Int. J. Biosci.

2014
International Journal of Biosciences | IJB |
ISSN: 2220-6655 (Print) 2222-5234 (Online)
http://www.innspub.net
Vol. 5, No. 7, p. 248-259, 2014

RESEARCH PAPER

OPEN ACCESS

Localization of quantitative trait loci (QTLs) controlling

drought tolerance in Barley
Leila Mehravaran1, Baratali Fakheri1, Javad Sharifi-Rad2, 3*
1

Department of Plant Breeding and Biotechnology, Faculty of Agriculture, university of Zabol,

Zabol, Iran
2

Zabol Medicinal Plants Research Center, Zabol University of Medical Sciences, Zabol, Iran

Department of Pharmacognosy, Faculty of Pharmacy, Zabol University of Medical Sciences, Zabol,

Iran
Key words: QTL, drought, tolerance, barley, agronomical traits.

http://dx.doi.org/10.12692/ijb/5.7.248-259

Article published on October 15, 2014

Abstract
In this study, quantitative trait loci (QTL) were mapped in 72 doubled-haploid lines derived from a cross
between Steptoe and Morex. They were planted in two augment designs with 3 replications at the research
farm of Agriculture Faculty of Zabol University were investigated under irrigated and dry conditions. In order to
easiness in marker assisted selection, number of QTLs ,their genomic location, additive effects and the nearest
markers were identified for agronomic traits and grain yield. There were significant differences among the lines
for all studied traits. Maximum correlation observed between grain yield and number of spike (r=0.696(. QTL
analysis was carried out using genetic linkage map derived from 327 molecular marker of RFLP and QTL
cartographer software with composite interval mapping method. In general we found 40 QTLs for the traits (9
QTLs in normal condition, 18 QTL in stress condition and 13 QTL in the mean of these two conditions).
Phenotypic variations that were explained by these QTLs changed from 11.97 to 64.49. The highest and lowest
phenotypic variations were related to length spike and grain yield QTLs in stress conditions, respectively. LOD
scores ranged from 2.5 to 8.6 the lowest and highest LOD scores were attained for the QTLs of Thousand grain
weight (Qtgw6Hn) in normal condition and length spike (Qspl3H) in stress condition. One QTL co-locations
confirmed the observed correlations among the traits. Therefore through marker and-assisted selection (MAS) in
this population would be unlimited.
* Corresponding

Author: Javad Sharifi-Rad Javad.sharifirad@gmail.com

248 Mehravaran et al.

Int. J. Biosci.

2014

Introduction

variance described by these QTLs changing from 11.9

Mainly agronomical important characters inherited

to 61.1%. Favorable QTL-alleles are useful as a

quantitatively and are known to be influenced by

breeding resource after they have been fixed in closely

environmental factors. The major goals for plant

isogenic lines. However, these favorable QTLs often

breeders are to improve genotypes with a high yield

lose their effects after purified into elite lines (Pillen

potential and the capacity to maintain yield across

et al., 2003).

environments. With the evolution of molecular

markers, breeders have a completing apparatus to

Several authors have concentrated their attempts on

traditional selection, and markers joined to the

the dissection of drought tolerance in traits that have

variation in a trait interesting could be used to help

a positive influence on yield permanence (Acevedo,

breeding programs. However, the recognition of

1987; Monneveux, 1991). Osmotic adaptation and

related markers linked to the variation of yield

carbon isotope discrimination are such traits (Morgan

components and permanence is difficult and time-

1983; Farquhar and Richard 1984; Blum 1988;

consuming. However, multiple studies reported

Craufurd et al., 1991; Acevedo 1993; Merah et al.,

quantitative trait loci (QTLs) for such traits in barley

2001). Investigation Teulat et al. (1998, 2001) traits

(Hayes et al., 1993; Thomas et al., 1995; Kjaer and

related to plant water status and OA were measured

Jensen 1996; Mather et al., 1997; Yin et al. 1999; Zhu

in controlled situations and QTLs recognized at two

et al. 1999).

soil-moisture contents. Experiments performed under

controlled conditions present benefit because the

The polygenic nature of numerous morphological

traits

characters has been described in many studies

environmental fluctuations (Price and Courtois,

(Mohammadi et al., 2005; Peighambari et al., 2005),

1999). They are particularly fitted for traits such as

which makes genetic inheritance complicated and

OA where measurements are troublesome and time-

difficult to use the molecular markers. Interval

consuming

mapping is a powerful approach that permits the

measurements

identification and genetic mapping of loci controlling

management, and timing of the water stress). On the

complex traits like grain yield and its components,

other hand, field experiments represent realistic

having

breeding.

situations (Price and Courtois, 1999). In such a

Knowledge of numbers and effects of quantitative

background, the co-location of QTLs for drought

trait loci (QTLs) can help breeders to understand the

tolerance-related traits such as OA or relative water

genetic control of these traits and design more

content, measured in controlled situations, and of

efficient selection strategies (Broman et al., 1999).

