Mineral Nutrition On Root Growth

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From: N. K. Fageria and A. Moreira, The Role of Mineral Nutrition on Root Growth
of Crop Plants. In Donald L. Sparks, editor: Advances in Agronomy, Vol. 110,
Burlington: Academic Press, 2011, pp. 251-331.
ISBN: 978-0-12-385531-2
Copyright 2011 Elsevier Inc.
Academic Press.
Author's personal copy

C H A P T E R
F O U R
The Role of Mineral Nutrition

on Root Growth of Crop Plants
N. K. Fageria* and A. Moreira
Contents
252
255
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312
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1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
Introduction
Root-Induced Changes in the Rhizosphere
Root Systems of Cereals and Legumes
Contribution of Root Systems to Total Plant Weight
Rooting Depth and Root Distribution
Root Growth as a Function of Plant Age
RootShoot Ratio
Root Growth Versus Crop Yield
Genotypic Variation in Root Growth
Root Oxidation Activity in Oxygen-Deficient Soils
Root Growth in Conservation Tillage Systems
Mineral Nutrition Versus Root Growth
12.1. Nitrogen
12.2. Phosphorus
12.3. Potassium
12.4. Calcium
12.5. Magnesium
12.6. Sulfur
12.7. Micronutrients
13. Management Strategies for Maximizing Root Systems
13.1. Soil management
13.2. Plant management
14. Conclusions
Acknowledgment
References
Abstract
Agriculture is going through a profound revolution worldwide due to increasing
world demand for food, higher costs of energy and other inputs, environmental
pollution problems, and instability of cropping systems. In this context,
* Rice and Bean Research Center of Embrapa, Santo Antonio de Goias, GO, Brazil
Western Amazon Research Center of Embrapa, Manaus, AM, Brazil
Advances in Agronomy, Volume 110
ISSN 0065-2113, DOI: 10.1016/B978-0-12-385531-2.00004-9
{
2011 Elsevier Inc.

All rights reserved.
251

252
N. K. Fageria and A. Moreira
knowledge of factors that affect root development is fundamental to improving

nutrient cycling and uptake in soilplant systems. Roots are important organs
that supply water, nutrients, hormones, and mechanical support (anchorage) to
crop plants and consequently affect economic yields. In addition, roots improve
soil organic matter (OM) by contributing to soil pools of organic carbon (C),
nitrogen (N), and microbial biomass. Root-derived soil C is retained and forms
more stable soil aggregates than shoot-derived soil C. Although roots normally
contribute only 1020% of the total plant weight, a well-developed root system
is essential for healthy plant growth and development. Root growth of plants is
controlled genetically, but it is also influenced by environmental factors. Mineral nutrition is an important factor influencing the growth of plant roots, but
detailed information on nutritional effects is limited, primarily because roots
are half-hidden organs that are very difficult to separate from soil. As a result, it
is difficult to measure the effect of biotic and abiotic factors on root growth
under field conditions. Root growth is mainly measured in terms of root density,
length, and weight. Root dry weight is often better related to crop yields than is
root length or density. The response of root growth to chemical fertilization is
similar to that of shoot growth; however, the magnitude of the response may
differ. In nutrient-deficient soils, root weight often increases in a quadratic
manner with the addition of chemical fertilizers. Increasing nutrient supplies
in the soil may also decrease root length but increase root weight in a quadratic
fashion. Roots with adequate nutrient supplies may also have more root hairs
than nutrient-deficient roots. This may result in greater uptake of water and
nutrients by roots well supplied with essential plant nutrients, compared with
roots grown in nutrient-deficient soils. Under favorable conditions, a major part
of the root system is usually found in the top 20 cm of soil. Maximum root
growth is generally achieved at flowering in cereals and at pod-setting in
legumes. Genotypic variations are often found in the response of root growth
to nutrient applications, and the possibility of modifying root system response
to soil properties offers exciting prospects for future improvements in crop
yields. Rooting pattern in crop plants is under multi- or polygenic control, and
breeding programs can be used to improve root system properties for environments where drought is a problem. The use of crop species and cultivars
tolerant to biotic and abiotic stresses, as well as the use of appropriate cultural
practices, can improve plant root system function under favorable and unfavorable environmental conditions.
1. Introduction
Roots are important plant organs. They absorb water and nutrients
from the soil and translocate them to plant tops (Merrill et al., 1996, 2002;
Sainju et al., 2005a; Stone et al., 2001). Roots also give mechanical support
to plants and supply hormones that affect many physiological and biochemical processes associated with growth and development. Roots exert control

The Role of Mineral Nutrition on Root Growth of Crop Plants
253
over whole-plant growth and development by controlling the uptake of

mineral nutrients (Zobel, 1986). Cytokinins produced in roots are translocated to shoots and participate in the control of leaf protein biosynthesis
(Oritani, 1995). In addition, cytokinins may control nitrogen assimilation in
the root itself. Oritani (1995) also reported that rice roots synthesize zeatin
glucoside, a bound cytokinin, in addition to isopentenyladenine (IP),
isopentenyl-adenosine (IPA), trans-zeatin riboside (tZR), and trans-zeatin
(tZ), which are free cytokinins in zeatin-related compounds. Zobel (2005a)
reported that root system dynamics are instrumental in the maintenance of
biological and chemical equilibrium within the soil and modulate changes
to soil quality. In addition, genotypes with inherently large root systems
have been associated with reduced lodging in cereals and legumes (Stoffella
and Kahn, 1986). Soil is knitted together by plant roots, which form
complex and structurally diverse reinforcing structures. Near the soil surfaces, fibrous plant roots hold soil aggregates together against the stresses of
water and wind.
Vigorous root systems are needed for the development of healthy plants
and consequently, higher yields. Roots that are left in the soil after crop
harvest improve soil organic matter (OM) content and contribute to the
nitrogen cycle and microbial activity (Sainju et al., 2005a). All these activities improve soil structure, soil water holding capacity, water infiltration
into the soil, as well as reduce soil bulk density and soil erosion, ultimately
leading to greater soil productivity.
Processes that are largely controlled or directly influenced by roots and
often occur in the vicinity of the root surface are often referred to as
rhizosphere processes (Cheng and Kuzyakov, 2005). These processes may
include root turnover, rhizodeposition, root respiration, and rhizosphere
microbial respiration that are a result of microbial utilization of rhizodeposits. Rhizosphere processes play an important role in the global C cycle.
Terrestrial ecosystems are intimately connected to atmospheric carbon
dioxide levels through photosynthetic fixation of CO2, sequestration of
CO2 in plant and soil biomass, and the subsequent release of C through
respiration and decomposition of organic matter (Cheng and Kuzyakov,
2005). Carbon cycling belowground is increasingly being recognized as one
of the most significant components of the ecosystem C fluxes and pools
(Cheng and Kuzyakov, 2005; Jackson et al., 1997; Zak and Pregitzer, 1998).
Roots improve soil aggregation, which controls biological and hydrologic properties of the soil. A soil aggregate is a group of primary soil
particles that adhere to one another more strongly than to surrounding
soil particles (Follett et al., 2009). Root materials remain mixed within the
soil as they decompose, providing a gum-like material that cements soil
particles into aggregates (Melillo and Gosz, 1983; Tresder et al., 2005). Root
exudation occurs when organic acids either are leaked from the root or are
released as a means of interacting with microbes in the rhizosphere. These

254
exudates can influence microbial activity and the subsequent metabolism of

root detritus. Mucilages released by the root cap and epidermis link particulate organic residues with mineral fragments. Microbes in the rhizosphere
use plant mucilage as a substrate for growth and secrete their own mucilage,
thereby producing mucigel, a mucilaginous material of mixed origin which
stabilizes finer aggregates. In addition, in many soils, fungal hyphae are
important for stabilizing larger structural units (Goss and Kay, 2005).
Finally, allocation of carbohydrates and other C-containing molecules
directly to mycorrhizal fungi forms another conduit of C into the soil as
these microbes secrete their own exudates and as their tissues senesce
(Tresder et al., 2005).
The amount of C and N supplied by roots can be significant for maintaining or improving soil organic matter (Sainju et al., 2005b). The organic
input from plant roots to the surrounding soil is the principal support of the
biological activity and abundance of organisms in the rhizosphere (Cheng
et al., 1994; Kirchner et al., 1993). As much as 743% of the total aboveground and belowground plant biomass can be contributed by roots (Kuo
et al., 1997a,b). Roots can supply from 400 to 1460 kg C ha 1 during a
growing season (Kuo et al., 1997a; Qian and Doran, 1996). Liang et al. (2002)
reported that roots contributed as much as 12% of soil organic C, 31% of
water soluble C, and 52% of microbial biomass C within a growing season.
Roots may play a dominant role in soil C and N cycles (Gale et al., 2000a;
Puget and Drinkwater, 2001; Wedin and Tilman, 1990). Roots may have
relatively greater influence on soil organic C and N levels than the aboveground plant biomass (Boone, 1994; Haider et al., 1993; Milchumas et al.,
1985; Norby and Cortrufo, 1998; Sanchez et al., 2002). Balesdent and
Balabane (1996) reported that corn roots contributed 1.6 times more C to
soil organic C than did stover. Root-derived C is retained and forms more
stable aggregates than does shoot-derived C (Gale et al., 2000a,b).
Rhizodeposition, such as root exudates, mucilages, and sloughed cells,
may be a significant source of soil organic C (Balesdent and Balabane, 1996;
Buyanovsky et al., 1986; Sainju et al., 2005a). Helal and Sauerbeck (1987)
estimated that the amount of C released from roots as rhizodeposit could be
more than 580 kg C ha 1. This rhizodeposition increases microbial activity
and influences N mineralization in the soil (Bakken, 1990; Texier and Biles,
1990). Carbon contribution from corn root biomass and rhizodeposition to
soil organic C can be as much as 1.73.5 times greater than from stover
(Allmaras et al., 2004; Wilts et al., 2004).
The environment is seldom optimum for extensive and effective root
growth. Canopy conditions that limit photosynthesis reduce shoot growth
and limit assimilate translocation to the roots, thus reducing root growth
(Miller, 1986). Root growth is under multi- or polygenic control and is also
influenced by environmental factors, including soil temperature, soil moisture content, solar radiation and soil physical, chemical, and biological

255
properties (Fageria, 2002c, 2009; Klepper, 1992; Merrill et al., 1996;

Russell, 1977; Sainju et al., 2005b; Zobel, 1991). Most of the root biomass
of annual crops is located in the 020 cm soil depth. This may be associated
with greater organic matter, nutrients, aeration, and water availability in the
top soil layer compared to lower soil depths (Merrill et al., 1996, 2002; Qin
et al., 2004; Sainju et al., 2005a; Stone et al., 2001).
Increased knowledge of root architecture and root development dynamics could help improve crop productivity in agroecosystems. Better understanding of root architecture and growth dynamics of annual crops may lead
to a more efficient use of applied nutrients and water. The study of plant
roots is one of the most promising, but least explored, areas of research
related to plant growth. The aerial portions of plant species have received
greater attention and study, probably because of their conspicuousness and
easy access, while the subterranean portions have been neglected because of
the difficulty of observing and sampling them and the disruption of root
systems when they are removed from soil. Many crop root studies have
relied on soil cores and more recently on minirhizotron observations (Box
and Ramseur, 1993; Zobel, 2005b). The data collected using these methods
may not be representative of the crop as a whole (Andren et al., 1991;
Hansson et al., 1992; Hoad et al., 2001; Parker et al., 1991). In addition,
information about annual field crop root growth dynamics as a function of
environmental factors is scattered and often not readily accessible.
The primary objectives of this chapter are to review the latest advances in
relation to the role of mineral nutrition in the growth and development of
roots of annual crops. To make the subject matter as practical as possible, most
of the discussion is supported by experimental results. Our approach should
enhance understanding on the contribution of roots to total dry matter of
crops, to assess the effects of root system size and form on overall crop growth,
and to relate the effects of root growth on the environment. This information
may be useful for agricultural scientists in the fields of plant nutrition, water
use, breeding, and plant physiology who are interested in conducting
research to manipulate plant root systems in favor of higher yields.
2. Root-Induced Changes in the Rhizosphere

Pinton and Varannini (2001) suggested that the soil layer surrounding
roots should be termed the ectorhizosphere and the root inside the layer
colonized by microorganisms should be designated as endorhizosphere. The
two areas are separated by the root surface known as rhizoplane (Fig. 1).
Growing roots release an appreciable amount of organic components into
the rhizosphere. Marschner (1995) reported that three major components
released by roots are low-molecular weight organic compounds (free

256
Ectorhizosphere
Root hair
Mucigel
(plant and bacteria)
Endorhizosphere
Stele (xylem, phloem)

Epidermis
Cortex
Endodermis
Root cap
Sloughed root cap cell
Figure 1 Root cross section showing ecto- and endorhizosphere (Fageria and Stone,
2006).
exudates), high-molecular weight gelatinous materials (mucilage), and

sloughed-off cells and tissues and their lysates (Fig. 1). The rhizosphere is
the soil zone adjacent to plant roots which is physically, chemically, and
biologically different than bulk or nonrhizosphere soil. Plants influence the
physical (temperature, water availability, and structure), chemical (pH,
redox potential, nutrient concentration, root exudates, Al detoxification,
and allelopathy), and biological properties (microbial association) in the
rhizosphere. Their effects include changes in nutrient solubility, transport,
and uptake of mineral nutrient, and ultimately plant growth. Major rhizosphere changes are synthesized in Fig. 2 and their influence on nutrient
availability is discussed in details by Fageria and Stone (2006).
3. Root Systems of Cereals and Legumes

Cereals as well as legume seeds contain relatively large reserves of
storage carbohydrates and nutrients which allow the initial root system to
grow rapidly to considerable depth (Marschner, 1998). Branching often

Physical changes
Temperature
Water holding capacity
Structure
257
Biological changes
Nitrogen fixation
PGPR bacterias
Mycorrhizal fungi
Harmful microorganisms
Chemical changes
pH
Redox potential
Release of organic compounds by roots
Nutrient concentration and availability
AI detoxification
Allelopathy
Figure 2 Major physical, chemical, and biological changes in the rhizosphere (Fageria
and Stone, 2006).
begins before the leaves have unfolded, with the result that the plant
establishes early contact with moist soil (Hoad et al., 2001). Generally,
roots are classified into four groups. These groups are the taproot, basal
roots, lateral roots, and shootborn or adventitious roots (Zobel, 2005a).
When plants produce secondary shoots (tillers) or shoot branches which
develop roots, these roots are commonly called adventitious roots. To
indicate the true origin of these adventitious roots, the term shootborn is
sometimes used (Zobel, 2005a). The primary function of the taproot, basal
roots, and adventitious roots is to establish the most optimum framework
from which to initiate small lateral roots to effect water and nutrient uptake
(Zobel, 2005b). The taproot penetrates relatively deeply to ensure an

