398, 401
Frankliniella spp. 287
Frost damage 48, 49, 52, 88, 396
Frying 412413
Fusarium solani (Black root rot) 277
Fungal pathogens 263275, 277
Fungicides 264, 267, 269, 273
Fusarium oxysporum (wilt pathogen)
272273
Genebanks
collection sources 64, 6768
locations 64, 66, 67, 140
management 6667, 6869, 318
trait evaluation 6970
uses of germplasm 7072, 104,
115116, 139
Genome sequencing 93, 157, 160167
Germination 35, 43, 86, 352
advanced by seed priming 221
see also Sprouting
Globe mutant 80
Graptostethus servus 284
Grasshoppers (Orthoptera) 283, 284,
294295
Index
451
Gravity separation 395
Green manures 241242
Greges 25, 26, 28
Grey mould disease 56, 266267
Growth habit 34, 35, 3940, 40, 77
adaptation to cropping conditions
4445
and water deficit 127, 172, 184, 257
and weed competition 57, 334
Gypsum application 202, 203
Harvest
area, world 5, 78
costs 111112
handling damage 394
index 175, 176, 180
interference by weeds 327
machinery 255257, 257, 258
importance of cleaning 332
methods 57, 223, 249, 253260, 254
timing of 353, 401
Height, plant 3839, 79, 148
Helicoverpa armigera (pod-borer) 285,
290, 291, 295296, 297
Helminthosporium sp. (Helminthosporium
leaf spot) 277
Herbicides
pre-harvest application 337, 393,
401
resistance in weeds 332333, 337,
338
resistant lentil cultivars 57, 146,
158159, 338, 347
weed control 57, 250, 336337,
346347
Heterodera ciceri (cyst nematode)
275276
History, lentil cultivation 1, 1314,
2930, 47, 103
Hoeing, hand 253, 334335
Hybridization barriers 2124, 294
Hypera postica 287, 293
ICARDA (International Centre for

Agricultural Research in Dry
Areas)
collaborative links 114, 115116,
151
collection sources, lentil 6466, 65
genetic diversity analysis 159

lentil improvement programme
103104, 105106, 445
supply of genetic resources 67, 112,
149, 425426, 427
wild Lens spp. collected 6466, 66
Immunoassays, for virus detection
316317
Imports of lentil 10, 1011
Improvement programmes, lentil
developed countries 142
Ethiopia 428430
ICARDA 103104, 105106, 445
South Asia 108109, 124, 131,
427428
Indent cleaning 395
Infrared drying 402
Inoculants, rhizobial 205, 206, 235,
236239, 355
Insect pests see Pests, insect
Insecticides 294, 296298, 299, 319
Intercropping 45, 122, 215216, 240241,
291
International Centre for Agricultural
Research in Dry Areas see
ICARDA
Iron
content, in lentil cultivars 112, 113,
150151
deficiency, soil 5455, 198, 203204
Irrigation
effect on crops 45, 51, 179, 222223
as weed source 332
Karyotypes, Lens spp. 21, 23, 9293

Khichri 103, 416417
Labour costs 56, 57, 248249, 255

Landraces 30, 4950
preservation 6768, 70
as source of cultivars 115, 115, 128
Laphygma exiqua 285
Laspreyresia (pod borer) 285
Leaf miners 283, 286, 291, 295, 297
Leafhoppers 308, 310, 315
Leaflets 41, 78
Leaves 4041, 44, 7778, 182
diseases of 263272, 277, 306314
pest damage 283, 288, 295
452
Index
Lectins 293, 383
Lens culinaris, regional varieties (greges)
25, 26, 28
Lentil, Black Gram and Mungbean
Development Pilot Project
(LBMDP) 427428, 444
Lentil yellows disease 289, 309
Leveillula taurica (powdery mildew
pathogen) 271272
Liming, effects on soil 201202, 202
Limiting factors, to lentil growth 2, 104,
122123
Linkage mapping 132, 151, 162,
162163, 164
disease resistance 90, 91, 145
see also Markers, genetic
Liriomyza sp. (see Leaf miners) 286, 295
Livestock, uses of lentil for 13, 138, 397
Lodging 57, 148, 248
Lygus bugs 283, 284, 288, 290, 297, 401
Macrophomina phaseolina (dry root rot)