QTLs for agronomic traits, measured in various

A few compared QTLs detected under water-stressed

Mediterranean field situations, would reinforce the

and irrigated conditions in cereals (Ribaut et al., 1996

value of such criteria. actually, once genetic markers

in corn and Lafitte and Courtois 1999 in rice, as

associated with field performance are identified, it

examples) and, except the study by Baum et al.

will become feasibility to study which adaptive traits

(1996). Various reports on the implementation of the

co-segregate with yield or yield components. The

QTL strategy are available for barley (Backes et al.,

comparison between loci included in these complex

1995; Marquez-Cedillo et al., 2001; Pillen et al.,

traits, as well as the study of their relationships and

2003; Mohammadi et al., 2005; Peighambari et al.,

their genetic basis could help in the identification of

2005) in which favorable exotic QTL alleles for

the pertinent drought tolerance criteria and of the key

important agronomic traits have been recognized. For

chromosomal regions involved. Favorable alleles at

example, Peighambari et al. (2005) by using 72

the closed markers could then be selected to improve

doubled haploid (DH) barley lines identified twenty-

the yield performance of barley in the Mediterranean

three QTLs controlling various traits. Phenotypic

area. The objectives of this work was to identify QTLs,

great

importance

in

249 Mehravaran et al.

plant

are

evaluated

in

theoretically

field

with

conditions

concurrently,

importance

limited

(multiple
of

the

Int. J. Biosci.

2014

their effects and locations on linkage groups for 1000

the North American Barley Genome Mapping Progect

grain weight, spike length, plant height, awn length,

from Steptoe/Morex F1s, and used for mapping of

number of grains per spike, number of spikes and

forage quality traits. This map comprises 327 RFLP

grain yield using SteptoeMorex doubled haploid

markers with and average density of 3.75 cM. QTL

(DH) lines of barley.

analysis

was

environment

conducted
and

for

separately

combining

for

each

environments.

Materials and methods

Analysis was performed using WinQTL cartographer

Seventy-two doubled haploid lines from the cross of

2.5. A series of 1000 permutation were run to

Steptoe (CI15229)Morex (CI15773) together with

determine the experiment-was significance level at

both parents and 4 control lines, were utilized to

P=0.05 of logarithm of the odds ratio (LOD) for the

determine

were

traits. The genome was scanned at 2 cM intervals and

developed through a modified Hordeum bulbosum

the window size was set at 10 cM. Cofactors were

technique, as described by Chen and Hayes, by the

chosen using the forward-backward method of

Oregon state University Barley Breeding Program and

stepwise regression. The percentage of phenotypic

were Kindly provided by Hayes (Department of Crop

variance explained by a specific QTL value (R2) was

and Soil Science, Oregon State University, Corvallis,

taken as the peak QTL position as determined by

USA). The experiment was carried out in Zabol

WinQTL cartographer 2.5. The LOD peaks were

University Agricultural Research Station, in 2013

considered to indicate the most likely position of QTL

year, in two augment designs with 3 replications

effects. 95% confidence intervals were calculated by

under two normal irrigation and drought stress

1000-bootstrap re-sampling (Lebreton et al., 1998),

conditions. Planting for both conditions were as

as proposed in the WinQTL cartographer 2.5 package.

drought

tolerance.

The

DHs

sowing in irrigated land. Irrigation methods were as

ridge planting and in total period of growth 6 stage,

Results and discussion

time observed dry visual in the ground. In stress

Analysis of variances of the 72 doubled haploid lines

conditions, point evaluate drought tolerance, once

and their parents (Steptoe and Morex) showed a

irrigation carried out and it even in during grain

highly significant (p<0.01) genotype effect for all

filling period, that field due to excessive dry have the

traits (Table 1). Environment effect on length awn,

loss. Measured Traits include: length of spike, length

plant height, thousand-grain weight, Number of

of awn, plant height, thousand grain-weights, number

grains per ear and grain yield was highly significant

of tiller, number of spike, number of grain in spike

(p<0.01) and for other traits was significant (p<0.05).

and grain yield.

Genotype environment (G E) interaction was also

highly significant (p<0.01) for the thousand-grain

Analysis of variance (ANOVA) was performed using

weight and Number of tillers in plant and significant

proc ANOVA procedure in SAS (SAS Inst. Inc., Cary,

(p<0.05) for Number of spike in plant (Table 1).

NC). Heritability of entry was estimated as narrow-

Traits of length spike, length awn, plant height,

sense heritability by:

Number of grains and grain yield werent affected

h2=

genotype environment interaction. Previous studies

1- (MSGS / MSG).

have also reported significant genotype and genotype

Where MSGS is the variance of genotype. Simple

environment interaction effect for different traits in

correlation analysis was carried out for all studied

this population. Hayes et al. (1994) and Gibson et al.

characteristics using entry means and proc CORR

(1994)

procedure in SAS.

population for grain quality traits. Morex showed

reported

alike

fluctuation

within

this

higher values for traits such as plant height,

A molecular marker linkage map (Current at

thousand-grain

http://barleygenomics.wsu.edu/) was developed by

compared with Steptoe, And for other traits showed

250 Mehravaran et al.

weight

and

grain

yield,

when

Int. J. Biosci.

2014

lower values (Table 2). The differences between the

transgrasive genotypes over each parent indicated the

means of parents were not significant for all the

wide spectrum variation of studied traits in offspring.

studied traits, indicating that variation between the

The phenomenon of transgressive variation could

parents for these traits were narrow (Table 2). The

be interpreted as favorable alleles being dispersed

difference between the mean of doubled haploids (

DHs)

and their midparent was not significant for all the

traits (P>0.05), indicating that the 72 DHLs in this

study are representative of the total possible DHLs
from the cross Steptoe/Morex and that the studied
traits are mostly controlled by additive gene effect.
Phenotypic distribution of DH population was
normal. Phenotypic

means

of parents showed

variation for most traits and fell within the ranges

exhibited by progeny individuals (Table 2), providing
evidence

for

directions

transgrecive

(positive

and

segregation

in

negative).

both

Positive

transgrasive means that a majority of DH lines were

superior

to

the

parental

lines

and

negative

transgrasive means that a majority of DH lines was

inferior to the parental lines. The best DH when
compared with the best parent showed higher values
for all studied traits, indicating Positive transgrasive
for these traits. For all studied traits, except
thousand-grain weight the worst DH when compared
with the worst parent showed lower values. Difference
between the worst DH when compared with the worst
parent for thousand-grain weight trait was highly
significant (p<0.01) and for other traits were not
significant that indicating negative transgresive for
mostly

studied

traits.