258
adequate supply of soil water, the basal roots spread out laterally to ensure a
structure for lateral roots that take up P and other nutrients that are less
abundant in the lower levels of the soil profile (Zobel, 2005b), and to
provide a degree of lodging resistance to the plant as it matures and produces
seed (Barlow, 1986; Stoffella et al., 1979). For many grasses and other species
in which root secondary thickening is not important, the shootborn roots
take over the role of the basal roots. The shootborn roots continue to build
the framework with larger and larger conducting roots as the plant increases
in size (Zobel, 2005b). The basal and shootborn roots probably provide little
direct uptake of nutrients and water (St. Aubin et al., 1986).
Besides length and weight, surface area is an important parameter of the
root system in crop plants. The form of root systems and their development
conditions greatly affect the surface area of roots. The surface area of roots
has a high positive correlation with the amount of nutrient absorption
(Takenaga, 1995). Various studies show that 9095% or more of the root
length of an intact plant is made up of roots <0.6 mm in diameter (Zobel,
2003, 2005b).
Monocots and dicots typically have different root system structures.
Root systems of monocots are fibrous, whereas dicots often have taproots.
The fibrous root systems of monocots consist of seminal, nodal, and lateral
roots. Seminal roots develop from primordia within seeds and nodal roots
develop adventitiously from lower stem nodes. All adventitious roots of
stem origin are called nodal roots to distinguish them from other adventitious roots that emerge from the mesocotyl or elsewhere on the plant.
Nodal roots are identified by the node number from which they originate.
Nodal roots may be functional or nonfunctional (Thomas and Kaspar,
1997). Functional nodal roots are defined as roots that have emerged from
stem nodes, entered the soil, and developed lateral roots and/or root hairs.
Nonfunctional nodal roots are defined as roots that have emerged from
aboveground stem nodes and have not entered the soil or produced lateral
roots (Thomas and Kaspar, 1997).
Initial seminal or nodal roots develop laterals that are classed as roots of
the first order, roots that develop from first-order roots are classed as
second-order roots, and additional roots that develop from these laterals
are classed as third-order roots, fourth-order roots, etc. (Yamauchi et al.,
1987a,b). Nodal roots are also known as adventitious, coronal, and/or
crown roots. Roots of cereals such as rice include mesocotyl, radical
(seminal), and nodal or adventitious roots (Yoshida, 1981). Mesocotyl
roots emerge from the axis between the coleoptile node and the base of
the radical, and they typically develop only when seeds are planted very
deep or are treated with chemicals (Yoshida, 1981). Until adventitious roots
develop, seedlings must rely on roots which initiate on the subcoleoptile
internodes above the seed or seminal roots below the seed. Adventitious
roots are important to seedling establishment because they can conduct

259
more water than smaller diameter seminal roots. Adventitious roots

may develop as early as 2 weeks after sowing. Seedling survival may increase
when seeds are sown at greater soil depths, where greater soil water availability may increase adventitious root development (Fageria et al., 2006).
Tiller roots do not form in cereals until tillers have two to three leaves
(Kleeper et al., 1984), and until these roots have developed, parent culms
must provide nutrients and water. Parent culms may also have to provide
hormonal control so essential for tiller survival. Delayed root production by
tillers may explain why late tillers often do not survive (Kleeper et al., 1984).
Figure 3 shows the radical and adventitious root system of upland rice
(Oryza sativa L.) (cereal) and Fig. 4 shows the tap root system of dry bean
(Phaseolus vulgaris L.) (legume).
In addition to their morphological differences, roots of cereals and
legumes have different physiochemical properties. The surface of plant
roots has a negative electric charge, mainly due to carboxyl groups in the
pectin of the root cell walls. The density of this negative charge is defined as
a cation exchange capacity (CEC; Takenaga, 1995). The CEC of cereals
3rd leaf
4th leaf
2nd leaf
1st tiller
1st leaf
Adventitious root
Radicle root
Figure 3 Root system of upland rice seedlings (Fageria, 2007).

260
First trifoliate
Gem
Primary leaf
Epicotyl
Hypocotyl
Adventitious roots
Basal roots
Lateral roots
Figure 4
Tap root
Root system of dry bean seedling (Fageria and Santos, 2008).
such as rice, barley, and corn is typically lower than the CEC of legumes like
dry bean and broad bean. Roots with high CEC absorb more divalent
cations like Ca2 and Mg2 than monovalent cations such as K and
NH4. On the other hand, roots with lower CEC absorb more monovalent
than divalent cations. Hence, in grasslegume mixtures, legumes generally
suffer with K deficiency due to large uptake of this element by grasses.
Essau (1977), Fageria et al. (2006), Klepper (1992), Leskovarant and Stofella
(1995), OToole and Bland (1987), and Zobel (1991, 2005a,b) have dealt
extensively with various types of monocotyledonous and dicotyledonous
roots and root hairs, and their growth and morphology.
4. Contribution of Root Systems to Total

Plant Weight
Crops can accumulate photosynthetic products in their stems, leaves,
grains, and roots, and the development of robust root systems is necessary to
produce good crop yields. A well-developed root system is needed to
absorb adequate amounts of water and nutrients, especially when plants

261
are under abiotic or biotic stress. The traditional view is that a large vigorous
root system, through avoidance of plant water deficits, is required for high
yields in water-limited environments (Ludlow and Muchow, 1990). In
addition, large root systems add more organic matter to the soil, creating
better environmental conditions for the growth and development of crops.
The distribution of photosynthetic products in the root, shoot, and grain is
determined genetically, but it also varies with environmental conditions
(Fageria, 1992, 2009; Sainju et al., 2005a). Because roots are so difficult to
separate from soils, little is known about the effects of soil and crop management practices on their growth and development (Sainju et al., 2005b).
In most annual plant species, only 1020% of mature plant biomass
consists of root tissue (Zobel, 1986). Similarly, Fageria (1989a) also reported
that the contribution of roots of food crops to the total plant weight varies
from 1020%, depending on species and cultivars within species. However,
Sainju et al. (2005b) reported that C accumulation in cotton and sorghum
roots ranged from 1% to 14%. Data in Table 1 show the contribution of the
root system of 20 upland rice genotypes grown at two N rates in a Brazilian
Oxisol. In this study, the nitrogen genotype interaction was significant
because some genotypes were highly responsive to the N application while
others were not. Thus, genotype selection is an important strategy for
upland rice production in Brazilian Oxisols. In the control treatment, the
contribution of the root system to total plant weight varied from 12% to
30%, with an average value of 22%. At the 300 mg N kg 1 soil treatment,
the contribution of root weight to total plant weight varied from 3% to
21%, with an average value of 14%. The proportionally lower root dry
weight at the higher N rate was associated with a significant increase in the
grain and shoot weight of rice genotypes with the addition of N fertilizer
(Fageria and Baligar, 2005).
The contribution of root systems to total plant weights of tropical
legume cover crops under three P levels is shown in Table 2. There was a
significant influence of P rate, cover crop species and P species interaction was significant, indicating different responses of cover crops at different
P rates. At 0 mg P kg 1 level, the contribution of roots to total plant weight
varied from 4.91% to 21.50%, with an average value of 12.98%. With
100 mg P kg 1, the root contribution to the total plant weight varied
from 11.30% to 25.16%, with an average value of 18.93%. At the
200 mg P kg 1 P rate, the contribution of roots to the total plant weight
varied from 6.36% to 25.91%, with an average value of 14.79%. Overall,
increase in root contribution to the total plant weight with the increase in P
rate may be associated with the response of legumes to P fertilization.
Significant differences have been reported among the crop species and
genotypes of the same species in the absorption and utilization of P
(Epstein and Bloom, 2005; Fageria, 2009; Marschner, 1995). Tian et al.
(1998) reported significant responses of legume cover crops grown on

262
Table 1 Contribution of upland rice roots in the total plant dry weight (%) as
influenced by N rates
Genotype
0 mg N kg 1 soil
300 mg N kg 1 soil
BRA01506
BRA01596
BRA01600
BRA02535
BRA02601
BRA032033
BRA032039
BRA032048
BRA032051
BRA042094
BRA042156
BRA042160
BRA052015
BRA052023
BRA052033
BRA052034
BRA052045
BRA052053
BRS Primavera
BRS Sertaneja
Average
F-test
N rate (N)
Genotype (G)
NG
CV(%)
18abcd
18abcd
23abcd
25abcd
23abcd
20abcd
14cd
12d
23abcd
25abcd
24abcd
28abc
17abcd
16bcd
22abcd
30a
27abc
29ab
28abc
26abc
22
3g
3g
6fg
14cde
19abcd
15bcde
21ab
19abc
14cde
12e
15cde
15cde
13de
21a
15bcde
12e
17abcde
11ef
12e
14de
14
*
**
**
17
Root dry weight

100:
Dry weight of root; shoot; and grain
*,** Significant at the 5% and 1% probability levels, respectively. Means followed by the same letter in
the same column are not significant at the 5% probability level by Tukeys test.
Contribute of roots in total plant dry weight%
Alfisols in Africa. Similarly, Ae et al. (1990) reported that pigeon pea as a

cover crop was more efficient in utilizing iron-bound P than several other
cover crops. These authors also reported that this ability of pigeon pea was
attributed to root exudates, in particular, piscidic acid and its p-O-methyl
derivative, which release P from FeP by chelating Fe3. These results also
show that P fertilization also improves the root weight of cover crops,
which may be beneficial in improving soil organic matter content and soil
microbial activities.

263
Table 2 Contribution of tropical legume cover crops root dry weight (%) in the total
plant weight as influenced by phosphorus rates
Cover crop
0 mg P kg 1 100 mg P kg 1 200 mg P kg 1
Crotalaria breviflora
Crotalaria juncea L.
Crotalaria mucronata
Crotalaria spectabilis Roth
Crotalaria ochroleuca G. Don
Calopogonium mucunoides
Pueraria phaseoloides Roxb.
Cajanus cajan L. Millspaugh
Cajanus cajan L. Millspaugh
Dolichos lablab L.
Mucuna deeringiana (Bort) Merr.
Mucuna aterrima (Piper & Tracy)
Holland
Mucuna cinereum L.
Canavalia ensiformis L. DC.
Average
F-test
P rate (P)
Cover crops (C)
PC
20.77a
12.28abc
4.91c
11.66abc
5.18c
11.66abc
7.22bc
21.50a
16.90abc
12.81abc
19.70ab
14.17abc
11.39ef
19.77abcdef
19.29abcdef
24.72ab
22.76abc
12.16def
17.17abcdef
21.12abcde
21.80abcd
22.22abc
15.08bcdef
25.16a
21.22ab
12.49bcd
21.96ab
9.26cd
17.38abc
25.91a
8.15cd
6.36d
13.77bcd
14.71bcd
13.36bcd
16.43abc
11.09abc
11.83abc
12.98
13.44cdef
11.30f
18.93
16.94abc
9.12cd
14.79
**
**
**
Root dry weight

100:
Dry weight of root and shoot
** Significant at the 1% probability level. Means followed by the same letter in the same column are not
significant at the 5% probability level by Tukeys test.
Contribute of roots in total plant dry weight%
5. Rooting Depth and Root Distribution

Rooting depth and distribution are important traits for absorption of
water and nutrients from the soil profile. Rooting depth, that is, the
maximum depth that roots reach, is difficult to ascertain in the field
(Hoad et al., 2001). Hsiao et al. (2009) reported that roots typically reach
maximum depth about the time when the canopy begins to senesce under
nonstress conditions. Gregory (1994) reported that rooting depth in cereals
increases until anthesis. Hoad et al. (2001) reviewed the rooting depth
literature for cereals and concluded that individual roots of cereal crops
can reach a depth of over 2.0 m under favorable conditions. Soil compaction reduces rooting depth (Lipiec et al., 1991; Unger and Kaspar, 1994).

264
Martino and Shaykewich (1994) reported that the proportion of roots

penetrating the soil is inversely related to the soil penetration resistance.
Ueda (1936) observed that wheat cultivars with relatively great cold
resistance had roots that penetrated rapidly into deeper soil layer at early
stages of growth. Similarly, Sanders and Brown (1978) reported that differences in the yields of indeterminate and determinate soybean varieties
mirrored the distribution of their root systems, although the dry weights
of their root systems were almost the same. Yamauchi et al. (1987b)
compared the root system distributions of 13 species of cereals and reported
that there was a significant difference among cereals in the distribution and
depth of penetration of root systems. Some of the cereals had concentrated root systems and others had scattered type root systems.
Large amounts of organic matter and immobile nutrients are generally
found in the upper soil layers. Hence, a major part of the roots of the most
crops is concentrated in the upper 020 cm soil depth (Gregory, 1994).
Application of nitrogen fertilizer to barley caused an accumulation of
9097% of the root mass in the top 30 cm soil layer (Hansson and
Andren, 1987). Similarly, Haberle et al. (1996) reported only a few
unbranched primary roots below a depth of 25 cm in fertilized wheat.
Differences in root lengths, dry weights of roots at different soil depths,
and the extent of rooting at the seedling stage were related to differences in
yield and the ability of wheat cultivars to escape drought (Hurd, 1974).
Upland rice cultivars, which are more drought tolerant than lowland
cultivars, have deeper and more prolific rooting systems (Steponkus et al.,
1980). When soil types did not restrict the rooting potential, deep rooting of
bean cultivars was positively associated with seed yield, crop growth, cooler
canopy temperature, and soil water extraction (Sponchiado et al., 1989).
Because about 90% of the total NH4, P, and K uptake and root length of
flooded rice cultivars occur within the surface 20 cm of soil, samples
collected for routine soil tests should be taken from the top 20 cm (Teo
et al., 1995). Lowland rice plants develop a surface mat of roots in the
oxygenated zone near the soil surface soon after application of flood waters
(University of Arkansas Cooperative Extension Service Rice Committee,
1990).
Durieux et al. (1994) reported that more than half of the root length of
maize was located in the surface 020 cm depth at all sampling times during
a season. Roots of the peanut (Arachis hypogaea L.) cultivar Florunner
penetrated to depths up to 280 cm when grown in a sandy soil, and the
most extensive root growth occurred in the top 30 cm (Boote et al., 1982).
Sharratt and Cochran (1993) reported that 85% and 95% of the root mass of
barley was located in interrows of the top 20 and 40 cm of soil, respectively.
Welbank and Williams (1968) also found that nearly 80% of barley roots
occupied the uppermost 15 cm of soil. A study conducted by Stone and
Pereira (1994a,b) of four common bean cultivars and three upland rice

265
cultivars to evaluate rooting depths in an Oxisol showed that 70% of the

roots were concentrated in the top 20 cm layer and about 90% were
concentrated in the top 40 cm soil depth of both crops.
The presence of these roots in surface soil layers may contribute to large
amounts of nutrients measured in the upper 20 cm of the soil. Using the
ClaassenBarber model to predict nutrient uptake by maize grown in silt
loam soil, >90% of K and P uptake occurred in the top 20 cm soil depth
(Schenk and Barber, 1980). Silberbush and Barber (1984) reported that
about 80% of P and 54% of K uptake by soybean was from 0 to 15 cm
depths.
Soybean cultivars differ in their rate of downward growth during specific
shoot development stages and in their maximum rooting depth on specific.
Cultivars selected for rapid taproot elongation rates in a greenhouse trial
were found to have greater rooting depths in rhizotron and field trials than
cultivars selected for slow taproot elongation (Kaspar et al., 1978, 1984).
6. Root Growth as a Function of Plant Age