277
Macrosiphum creelii 284
Malnutrition 1112, 377
Manures
farmyard 200, 273
green 241242
weed seed elimination 331332
Markers, genetic
molecular 88, 159163, 165, 299
expressed sequence tag (EST) 157,
160, 162
SSR (microsatellite) 70, 132,
159160, 161
morphological 163164
in taxonomy 1821
use in selection 165166, 167
Marketing
in developed countries 356,
404406
enhanced cultivars 112
farmer involvement 360, 363364
lentil type classes 138139, 351
see also Consumption of lentils
Masur (lentil) 415, 416, 418
Maturity, time to 34, 4849, 116,
181182
Mechanization
field preparation 249251
harvesting 57, 111112, 249,

255260, 257
processing 398400
seed cleaning 351, 392, 394396
seeding 251252
weed control 252253, 253, 334336
worldwide extent 248, 356, 357, 359
Medicine, uses of lentil in 13, 421422
Mediterranean environments 50, 52,
176177
Melanoplus sp. 283, 284
Meloidogyne spp. (root-knot nematodes)
275276
Microarrays, in gene expression study
156157, 157159
Microbes, soil 236, 241, 273
Micronutrients, soil 197198, 199, 234
Micropropagation 158
Microsatellite markers see Simple
sequence repeat (SSR) markers
Mildew, powdery 91, 271272
Milling effiency 398, 401
Minerals, essential (in diet) 374, 376,
376377, 377
Mixed cropping 215
Mjeddarah (mujaddarah) 2, 103, 417
Moisture content, seed 353, 354,
393394, 400
Molybdenum 198, 201
Monocropping 214
Morphological characteristics of Lens
spp. 1516, 17, 3435
Mujaddarah (mjeddarah) 2, 103, 417
Mutations, induced 7677, 80
for breeding 108, 131, 145146, 294,
347
Mycorrhizas, arbuscular (AM) 204, 206,
236
Myzus persicae 284, 289, 298
Nematode pathogens 275276, 277

Nezara viridula 284
Nitrogen
fixation 175, 184, 229234, 231, 242
requirements 195196, 199
see also Rhizobia, symbiosis with
lentil
No-till systems 218, 239240, 249, 250,
336
Nodules, root 37, 38, 175, 235236, 239
Index
453
Non-governmental organizations
(NGOs) 363, 428
Nucleic acid-based assays, for virus
detection 317
Nutritional value of lentils (human) 10,
12, 369, 404405
after food preparation 409, 410413
carbohydrates 371, 373, 373374
dietary fibre 374, 379380, 403, 421
energy 369, 374
fats (lipids) 374, 375
minerals 112, 374, 376, 376377, 377
proteins 8788, 369371, 370, 372,
408
vitamins 377379, 378, 412
Oligosaccharides 371, 373, 374, 379

Ophiomyia phaseoli 286
Origins, of cultivated lentil 2531
Orobanche spp. (broomrapes) 344345,
346348, 355
Osmotic adjustment (OA) 184185, 185
Outreach programmes 427428, 436,
437, 444445
Packaging 397, 402