Positive

and

negative

between the two parental lines. Bregitzer and

Campbell (2001), in a study to determine the QTLs
associated with plant regeneration in this population
also reported that transgressive segregation occurred.
Phenotypic coefficient variation for all studied traits
was most of genotypic coefficient variation. High
phenotypic and genotypic coefficient variation for
trait of number of spike, number of grain, number of
tiller and grain yield showed that these traits have
role effective in phenotypic and genotypic variation.
In selection intensity of 5 percent, the expected rate of
genetic output as a percentage of the mean is
expressed is varied from 1.13 for awn length trait to
46.11 for the number of grain trait. Narrow-sense
heritabilitys (h2) presented in Table 2 showed high
values ranging from 25.07 to 64.57 %. Therefore, all
studied traits were moderately heritable. Heritability
provides an estimate of how much variability is due to
genetic factors. However, the efficiency of selection
depends on the amount of heritability and expected
genetic advance. Traits that have high heritability and
genetic advances may be controlled by genes that
have act together. Furthermore, high heritability
estimates and genetic output may be as a result of
lowest environmental variance of traits (Orf et al.,
1999).

Table 1. Results of analysis of variance of 9 salt tolerance traits in 72 Steptoe/Morex doubled haploid lines
(DHLs) barley population and their two parents.
S.O.V

D.F

Mean Squares
SPL

AWL

PHT

NOS

NOG

Condition

30.13*

113.04**

12933.66** 888.8** 36.33*

29.18*

13022.78** 15459**

Blook(Condition)

2.38ns

4.93ns

300.697*

9.06ns

3.66**

1.66*

204.81*

705.57**

Genotype

77

3.20**

6.90**

179.93**

68.52**

3.48**

2.29**

207.55**

368.06**

Genotype Condition

77

1.13ns

5.17ns

94.59ns

38.57**

1.93**

0.95*

94.56ns

259.53ns

Error

308

1.28

4.18

95.71

18.40

1.063

0.67

80.77

197.38

C.V

22.49

22.29

15.64

11.75

29.97

34.80

37.01

74.16

R2

0.48

0.45

0.54

0.62

0.59

0.58

0.59

0.52

TGW

NOT

GYL

* and **, significant at 0.05 and 0.01 probability level, respectively; ns, non significant.
SPL: spike length; AWL: Awn Length; PHT: Plant height; TGW: Thousand grain weight; NOT: Number of tiller;
NOS: Number of spike; NOG: Number of grain; GYL: Grain yield.

251 Mehravaran et al.

Int. J. Biosci.

2014

Simple correlation analysis indicated that there was a

exist significant positive correlation (Table 3). Thus in

Positive highly significant relationship between yield

breeding program of barley for resistance to drought

and all studied traits except thousand-grain weight.

selection of each one of these traits can be useful and

This result is alike with results peighambari et al.

effective in increase yield. Correlations between traits

(2005) except in here there is not correlation between

may be to QTLs co-location with positive or negative

the yield and thousand-grain weight. Mohammadi

allelic effects. Moreover, variation of one trait may

and Baum (2005) and Blum and Penuel (1990) also

illustrate variation of others. Negative correlation

observed

between number of grain in spike and thousand-grain

highly

significant

positive

correlation

between number of grain trait and grain yield. In this

weight

can

be

due

to

florets

contest

for

results observed very high correlation between plant

photosynthesis material that cause reduced grains

height with spike length and number of grain in spike

weight. Golparvar et al. (1381) also obtained similar

and high correlation between plant heights with awn

results for these two traits.

length. Between spike length and number of tiller

Table 2. Simple statistics, genetic gain and heritability of 8 studied traits in a population of 72 Steptoe/Morex
DHLs barley and their two parents.
Item
Steptoe (P1)
Morex (P2)
P1- P2
=( P1+ P2)/2
WorstDHs
BestDHs
Range
DHs

SDDHs
CVDHs

X DHs- X
GGN=WDH-Wp
GGp=BDH-BP
P

SPL
5.01
3.71
1.30ns
4.36

AWL
9.22
8.66
0.55 ns
8.94

PHT
59.93
61.73
-1.80 ns
60.83

TGW
27.60
40.45
12.85 ns
34.02

NOT
4.01
2.88
1.13 ns
3.45

NOS
2.16
1.56
0.60 ns
1.86

NOG
24.78
12.06
12.71 ns
18.42

GYL
15.44
7.39
8.04 ns
11.42

3.51
6.72
3.21
5.04

7.33
15.32
7.99
9.14

49.87
74.70
24.82
62.84

30.23
45.81
15.58
36.49

2.25
5.16
2.91
3.38

1.38
3.76
2.38
2.32

11.59
36.92
25.33
24.41

6.32
58.77
52.45
18.92

0.70
13.93
0.677

1.06
11.58
0.202

5.52
8.78
2.01

3.00
8.23
2.472

0.59
17.50
-0.071

0.50
21.57
0.454

5.84
23.93
5.985

7.61
40.23
7.50

-0.19 ns
1.71 ns

-1.33 ns
6.10 ns

-10.05 ns
12.96 ns

2.63**
5.36 ns

-0.63 ns
1.15 ns

-0.18 ns
1.60 ns

-0.47 ns
12.14 ns

-1.07 ns
43.33

20.05
42.15
1.20
58.51

17.87
42.03
11.46
54.45

ns

GCV(%)
11.62
5.85
6.03
6.12
14.77
PCV5%
24.88
23.86
16.73
15.04
36.87
GC5%
1.67
1.13
10.24
4.95
1.16
2
h
64.57
25.07
47.42
43.72
44.54
**, significant at 0.01 probability level, respectively; ns, non significant.