Root development varies with stages of plant growth and development. The most rapid development of corn (Zea mays L.) roots occurs
during the first 8 weeks after planting (Anderson, 1987). As corn plants age,
growth of roots generally increases at slower rates than shoots (Baligar,
1986). After silking, corn root length declines (Mengel and Barber, 1974).
This decline in root length after silking presumably is due to the high C
demand of grain resulting in enhanced translocation of C and N to grain,
including some C and N that roots would normally obtain (Wiesler and
Horst, 1993).
Peanut (A. hypogaea L.) root length density and root weight density
increased at each soil depth increment from planting to 80 days after
planting (Ketring and Reid, 1993). These authors reported that roots had
penetrated to depths of 120 cm 4045 days after planting and spread laterally
to 46 cm in mid-furrow. The 015 cm depth increment had the highest
mean root length density, which increased to a maximum of 2.1 cm cm 3 at
80 days after planting (Ketring and Reid, 1993). This meant that peanut
roots were established both deeply and laterally in the soil profile early in the
growing season. This would be advantageous in drought environments and
helpful for water management.
Sunflower (Helianthus annuus L.) rooting depth reached 1.88 m at the
beginning of disk flowering and 2.02 m at the completion of disk flowering
( Jaffar et al., 1993). In a review of depth development of roots with time for
55 crop species (Borg and Grimes, 1986), it was shown that maximum
rooting depth for most crop species was generally achieved at physiological

266
Root dry weight

(g plant1)
maturity. Kaspar et al. (1984) noted that the rate of soybean (Glycine max
L. Merr.) root depth penetration reached a maximum during early flowering and declined during seed fill. However, some root growth was observed
throughout the reproductive stage until physiological maturity (Klepper and
Kaspar, 1994).
Slaton et al. (1990) studied root growth dynamics of lowland rice and
found that maximum root growth rates were reached between active
tillering and panicle initiation, and maximum root length was reached by
early booting. Beyrouty et al. (1987) noted that the most rapid rate of root
and shoot growth in flooded rice occurred before panicle initiation, which
corresponds to the plant transition between vegetative and reproductive
growth. Approximately 77% and 81% of total shoot and root biomass,
respectively, was achieved before panicle initiation. Following panicle
initiation, the length of roots and shoots increased only slightly until harvest
(physiological maturity). Beyrouty et al. (1988) also reported that lowland
rice root growth was most rapid during vegetative growth, with maximum
root length occurring at panicle initiation. Root length either plateaued or
declined during reproductive growth.
Fageria and Santos (2011) studied the root and shoot growth of lowland
rice during its growth cycle (Fig. 5). Root dry weight increased in a
quadratic fashion with the advancement of plant age from 19 to 120 days,
but shoot dry weight increased linearly during the growth cycle. Development of the root system was slow during the first 40 days after sowing and
then it increased almost linearly until physiological maturity. The slow
40
Y = 2.4581 + 0.0952X + 0.00098X 2

R 2 = 0.9063**
30
20
10
Shoot dry
weight (g plant1)
0
60
Y = 12.5474 + 0.5287X
R 2 = 0.9702**
50
40
30
20
10
0
20
40
60
80
100
120
140
Plant age (days after sowing)
Figure 5 Root and shoot dry weight of lowland rice as a function of plant age (Fageria
and Santos, 2011).

267
Maximum root
length (cm)
increase in root dry weight early in the growth cycle may be associated with
low translocation of photosynthetic materials due to low leaf area (Fageria,
2007). When there is a low amount of photosynthetic product, a major part
goes to the shoot, and very little is translocated to the roots (Fageria, 1992).
Root growth generally parallels shoot growth in crop plants. When a
large amount of nutrients, especially N, is supplied to leaves from roots,
photosynthesis remains high during maturation, which secures the supply of
carbohydrates to roots. Hence, the activities of roots and shoots are mutually
dependent (Osaki et al., 1997). Figure 6 shows maximum root length and
root dry weight of dry bean during the growth cycle of a Brazilian cultivar
BRS Valente under greenhouse conditions. Maximum root length was
40
30
20
10
Root dry weight

(g plant1)
0
4
Y = 6.0778 + 0.7862X 0.0049X 2)

R 2 = 0.8353**
Y = 0.0267EXP(0.0824X 0.00032X 2)
R 2 = 0.8545**
Number of
trifoliates (plant1)
0
20
Y = 0.4461EXP(0.0787X 0.00041X 2)
R 2 = 0.9216**
10
Shoot dry weight

(g plant1)
0
20
Y = 0.0297EXP(0.1161X 0.00054X 2)
R 2 = 0.9430**
10
20
40
60
80
100
Plant age (days after sowing)
Figure 6 Relationship between plant age and dry bean growth parameters (Fageria
and Santos, 2008).

268
achieved at 80 days after sowing. However, root dry weight increased

quadratically with plant age, much like shoot growth. The youngest
growing portions of the roots are most active in water and mineral nutrient
uptake; therefore, the root system must continually grow to provide
adequate nutrition for the plant (Brown, 1984).
7. RootShoot Ratio
The partitioning of photoassimilate between roots and shoots has
frequently been analyzed as a balance between root and shoot activity
(Brouwer, 1966; Davidson, 1969a; Werf, 1996). Different plant species
may have different patterns for photosynthate transportation and allocation
to shoot and root (Dyer et al., 1991; Freckman et al., 1991). There is an
interdependence of shoot and root for growth and development. The shoot
relies on the root for water and nutrients, while the roots depend on the
shoot for carbohydrates (Hoad et al., 2001). The terms shoot and root
are used here in a botanical sense and refer, respectively, to the entire aerial
and subterranean portions of higher seed plants (Aung, 1974). In the early
part of the twentieth century, shootroot ratios were used rather extensively
to characterize plant response to imposed nutritional changes. Root growth
is closely related to the whole plant growth. This relationship is called
allometry or relative growth. Root dry weight is related to the total dry
weight of a plant using the following equation (Yoshida, 1981):
WR HWTh ;
where WR is the root dry weight, WT is the total dry weight (shoot
dry weight root dry weight), and H and h are constants. The above
relationship has been tested for different rice cultivars grown under various
environmental conditions, and can be expressed by the following equation
(Yoshida, 1981):
WR 0:212WT0:936 :
When plants are small (substitute 1 for WT), WR/WT is 0.2; WR/WT
values approach 0.1 as plants grow larger (substitute 105 for WT). In other
words, ratio of root dry weight to total dry weight ranges from 0.2 at the
seedling stage to 0.1 at the reproductive stage (heading) for rice (Yoshida,
1981).
The above relationship between root and total dry weights gives an
estimate of root mass that remains in soil if shoot weight is known. For
example, when plants produce shoot dry weights of 3 Mg ha 1 at heading,

269
root dry weights remaining in soil should be 330 kg ha 1. Partitioning of

dry matter in roots relative to shoots is high during the seedling stages of
growth and steadily declines throughout development (Evans and Wardlaw,
1976). The shoot:root biomass ratio changes during ontogeny, generally
becoming high as the plant approaches flowering, and stabilizing after
flowering (Zobel, 1986). Shoot-to-root ratios of common bean, rice,
wheat, and cowpea increased as plants advanced in age (Fageria, 1992).
Increases in shoot-to-root ratios indicate that shoots have a higher priority
for photosynthate accumulation than roots. If shootroot ratios decrease
with time, roots have preferential utilization of photosynthates under the
existing plant growth conditions. A relatively high conservation of photoassimilate in shoots may increase the plants photosynthetic leaf area while
decreasing root biomass and the plants capacity for water and nutrient
uptake (Werf, 1996).
Environmental stresses increase the relative weights of roots compared to
shoots (Eghball and Maranville, 1993). Decrease in the availability of N, P,
or water increased rootshoot ratios of perennial ryegrass (Lolium perenne L.)
(Davidson, 1969b). Although deficiencies of many mineral elements influence plant growth and rootshoot relationships, invariably water and N
deficiency limit shoot growth the most. Rootshoot ratios of 28-day-old
maize plants were 0.27, 0.15, and 0.18 at volumetric soil moisture contents
of 0.22, 0.27, and 0.32 m3 m 3, respectively (Mackay and Barber, 1985).
When plants are N-deficient, relatively more photosynthate is used by roots
as they develop greater length to aid the plant in obtaining more N. In
general, when low nutrient levels do not reduce maize grain yield by more
than 20%, addition of N will reduce total root weights even though shoot
weights increase (Barber, 1995). Champigny and Talouizte (1981) reported
that under N deprivation, translocation of photoassimilates from shoots to
roots increased because of increased sink strength of roots compared to
shoot sinks. In an experiment with 18-day-old wheat seedlings deprived of
N for 7 days, soluble sugar contents in roots were higher than in the
corresponding roots of seedlings grown continuously with complete nutrient solutions (Talouizte et al., 1984). Similarly, rootshoot ratios of maize
plants were higher when grown with low soil N compared to adequate N
(Eghball and Maranville, 1993).
Soil salinity is another important soil chemical property that influences
shootroot ratios. The depressing effect of salinity on root growth is
generally less severe than its effect on shoot growth. Shalhevet et al.
(1995) summarized the results of 10 experiments relating shoot and root
growth to salinity. In all the experiments, the root and shoot responses were
evaluated by measuring fresh or dry weights at the end of the experimental
periods. All the 10 experiments produced either the same or stronger
growth responses of shoots than roots because of the imposed osmotic
potential. However, Slaton and Beyrouty (1992) observed shootroot ratios

270
of rice remained constant as a result of a functional equilibrium in which

shoot growth was proportional to root growth.
Partitioning of photosynthates and their effects on dry matter distribution is influenced by several environmental factors such as low temperature,
drought, and mineral nutrient deficiency (Wardlaw, 1990). The mineral
nutrients P and N exerted pronounced influences on photosynthate and dry
matter partitioning between shoots and roots (Costa et al., 2002). Phosphorus and N-deficient plants usually produce proportionately more dry matter
to roots than shoots, compared with unstressed plants. This probably results
from higher export rates of photosynthate to roots in deficient plants. Leaf
expansion is highly sensitive to low tissue P concentrations, producing
higher concentrations of sucrose and starch in P-deficient leaves because
of reduced demand (Fredeen et al., 1989). Thus, roots become more
competitive for photosynthates than shoots, which leads to higher export
of carbohydrates to roots with correspondingly lower shootroot ratios
(Rufty et al., 1993). Cakmak et al. (1994) reported that dry matter partitioning between shoots and roots of common bean was affected differently by
low supplies of P, K, and Mg. Although total dry matter production was
somewhat similar in P-, K-, and Mg-deficient plants, K- and especially Mgdeficient plants had greater than normal shootroot ratios, while P-deficient
plants had smaller than normal shootroot ratios (Cakmak et al., 1994).
Shootroot dry weight ratios were 1.8 in P-deficient, 4.9 in control, 6.9 in
K-deficient, and 10.2 in Mg-deficient plants. Upland rice usually has high
root/shoot weight ratios than lowland rice, an adaptation improving access
to soil water (Dingkuhn and Kropff, 1996).
8. Root Growth Versus Crop Yield

Roots are responsible for absorption of water and nutrients which are
important resources affecting crop yields. In addition, roots improve soil
organic matter content and biological activity in the rhizosphere. Root
length and root dry weight are standard root parameters that are measured
in many studies, largely because they are more easily determined than other
root system properties (Gregory, 1994). Barber and Silberbush (1984)
studied the relationship between root length and soybean yield and concluded that yield was significantly related to total root length at the R6 (full
seed) stage. These authors concluded that root growth is important in
determining the nutrient supply to the shoot which, in turn, affects crop
yield. Similarly, Thangaraj et al. (1990) reported that root length density of
lowland rice at flowering was directly proportional to grain yield. Leon and
Schwang (1992) used the grid intercept method (Newman, 1966) to evaluate differences in total root length between cultivars of oats and barley and

271
Grain yield
(g plant1)
Y = 3.6178EXP(0.9326X 0.0109X
20
R = 0.5101**
10
20
30
40
Maximum root length (cm)
Figure 7 Relationship between maximum root length and grain yield of upland rice.
Not the root dry weight.
Grain yield
(g plant1)
20
10
Y = 1.0257EXP(0.9397X 0.0718X
R 2 = 0.8656**
0
Root dry weight (g plant1)
Figure 8
Relationship between root dry weight and grain yield of upland rice.
found that yield stability was correlated with root system length. The first
author studied the relationship between maximum root length and root dry
weight and grain yield of upland rice (Figs. 7 and 8). Grain yield increased in
a quadratic fashion with increasing root length or root dry weight, and root
dry weight was a better predictor than root length of yield. Similarly, the
author studied the relationships between root length and root dry weight
and shoot dry weight of tropical legume cover crops (Figs. 9 and 10). There
was a significant increase in shoot dry weight of legume cover crops with
increasing root length and dry weight, and as with upland rice, root dry
weight was a better predictor than root length of shoot dry weight.
9. Genotypic Variation in Root Growth

Variability in root growth among crop species and among genotypes
of the same species is widely reported in the literature (Fageria. 2009;
Gregory, 1994; Kujira et al., 1994; Marschner, 1998; OToole and Bland,
1987). This variability can be used in improving the yield of annual crops by

272
Y = 2.9808 0.2634X + 0.0076X 2
Shoot dry weight (g plant1)
R 2 = 0.8783**
7
6
5
4
3
2
1
0
10
20
30
40
50
Shoot dry weight (g plant1)
Figure 9 Relationship between maximum root length and shoot dry weight of
tropical legume cover crops. Values are averages of 14 tropical legume cover crops.
Y = 0.3422EXP(5.5187X 2.7205X
8
7
R 2 = 0.9712**
6
5
4
3
2
1
0
0.2
0.4
0.6
0.8
1.0
1.2
Figure 10 Relationship between root dry weight and shoot dry weight of tropical
legume cover crops. Values are averages of 14 tropical legume cover crops.
incorporating vigorous root growth into desirable cultivars. Vigorous root

growth is especially important when nutrient and water stress are significant
(Gregory, 1994). Ludlow and Muchow (1990), in their review of traits
likely to improve yields in water-limited environments, place a vigorous
rooting system high in their list of properties to be sought.
OToole and Bland (1987) reviewed genotypic variation in root growth
of annual crops and reported significant differences in rooting depths,
maximum root length, and distribution pattern in the soil profile. Hurd
(1974) and Yoshida and Hasegawa (1982) reported rooting depth differences among genotypes of wheat (Triticum aestivum L.) and rice,