Parasites, lentil
bacteria 277
fungi 263275, 277
nematodes 275276, 277
viruses 306314, 307, 308
weed plants 253, 329, 333, 343348,
351
Parasitoids 294296, 296
Partitioning, of assimilate to seeds 44,
184
PCR (polymerase chain reaction) 317
Pea aphid (Acyrthosiphon pisum) 290,
294, 314
Pea enation mosaic virus-1 (PEMV-1) 306,
307, 308, 312313, 314, 315
Pea seed-borne mosaic virus (PSbMV) 91,
289, 306, 307, 308, 313, 315
Pea streak virus (PeSV) 306, 307, 308,
313314, 315
Peanut stunt virus (PSV) 307
Pedigree selection 145
Peronospora lentis (downy mildew) 277
Pesticides see Insecticides
Pests, insect
biological control 294296, 296, 299
chemical control 294, 296298, 299,
319
host resistance 291294, 298299
monitoring methods 290
species 282288, 284287
as virus vectors 288289, 307, 314,
315, 319
Phenolics 380, 380381, 383
Phoma medicaginis (Phoma leaf spot) 277
Phosphates
deficiency, and nitrogen fixation
199, 234
fertilizers 195, 196197, 204205
Photoperiod, flowering response 4950,
123, 181
Photosynthesis, rate of 183184
Phyllotreata chotanica 287
Phytate 382, 409
Phyto-oestrogens 381382
Piezodorus rubsofasciatus 284
Pod borers 283, 284, 288, 291, 298
see also Etiella zinckenella; Helicoverpa
armigera
Pods 36, 42, 44, 7778, 82
dehiscence 35, 82
effect of water availability 173, 173
Polishing 401, 416
Pollination 4142, 141
Polysaccharides 373374, 409
Potassium 197, 203, 206
Powdery mildew disease 91, 271272
Pratylenchus spp. (root-lesion
nematodes) 275276
Precoz 3940, 49, 89, 108109, 131
Priming, seed 221, 354
Processing, lentil
by-products 397, 405, 409
for consumption 383, 391, 420421
post-harvest 224, 392
primary 391, 392397, 393
secondary 391, 397403, 400
tertiary 391, 403
Production of lentil
costs 205207, 248, 260, 359, 440
geographical distribution 58, 6,
78, 350, 355356
global amounts 4, 5, 103, 121122,
403
454
Index
Production of lentil continued
integrated management practices
223, 263264, 265, 267, 319320
see also Cropping systems
Productivity see Yield, lentil crop
Profitability, lentil crop 59, 356357,
398, 403, 433434
Proteins 8788, 369371
see also Amino acids
Pseudomonas radiciperda (Bacterial root
rot) 277
Pubescence 16, 38, 41, 79
Pulses, global production 4, 5, 9, 404
Pure line selection 106, 128, 142
Pythium ultitmum (Pythium root and
seedling rot) 277
Quail pea mosaic virus (QPMV) 307

Quality control, seed 350354, 355
Quantitative trait loci (QTL) 70, 132,
162, 164, 167
Rabi crops see Winter crops

Rainfall 5051, 176177
Raised bed planting 220221, 222
Recipes, lentil 413420
Recombination breeding 128, 131
Red clover vein mosaic virus (RCVMV)
307
Relay cropping 122, 216, 218, 220, 221
Research
benefits 425426, 439, 442443, 443
funding sources 152, 356
Residues, crop 200201, 240
Rhizobia, symbiosis with lentil 55, 229,
233, 235239
Rhizoctonia solani (wet root rot) 277
Rice, cropping patterns with lentil 214,
215, 216, 218
Rolling 251252
Root-knot nematodes 275276
Root-lesion nematodes 275276
Roots
anatomy 43
diseases of 272273, 275276, 277
growth patterns 3637, 37, 182183
induction 158
nodulation 37, 38, 175, 235236,
239
parasitic weed infection 344345,

347
Rotation, crop
disease control 269, 276
soil nitrogen benefit 241242
and weed growth 333, 334
Rotylenchulus reniformis (reniform
nematode) 277
Row spacing, lentil crop
and mechnical weed control
334335
optimum 215216, 221
Rust disease 89, 264266, 440
Salad, lentil 403, 414