22.45
81.12
9.35
29.48

SPL: spike length; AWL: Awn Length; PHT: Plant height; TGW: Thousand grain weight; NOT: Number of tiller;
NOS: Number of spike; NOG: Number of grain; GYL: Grain yield.
In general we found 40 QTLs for all studied traits

were attained for the QTLs of spike length (Qspl3H)

those 9 QTLs for normal condition, 18 QTLs for

on chromosome 3H and Thousand grain weight

drought stress condition and 13 QTLs for the mean of

(Qtgw6Hn) on chromosome 6H in stress condition.

two conditions. Phenotypic variations that were

QTL models explained about 38.5, 64.49 and 44.03 of

explained by these QTLs, changed from 8.02 to 30.87.

the total variation of spike length in normal and stress

The highest and the lowest phenotypic variances were

conditions and means of them, respectively. Five

related to spike length in mean and stress condition

QTLs were identified were detected on chromosomes

(Qspl3Hm, Qspl4Hs), respectively. LOD was ranged

3H, 4H and 7H, respectively. These QTLs (Qspl3H,

in 2.5 to 8.6. The highest and the lowest LOD scores

Qspl3Hn, Qspl3Hm, Qspl4Hs, Qspl7Hsa, Qspl7Hm

252 Mehravaran et al.

Int. J. Biosci.

2014

and Qspl7Hsb) were located near ABC323, Dor4A,

markers, respectively. Plant height-QTLs Qpht4Hsa,

WG114, ABC154A and AmY2 markers, respectively.

Qpht4Hsb, Qpht7Hsa and Qpht7Hsb were located on

Qspl3H was stable in two conditions and means of

chromosomes 4H and 7H near Adh4, ABG500b,

them. Mohammadi and Baum (2008) also in this case

ABG320 and AmY2 markers and controlled about

QTLs identified on chromosomes 2H, 3H, 4h and 7H.

53.38% of the total variation in stress conditions.

Peighambari et al. (2005) reportd existence QTLs

Mohammadi and Baum (2008) also in this case QTLs

controller spike length on chromosomes 2 and 3.

identified on chromosomes 2H, 4H, 6H and 7H in

There were three QTLs identified on chromosomes

Telhadia place. Peighambari et al. (2005) reported

2H and 4H for awn length controlling nearly 33.04

existence

and 14.9 of the total phenotypic variation for the trait,

chromosome 4H, they in general found 3 QTLs for

in stress conditions and means of them, respectively.

plant height in barley that explained 16 to 30% of

This

phenotypic variation. Kandemir et al. (1999) found a

three

QTLs,

Qawl2Hs,

Qawl2Hm

and

Qawlh4Hs near KsuF15, KsuF15 and WG1026B

QTLs

controller

plant

height

on

PHT QTL on chromosome 3H.

Table 3. Simple correlations for salt tolerance traits in the 72 Steptoe Morex doubled haploid (DH) line barley
population and their two parents.
SPL

AWL

PHT

TGW

NOT

NOS

AWL

0.208ns

PHT

0.513**

0.261*

TGW

0.010 ns

0.192 ns

0.212 ns

NOT

0.277*

0.220 ns

-0.009 ns

-0.168 ns

NOS

0.394**

0.339**

0.176 ns

-0.074 ns

0.847**

NOG

0.381**

0.145 ns

0.413**

-0.202 ns

0.171 ns

0.412**

GYL

0.337**

0.594**

0.322**

0.001 ns

0.493**

0.696**

NOG

0.650**

**, significant at 0.01 probability level, respectively; ns, non significant.

SPL: spike length; AWL: Awn Length; PHT: Plant height; TGW: Thousand grain weight; NOT: Number of tiller;
NOS: Number of spike; NOG: Number of grain; GYL: Grain yield.
QTLs for thousand grain weight according about

Rrn2, ABC154A and ABC465 markers, respectively.

33.99 and 20.78 % of the total variation for this trait

Six QTL affecting number of spike in bush were found

in normal conditions and means of two conditions,

on chromosomes 1H, 2H, 3H and 7H controlling

respectively. Qtgw2Hm, Qtgw2Hn, Qtgw6Hn and

approximately 13.02, 36.39 and 42.58 in normal and

Qtgw7Hn were located on chromosomes 2H, 6H and

stress

7H near MWG557, CDO537, Nar7 and ABC156d

Qnsp2Hn, Qnsp2Hs, Qnsp2Hm, Qnsp3Hm and

markers, respectively. Peighambari et al. (2005)

Qnsp7Hs. QTLs were located near Hor1, ABC306,

found 3 QTLs for 1000 grain weight on chromosomes

CDO537, WG996, ABG396 and ABC154A markers,

1, 5 and 7H that explained variation ranging from 14

respectively.

conditions

and

their

mean.

Qnsp1Hm,

to 20% of the total phenotypic variation.

Number of grain mapped to a region of chromosome
Five QTL affecting number of tiller were found on

2H, 4H and 7H, near CDO588, Adh8, ABG394,

chromosomes 2H, 5H, 7H controlling approximately

ABG500b,

33.19 and 25.04 % of the total variation in stress

respectively. These QTLs (Qnog2Hs, Qnog2Hm,

condition and means of two conditions, respectively.