273
respectively. Similarly, Brown et al. (1987) and Pan et al. (1985) also
reported differences in rooting depths of corn and barley (Hordeum vulgare
L.), respectively. Similar variability exists in dicotyledonous species where
most work has been undertaken on legumes, including dry bean (Fageria,
2002a; White and Castillo, 1989), soybean (Taylor et al., 1978; Zobel, 2005a),
chickpea (Cicer arietinum L.; Vincent and Gregory, 1986), peanut (Mathews
et al., 1988), and white clover (Trifolium repens L.; Caradus, 1990). Hamblin and
Tennant (1987) compared root growth and water uptake of wheat and lupin
(Lupinus albus L.) and Gregory and Brown (1989) did similar work with barley
and chick pea. These authors concluded that there were differences between
the species in the root length necessary to extract water, and the rate of water
extraction was greater in legumes than in cereals. Fageria (1991) studied root
dry weight of the Brazilian upland rice cultivars IAC 47 and the International
Rice Research Institute (IRRI) cultivar IR 43 and concluded that root dry
weight of Brazilian cultivars was almost double that of the IRRI cultivar
during the entire growth cycle. Xiaoe et al. (1997) reported that hybrid rice
has a more vigorous root system, larger panicle, and more grains per panicle
than traditional rice cultivars.
The genotypic variability in the root growth of annual crops has been
used to identify superior genotypes for drought-prone environments
(Gregory, 1994; Hurd et al., 1972). Gregory and Brown (1989) reviewed
the role of root characters in moderating the effects of drought and concluded that roots may have a direct effect, by increasing the supply of water
available to the crop, or an indirect effect by changing the rate at which the
supply becomes available. Where crops are grown on deep soils and water is
stored throughout the whole soil profile, the depth of rooting has a major
influence on the potential supply of water (Gregory, 1994). Rain may
replenish the upper soil during the season, but later growth and grain filling
in many crops are accomplished during periods of low rainfall when soil
moisture stored deep in the profile must be utilized. Sponchiado et al. (1989)
reported that in dry bean, drought avoidance results from root growth and
soil water extraction deep in the profile.
Atkinson (1990) reported significant variation in the speed of root
penetration, specific root length, branching pattern, root density, total
root mass, and root hair development of 25 spring barley cultivars. Information reported in the literature on old and new cereal cultivars indicates
that more modern cultivars are more responsive than older cultivars to high
nutrient availability (Haberle, 1993; Haberle et al., 1995), although they
tend to have a lower root fraction (Wahbi and Gregory, 1995). Root
physiological characteristics also differ among cultivars and can affect processes like nutrient acquisition (Hoad et al., 2001; Marschner, 1998). The
rate of uptake of nutrient per unit root length depends on the nutrient
availability but also varies considerably among cultivars (Hoad et al., 2001;
Romer, 1985).

274
10. Root Oxidation Activity in Oxygen-Deficient

Soils
Roots require O2 for respiration, water and mineral absorption, and
other metabolic activities. Brailsford et al. (1993) reported that roots asphyxiate within a few hours or days at normal growing temperatures unless at
least a small amount of oxygen (0.1 kPa in a flowing gas phase) is available.
The O2 used by roots is replaced largely by molecular diffusion between soil
air and the aboveground atmosphere (Miller, 1986). Waterlogging, defined
as the soil saturated or nearly saturated with water, significantly reduces
oxygen availability to roots, causes the soil to become reduced, results in
the production of toxic compounds, and generally reduces root growth
(Ellis, 1979; Hoad et al., 2001). Cereals are generally more tolerant to
waterlogging than legumes. Hoad et al. (2001) reported that cereal roots
can survive short periods of waterlogging without adverse effects because
they can use the small quantities of oxygen dissolved in the soil water and
are capable of anaerobic metabolism when oxygen demand exceeds supply.
Oxygen requirements are higher for growing root tips than for the older
parts of the root system. The effect of waterlogging is least at low temperatures when more oxygen can be dissolved in the water and biological
activity is depressed (Hoad et al., 2001). The effect of waterlogging is
reduced with the application of chemical fertilizers, especially K, which
can improve the respiration capacity of roots (Fageria, 2009).
Most agricultural crops are mesophytes, which, for maximum growth,
require an environment that is neither too wet nor too dry (Fageria, 1992).
However, some plant species grow well under anaerobic conditions. It has
long been known that marsh plants or hydrophytes, such as rice, are
genetically adapted to grow in reduced soil environments (Horiguchi,
1995). Adaptation to waterlogging in hydrophytes is the result of their
unique ability to translocate O2 from the shoot to the root system. Ando
et al. (1983) reported that in rice oxygen absorbed from the atmosphere by
the shoots can be translocated to the root system and released into the
rhizosphere within 5 min. Generally, flooded rice roots have a reddish
brown color due to oxidation of the Fe ions by oxygen release from the
roots and their deposition on the root surface (Fageria et al., 2008a,b).
Flooding induces many changes in plant roots, of which formation of
aerenchyma (large interconnected intercellular spaces) is an important adaptive mechanism (Laan et al., 1989). Some species like corn (Z. mays L.)
develop aerenchyma as a response to flooding or anoxia (Armstrong and
Drew, 2002), a facultative development, while others develop them routinely regardless of the environmental conditions (constitutive development; Barlow, 2002; Zobel, 2005a). The development of aerenchyma in

275
rice (O. sativa L.), and other species and crops growing in wet soils, allows
the roots to grow into saturated soils and still extract nutrients (Zobel,
2005b). In corn, the leaf bases also form aerenchyma when submerged,
while the enclosed stem bearing aerenchymatous nodal (adventitious) roots
may elongate more rapidly ( Jackson, 1994). Aerenchyma formation in corn
roots appears to be triggered by the loss of tonoplast integrity (Campbell and
Drew, 1983), although cell wall disintegration is also involved at an early
stage, at least in rice ( Jackson, 1994; Webb and Jackson, 1986). Aerenchymatous roots are formed either by some cell wall separation and cell
collapse (lysigeny) or by cell separation without collapse (schizogeny).
Figure 11 shows cross sections of rice and wheat roots, and lysigenous and
schizogenous intercellular space. Both forms result in large longitudinal
channels in root cortices, and such structures enhance diffusion of atmospheric or photosynthetic oxygen from shoots to roots so that aerobic
respiration and growth can be maintained (Armstrong, 1979). Changes in
root morphology occur after flooding for both wetland and nonwetland
plant species. Flooding may also increase branching of roots, development
of new adventitious roots, and superficial rooting (Laan et al., 1989).
Cellular spaces that exist in roots facilitate oxygen diffusion; however,
the amount of aeration varies greatly among plant species. In the case of
certain marsh plants like rice, root cortical cells are arranged in columns, and
when channels form, the spaces become large and continuous, facilitating
diffusion. However, in the case of certain terrestrial plants, the cellular
arrangement is oblique and the spaces formed are small (Horiguchi, 1995).
There are marsh plants, however, that do not display columnar forms, while
some terrestrial plants do (Horiguchi, 1995). In the case of rice, large
cortical aerenchyma spaces develop schizogenously and lysigenously even
in well-aerated soils. Horiguchi (1995) reported that in rice and other
graminaceous plants, aerenchymous cells are well developed in both the
nodes and the internodes. The oxidizing capacity of roots of crop plants also
depends on the soil fertility level. Higher soil fertility improves the oxidation activity of rice roots (Horiguchi, 1995). Application of silicon to rice
A
Rice
B
Lysigenous
intercellular
space
Wheat
Schizogenous
intercellular
space
Figure 11 Cross section of a rice (A) and wheat (B) root showing lysigenous and
schizogenous intercellular spaces (from Horiguchi, 1995).

276
crops increases the diffusion of oxygen from the tops to the roots, strengthening oxidation activity and the deposition of ferrous iron on root surfaces,
and restricting the transport of excess iron to the top (Horiguchi, 1995).
Development of adventitious roots in some crops is related to plant
tolerance of soil waterlogging. Such roots developed by corn after one
day of flooding were short and porous enough to allow significant O2
movement from the shoots to the root meristems (Miller, 1986; Wenkert
et al., 1981). Despite this adventitious root formation and increased root
porosity, corn and most other crop plants soon perish under such conditions
(Miller, 1986).
11. Root Growth in Conservation

Tillage Systems
Conservation tillage is defined as any tillage sequence, the object of
which is to minimize or reduce loss of soil and water; operationally, a tillage
or tillage and planting combination that leaves a 30% or greater cover of
crop residue on the surface (Soil Science Society of America, 2008). The
benefits of conservation tillage are reducing soil erosion, conserving soil
moisture, avoiding fluctuations of soil temperature in the arable soil depth,
and reducing the costs of soil preparation. In addition, the use of conservation tillage is being encouraged as part of a strategy to reduce C loss from
agricultural soils (Kern and Johnson, 1993). Decomposition rates are generally slower in no-till than conventional tillage in which the decomposition
of soil organic matter is promoted by the stirring of the soil and alterations in
the soil microclimate (Parton et al., 1996). Holland and Coleman (1987)
suggested that C retention is increased in no-till because the surface residue
is primarily decomposed by fungi, which have higher assimilation efficiency
than the bacteria, which dominate the decomposition processes of residue
mixed into the soil. Gale and Cambardella (2000) reported that there was a
clear difference in the partitioning of surface residue and root-derived C
during decomposition and imply that the beneficial effects on no-till on soil
organic C accrual are primarily due to the increased retention of rootderived C in the soil. Knowledge of how plant root systems grow under
conservation tillage is important because this practice is widely adopted in
many countries around the world, most notably, in countries such as United
States, Brazil, Argentina, Canada, and Australia (Bolliger et al., 2006). Fortyfive percent of the total cultivated land in Brazil is now estimated to be
managed with conservation tillage, although in southern Brazil, this figure is
reported to exceed 80% (Bolliger et al., 2006).
Conservation tillage reduces soil erosion, conserves soil moisture, conserves energy, increases soil organic matter content, and consequently, soil

277
quality. However, conservation tillage may compact surface soil horizons and
may lead to poor root growth. Adverse effects of soil compaction on crop
production have been recognized for many years. Cato the Elder (234149 B.C.)
wrote that the first principle of good crop husbandry is to plow well and the
second principle is to plow again, presumably to provide a mellow (wellaerated) seedbed (Unger and Kaspar, 1994). Conservation tillage increases
soil bulk density (Martino and Shaykewich, 1994) which may inhibit root
growth in the upper part of the soil profile (Cannell, 1985; Lampurlanes
et al., 2001), reducing nutrient uptake and plant growth (Peterson et al.,
1984). Qin et al. (2004) reported that wheat root length density, mean root
diameter, and percentage of small-diameter roots were lower in no-till than
conventional tillage. Root diameter may be indicative of the effects of soil
strength on root growth and affects the utilization of nutrients in the soil.
Sidiras et al. (2001) reported thicker barley roots under conventional tillage
than under no-till. In general, bulk densities that impede root growth are
1.55 Mg m 3 for clay loams, 1.65 Mg m 3 for silt loams, 1.80 Mg m 3 for
sandy loams, and 1.85 Mg m 3 for loamy fine sands (Miller, 1986).
Tillage-induced differences in the soil nutrient status may also have a
significant impact on root growth (Qin et al., 2004). Conservation tillage
often results in the stratification of soil nutrients, especially of immobile
nutrients like P (Crozier et al., 1999; Holanda et al., 1998; Logan et al.,
1991). This produces greater soil fertility near the soil surface which, in
contrast to the effects of compaction described above, causes an increase in
root length density near the soil surface under conservation tillage (Cannell
and Hawes, 1994; Gregory, 1994). Frequently, root growth is greater from
0 to 5 cm in conservation and no-tillage systems than in conventional tillage
systems (Chan and Mead, 1992; Rasmussen, 1991; Wulfsohn et al., 1996).
Radial root swelling has been reported for lupins (Lupinus angustifolius
L.) grown in compacted soil (Atwell, 1989), for barley (H. vulgare L.) under
mechanical impedance (Wilson et al., 1977), and for mustard (Brassica sp.) in
drying soil (Vartanian, 1981). Studies on root elongation of cotton as a
function of soil strength and soil water content showed that root elongation
is more sensitive to soil strength than water content (Taylor and Ratliff,
1969). Root volumes were less affected than length, indicating an increase
in root diameter (Ball et al., 1994). Chassot et al. (2001) reported that
conservation tillage decreases soil temperature, and this may be the main
reason for the poor growth of the roots and shoots of corn seedlings
compared with conventional tillage under temperate humid conditions.
Considering the many advantages of conservation tillage compared to
conventional tillage, the effects of soil compaction produced by conservation tillage can be minimized. Unger and Kaspar (1994) reported that
growing deep-rooted crops in rotation will help avoid or alleviate compaction, improving root distribution and increasing rooting depth. These
authors also reported that weather conditions and soil moisture can enhance

278
or diminish the effects of compaction on root growth. Even if compaction

limits root growth, subsequent weather events may either enhance or
diminish the effect of the root limitation on crop growth. The first author
has conducted field experiments on an upland rice-dry bean rotation using
conservation tillage on Brazilian Oxisols. The upland rice root system was
adversely affected, and rice yields were low (around 2000 kg ha 1), about
half that expected in a field experiment with conventional soil preparation
and favorable environmental conditions. However, dry bean yields were
greater than 3000 kg ha 1 (Fageria, 2008; Fageria and Stone, 2004). Hence,
selecting the appropriate crop is important.
12. Mineral Nutrition Versus Root Growth

There are 17 nutrients essential for plant growth and development and
these are carbon (C), hydrogen (H), oxygen (O), nitrogen (N), phosphorus
(P), potassium (K), calcium (Ca), magnesium (Mg), sulfur (S), zinc (Zn),
copper (Cu), manganese (Mn), iron (Fe), boron (B), molybdenum (Mo),
chlorine (Cl), and nickel (Ni). Among these nutrients, plants take C, H, and
O from air and soil water and the remaining from the soil solution. About
95% of the plant weight is C, H, and O, and the remaining 5% is the other
14 nutrients. The C, H, O, N, P, K, Ca, Mg, and S are required by plants in
large amounts and for this reason are known as major or macronutrients.
The remaining nutrients, Zn, Cu, Mn, Fe, B, Mo, Cl, and Ni, are classified
as micronutrients because they are required in small amounts by plants
(Fageria et al., 2002). Among the micronutrients, Cl is also absorbed by
plants in large amounts, but it is needed in only small amounts, and Cl
deficiency is rarely observed in crop plants.
All the 17 nutrients are equally important for plant growth. If any of
these nutrients is limiting in the growth medium, plant growth will be
reduced. An example is given in Fig. 12, where at an adequate fertility level,
the relative dry weight of dry bean was 100%. When essential nutrients were
omitted from the soil or were not applied, growth was reduced, but the
amount of reduction depended on the nutrient. The impacts of deficiencies
on growth were in the order of P > Ca > Mg > N K > S among
macronutrients and B > Zn > Cu > Fe > Mn > Mo among micronutrients. Similarly, the influence of N, P, and K on shoot and root growth of
dry bean is shown in Fig. 13. Both shoot and root growth were significantly
reduced when N, P, or K were omitted from the growth medium.
Figures 14 and 15 show root growth of wheat, upland rice, and corn with
the addition of N P K and with the omission of these nutrients from
the growth medium. Root growth of these three crop species was decreased
in the absence of N, P, and K in the soil. A significant amount of variation