Salinity, soil 5354, 110, 202203, 232
Saponins 382
Sclerotinia sclerotiorum (stem rot
pathogen) 273275
Sclerotium rolfsii (collar rot) 277
Screening, field (of genotypes) 90, 111,
126, 144, 147
Seedbed preparation 217218, 239,
249251
Seedcoat
chemical composition 8485, 380
colour 9, 43, 8284
compared to cotyledons 408
hardness 86, 353
pattern (spotting) 85, 139
physical damage to 353, 394
thickness/structure 44, 398, 399
see also Dehulling
Seeding see Sowing
Seedlings, diseases of 266267, 275276,
277, 352
Seeds 36
chemical treatment 355
cleaning 391, 392396
disease transmission 263264,
272273, 316, 316
effect of water availability 173,
173175, 174
inoculation with rhizobia 202, 203,
237239, 355
moisture content 353, 354, 393394,
400
non-nutritive components 379383
nutritional composition 368379,
369
Index
455
of parasitic weeds 333, 345, 346
pest damage 224, 283, 288, 395, 401
priming 221, 354
quality 350354, 355
genotype purity 351, 352353
health 351, 352
vigour 351, 352, 353, 354
weed seed contamination 327,
331, 351
sizes, variation in 35, 4243, 8586,
187
supply 358
farmer-based 360, 361364, 362,
430, 434
formal/commercial 152, 356, 358,
360
informal/non-commercial 360
see also Cotyledons; Seedcoat
Selection, marker-assisted (MAS) 165
166, 167
Simple sequence repeat (SSR) markers
70, 132, 159160, 161
Single seed descent (SSD) 145
Sitona sp. (weevil) 116, 282, 283, 287, 288,
289, 292, 293, 296
nodule damage 37, 236
Sizing, seed 395, 396
Smynthurodes betae (root aphid) 284
Snacks, lentil 402, 418420
Soaking (pre-cooking) 409, 411
Sodicity, soil 54, 203204
Soil
diseases carried by 272276
evaporation 176, 177
health, contribution of lentils to
214, 232242, 233
inoculation with rhizobia 234, 235,
236239
nitrogen content 230234, 241242
nutrients
availability, and pH 201202, 202,
217
deficiencies 53, 5455, 198,
201204, 234
removal by seed crop 195, 197
status improvement 198201,
204206, 214
toxicities 5254, 110111, 201204,
232
types 3637, 5255, 8889, 217
Soup, lentil 2, 13, 414415, 415
South Asia
cropping systems 122, 213214,
215216
pilosae landrace 108, 121
recent cultivars released 129130
yield constraints 122123
see also Bangladesh
Sowing
depth 221, 334
equipment 251
rates 178, 220, 251, 334
row spacing 42, 215216, 221, 334
timing of 56, 123124, 177, 218220
to avoid infestations 291, 333
see also Seedbed preparation
Soybean dwarf virus (SbDV) 307, 308,
314, 315, 318
Species of Lens
geographical distribution 22,
2829, 68
in ICARDA collections 6466, 66
taxonomic status 1624, 1920
Splitting 399, 400, 401
Spodoptera sp. 280, 285
Sprouting 409, 411412, 418, 419420
Starch 44, 369, 373374, 403
Stem nematodes 275276
Stem rot disease, Sclerotinia 273275
Stemphylium botryosum (blight pathogen)
269271
Stems 3840, 44, 80
branching pattern 3940, 40, 4445
diseases of 263264, 268269,
273275, 277
height 3839, 79, 148
see also Growth habit
Stipules 24, 40, 80
Stomata 44, 183, 270
Storage
conditions 6667, 392394
pests 224, 283, 395, 401
Straw, lentil 248, 255, 259
Stubble
burning 335
standing, protective function 149,
336
Subsistence crop, lentil as 195, 356, 357
Subspecies of Lens culinaris
geographical distribution 26,
2829
taxonomic status 2425
456
Index
Subterranean clover stunt virus (SCSV)
307
Sugars 371
Summer crops 48, 219
Swathing 257258, 337, 401
Tagine, lentil 414

Tannins 85, 381, 408
Taxonomy 1525
Technology transfer see Outreach
programmes
Temperature
effects on seed yield 5152
flowering response 4950, 124, 181
Tendrils 31, 40, 7879
Testa see Seedcoat
Thielaviopsis basicola (black streak root
rot) 277
Threshing 255, 259, 260
Thrips 41, 282, 283, 287
Thysanoplusia oricholcea 285
Tillage
choice of techniques 218, 249250
effect on weeds 335, 336
effects on soil health 239240
Tobacco streak virus (TSV) 307
Tomato black ring virus (TBRV) 307
Tomato spotted wilt virus (TSWV) 307,
308, 314
Tomato yellow leaf curl virus (TYLCV)
307
Toxins, seed 383
Traits
for agroecological regions 4445,
107
correlation with yield 126127
evaluation, for genebank database
7072
selection, in breeding programmes
114, 143144, 147151
Transformation, genetic 116117,
157159
Transpiration 176, 181, 185186
Trichoplusia ni 285
Turnip mosaic virus (TuMV) 307
Tychius quinquepunctatus 287, 288
Uromyces viciae-fabae (rust pathogen)