Qnog4Hs, Qnog4Hm, Qnog7Hn and Qnog7Hm)

These QTL (Qnot2Hs, Qnot2Hm, Qnot5Hs, Qnot7Hs,

according about 17.42, 34.8 and 45.68 % of the total

and Qnot7Hm) were near CDO474B, CDO474B,

variation in normal and stress condition and their

253 Mehravaran et al.

ABC308

and

VAtp57A

markers,

Int. J. Biosci.

2014

mean. Mohammadi and Baum (2008) recognize two

Four QTL affecting yield grain were found on

QTL for number of grain trait on chromosomes 2H

chromosomes

and 4H in Telhadia and Barda two place. Teulat et al,

approximately 27.76, 11.97 and 23.78 in normal and

(2001) and Spagnoletti et al, (1985) recognized QTLs

stress

on chromosomes 3H and 4H for this trait.

Qgyl2Hm, Qgyl4Hs and Qgyl5Hn QTLs were located

2H,

conditions

4H
and

and
their

5H

controlling

mean.

Qgyl2Hn,

near ABG358, Adh8, WG114 and MWG514B markers,

respectively.
Table 4. Quantitative trait loci (QTL) for 9 salt tolerance traits identified by composite interval mapping for 72
Steptoe/Morex doubled haploid (DH) barley lines population. (first salinity level).
Trait

QTL name

R2

Total R2

SPL

Qspl3Hm

3H

ABC323

60.8

53.7-69.8

8.3815

-0.3918

30.87

44.03

Qspl7Hm

7H

ABC154A

46.2

44.5-52.5

4.1497

-0.2797

13.16

Qawl2Hm

2H

KsuF15

100.8

97.3-111.2

4.6874

0.4262

14.9

AWL

Chr. name Nearest marker

QTL pos.

QTL interval LOD score Allelic effect

14.9

PHT

TGW

Qtgw2Hm

2H

MWG557

66.8

61.6-69.2

5.0011

1.4047

20.78

20.78

NOT

Qnot2Hm

2H

CDO474B

70.3

68.2-73

3.6032

-0.2279

13.95

25.04

Qnot7Hm

7H

ABC465

60.2

48.1-64.3

2.9622

-0.2089

11.09

NOS

NOG

GYL
aQTL

Qnsp1Hm

1H

Hor1

15.3

8.6-30.6

3.3104

0.1756

11.88

Qnsp2Hm

2H

WG996

72.8

71.6-76.3

5.3776

-0.2315

20.69

Qnsp3Hm

3H

ABG396

73

71.9-78.2

2.8430

-0.1966

10.01

Qnog2Hm

2H

Adh8

62

55.2-66.1

3.4164

-2.2045

13.86

Qnog4Hm

4H

ABG500b

118.3

109.4-121.7

4.4843

-2.4547

16.34

Qnog7Hm

7H

VAtp57A

94

89.8-97.5

4.2028

-2.3284

15.48

Qgyl2Hm

2H

Adh8

65

59.6-66.7

5.3403

-3.7675

23.78

position expressed in cM, from origin of the linkage group ( end of short arm).

cProportion

of phenotypic variance explained by the QTL.

dTotal

bPeak

42.58

45.68

23.78

value of the LOD.

phenotypic variance explained by the model.

SPL: spike length; AWL: Awn Length; PHT: Plant height; TGW: Thousand grain weight; NOT: Number of tiller;
NOS: Number of spike; NOG: Number of grain; GYL: Grain yield.
Several QTL co-locations, confirm the observed

These co-locations could be either because of linkage

correlations among the traits. For example, in stress

between two genes or the pleiotropy effect of one

conditions, Qnot7Hs and Qnsp7Hs QTLs conceded in

gene. In the later case, the correlation between traits

their genomic locations with each other. The positive

will never be broken. Pleiotropy controls the common

correlation between number of tiller and number of

sub-fraction of the traits, and therefore, would result

spike would come from these common QTLs with

in the concurrent increase or decrease of the

congruent effects. QTLs associated with thousand-

correlated traits when we select for only one trait. The

grain weight and number of spike (Qtgw2Hn and

large effect of QTL near the ABC154A marker on

Qnsp2Hs) were also co-localized. Han and Ullrich

chromosome 7H on number of tiller and number of

(1994), reported several

QTL co-locations, for

spike traits, raises an intriguing question about

different traits, For example, QTLs for kernel weight

whether a single gene or a cluster of genes for drought

and grain protein were located in similar region on

tolerance traits is located on these chromosomes. The

2H. siahsar et al, (2009) identified several QTL co-

answer to this question requires a detailed study

locations effective for difference trait in forage

using a more densely saturated map. Because, there is

qualification in same population. Siahsar and Narouie

one difficulty that must be faced in attempting to

(2010) (Rosiellen et al., 1981) distinguish several QTL

discern the nature of QTL governing of agronomical

co-locations for difference physiological trait relation

traits. Mansur et al (1993) and orf et al (1999),

with salt stress in this population.

reported cluster of QTLs with various effects on

254 Mehravaran et al.

Int. J. Biosci.

2014

flowering, maturity, plant height and lodging. The

probably only to the closer one hundred genes. This is

best confidence interval with which mapping can

an impossible wide window in which to search for a

localize QTL, regardless of the size of the mapping

gene without some obviously idea of what it might be.

population, is perhaps 10 cM. With a mapping

Overall, this region of chromosome 2H and 7H is

population

of

important for drought tolerance in barley, and

phenotyping it may be 20-30 cM. Under the most

therefore the region may be used as an important

hopeful conditions the best map location of a QTL is

target for improving drought tolerance of barley.

constrained

by

the

difficulties

Table 5. Quantitative trait loci (QTL) for 9 salt tolerance traits identified by composite interval mapping for 72
Steptoe/Morex doubled haploid (DH) barley lines population. (first salinity level).
Trait

SPL

AWL
PHT

TGW

NOT

NOS

NOG

GYL

aQTL

QTL name Chr. name Nearest marker QTL pos.