279
Relative shoot dry weight (%)
120
100
80
60
40
20
u
C
F
e
g
M
a
C
AF
Figure 12 Relative shoot dry weight of dry bean as influenced by adequate fertility
level (AFL) and other nutrients were not applied or omitted from the Oxisol.
120
Shoot
Root
Relative dry weight (%)
100
80
60
40
20
0
N+P+K
Figure 13 Relative dry weight of shoot and root of dry bean as influenced by N, P, and
K fertilization.
exists, both within and among crop species, in nutrient acquisition and use.
This variability reflects differences in root morphology and mechanisms that
either aid or prevent ion movement into the root (Gabelman et al., 1986).
12.1. Nitrogen
Nitrogen is one of the most yield-limiting nutrients in crop production in
most agroecosystems. Nitrogen plays numerous key roles in plant biochemistry, including being an essential constituent of enzymes, chlorophyll,

280
N+P+K
Wheat
Upland
rice
Figure 14 Root growth of wheat at the top and upland rice at the bottom grown at
N P K and N, P, and K levels.
N+P+K
Corn
Figure 15 Corn root growth at N P K and N, P, and K levels.
nucleic acids, storage proteins, cell walls, and a vast array of other cellular
components (Harper, 1994). Consequently, a deficiency of N in crop plants
profoundly influences plant growth, development, and yield. The recovery

281
of applied N with chemical fertilizers is lower than 50% for most annual
crops. The low recovery of N is associated with loss of this element
by leaching, denitrification, volatilization, incorporation into soil microorganisms, and soil erosion (Fageria and Baligar, 2005). To improve
the efficiency of N uptake and use by crop plants, root systems play an
important role.
Root morphology is influenced by the amount of N fertilizer applied
(Eghball et al., 1993) and factors such as temperature (Feil et al., 1991) and
soil mechanical impedance (Bengough and Mullins, 1990). Eghball et al.
(1993) showed that N stress in corn reduced root branching. Similarly,
Maizlish et al. (1980) showed greater root branching in corn with increasing
levels of applied fertilizer N. Costa et al. (2002) reported that greater root
length and root surface area were obtained at an N fertilizer rate of
128 kg N ha 1 compared with either the absence of fertilizer N or the
higher rate of 255 kg N ha 1. Nitrogen fertilizer improves root growth in
soils having low-OM content (Gregory, 1994; Robinson et al., 1994).
Nitrogen fertilization may increase crop root growth by increasing soil N
availability (Garton and Widders, 1990; Weston and Zandstra, 1989). Sainju
et al. (2001) observed that tomato (Lycopersicon esculentum Mill.) root growth
was greater with hairy vetch and crimson clover cover crops and
90 kg N ha 1 than with no cover crops or N fertilization. Nitrogen also
improves production of lateral roots and root hairs, as well as increasing
rooting depth and root length density deep in the profile (Hansson and
Andren, 1987). Hoad et al. (2001) reported that surface application of
nitrogen fertilizer increases root densities in the surface layers of the soil.
Nitrogen fertilization can increase root length and root surface area and
decrease root mass per unit area of corn (Anderson, 1987; Costa et al., 2002).
It is well known that roots tend to proliferate in nutrient-enriched soil zones
(Drew et al., 1973; Qin et al., 2005). Russell (1977) refers to this as a
compensatory response. The results of pot experiments showed that corn
roots were longer and thinner in zones that were rich in N (Durieux et al.,
1994; Zhang and Barber, 1992, 1993).
Root mass was less affected by N than root length, but the effect may
depend on the stage of maturity of the crop (Baligar et al., 1998). Higher
rates of application of N reduced root growth and depth of rooting in wheat
(Comfort et al., 1988) and reduced root:shoot ratio in rye (Brouwer, 1966).
In corn, the primary root system was 16% thicker when NH4N was
applied rather than NO3N. The NH4N treatment also increased the
diameters of lateral and first- and second-order nodal roots (Anderson et al.,
1991). Baligar et al. (1998) reported that relative dry weights of roots of rice,
dry bean, corn, and soybean were reduced by 38%, 56%, 35%, and 11%,
respectively, when N was omitted from a complete fertilizer. Nitrogen
deficiency also reduces branching and root hairs in cereals and legumes
(Baligar et al., 1998).

282
Figure 16 shows that increasing the N fertilizer rate from 0 to

400 mg kg 1 increased root dry weight of upland rice in a linear fashion
in Brazilian Oxisol, with the rate of N fertilizer explaining 59% of the
variation in root weight. Nitrogen fertilization increases production of
thinner roots with fine root hairs (personal visual observation).
Data in Table 3 show root length and root dry weight of 20 upland rice
genotypes grown on a Brazilian Oxisol. These two traits were significantly
influenced by N rate and genotype treatments. A significant interaction
between genotypes and N rates was found for root growth because some
genotypes were highly response to the N application while others were not.
Figures 1720 show root growth of upland rice cultivars/genotypes at two
N rates. Root growth of all the cultivars/genotypes improved with the
addition of N; however, there were differences among genotypes for root
development. Thus, selecting genotypes for N use efficiency may be an
important aspect of improving root growth and consequently, the yield of
upland rice in Brazilian Oxisols.
Nitrogen sources also affect root growth in upland rice (Fig. 21). Root
dry weight increased in a quadratic exponential fashion with the application
of N in the range of 0400 mg kg 1 of soil. In the case of urea, maximum
root dry weight was obtained with 281 mg N kg 1 of soil. Figures 22 and
23 show how the root growth of upland rice is affected by application of
urea and ammonium sulfate in the Brazilian Oxisol. Ammonium sulfate
produced more vigorous root systems, especially at higher N rates, than
urea, perhaps because upland rice is highly tolerant to soil acidity and
ammonium sulfate reduces soil pH more than urea. Fageria (2009) reported
that upland rice can tolerate up to 70% Al saturation in the soil. Fageria
2
Y =1.6407 + 0.0053X
R 2 = 0.5944**
50
100
150
200
250
300
350
400
Rate of N application (mg kg1)
Figure 16 Influence of nitrogen on root dry weight of upland rice. Values are averages
of 20 upland rice genotypes.

283
Table 3 Root length and root dry weight of 20 upland rice genotypes as influenced
by nitrogen fertilization
Root length (cm)
Root dry weight (g plant 1)
Genotype
0 mg N kg 1 300 mg N kg 1 0 mg N kg 1 300 mg N kg 1
BRA01506
BRA01596
BRA01600
BRA02535
BRA02601
BRA032033
BRA032039
BRA032048
BRA032051
BRA042094
BRA042156
BRA042160
BRA052015
BRA052023
BRA052033
BRA052034
BRA052045
BRA052053
BRS Primavera
BRS Sertaneja
Average
F-test
N rate (N)
Genotype (G)
NG
CV(%)
34.67ab
30.00b
27.00ab
31.67ab
28.00b
30.00a
43.00a
28.50b
30.67ab
30.33b
29.33b
29.67b
31.00ab
29.67b
30.33b
29.00b
28.67b
29.00b
29.33b
30.00b
30.49
NS
**
**
14.25
21.00cd
15.67d
25.33bcd
30.00abcd
32.67abc
28.00abcd
34.67abc
32.00abc
35.67ab
38.00ab
33.00abc
32.50abc
35.00abc
26.50abcd
31.33abc
37.67ab
37.67ab
36.50ab
36.33ab
40.33a
31.99
0.92a
0.87a
1.14a
1.33a
1.11a
1.12a
1.24a
1.05a
1.31a
1.78a
1.19a
1.51a
1.67a
1.48a
1.56a
1.78a
1.75a
1.65a
1.68a
1.45a
1.38
0.40f
0.45f
1.03ef
3.25abcd
3.73ab
2.41cd
3.77ab
3.62abc
2.82bcd
2.31de
2.84bcd
3.33abcd
2.83bcd
4.14a
3.38abcd
2.58bcd
3.66abc
2.72bcd
2.49bcd
2.68bcd
2.72
*
**
**
15.86
Source: Fageria (2011).

*,**Significant at the 5% and 1% probability levels, respectively. Means followed by the same letter in
the same column are not significant at the 5% probability level by Tukeys test.
(2009) also reported that rice growth was better in 10 mg L 1 Al than with
0 mg Al L 1 in nutrient solution. Brazilian rice cultivars (lowland as well as
upland) are highly tolerant to soil acidity (Fageria et al., 2004). Another
possible explanation is that ammonium sulfate has about 24% S, which may
improve root growth if the extractable soil S level is lower than 10 mg kg 1.
The timing of nitrogen application can also influence the root growth of
crop plants (Table 4). The treatment T3, which produced maximum grain
yield, also produced minimum root length; treatment T2 which produced
minimum grain yield produced maximum root length. There was a

BRS Sertaneja
N0
N300
Figure 17 Root growth of cultivar BRS Sertaneja at 0 and 300 mg N kg 1 soil
(Fageria, 2011).
BRS Primavera
N0
N300
Figure 18 Root growth of cultivar BRS Primavera at 0 and 300 mg N kg 1 soil
(Fageria, 2011).

285
BRA052053
N0
N300
Figure 19 Root growth of genotype BRA052053 at 0 and 300 mg N kg 1 soil

(Fageria, 2011).
negative association between root length and grain yield (Y 31.2041

0.2718X, R2 0.7396**). Root dry weight was significantly related to
grain yield (Y 7.6345 5.8030X 0.4081X2, R2 0.8747**;
Fageria, 2011). Figure 24 shows root growth of upland rice under different
N timing treatments. Root dry weight was least (treatment T2) when all N
fertilizer was applied at planting. Root and shoot yields were better in
treatments T3 and T4, when N was applied later in the growth cycle
(Fageria, 2011). Better root growth may be responsible for higher absorption of nutrients and water in the T3 and T4 treatments which resulted in
higher grain and straw yields.
The depth of N placement can influence NO3N distribution in the soil
and root growth of wheat (Sharma and Chaudhary, 1984). Root length
density decreased abruptly below the 15 cm depth when N was surface
applied, whereas root length density decreased more gradually below 15 cm
when N was placed at the 10 cm depth. Drew (1975) reported that barley
root weight increased in the zone of nutrient localization and decreased in
the deficient zone. Murphy and Zaurov (1994) reported that N fertilization
at the 5, 10, and 15 cm soil depths produced greater root mass than N
fertilization at 0 cm soil depth or surface fertilization.

286
BRA052045
N0
N300
Figure 20 Root growth of genotype BRA0522045 at 0 and 300 mg N kg 1 soil

(Fageria, 2011).
5
(NH4)2SO4 (Y ) = 1.9132EXP(0.00093X + 0.000031X
2)
R 2 = 0.5296**
3
2
CO(NH2)2 (Y ) = 1.1194+0.0135X 0.000024X 2
R 2 = 0.4122**
50
100
150
200
250
300
350
400
Nitrogen application rate (mg kg1)
Figure 21 Relationship between nitrogen application rate by ammonium sulfate and

urea and root dry weight of upland rice (Fageria et al. 2011).

287
CO(NH2)2
0N
50 N
300 N
Figure 22 Root growth of upland rice at 0 (left), 50 (center), and 300 (right)
mg N kg 1 soil supplied with urea.
The effects of two N rates on root length and root dry weight of six
tropical legume cover crops are showed in Table 5. Root length as well as
root dry weight were significantly increased with the addition of N fertilizer. Similarly, root length as well as root dry weight were significantly
different among cover crop species. Root growth of four legume cover
crops is shown in Figs. 2528. In all cases, root growth was better at the
higher N rate compared to low N rate.
12.2. Phosphorus
Phosphorus is one of the most yield-limiting nutrients in tropical highly
weathered soils (Fageria, 2009; Fageria and Baligar, 2003, 2008). The
deficiency of P in these soils may be related to the low natural level of this
element in these soils as well as to the immobilization of P in these soils
(Fageria and Baligar, 2008). Phosphate plays many roles in the physiology
and biochemistry of pants. It is a component of important compounds like

288
(NH4)2SO4
0N
300 N
400 N
Figure 23 Root growth of upland rice at 0 (left), 300 (center), and 400 (right)
mg N kg 1 soil with ammonium sulfate.
Table 4 Root length and root dry weight of upland rice as influenced by nitrogen
timing treatments
Nitrogen timing treatmenta
Root length (cm)
T1
T2
T3
T4
T5
Average
F-test
CV(%)
28.75b
35.00a
25.75b
27.00b
28.25b
28.95
**
8.7
4.21bc
2.56c
5.74ab
7.70a
4.98b
5.04
**
20.9
Source: Fageria (2011).

** Significant at the 1% probability level. Means followed by the same letter in the same column are not
significant at the 5% probability level by the Tukeys test.
a
T1 (1/2 N applied at sowing 1/2 applied at panicle initiation), T2 (total N applied at sowing),
T3 (1/3 N applied at sowing 1/3 N applied at active tillering 1/3 N applied at the panicle
initiation), T4 (1/2 N applied at initiation of tillering 1/2 N applied panicle initiation), and T5 (2/3
N applied at sowing 1/3 N applied at panicle initiation).

289
T1
T3
T2
T5
T4
Figure 24 Root growth of upland rice at harvest under N timing treatments.

T1 (1/2 N applied at sowing 1/2 applied at panicle initiation), T2 (total N applied
at sowing), T3 (1/3 N applied at sowing 1/3 N applied at active tillering 1/3 N
applied at the panicle initiation), T4 (1/2 N applied at initiation of tillering 1/2 N
applied panicle initiation), and T5 (2/3 N applied at sowing 1/3 N applied at panicle
initiation). Source: Fageria (2011).
Table 5 Influence of nitrogen rate on root length and root dry weight of six tropical
legume cover crops
Root length (cm)
Cover crops
0 mg N kg 1 100 mg N kg 1 0 mg N kg 1 100 mg N kg 1
Showy
crotalaria
Calopogonio
Pueraria
Pigeon pea
Lablab
Gray mucuna
bean
Average
F-test
N rate (N)
Cover crops
(C)
NC
CV(%)
22
29
0.48
0.77
28
23
30
27
28
34
24
34
32
31
0.30
0.16
0.73
0.74
2.01
0.46
0.15
0.48
1.72
1.84
26b
31a
0.74b
0.90a
**
**
**
**
NS
6.47
**
16.27
**, NS Significant at the 1% probability level and nonsignificant, respectively. Means in the same line for
each growth parameter, followed by the same letter are not significantly different by Tukeys test
at the 5% probability level.

