qtl analysis for yield and other agronomic traits in barley

QTL ANALYSIS FOR YIELD AND QTL ANALYSIS FOR YIELD AND

OTHER AGRONOMIC TRAITS IN OTHER AGRONOMIC TRAITS IN

BARLEYBARLEY

INTRODUCTIONINTRODUCTION

Production Statistics, Economics and MarketsProduction Statistics, Economics and Markets

Barley ranks fourth among the cereals in terms of total world production. The Barley ranks fourth among the cereals in terms of total world production. The

average productionaverage production is is around around 136 136 million million metric tonnes. The map below shows metric tonnes. The map below shows

the distribution of production by country.the distribution of production by country.

Source: FAOSTAThttp://oregonstate.edu/instruct/css/330/five

Countries that Produced the Most Barley (3-year averages, 2000-2002)

CountryArea

(ha x 1,000)Yield

(kg/ha)Production

(Mt x 1,000)

Russian Federation 8061213417200

Germany 2052594012192

Canada 4050260910565

France 1627 6240 10154

Ukraine 397422538953

Spain 313827578652

Turkey 360621177633

United Kingdom1159 55106388

Australia 34201719 5879

United States1840 3136 5770

Denmark 7745244 4061

Leading Barley Importer and Exporter Countries (Averages for 1998-2000)

CountryImports

(Mt x 1,000)Country

Exports(Mt x 1,000)

Saudi Arabia4447France6758

Japan2531Germany4040

China2160Australia3973

Belgium-Luxembourg1459Canada2196

Russian Federation1005United Kingdom1898

Germany941Belgium-Luxembourg1223

Brazil880United States985

Netherlands823Denmark960

Morocco794Ukraine841

Italy790 Turkey659

United States778 Sweden606

Production Statistics, Economics and Production Statistics, Economics and

MarketsMarkets

Barley (Hordum vulgare L.), the oldest among cereals, is an autogamous diploid (2n=14) with an average genome size of ~5000 Mbp. It stands fourth among cereals as far as nutritive value and area under cultivation are concerned, but it is in first standing with regard to easily getting adapted and being cultivable in varied climatic conditions as well as the extent to which it is cultivated.

Barley, with an area under cultivation of 1.5 million hectares and 2.5 million tons of grain yields is cultivated in most provincial areas in Iran

Determination of genes controlling quantitative traits (QTL), such as yield and yield components is of paramount importance to plant breeders.

QTL can help breeders to be aware of genetic control of quantitative traits and to improve such traits through an implementation of more efficacious selections.

Mapping of quantitative traits controlling genes (QTLs) in a plant is carried out by use of suitable genetic population (AILs, BC, RIL, DH, F3, F2) in which there exists enough diversity for different traits.

QTL analysis can answer a variety of questions including:

How many zones in a genome control a quantitative trait?

Are all these zones equally effective?Is the effect of some zones more and some

others less?How are the QTLs distributed throughout

the genome?Are they distributed throughout the whole

genome or are they concentrated in a particular zone in the genome?

Different markers such as isozymes, SSR, AFLP, RAPD and others are employed in such research activities.

To locate QTLs in a genome after their recognition, there would be a need for a genetic map (a base map).

Nowadays such maps have been prepared for many crop plants including barley. These maps get more and more saturated and more accurate with more research works. In preparation of these maps land marking markers such as RFLP are used.

For determination of QTLs and their locations, different traits are measured in a given population to ascertain the existance of sufficient diversity for the traits measured. Then through the use of molecular markers the genome of the genotypes are being searched to obtain a saturated map. Using this map, then the markers linked to QTLs are being designated. In this way and by the help of the base map the location of QTLs in genome are detected. There are different methods of searching a genome for QTLs, such as regression and methods of maximum likelihood. There has been different softwares such as MULTIQTL, JOINMAP, Mapmaker QTL, etc. designed for this purpose.

This study was carried out to identify genes controlling QTLs related to agronomic traits as well as genes controlling traits related to drought tolerance and also to find the effect of drought stress on yield and yield components in barley.

LITERATURE REVIEWLITERATURE REVIEW

Summary of the QTLs detected from previous studies using the Steptoe Summary of the QTLs detected from previous studies using the Steptoe