LOD score

Allelic effect

normal

stress

normal

stress

R2
normal

stress

Qspl3H

3H

ABC323

60.8

3.7834

8.6785

-0.5995

-0.3928

16.65

25.56

Qspl3Hn

3H

Dor4A

77

4.6101

-0.3796

21.85

Qspl4Hs

4H

WG114

121.4

3.3218

-0.2254

8.02

Qspl7Hsa

7H

ABC154A

46.2

6.7170

-0.3502

18.18

Qspl7Hsb

7H

AmY2

113.3

Qawl2Hs

2H

KsuF15

100.8

2.8747

0.2966

10.98

Qawl4Hs

4H

WG1026B

55.8

5.7548

0.5084

22.06

Qpht4Hsa

4H

Adh4

58.9

3.8289

1.9842

13.79

Qpht4Hsb

4H

ABG500b

120.3

3.2816

-1.874

11.97

Qpht7Hsa

7H

ABG320

14.7

3.1892

-1.7927

11.24

4.0705

0.2864

12.73

Qpht7Hsb

7H

AmY2

115.3

4.1202

2.1431

16.38

Qtgw2Hn

2H

CDO537

68.8

3.3398

1.2161

11.74

Qtgw6Hn

6H

Nar7

76

2.5416

-1.0367

8.71

Qtgw7Hn

7H

ABC156d

65.6

3.4516

1.3591

13.54

Qnot2Hs

2H

CDO474B

70.3

2.9012

-0.3075

11.17

Qnot5Hs

5H

Rrn2

48.2

3.1130

0.3110

11.85

Qnot7Hs

7H

ABC154A

46.2

2.7084

-0.2902

10.17

Qnsp2Hn

2H

ABC306

71.8

2.8531

-0.2087

13.02

Qnsp2Hs

2H

CDO537

68.8

3.7429

-0.2909

15.74

Qnsp7Hs

7H

ABC154A

46.2

2.8253

-0.2492

11.51

Qnog2Hs

2H

CDO588

82.2

2.7133

-2.2124

9.14

Qnog4Hs

4H

ABG394

129.3

6.3223

-3.6809

25.66

Qnog7Hn

7H

ABC308

85.3

3.6833

-2.8972

17.42

Qgyl2Hn

2H

ABG358

44.3

3.8412

-3.5703

15.67

Qgyl4Hs

4H

WG114

122.4

3.1143

-2.1523

11.97

Qgyl5Hn

5H

MWG514B

141.7

2.7436

3.2492

12.09

position expressed in cM, from origin of the linkage group ( end of short arm).

cProportion

bPeak

value of the LOD.

of phenotypic variance explained by the QTL. dTotal phenotypic variance explained by the model.

SPL: spike length; AWL: Awn Length; PHT: Plant height; TGW: Thousand grain weight; NOT: Number of tiller;
NOS: Number of spike; NOG: Number of grain; GYL: Grain yield.
One of the major goals of QTL mapping is to select

constituent for MAS. Coincident QTLs provide

markers that linked to genes contributing to change

relative permanence of genetic control which would

in the trait of interest. QTL persistence across

overcome the problem associated with the interaction

different environments and background is important

of QTL by environment. In this study moreover 40

255 Mehravaran et al.

Int. J. Biosci.

2014

QTLs were detected for agronomical traits and grain

Program Annual Re-port 1987, ICARDA, Aleppo,

yield associated with drought tolerance and some of

Syria, 101116.

these QTLs appear to be quite stale between two

conditions,

thus

gain

through

marker-assisted

Acevedo E. 1993. Potential of carbon isotope

selection (MAS) in this population would be

discrimination as a selection criterion in barley

unlimited. In previous studies the utility of marker-

breeding. In: Ehleringer JR, Hall AE, Farquhar GD

assisted selection for traits such as yield and quality

(eds)

in barley has been demonstrated.

relations. Academic Press, New York, 399417.

Stable

isotopes

and

plant

carbon-water

http://dx.doi.org/10.1016/B978-0-08-091801We detected a number of variable QTLs for the same

3.50035-0

trait in different environment. Thus suggesting the

presence of QTLenvironment interaction variation.

Backes G, Granner A, Foroughi-Wehr B,

The interaction of QTLs and environment were

Fischbeck G, Wenzel G, Jahoor A. 1995.

observed in many studies (Golparvar et al., 2002;

Localization of quantitative trait loci (QTLs) for

Lafitte et al., 2006; Zhu et al., 1999). Since

agronomic important characters by the use of a RFLP

environment plays a enormous role in the phenotypic

map in barley (Hordeum vulgare L.). Theoretical and

expression of traits associated with drought tolerance,

Applied Genetics 90, 294302.

therefore QTL analysis must be repeated in different

http://dx.doi.org/10.1007/BF00222217

environments, because a QTL may be expressed in

one environment but not in another, or expressed

Baum M, Sayed H, Araus JL, Grando S,

strongly in one environment but weakly in another,

Ceccarelli S, Backes G, Molher V, Jahoor A,

and or expressed very differently and opposite effects

Fischbeck G. 1996. QTL analysis of agronomic

in different environments.

important characters for dryland conditions in barley.