Black mucuna bean
100 N
0N
Figure 25 Root growth of black mucuna bean tropical legume cover crop at 0 (left)
and 100 (right) mg N kg 1 soil.
0N
Gray mucuna bean

100 N
Figure 26 Root growth of gray mucuna bean tropical legume cover crop at 0 (left)
and 100 (right) mg N kg 1 soil.

291
Lablab
0N
100 N
Figure 27 Root growth of Lablab tropical cover crops at two N levels: 0 (left) and 100
(right) mg N kg 1 soil.
phospholipids, phosphorylated sugars and proteins, DNA (deoxyribonucleic acid), and RNA (ribonucleic acid). It is also a component of ATP
(adenosine 5-triphosphate), PEP (phosphoenolpyruvate), NADPH (nicotinamide adenine dinucleotide phosphate, reduced), and other biochemicals
that use the phosphate bond in energy utilization and storage (Blevins,
1994).
Phosphorus is a key nutrient essential for root development in highly
weathered tropical soils. Baligar et al. (1998) reported that P increased the
root weight of wheat, dry bean, and cowpea in a quadratic fashion with
increasing P rate from 0 to 200 mg kg 1 of soil. The regression equations
related to P rates versus root dry weight were Y 0.4019 0.094X
0.00031X2, R2 0.74* for wheat, Y 0.4813Exp. (0.019X 0.000071
X2), R2 0.63* for dry bean, and Y 0.7351 0.0232X 0.000073
X2, R2 0.80** for cowpea. Based on these regression equations, maximum root dry weight for wheat was achieved at 152 mg P kg 1, whereas
maximum root dry weight for common bean and cowpea was achieved at
134 and 159 mg P kg 1 soil, respectively. These results indicate that
increasing P levels increased root growth, but root growth was reduced at

292
Crotalaria spectabilis
0N
100 N
Figure 28 Root growth of Crotalaria spectabilis tropical legume cover crops at two N
levels: 0 (left) and 100 (right) mg N kg 1 of soil.
higher P levels, and the crops had different P requirements to achieve

maximum growth. Overall, the root growth of cereals and legume crops
was reduced if P was deficient. Most studies indicate that, within certain
limits, both root and shoot growth vary similarly as P level increases. Above
certain levels, further increases in P supply do not affect root or shoot
growth (Troughton, 1962). Fageria et al. (2006) reported that root dry
weight was reduced 62% in rice, 74% in common bean, 50% in corn, and
21% in soybean without added soil P, compared to adequate P in a Brazilian
Oxisol.
Fageria et al. (2011) studied the influence of phosphorus on root dry
weight and root length of 20 upland rice genotypes grown on a Brazilian
Oxisol. Phosphorus level and genotype interactions for root dry weight and
root length were significant, indicating different responses of genotypes to
varying P levels (Table 6). Root dry weight of 20 upland rice genotypes at
low P level varied from 2.00 to 5.68 g plant 1, with an average value of
3.41 g plant 1. At the high P level, root dry weight varied from 2.43 to
8.55 g plant 1, with an average value of 4.01 g plant 1. However, the effect

293
Table 6 Root dry weight and root length of 20 upland rice genotypes as influenced
by P levels
Genotype
BRA01506
BRA01596
BRA01600
BRA02535
BRA02601
BRA032033
BRA032039
BRA032048
BRA032051
BRA042094
BRA042156
BRA042160
BRA052015
BRA052023
BRA052033
BRA052034
BRA052045
BRA052053
BRS
Primavera
BRS
Sertaneja
Average
F-test
P level (P)
Genotype
(G)
PG
Root length (cm)
Low P
(25 mg kg 1)
High P
(200 mg kg 1)
Low P
(25 mg kg 1)
High P
(200 mg kg 1)
3.92ab
2.78ab
2.81ab
3.12ab
4.42ab
3.70ab
2.91ab
3.96ab
2.00b
2.82ab
2.50b
5.68a
3.91ab
4.69ab
2.23b
3.18ab
3.07ab
2.57ab
3.56ab
3.22c
2.73c
3.03c
4.30c
3.20c
3.62c
4.36c
3.91c
2.58c
3.92c
2.91c
8.32ab
2.98c
8.55a
2.43c
3.99c
3.08c
3.87c
5.21bc
26.00ab
35.67ab
36.00ab
28.67ab
31.33ab
23.00b
27.67ab
37.00a
36.00ab
30.00ab
29.00ab
32.67ab
27.00ab
29.67ab
27.00ab
31.00ab
38.33a
28.33ab
29.67ab
26.67a
28.00a
29.00a
33.33a
27.00a
29.67a
27.00a
33.33a
30.67a
27.33a
27.00a
33.00a
27.00a
34.33a
29.00a
24.67a
24.67a
23.67a
25.00a
4.36ab
3.92c
34.00ab
23.67a
3.41
4.01
30.9
28.20
NS
**
*
**
**
Source: Fageria et al. (2011).

*,**, NS Significant at the 5% and 1% probability level and nonsignificant, respectively. Means in the
same column followed by the same letter are not significantly different at the 5% probability
level by the Tukeys test.
of P level on root system dry weight was not significant. Root length varied
from 23.00 to 38.33 cm with an average value of 30.9 cm at low P level.
At high P level, root length varied from 23.67 to 34.33 cm, with an average
value of 28.20 cm. There was a significant 10% decrease in root length at the
high P level compared to the low P level. However, higher P level roots had
more fine hairs compared to lower P level (personal observations).

294
Figures 2931 show root growth of upland rice under different P rates.
Unlike the data cited above, root growth of all the cultivars/genotypes
increased with increasing P levels.
The first author studied the influence of P levels on the root growth of
14 tropical legume cover crops grown on a Brazilian Oxisol. Root dry
weight was significantly influenced by P; crop species and P crop species
interactions were significant (Fig. 32). The significant P crop species
interaction indicates significant variation in shoot dry weight with the
variation in P levels. At the lowest P level (0 mg kg 1), maximum root
dry weight of 0.77 g plant 1 was produced by white jack bean (Canavalia
ensiformis) and minimum root dry weight of 0.01 g plant 1 was produced by
crotalaria (Crotalaria mucronata) and pueraria (Pueraria phaseoloides). At the
medium P level (100 mg P kg 1), maximum root dry weight of
1.91 g plant 1 was produced by black mucuna bean (Mucuna cinereum)
and minimum root dry weight of 0.07 was produced by crotalaria (Crotalaria
0P
50 P
175 P
Figure 29 Root growth of upland rice under 0 (left), 50 (center), and 175 (right)
mg P kg 1 soil (Fageria and Stone, 1999).
0P
25 P
50 P
75 P 100 P 200 P
IAC 164
Figure 30 Root growth of upland rice cultivar IAC 164 at different P levels.

295
BRA 01596
25 P
200 P
Figure 31
Root growth of upland rice genotype BRA01596 at two P levels.
breviflora), with an average value of 0.63 g plant 1. At the highest P level

(200 mg P), maximum root dry weight of 1.42 g plant 1 was produced by
gray mucuna bean (Mucuna cinereum) and minimum root dry weight of
0.09 g plant 1 was produced by calopogonio (Calopogonium mucunoides)
and pueraria (Pueraria phaseoloides). The variation in root dry weight is
genetically controlled and also influenced by environmental variables, like
the supply of mineral nutrition (Baligar et al., 2001; Caradus, 1990; Fageria
et al., 2006).
Maximum root length of tropical legume cover crops varied from 15.5
to 36 cm at the low P level, from 20.5 to 50.33 cm at the medium P level
and 18.33 to 52.33 cm at the high P level (Fig. 33). Overall, root length also
increased with increasing P level. Figures 3436 show root growth of
tropical legume cover crops as influenced by P levels. The improvement
in root length by improved P nutrition has been reported by Fageria (2009)
in various crop species. Barber (1995), Fageria et al. (2006), Marschner
(1995), and Mengel et al. (2001) reported that mineral nutrition has tremendous effects on root growth, development, and function and, subsequently,
the ability of roots to absorb and translocate nutrients. These authors further
reported that mineral deficiencies induce considerable variations in the
growth and morphology of roots and such variations are strongly influenced
by plant species and genotypes.

296
1.6
a
200 mg P kg1
1.4
1.2
b
1.0
0.8
0.6
0.4
0.2
b
b
de
de
d
de
de
Root dry weight (g plant )
0
2.0
a
100 mg P kg1
1.8
1.6
1.4
1.2
1.0
bc
bc
bcd
0.8
cde
0.6
ef
0.4
0.2
ef
ef
df
0
0.8
a
0 mg P
0.6
kg1
0.4
c
cd
Figure 32
de
Canavalia ensiformis
Mucuna cinereum
Mucuna aterrima
e
Mucuna deeringiana
Crotalaria ochroleuca
Calopognium
mucunoides
de
cde
Dolichos lablab
Cajanus cajan (black)

Cajanus cajan
(mixed color)
de
Pueraria phaseoloides
Crotalaria spectablis
Crotalaria juncea
de
cd
0.2
Root dry weight of 14 tropical legume cover crops as influenced by P rates.
The influence of P fertilization on dry bean root dry weight was studied
by Fageria (1989b). Root dry weight of three bean genotypes increased
significantly in a quadratic fashion but differed from genotype to genotype
(Table 7). The variability in root dry weight due to P fertilization was about

297
70
200 mg P kg1
60
a
ab
ab
ab
50
40
30
c
cd
20
cd
d
c
cd
cd
cd
10
0
60
100 mg P kg1
50
bc
40
cde cde
30
20
def def
f
def
cde
ef
bcd
def
10
0
40
0 mg P kg1
ab
ab
de
de cde
cd
cd
Cajanus cajan
(mixed color)
cde
20
cd
Cajanus cajan (black)
30
cd
bc
de
Canavalia ensiformis
Mucuna cinereum
Mucuna aterrima
Mucuna deeringiana
Dolichos lablab
Pueraria phaseoloides
Calopogonium
mucunoides
Crotalaria ochroleuca
Crotalaria spectablis
Crotalaria juncea
10
Figure 33 Maximum root length of 14 tropical legume cover crops at three P levels.
52% in genotype Carioca, 35% in genotype CNF10, and 70% in genotype

CNF4856. Such information may contribute to the selection of cultivars
specific to soil type and management systems, resulting in increasing yields
on the soils of variable P fertility (Fageria, 1989b).

0P
200 P
Figure 34 Root growth of tropical legume cover crop Crotalaria mucronata at 0 and
200 mg P kg 1 of soil.
Gray mucuna bean

0P
200 P
Figure 35 Root growth of tropical legume cover crop gray mucuna bean at 0 (left) and
200 (right) mg P kg 1 of soil.

299
0P
200 P
Figure 36 Root growth of tropical legume cover crop Crotalaria breviflora at 0 and
200 mg P kg 1 of soil.
12.3. Potassium
Potassium plays an important role in the growth and development of plants,
including the root system. Many enzymes are activated in plants by potassium, and it is also required for photosynthesis, transport of photosynthate,
and protein synthesis (Blevins, 1994). Potassium plays a role in cell growth
following cell division by serving as a major component in cell turgor. It also
maintains ionic balance and electrical neutrality in plants. Crops that produce large quantities of protein per unit area of land require more K than
those that produce less protein (Blevins, 1994). Potassium also plays an
important role in opening and closing the stomata.
Tennant (1976) reported that potassium deficiency stops root growth
completely within 1012 days of planting in wheat. Inadequate K reduces
root growth and consequently, crop yields (Baligar et al., 1998). A deficiency of K in an Inceptisol reduced root growth by 23% in lowland rice, by
30% in dry bean, by 12% in corn, and by 11% in soybean (Baligar et al.,

300
Table 7 Influence of phosphorus fertilization on the root growth of three dry bean
genotypes at harvest
Root dry weight (g per 2 plants)
1
P rate (mg kg
Carioca
CNF10
CNF4856
0
0.29
0.45
0.23
25
1.13
1.79
1.38
50
1.66
1.28
1.83
75
1.31
1.29
1.52
100
1.34
1.60
1.59
125
1.62
1.33
1.78
150
1.31
1.75
1.82
175
1.18
1.29
1.48
200
1.79
1.50
1.67
Regression analysis
P rate versus Carioca (Y) 0.6459 0.0128X 0.000044X2,
R2 0.5179**
P rate versus CNF10 (Y) 0.8704 0.0109X 0.000042X2,
R2 0.3509*
P rate versus CNF4856 (Y) 0.6145 0.0193X 0.000075X2,
R2 0.6953**
Source: Adapted from Fageria (1989b).
*,** Significant at the 5% and 1% probability levels, respectively.
1998). On an Oxisol, a 35% lower root dry mass was observed in 13 corn
genotypes when K levels were 0 versus 200 mg kg 1 of soil (Baligar et al.,
1998). Data in Table 8 show that at 0 mg K kg 1 of soil root dry weight of
common bean (P. vulgaris L.) genotypes varied from 1.54 to 3.14 g per
3 plants, a variation of twofold. At the 200 mg K kg 1 level, root dry weight
varied from 1.50 to 2.30 g per 3 plants, a variation of 1.5-fold. Similarly,
maximum root length varied from 42 to 46 cm at low K level and 3244 cm
at higher K level. At the higher K level, there was a slight decrease in the
root length of all the genotypes, and the root weight of three genotypes also
decreased at the higher K level. However, at the higher K level, there were
more root hairs than at the low K level (visual observations).
There is widespread evidence for genotype diversity in the root characteristics of many crops in response to the environment and increasing
interest in using this diversity to improve agricultural production and
consequently, nutrient-use efficiency (Barber, 1994; Gregory, 1994).
Mullins et al. (1994) studied K placement effects on the root growth of
cotton grown on a fine sandy loam soil. Root density measurements taken
in-row showed that root growth at depths >20 cm was improved with inrow subsoil additions of K. Tupper (1992) also observed increased cotton
taproot length when K fertilizer was band-applied in the subsoils of

301
Table 8 Root dry weight and maximum root length of six common bean genotypes as
influenced by potassium levels applied to a Brazilian Oxisol
0 mg K kg 1
200 mg K kg 1
Maximum
Root dry weight root length
Genotypes (g per 3 plants) (cm)
Maximum
Root dry weight root length
(g per 3 plants) (cm)
Apore
Perola
Ruda
IAC
Carioca
Jalo
Precoce
Safira
Average
1.54
1.97
1.94
3.14
45
42
44
45
1.67
2.04
2.30
1.70
32
39
35
38
2.24
42
1.67
36
1.77
2.10
46
44
1.50
1.81
44
37
Source: Fageria et al. (2008a).