264266
Uses of lentil 12, 213, 410

animal feed 2, 13, 138, 397
food (human) 102103, 402403,
409410, 413421
medicinal 13, 421422
processing by-products 397, 405,
409
Utera cropping see Relay cropping
Variation
environmental 3839, 4041, 58
genetic 6970, 121, 125, 127
quantitative 86, 88, 162
see also Diversity, genetic
Varieties, historical records 1314
Vectors, of virus diseases 288289, 307,
314, 315, 319
Vegetable, lentil as 417418
Vicieae, genera 1416, 68
Village-based seed enterprises (VBSEs)
361364, 362
Viruses, lentil
control 317320
detection 316317
disease transmission 288289,
314316, 315, 316
geographical distribution 308
species 306314, 307, 320
Vitamins 377379, 378, 412
Water availability
deficit effects 127, 172175, 173,
174, 180186
management of 222223
see also Irrigation; Rainfall
Water use efficiency (WUE) 5051,
176180, 178, 179
Waterlogging 51, 54, 217
Weeds
control methods 253
biological 331
chemical 57, 250, 336337, 346347
cultural 5657, 333334
integrated 337338, 346
mechanical 252253, 334336
preventive 331333
effect on yield 223, 326328, 327,
328, 329
Index
457
as pest hosts 291
seed viability 331, 333, 335, 336
sources 331333
species 328329, 330, 343345
Weevils see Sitona crinitus (pest weevil)
Wild Lens taxa 1624, 6466, 68, 82
Wilt disease, Fusarium 90, 272273
Winter crops
post-monsoon 4748, 116, 214216
temperate zone 34, 47, 109110
Winter hardiness 52, 88, 109110, 116
effect of weeds 223, 326328, 327,

328, 329
global and regional 6, 68, 78
limiting factors 2, 104, 122123
new and local cultivars compared

439440, 440
pest infestation losses 283, 288
and soil nutrient status 54, 195, 197
stability 58, 123
and water use efficiency 5051,
176178, 178, 180
Xanthomonas sp. (Bacterial leaf spot) 277
Yield, lentil crop

effect of inoculation 237
effect of seed priming 221, 354
effect of sowing time 218219
Zero tillage see No-till systems

Zinc
content, in lentil cultivars 112, 113,
150151
deficiency, soil 55, 198, 203

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The Lentil

Botany, Production and Uses

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William Erskine,1 Fred J. Muehlbauer,2 Ashutosh Sarker3

for Legumes in Mediterranean Agriculture (CLIMA), The University of

CABI Head Ofce

CABI is a trading name of CAB International

Tel: +44 (0)1491 832111

Tel: +1 617 395 4056

CAB International 2009. All rights reserved. No part of this publication

978 1 84593 487 3

Typeset by AMA Dataset Ltd, Preston, UK.

2. Global Production, Supply and Demand

3. Origin, Phylogeny, Domestication and Spread

4. Plant Morphology, Anatomy and Growth Habit

5. Agroecology and Crop Adaptation

6. Genetic Resources: Collection, Characterization, Conservation

8. Genetic Enhancement for Yield and Yield Stability

9. Breeding for Short Season Environments

10. Improvement in Developed Countries

Advances in Molecular Research

13. Soil Nutrient Management

14. Cropping Systems and Production Agronomy

15. Biological Nitrogen Fixation and Soil Health Improvement

17. Diseases and their Management

18. Insect Pests and their Management

19. Virus Diseases and their Control

20. Weed Management

21. Parasitic Weeds

22. Seed Quality and Alternative Seed Delivery Systems

23. Nutritional and Health-benecial Quality

24. Postharvest Processing and Value Addition

25. Food Preparation and Use

The colour plate section can be found following p.224

This page intentionally left blank

W. Erskine,1 F.J. Muehlbauer,2 A. Sarker3 and B. Sharma4

University of Western Australia, Crawley, Western Australia, Australia;

The lentil (Lens culinaris Medikus subsp. culinaris) is a lens-shaped grain

Adequate variability for many of the crops genetic constraints exists

Global Production, Supply and Demand

2.2. Production, Productivity and Area

CAB International 2009. The Lentil: Botany, Production and Uses

Global Production, Supply and Demand

Area harvested (million ha)

Global lentil productivity was only 560 kg/ha in 19611963, while it

Mean yield (kg/ha)

Global Production, Supply and Demand

Area (1000 ha)

Table 2.2. continued

Area (1000 ha)

North and Central

Global Production, Supply and Demand

Global Production, Supply and Demand

Table 2.5. Extract from FAO lentil consumption data

when on occasion national production is insufcient to meet export

de Agricultura Sostenible (CSIC), Crdoba, Spain; 2Universidad de Len,

3.1. Historical Data

J.I. Cubero et al.

As in the time of Columella, lentils were also used to feed pigeons.

3.2. The Genus Lens and the Vicieae

Origin, Phylogeny, Domestication and Spread

J.I. Cubero et al.