In: Proc Vth IOC-VII IBGS meeting, 30 July6

This study is one of few reports of QTL analysis in

August 1996, Saskatoon, Saskatchewan, Canada, 241

barley evaluated in Iran. The means of all DH lines

243.

were close to the mid-parental values for most traits.

Although phenotypic distribution of DH population

Blum A. 1988. Drought resistance. In: Plant

was normal, transgressive segregation was also

breeding for stress environments. CRC Press, Boca

observed in both directions for all traits. Significant

Raton, Florida, 4376.

variation and normal distribution of all traits

measured in this study suggest the suitability of the

Blum A, Penue l. 1990. Physiological of attributes

DH population for QTL analysis. In spite of, the small

associated with drought resistance of wheat cultivars

sample size used for technical reasons, a total of 50

in a Mediterranean environment. Australian Journal

significant QTLs were found for the 8 studied traits.

Agriculture research 41, 799-810.

One of detected QTLs appears to be quite stale

http://dx.doi.org/10.1071/AR9900799

between two conditions. QTL co-locations approve

the observed correlations among the traits. Although

Bregitzer P, Campbell RD. 2001. Genetic markers

the detected regions need to be more accurately

associated with green and albino plant regeneration

mapped, the information obtained should help in

from embryogenic barley callus. Crop Science 41,

marker-assisted selection.

173-179.
http://dx.doi.org/10.2135/cropsci2001.411173x

References
Acevedo E. 1987. Gas exchange of barley and wheat

Broman KW, Speed TP. 1999. A review of

genotypes under drought. In: Cereal Improvement

methods for identifying QTLs in experimental

256 Mehravaran et al.

Int. J. Biosci.

2014

crosses, In:Seillier- Moiseiwitsch, F. (ed.), Statistics in

January 17-21.

Molecular Biology and Genetics 33, 11442 P. IMS

Lecture Notes-Monograph Series.

Kjaer B, Jensen J. 1996. Quantitative trait loci for

grain yield and yield components in a cross between a

Craufurd PQ, Austin RB, Acevedo E, Hall MA.

six-rowed and a two-rowed barley. Euphytica 90,

1991. Carbon iso-tope discrimination and grain yield

3948.

in barley. Field Crops Res 27, 301313.

http://dx.doi.org/10.1016/0378-4290(91)90038-W

Lafitte R, Courtois B. 1999. Genetic variation in

performance under reproductive-stage water deficit

Farquhar GD, Richards RA. 1984. Isotopic

in a doubled-haploid rice population in upland fields.

composition of plant carbon correlates with water-

In: Ribaut JM, Poland D (eds) Molecular approaches

use-efficiency

for genetic improvement of cereals for stable

of

wheat

geno-types.

Australian

Journal Plant Physiology 11, 539552.

production in water-limited environments. CIMMYT,

http://dx.doi.org/10.1071/PP9840539

El Batan, Mexico 2125 June 1999. 97102.

Gibson LA, Bowman JGP, Oberthur LE, Blake

Lebreton

TK.

Semikhodskii

1994.

Determination

of

genetic

marker

CM,

Visscher
A,

PM,

Quarri

Halley

SA.

1998.

CS,
A

associated with ruminant digestion of barley. Proc.

nonparametric bootstrap method for testing close

West. Sect. Am. Soc. Anim. Sci. 45, 317-320.

linkage vs. Pliotropy on coincident quantitative trait

loci. Genetics 150, 931-943.

Golparvar

A.

2002.

Evaluation

some

of

morphological traits as selection criteria in wheat

Mansur LM, Lark KG, Kross H, Oliveira A.

breeding. Journal Iran Agronomic Sciences 4, 202-

1993. Interval mapping of quantitative trait loci for

207.

reproductive, morphological and seed traits of

Han F, and Ullrich SE. 1994. Mapping of

soybean (Glycine max L.). Theoretical and Applied

quantitative trait loci for malting quality traits in

Genetics 86, 907-913.

barley. Barley Genet. Newsl. 23, 84-97.

http://dx.doi.org/10.1007/BF00211040

Hayes PM, Lui BH, Knapp SJ, Chen F, Jones

Marquez-Cedillo LA, Hayes PM, Kleinhofs A,

B, Blake T, Franckowiak J, Rasmusson D,

Legge WG, Rossnagel BG, Sato K, Ullrich SE,

Sorrells

D,

Wesenberg DM. 2001. QTL analysis of agronomic

Kleinhofs, A. 1993. Quantitative trait locus effects

M,

Ullrich

SE,

Wesenberg

traits in barley based on the doubled-haploid progeny

and environmental interaction in a sample of North

of two elite North American varieties representing

American barley germplasm. Theoretical and Applied

different.

Genetics 87, 392401.

http://dx.doi.org/10.1007/BF01184929

Mather D, Tinker NA, LaBerge DE, Edney M,

Jones BL, Rossnagel BG, Legge WG, Briggs

Hayes PM, Iyambo OE. 1994. Summery of QTL

KG, Irvine RB, Falk DE, Kasha KJ. 1997.

effects in the SteptoeMorex population. Barley

Regions of the genome that affect grain and malt

Genetic. Newsl. 23, 98-143.

quality in a North American two-row barley cross.

Crop Science 37, 544554.

Kandemir N, Kleinhofs D, Kudrna A, Ullrich

http://dx.doi.org/10.2135/cropsci1997.0011183X003

SE. 1999. Molecular Marker Assisted analysis of two

700020039x

yields QTL in barley, in: Plant and Animal Genom VII

Conference. Town and Country Hotel, San Diego, CA,

257 Mehravaran et al.

Merah O, Delens E, Souyris I, Monneveux P.