Values were determined at physiological maturity.
Mississippi soils with low soil-test K. On the other hand, Hallmark and
Barber (1984) and Yibrin et al. (1993) reported that localized applications of
K did not promote root growth. However, K has been shown to promote
the root growth of some vegetable crops (Zhao et al., 1991). Fageria (1992)
determined the root growth of rice grown in nutrient solution as well as in
an Oxisols at the stress and nonstress levels of K. At the stress levels of K, rice
root growth was reduced compared with nonstress levels. Alfalfa herbage
yield, root weight, and root total nonstructural carbohydrates increased
with increasing K fertilizer (Kitchen et al., 1990).
12.4. Calcium
Absolute Ca deficiency is difficult to identify on plants grown in acidic soils
(Kamprath and Foy, 1985). Most acidic soils contain adequate total Ca for
most plants, and Ca-deficiency symptoms are rarely observed in the field.
Only in highly leached, acidic, low-cation exchange soils (Oxisols and
Ultisols) would absolute deficiencies be likely to occur (Garrity et al.,
1983). Levels of Ca required for essential growth functions are so low as
to approach those of micronutrients. Hence, the major role of Ca in soils
and in plants is to exclude or detoxify other elements such as Al, Mn, and
heavy metals that might otherwise become toxic (Garrity et al., 1983).
Gonzalez-Erico et al. (1979) evaluated the maize response to deep
incorporation of limestone on an Oxisol. They reported that incorporation

302
of limestone to depths of 30 cm improved root growth and increased water

utilization and grain yield of maize. Similar results were obtained for maize
and cotton when limestone was incorporated to depths up to 45 cm (Doss
et al., 1979).
Suitable diagnostic indices for prediction of Ca limitations on root
growth are either Ca saturation of the effective CEC or Ca activity ratio
of the soil solution, which has been defined as the ratio of Ca activity to the
sum of the activities of Ca, Mg, K, and Na (Bruce et al., 1988). Values
corresponding to 90% relative root length (RRL) of soybean were 0.05 for
the Ca activity ratio and 11% for Ca saturation. Calcium activity and Ca
concentrations in soil solutions and exchangeable Ca were less useful for
diagnostic indices (Bruce et al., 1988). The root growth of soybean was
improved with the addition of 12 Mg lime per hectare compared to control
treatment (Fig. 37).
12.5. Magnesium
Magnesium is an essential macronutrient for plant growth. The most wellknown and important role of Mg is its occurrence in chlorophyll molecules.
In addition to this, Mg is required for many essential physiological reactions,
especially phosphorylation reactions (Mengel and Kirkby, 1978).
Fageria and Souza (1991) determined the effects of Mg levels on root
weights of rice, common bean, and cowpea grown in an Oxisols of Central
Brazil (Fig. 38). Dry weights of rice roots were higher at the lowest Mg
concentration compared with the highest soil Mg concentration. Initial
exchangeable Mg levels of surface soils were 0.1 cmol kg 1. They increased
to 0.3 cmol kg 1 within 3 days after liming and to 0.75 cmol kg 1 at harvest
time (33 days after sowing). The lack of growth responses to applications of
Mg indicated that this level of exchangeable Mg was adequate to meet Mg
0 Mg lime ha1
Figure 37
12 Mg lime ha1
Root growth of soybean at two lime rates.

303
3
Upland rice
2
Y = 1.0752 0.08852X
R 2 = 0.85*
Root dry weight (g/3 plants)
0
Common bean
4
3
2
Y = 2.4497exp(0.3009X 0.05047X 2)
R 2 = 0.88*
1
0
Cowpea
4
3
2
Y = 2.6658exp(0.3007X 0.05983X 2)
R 2 = 0.80**
1
0
0
Extractable Mg in soil (cmolc kg1)
Figure 38 Root dry weights of upland rice, common bean, and cowpea grown with
different Mg levels in an Oxisol (adapted from Fageria and Souza, 1991).
requirements of upland rice grown in this limed soil. Dry weight of roots of
common bean increased with Mg application up to 3 cmol kg 1 of soil.
Similarly, significant responses of cowpea root growth to soil Mg levels
were observed, and maximum root weight was achieved at
2.5 cmol Mg kg 1 of soil.
12.6. Sulfur
Sulfur has long been recognized as an essential element for plant growth and
development and classified as a macronutrient. Crop responses to applied
sulfur have been reported in a wide range of soils in many parts of the world
(Fageria, 2009). Sulfur plays many important roles in the growth and
development of plants. Fageria and Gheyi (1999) summarized important
functions of the sulfur in the plant. It is an important component of two
amino acids, cysteine and methionine, which are essential for protein
formation. Since animals cannot reduce sulfate, plants play a vital role in

304
supplying essential S-containing amino acids to them. Sulfur plays an

important role in enzyme activation. It promotes nodule formation in
legumes. Sulfur is necessary in chlorophyll formation, although it is not a
constituent of chlorophyll. Maturity of seeds and fruits is delayed in the
absence of adequate sulfur. Sulfur is required by the plants in the formations
of nitrogenase. It increases the crude protein content of forages and
improves the quality of cereals for milling and baking. Sulfur increases the
oil content of oilseed crops and increases winter hardiness in plants.
It increases drought tolerance in plants, controls certain soil-borne diseases,
and helps in the formation of glucosides that give characteristic odors and
flavors to onion, garlic, and mustard. Sulfur is necessary for the formation of
vitamins and synthesis of some hormones and glutathione, and it is involved
in oxidationreduction reactions. Sulfur improves tolerance to heavy metal
toxicity in plants, and it is a component of sulfur contain sulfolipids. Organic
sulfates may serve to enhance the water solubility of organic compounds,
which may be important in dealing with salinity stress, and fertilization with
sulfate decreases fungal diseases in many crops. Few studies have assessed the
impacts of sulfur on root growth and function; however, the effects of sulfur
on root growth may be similar to those of N. Zhao et al. (2008) reported
that S application increased the root number and root dry weight of soybean
compared to control treatment.
12.7. Micronutrients
Micronutrients have also been called minor or trace elements, indicating
that their required concentrations in plant tissues are small compared to the
macronutrients (Fageria et al., 2002; Mortvedt, 2000). Based on physicochemical properties, except B and Cl, the essential micronutrients are
metals. Even though micronutrients are required in small quantities by
field crops, their influence can be as great as that of macronutrients in
crop production. Micronutrients are normally constituents of prosthetic
groups that catalyze redox processes by electron transfer (such as with
the transition elements Cu, Fe, Mn, and Mo) and form enzymesubstrate
complexes by coupling enzymes with substrates (Fe and Zn) or enhance
enzyme reactions by influencing molecular configurations between
enzyme and substrate (Zn) (Fageria et al., 2002).
Micronutrient deficiencies in crop plants are widespread because of (i)
increased micronutrient demands from intensive cropping practices and
adaptation of high-yielding cultivars which may have higher micronutrient
demand, (ii) enhanced production of crops on marginal soils that contain
low levels of essential micronutrients, (iii) increased use of high-analysis
fertilizers with low amounts of micronutrients, (iv) decreased use of animal
manures, composts, and crop residues, (v) use of many soils that are inherently low in micronutrient reserves, (vi) use of liming in acid soils, and (vi)

305
involvement of natural and anthropogenic factors that limit adequate

supplies and create elemental imbalance (Fageria et al., 2002).
12.7.1. Zinc
Deficiency of Zn in crop production is spread worldwide (Alloway, 2008).
Graham (2008) reported that half of the worlds soils are intrinsically
deficient in Zn. Zinc deficiency in annual crops is reported in Brazil
(Fageria and Stone, 2008), Australia (Graham, 2008), India (Singh, 2008),
China (Zou et al., 2008), Turkey (Cakmak, 2008), Europe (Sinclair and
Edwards, 2008), USA (Brown, 2008), and Africa (Waals and Laker, 2008).
Micronutrient deficiencies are also a worldwide problem in human health
(Welch, 2008). Zinc deficiency is the highest priority among micronutrients for agriculture to address (Graham, 2008).
In the Brazilian Cerrado region, Zn deficiency is very common in
annual crops, especially upland rice and corn (Fageria, 2009). Figures 39
and 40 show Zn-deficiency symptoms in upland rice and corn grown on
Brazilian Oxisols. Data in Table 9 show that Zn application of up to
120 mg kg 1 improved the root growth of upland rice and wheat significantly. Similarly, Figs. 41 and 42 show improvement in the root growth of
soybean and dry dean, respectively, with the addition of Zn in Brazilian
Oxisols.
12.7.2. Boron
Boron deficiency is common for plants grown in arid, semiarid, and heavy
rainfall areas in calcareous, sandy, light textured, acid, and low-OM soils
(Gupta, 1993). Differences between B sufficiency and toxicity are narrow
Figure 39
Zinc deficiency in upland rice grown on Brazilian Oxisol.

306
Figure 40
Zinc deficiency in corn grown on Brazilian Oxisol.
Table 9 Root dry weight of upland rice and root length of corn as influenced by zinc
fertilization
Zn rate (mg kg 1)
Upland rice root dry weight

(g per 4 plants)a
Wheat maximum root

length (cm)a
0
5
10
20
40
80
120
R2
1.08
1.33
1.48
1.55
1.73
1.45
1.18
0.75**
55
54
52
55
57
58
52
0.24*
Source: Adapted from Fageria (2002a).

*,** Significant at the 5% and 1% probability levels, respectively.
a
Upland rice plants were harvested 6 weeks after sowing and corn plants were harvested 4 weeks after
sowing.
(Marschner, 1995), and soils supplied with high amounts of municipal

compost, sludge, and biosolids tend to accumulate high amounts of B,
which may result in B toxicity. Boron is essential for pollen germination
and pollen tube growth in crop plants (Blevins, 1994). Boron requirements

10
40
307
80 mg Zn kg1
Soja
Figure 41 Root growth of soybean at different Zn levels.
120
5
mg Zn kg1
Dry
bean
Figure 42
Root growth of dry bean at three Zn levels.
of dicots are generally higher than monocots (Fageria, 2000). Fageria (2000)
reported that maximum root growth of upland rice can be achieved with
the application of 0.4 mg B kg 1 soil, whereas maximum root growth of
dry bean required 1.9 mg B kg 1 of soil. Figure 43 shows that B requirements for root growth varied among the crop species. Application of
24 mg B kg 1 soil decreased the root dry weight of upland rice and corn
in greenhouse studies. However, application of B at lower rates to the same
soil increased the root dry weight in dry bean, soybean, and wheat (Fig. 43).

308
0.3
Upland rice
0.2
0.1
Y = 0.26810.00806X
R 2 = 0.83*
0.0
Corn
2
Root dry weight (g/4 plants)
1
Y = 2.0432exp (0.05957X+0.00105X 2)
R 2 = 0.93**
Common bean
Y = 1.3058exp (0.03466X0.00961X 2)
R 2 = 0.98**
2
1
0
Soybean
0.6
0.3
Y = 0.7473exp (0.0991X0.00652X 2)
R 2 = 0.91*
0.0
Wheat
0.3
0.2
Y = 0.2793exp (0.01993X0.00328X 2)
R 2 = 0.90*
0.1
0.0
10
15
20
Boron applied (mg kg1)
25
Figure 43 Root dry weights of upland rice, maize, common bean, soybean, and wheat
grown with different B levels on an Oxisols of central Brazil (adapted from Fageria,
2000).

309
12.7.3. Copper
Copper deficiency is often observed on plants grown in soils inherently low
in Cu (coarse-textured and calcareous soils) and in soils high in OM, where
Cu complexes with organic substances (Fageria et al., 2002). Higher-thannormal Cu supplies usually inhibit root growth more than shoot growth
(Lexmond and Vorm, 1981). Use of Cu-containing fungicides and anthelmintic compounds in agriculture has resulted in Cu toxicity in some plants,
but naturally occurring Cu toxicity is relatively uncommon (Welch et al.,
1991). Root dry weight of wheat and root length of dry bean were significantly increased by the application of copper fertilization (Table 10).
12.7.4. Iron
Iron deficiency is a worldwide problem and occurs in numerous crops
(Fageria et al., 2002; Marschner, 1995). Iron deficiency occurs not because
of Fe scarcity in soil or plants, but because various soil and plant factors affect
Fe availability, inhibit its absorption, or impair its metabolic use (Marschner,
1995; Welch et al., 1991). Plant species that commonly become Fe-deficient
are peanut (A. hypogaea L.), soybean (G. max L.), sorghum (Sorghum bicolor
L. Moench), and upland rice (O. sativa L.). Iron deficiency reduces root
growth (Table 11). Iron toxicity (indicated by leaf bronzing) can be serious
for production of crops in waterlogged soils. For wetland rice, Fe toxicity is
the second most severe yield-limiting nutrient disorder, and it has been
reported in South America, Asia, and Africa (Fageria et al., 2008b). Iron
toxicity decreases the root growth of lowland rice (Fageria et al., 2008b);
however, genotypic differences exist (Fig. 44).
Table 10 Root dry weight of wheat and root length of dry bean as
influenced by copper fertilization
Cu rate (mg kg 1) Wheat (g per 4 plants)a Dry bean (cm)a
0
2
4
8
16
32
64
96
R2
0.53
0.60
0.50
0.48
0.47
0.47
0.43
0.17
0.88**
25
30
27
28
28
24
30
14
0.42**
Source: Adapted from Fageria (2002a).

** Significant at the 1% probability levels.
a
Wheat and dry bean plants were harvested 5 weeks after sowing.

310
Table 11 Root dry weight of lowland rice as influenced by iron

concentration in nutrient solution
Fe concentration (mg L 1)
20 days age
60 days age
0.0
2.5
5.0
10.0
20.0
40.0
80.0
160.0
Average
0.03
0.09
0.13
0.10
0.08
0.11
0.07
0.02
0.08
0.64
1.19
1.05
1.07
1.18
1.24
0.84
0.12
0.92
Source: Fageria et al. (1981).
CNA810208
CNA808951
Fe100 mg L1
Figure 44 Root growth development of two lowland rice genotypes at 100 mg Fe L 1

in nutrient solution. Source: Fageria et al. (2008a,b).

311
12.7.5. Other micronutrients

In addition to zinc, boron, copper, and iron, the other micronutrients that
are essential for the growth of higher plants are manganese, molybdenum,
chlorine, and nickel. Manganese deficiency has been reported for plants
grown in the coarse-textured and poorly drained coastal plain soils of the
United States (Reuter et al., 1988) and in the soils of Central America,
Bolivia, and Brazil (Leon et al., 1985). In Europe, Mn deficiency has been
reported on plants grown in peaty (England and Denmark), coarse-textured
(Sweden and Denmark), coarse/fine textured (Netherlands), podzolic, and
brown forest (Scotland) soils (Welch et al., 1991). Manganese deficiency has
also been reported on plants grown in the semiarid regions of China, India,
southeast and western Australia, Congo, Ivory Coast, Nigeria, and other
western African countries (Fageria et al., 2002). Manganese toxicity on crop
plants grown in many parts of the world has been reported to be more
important than Mn deficiency (Foy, 1984; Welch et al., 1991).
Molybdenum is the least abundant of the micronutrients in the lithosphere (Mortvedt, 2000), and soil concentrations range from 0.2 to
5 mg kg 1 (mean of 2 mg kg 1). Mo deficiency usually occurs on plants
grown in the broad areas of well-drained acid soils and in soils formed from
parent materials low in Mo. In Australia, Mo deficiency occurs on crops
grown in soils derived from sedimentary rocks, basalts, and granites
(Anderson, 1970). Peaty, alkaline, and poorly drained soils commonly
have high Mo. Iron oxides adsorb more Mo than Al oxides (Fageria et al.,
2002), and clay mineralogy can affect Mo adsorption, in the order montmorillonite > illite > kaolinite (Goldberg, 1993). Hydrous ferric oxides or
ferric oxide molybdate complexes and insoluble ferric molybdates may form
in well-aerated soils so that Mo solubility and availability to plants is low
(Welch et al., 1991). In poorly drained soils, the formation of soluble ferrous
molybdates or molybdites may lead to high Mo availability to plants. Plants
grown in high Mo soils of the intermountain valleys of western United
States have been reported to accumulate high Mo which has induced
molybdenosis (Cu deficiency) in cattle (Welch et al., 1991).
Chloride is essential to higher plants and is required for the watersplitting reactions in photosystem II (Kelley and Izawa, 1978). Nickel has
been shown to be essential for soybean (Eskew et al., 1983), and it is known
to be a constituent of urease (Blevins, 1994; Klucas et al., 1983). Since urea is
a widely used fertilizer in crop plants worldwide, Ni nutrition could be
important (Blevins, 1994). Information on the influence of these micronutrients on the root growth of crop plants is not available, and this aspect is
not discussed here.