Int. J. Biosci.

2014

2001. Ash content might predict carbon isotope

backcross QTL analysis in barley (Hordeum vulgare

discrimination and grain yield in durum wheat. New

L.). Theoretical and Applied Genetics 107, 340352.

Phytol 149, 275282.

http://dx.doi.org/10.1007/s00122-003-1253-9

http://dx.doi.org/10.1046/j.1469-8137.2001.00012.x
Price A, Courtois B. 1999. Mapping QTLs
Mohammadi M, Baum M. 2008. QTL Analysis for

associated with drought resistance in rice: progress,

morphological traits in doubled haploid population of

problems and prospects. Plant Growth Reg 29, 123

barley. Science and Technology of Agriculture and

133.

Natural Resources 45 (A), 111-120.

http://dx.doi.org/10.1023/A:1006255832479

Mohammadi M, Taleei A, Zeinali H, Naghavi

Ribaut JM, Hoisington DA, Deutsch JA, Jiang

MR, Ceccarelli S, Grando S, Baum M. 2005.

C, Gonzalez-de-Leon D. 1996. Identification of

QTL analysis for phenologic traits in doubled haploid

quatitative trait loci under drought conditions in

population of barley. Int. J. Agric. Biol 7, 820830.

tropical maize. 1. Flowering parameters and the

anthesis-silking interval. Theoretical and Applied

Monneveux P. 1991. Quelles stratgies pour

Genetics 92, 905914.

lamlioration gntique de la tolrance au dficit

http://dx.doi.org/10.1007/BF00221905

hydrique des cereals dhiver? In: Demarly Y, Chalbi N

(eds) Lamlioration des plantes pour ladaptation aux

Siahsar BA,

milieux arides. John Libbey Eurotext Paris, 165185.

Naghavi MR, Nabipour A, Sarrafi A. 2009. QTL

Peighambari

SA, Taleii

AR,

analysis of forage quality traits in barley (Hordeum

Morgan JM. 1983. Osmoregulation as a selection

vulgare L.). Cereal Res. Commun 37(4), 479-488.

criterion for drought tolerance in wheat. Aust J. Agric

http://dx.doi.org/10.1556/CRC.37.2009.4.1

Res. 34, 607614.

http://dx.doi.org/10.1071/AR9830607

Siahsar BA, Narouei M. 2010. Mapping QTLs

physiological traits associated with salt tolerance in

Orf JH, Chase K, Jarvik T, Mansur LM, Cregan

SteptoeMorex doubled haploid lines of barley at

PB, Adler FR, Lark KG. 1999. Genetics of soybean

seedling stage. Journal Food Agriculture Environ

agronomic traits: I. Comparison of three related

8(2), 751759.

recombinant inbred populations. Crop Science 39,

1642-1651.

Spagnoletti Z, Qualest CO. 1985. Geographical

http://dx.doi.org/10.2135/cropsci1999.3961642x

divercity for quantitative spike characters in a world

collection of durum wheat. Journal Crop Science 27,

Panes VG. 1957. Genetics of quantitative characters

235-241.

in relation to plant breeding. Indian Journal Genetic

17, 317-328.

Teulat B, This D, Khairallah M, Borries C,

Ragot C, Sourdille P, Leroy P, Monneveux P,

Peighambari SA, Yazdi Samadi B, Nabipour A,

Charrier A. 1998. Several QTLs involved in osmotic

Charmet G, Sarrafi A. 2005. QTL analysis for

adjustment trait variation in barley (Hordeum

agronomic

vulgare L.). Theoretical and Applied Genetics 96,

traits

in

barley

doubled

haploids

population grown in Iran. Plant Science 169, 1008-

688698.

1013.

http://dx.doi.org/10.1007/s001220050790

http://dx.doi.org/10.1016/j.plantsci.2005.05.018
Teulat B, Borries C, This D. 2001. New QTLs
Pillen K, Zacharias A, Lon J. 2003. Advanced

258 Mehravaran et al.

identified for plant water status, water-soluble

Int. J. Biosci.

2014

carbohydrates, osmotic adjustment in a barley

spring barley (Hordeum vulgare L.). Theoretical and

population grown in a growth-chamber under two

Applied Genetics 91, 10371047.

water regimes. Theoretical and Applied Genetics 103,

160-170.

Yin X, Stam P, Johan Dourleijn C, Kropff MJ.

http://dx.doi.org/10.1007/s001220000503

1999. AFLP mapping of quantitative trait loci for

yield-determining physi-ological characters in spring

Teulat B, Merah O, Souyris I, This D. 2001.

barley. Theoretical and Applied Genetics 99, 244

QTLs for agronomic traits from a Mediterranean

253.

barley progeny grown in several environments.

http://dx.doi.org/10.1007/s001220051230

Theoretical and Applied Genetics 103, 774787.

http://dx.doi.org/10.1007/s001220100619

Zhu H, Briceo G, Dovel R, Hayes PM, Liu BH,

Liu CT, Toojinda T, Ullrich SE. 1999. Molecular

Thomas WTB, Powell W, Waugh R, Chalmers

breeding for grain yield in barley: an evaluation of

KJ, Barua UM, Jack P, Lea V, Forster, BP,

QTL effects in a spring barley cross. Theoretical and

Swanston JS, Ellis RP, anson PR, Lance RCM.

Applied Genetics 98, 772779.

1995.

http://dx.doi.org/10.1007/s001220051134

Detection

of

quantitative

trait

loci

for

agronomic, yield, grain, and disease characters in

259 Mehravaran et al.