312
13. Management Strategies for Maximizing

Root Systems
Root growth of crop plants can be improved by adopting management practices that modify soils to fit crops, as well as to modify the plants to
fit the soil. Both of these strategies can be combined to get maximum
economic results.
13.1. Soil management

Soil management practices that can improve root growth of crop plants
include liming acid soils, use of gypsum, maintenance of organic matter,
use of adequate rates, sources and methods of fertilizer application, and deep
plowing.
13.1.1. Liming acid soils
Liming is the most common and effective practice to reduce soil acidity.
Lime requirements of crops grown on acid soils are determined by the quality
of liming material, status of soil fertility, crop species and cultivar within
species, crop management practices, and economic considerations. Soil pH,
base saturation, and aluminum saturation are important acidity indices that
are used to determine liming. Liming improves soil pH, Ca, and Mg contents
and reduces Al concentrations in the soil solution. In addition, liming
improves beneficial microbe populations in the soil. Furthermore, liming
improves P concentration in the soil solution by reducing P immobilization
by Fe and Al in acid soils (Fageria and Baligar, 2008). All these beneficial
effects of liming improve the root growth of crop plants. Nurlaeny et al.
(1996) found that liming increased shoot dry weight, total root length, and
mycorrhizal colonization of roots in soybean and corn grown on tropical acid
soils. Gonzalez (1976) reported that incorporation of lime to a 30 cm soil
depth allowed the corn roots to penetrate and use stored water throughout
the lime layer. Data in Table 12 show that liming increases soil pH and
consequently, the root dry weight of dry bean grown on a Brazilian Oxisol.
Similarly, Figs. 45 and 46 show how the root growth of dry bean and soybean
is influenced by soil pH. The root growth of both the legumes was significantly influenced by increasing soil pH. Dry bean root growth was maximum
at pH 5.9 and soybean produced vigorous root systems at pH 6.4.
13.1.2. Use of gypsum
Gypsum (CaSO4 2H2O) or phosphogypsum (e.g., byproducts of phosphoric acid manufacturing processes) applications are used to leach Ca deeper
into soil profiles where Ca can replace Al on cation exchange complexes.

313
Table 12 Influence of soil pH on root growth of dry bean grown on

a Brazilian Oxisol
Soil pH in H2O
Root dry weight (g per 4 plants)
4.1
4.7
5.3
5.9
6.6
7.0
R2
1.90
3.58
4.67
5.40
4.73
3.80
0.99**
Source: Adapted from Fageria (2002b).

** Significant at the 1% probability level.
pH 4.9 pH 5.9
pH 6.4 pH 6.7 pH 7.0
Dry
bean
Lowland soil
Figure 45 Root growth of dry bean grown on a Brazilian lowland soil (Inceptisol;
Fageria and Stone, 1999).
Much of the Al displaced by Ca can be leached from the root zones. This
practice works well in sandy soils or Oxisols with clay loam aggregates
which behave hydrologically like sands (Foy, 1992). Poor root growth of
crop plants has been frequently observed in highly weathered acid soils in
various countries (Alcordo and Rechcigl, 1993). The chemical factor identified as most responsible for poor root growth is excess soluble Al (Alcordo
and Rechcigl, 1993; Foy, 1992). Excess Al3 has been reported to inhibit
root growth by binding to the PO4 portion of DNA in the root cell nuclei,
reducing template activity and thus cell division (Matsumoto and
Morimura, 1980). In legumes, it has been shown to impair the growth of
root hairs and rhizobia, reducing root nodule initiation and function
(Munns and Franco, 1982). Excess Al may also adversely affect the root
and overall plant growth in nonphytotoxic ways by competing with Ca and

314
Lowland soil
pH 4.9
pH 5.9 pH 6.4
Soybean
pH 6.7 pH 7.0
Figure 46 Root growth of soybean in a Brazilian lowland soil (Inceptisol) at different

pH.
Mg for uptake by plants (Rengel and Robinson, 1989). The use of gypsum
can neutralize subsoil acidity by leaching CaSO4 and forming AlSO4
which is not toxic for root growth (Alva and Sumner, 1989). Ritchey
et al. (1980) reported that application of gypsum increased subsoil Ca and
Mg while decreasing Al and improving the root growth of corn in Brazilian
Oxisol.
13.1.3. Maintenance of adequate amounts of organic matter
The benefits of organic matter addition to soils include improving nutrient
cycling and availability to plants through direct additions as well as through
modification in soil physical and biological properties. The complementary
use of organic manures and chemical fertilizers has proved to be the best soil
fertility management strategy in the tropics (Fageria and Baligar, 2005).
Enhanced soil organic matter increases soil aggregation and water-holding
capacity, provides an additional source of nutrients, and reduces P fixation,
toxicities of Al and Mn, and leaching of nutrients (Baligar and Fageria,
1999). Build-up of organic matter through additions of crop and animal
residues increases the population and species diversity of microorganisms
and their associated enzyme activities and respiration rates (Fageria, 2002c).
The use of organic compost may result in a soil that has greater capacity to
resist the spread of plant pathogenic organisms. The improvement in the
overall soil quality may produce more vigorous root systems and higher
crop yields (Fageria, 2002c).

315
13.1.4. Use of adequate nutrient rates, sources, and methods of

application
Use of adequate rates and effective sources of nutrients are important
management practices to improve crop yield and plant root systems.
In addition, appropriate methods of fertilizer application are also important
for the development of vigorous root systems. Immobile nutrients like P
and K should be applied in bands to improve their uptake by plant roots.
13.1.5. Deep plowing
Deep plowing improves soil conditions for root growth by breaking compacted layers that roots cannot readily penetrate. If water tables are near the
soil surfaces, drainage can also be useful. When depths to root-restricting
hardpans are relatively shallow (<0.25 m), chisel plowing can be effective
for disrupting compacted layers.
13.1.6. Mulching, greater sowing depth, and sowing larger seeds
Root growth may be manipulated through cultural practices like mulching,
which can affect soil temperatures. Warmer soil temperatures generally
produce larger root systems. In addition, crop residues on soil surfaces
decrease soil evaporation and improve water-use efficiency. Greater sowing
depths may decrease seedling emergence rate, but it can also increase the
survival of emergent seedlings by increasing water availability. Larger seeds
generally produce seedlings with more extensive root systems. Sowing good
quality seeds could also improve root systems.
13.1.7. Leaching salts from soil profiles
Leaching may be required to prevent harmful accumulation of salts in the
root zone.. Steady-state leaching (Lr) requirements may be estimated
(Reichman and Trooien, 1993) as
Lr
Dd ECa
;
Da ECd
where Dd and Da are depths of drainage water and applied water, respectively, and ECa and ECd are electrical conductivity of applied water and
drainage water, respectively.
13.1.8. Integrated cropping systems and pest management
Reduced tillage, N side-dressing, and early planting can be included in
integrated pest management programs with no risk of increasing potential
for root damage from western maize rootworm (Roth et al., 1995). Miltner
et al. (1991) reported that cyst nematodes suppressed soybean root growth
on susceptible cultivars, whereas root growth of tolerant cultivars was

316
stimulated by the presence of soybean cyst nematodes. The use of tolerant

or nutrient-use-efficient cultivars may be an important practice to improve
root growth in stress environments. Stress-tolerant genotypes are being
identified and bred worldwide to solve some of the most difficult problems
of soil fertility such as subsoil acidity, salinity, low plant availability of Fe in
calcareous soils, and low P availability in acidic soils (Foy, 1992). Rhizobial
species/strains differ markedly in tolerance to low pH, and toxicities of Al
and Mn in tropical soils have been identified. The use of improved strains
can improve N2 fixation and the root growth of legumes grown in acidic
soils.
13.2. Plant management

Plant management is another important strategy in improving root systems
of crop plants. Adequate plant density and spacing can improve the root
growth of crop plants.
13.2.1. Genetic variability
An important plant management strategy is to exploit root system genetic
variability of crop species or cultivars within species. Plant genetic variability
can be defined as the heritable characters of a particular crop species or
cultivar that produce differences in the growth or production among
species, or cultivars of the same species, under favorable or unfavorable
growth conditions (Fageria, 1989a). Cultivar differences in root size are
quite common and have been related to differences in nutrient uptake
(Baligar et al., 1998; Caradus, 1990; Fageria et al., 2006). Differences
between white clover (T. repens L) populations and cultivars in P uptake
per plant at low levels of P have been related to differences in root size and
absolute growth rate (Caradus and Snaydon, 1986). There is widespread
evidence for genotype diversity in the root characteristics of many crops in
response to environment and increasing interest in using this diversity to
improve agricultural production and consequently, nutrient use efficiency
(Barber, 1994; Gregory, 1994). Mineral deficiency and toxicity, mechanical
impedance, moisture stress, oxygen stress, and temperature have tremendous effects on root growth, development, and function and, subsequently,
the ability of roots to absorb and translocate nutrients (Arkin and Taylor,
1981; Baligar et al., 1998; Barber, 1995; Marschner, 1995; Mengel et al.,
2001). Mineral deficiency induces considerable variations in growth and
morphology of roots, and such variations are strongly influenced by plant
species and genotypes. Overall, the growth of the main axis is little affected
by nutrient deficiency, but growth of lateral branches and their elongation
rates appear to be substantially reduced. Baligar et al. (1998) summarized the
effects of various essential elements as follows: nitrogen deficiency increases
root hair length, increases or has no effect on root hair density, and reduces

317
branching. Phosphorus deficiency increases the overall growth of roots and

root hair length, increases the number of second-order laterals, and either
increases or does not affect root hair density. K and Ca deficiencies reduce
root growth; however, high Mg levels reduce the dry mass of roots. The
effects of these nutrient stress factors on nutrient-use efficiency have not
been well explored. Baligar et al. (1998) states that low pH reduces root
mass, length, and root hair formation. In alkaline soils, ammonium toxicity
causes severe root inhibition and in general, salinity leads to reduction in the
mass and length of roots and dieback of laterals.
14. Conclusions
The study of plant roots is one of the most promising, but least
explored, areas of research related to mineral nutrition. Roots are poorly
studied primarily because of their physical location and growth habit.
Understanding plant root growth is important for improving productivity
of annual crops in agroecosystems. Root systems are important plant organs
because they absorb water and nutrients and also provide mechanical
support to the plant. In addition, roots synthesize growth substances and
hormones such as cytokinins that are important in leaf function and possibly, grain development. In plant growth analysis, the role of roots is
generally ignored due to the difficulty in getting accurate root growth
data under field condition. The principal structure of the root system
includes four developmentally distinct classes of roots. These are taproots,
basal roots, lateral roots, and shootborn roots. Current evidence suggests
that the four root classes that make up the primary and secondary root
systems are physiologically distinct from each other. Rooting depth and
spreading capacity are important traits for uptake of water and nutrients.
Root growth varies among plant species and cultivars within species and can
be modified by environmental factors. Genotypic differences in root growth
among crop species and genotypes of the same species under similar and
variable environmental conditions are now well demonstrated, and the
possibility of developing genotypes of desirable root systems to soil properties offers exciting prospects for the future. Root number, maximum length,
and root dry weight increase with increasing levels of macro- and micronutrients in the soil to a point beyond which root growth is suppressed.
Plant roots together with their associated bacteria and fungi play an important role in the formation, maintenance, and turnover of soil aggregates.
Currently, the techniques available to measure root systems are laborious
and time consuming, and this limits their use in plant physiological research.
Hence, it is necessary to develop root measurement techniques that are
simple, cheap, and less time consuming.

318
ACKNOWLEDGMENT
The authors thank Dr. Charles Allan Jones, Professor, Texas-A&M University, Texas
AgriLife Research and Extension Service, Dallas, Texas for their peer review of the chapter
and for giving valuable suggestions for its improvement.
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Mineral Nutrition On Root Growth

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The Role of Mineral Nutrition

2011 Elsevier Inc.

Author's personal copy

N. K. Fageria and A. Moreira

knowledge of factors that affect root development is fundamental to improving

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over whole-plant growth and development by controlling the uptake of

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N. K. Fageria and A. Moreira

exudates can influence microbial activity and the subsequent metabolism of

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properties (Fageria, 2002c, 2009; Klepper, 1992; Merrill et al., 1996;

2. Root-Induced Changes in the Rhizosphere

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N. K. Fageria and A. Moreira

Stele (xylem, phloem)

Sloughed root cap cell

exudates), high-molecular weight gelatinous materials (mucilage), and

3. Root Systems of Cereals and Legumes

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N. K. Fageria and A. Moreira

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more water than smaller diameter seminal roots. Adventitious roots

Figure 3 Root system of upland rice seedlings (Fageria, 2007).

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N. K. Fageria and A. Moreira

Root system of dry bean seedling (Fageria and Santos, 2008).

4. Contribution of Root Systems to Total

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N. K. Fageria and A. Moreira

300 mg N kg 1 soil

Root dry weight

Alfisols in Africa. Similarly, Ae et al. (1990) reported that pigeon pea as a

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The Role of Mineral Nutrition on Root Growth of Crop Plants

0 mg P kg 1 100 mg P kg 1 200 mg P kg 1

Root dry weight

5. Rooting Depth and Root Distribution

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N. K. Fageria and A. Moreira

Martino and Shaykewich (1994) reported that the proportion of roots

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cultivars to evaluate rooting depths in an Oxisol showed that 70% of the

6. Root Growth as a Function of Plant Age

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N. K. Fageria and A. Moreira

Root dry weight

Y = 2.4581 + 0.0952X + 0.00098X 2

Plant age (days after sowing)

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The Role of Mineral Nutrition on Root Growth of Crop Plants

Root dry weight

Y = 6.0778 + 0.7862X 0.0049X 2)

Shoot dry weight

Plant age (days after sowing)

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N. K. Fageria and A. Moreira

achieved at 80 days after sowing. However, root dry weight increased