MorexMorexChromosometrait references

1H

acid detergent fibre contentHan et al. 2003

enzyme activity and thermostability of beta-amylaseKaneko et al. 2001

grain yieldKandemir et al. 2000

containing malting-qualityHan et al. 1997

resistance to net blotch and spot blotchSteffenson et al. 1996

2H


resistance to leaf stripeArru et al. 2003

enzyme activity and thermostability of beta-amylaseKaneko et al. 2001

starch granulesBorem et al. 1999

grain yieldKandemir et al.2000

grain yieldZhu et al. 1999

grain yieldRomagosa et al. 1999

genetic variation in barley of crossability with wheatTaketa et al. 1995

malt beta-glucan & beta glucanase activityHan et al. 1995


Chromosometrait references

3H


head shattering, plant height & grain yieldKandemir et al. 2000

grain yieldRomagosa et al. 1999

seed dormancyLarson et al. 1996


partial resistance to bacterial leaf streakEl-Attari et al.1998


4H


malting quality Han et al. 1997

salt tolerane at germination and the seedling stageMano et al. 1997

powdery mildew resistance Iwasa et al, 1997


green malt beta-glucanse activityHan et al. 1995

Chromosometrait references

5H

alpha-amylase activity of maltAyoub et al. 2003


seed dormancyGao et al. 2003

endosperm-textureBeecher etal.2002

deep-seeding toleranceTakahashi et al. 2001

seed dormancyFeng Han et al. 1999

seed dormancyHayes et al. 1993


salt tolerance at germination and the seedling stageMano et al. 1997

finished malt beta-glucanse & grain yieldHan et al.1995

seed dormancy during seed developmentRomagosa et al. 1999


partial resistance to bacterial leaf streakEl-Attari et al.1998

Chromosometraitreferences

6H

salt tolerance at germination and the seedling stageMano et al. 1997


7H



green malt beta-glucanseHan et al.1995

containing malting quality Han et al. 1997

polymorphic microsatellite in the limit dextrinase gene


MATERIALS AND METHODSMATERIALS AND METHODS

Seeds from 72 doubled haploid (DH) lines of barley, the result of a cross between:Variety Steptoe (six rowed, high yielding, drought susceptible, highly dormant, susceptible to leaf stripe bacterial disease, used as forage) and variety Morex (six rowed, drought resistant, high quality malt and amylase, low dormancy and resistant to leaf stripe bacterial disease), received from University of Tulose, France, were vase planted in a greenhouse in autumn 2000. A sufficient number of seeds were obtained from the cross. The resulting seeds were planted in a randomized complete block design with three replications in the agricultural research farm of University of Tehran in the crop years 2001-2002 and 2002-2003. The planting dates were Nov. 14 and Oct. 28, respectively. A number of 40 seeds were planted in 2-m rows. Rows were 25cm apart. The experimental land was irrigated immediately after cultivation. To test the effect of drought tolerance (as compared to normal irrigation) on yield and yield components and also to identify the genes that control the traits related to drought tolerance, the 1st and 2nd replications were irrigated once more at spike initiation, while the 3rd replication was irrigated for 5 more times as practiced locally. Precipitation amounted to 229 and 221 mm (from planting to harvest date) for the two crop years, respectively.

Traits studied were:

X1: date of flowering (days)X2: date of spike initiation (days)X3: date of maturity (days)X4: plant height (cm)X5: spike length (mm)X6: tiller per plantX7: seed per spikeX8: 1000-seed weight (gr)X9: protein percentage (%)y: seed yield (gr)

A multiple linear regression model was selected to describe the traits with highest effects on variability. Heritability was calculated on the basis of mean of two- year data by use of the formula;2

/222

/ nreg negh

Where: is the variance component of genotype year and n stands for the number of environments. Additive variance and enviromental variance (Mse) were also determined. Diversity and genetic gain (GG= best DH-best P) for each trait were also determined.

neg /

2

)2

1V( DH

2A

Values obtained for each trait, for the irrigated replication and mean of the two stressed replications, were compared with t-test for each year as well as for the two year means. Analysis of variance was employed to test the effect of year and the effect of stress/year for all traits.

Genetic map of the population under study consisted of 327 RFLP markers with the mean distance of 3.75 cM between any two markers (NABGMP)*.

Considering field measurement results and markers information obtained from 72 DH lines a new map was constructed using 'Grain Genes' information of 150 doubled haploid lines. Then QTL analysis was performed for each trait and each chromosome in each year; and a combination of the above information was employed to determine the interaction of genotype environment (year and drought stress). Analysis of QTLs was performed on the basis of the average for the two years, using MULTI QTL software. Significant QTL effects (P= 0.027) with the LOD value of 2.33 for a nonsaturated map of studied traits was considered. Significant LODs were found by permutation test using MULTI QTL software.

LOD peaks with the probability of 95% were considered as the most probable QTL effects. Confidence intervals were assessed through 1000- time bootstrap replicated samplings, using MULTI QTL software.

* North American Barley Genome Mapping Project

Genetic map of barley Steptoe Genetic map of barley Steptoe Morex from Anderis Kleinhofs, updated 1/94 from the map referenced. Morex from Anderis Kleinhofs, updated 1/94 from the map referenced.

Reference : A. Kleinhofs et al. 1993. A molecular, isozyme and morphological map of the

barley

(Hordeum vulgare L.) genome, Theor.Appl.Genet. 86 : 705-712 Map position in

(cM)Kosambi

ChromosomeLocusMap Position

1HAga60.0

1HHor51.4

1HAct82.1

1HHor22.8

1HMWG036A8.6

1HMWG83713.3

1HHor115.3

1HABA00422.5

1HCDO9931.6

1HABG05337.5

1HIcal43.0

1HABG07444.3

1HABG500A49.6

1HABC16457.7

1HksuF2A59.0

1HABC152B60.3

1HRisBPP161C61.6

1HBCD351C62.3

1HPcr263.1


1HABR33767.0

1HCDO105B71.3

1HGlb172.4

1HABC16080.7

1HHis4A93.3

1HHis3B100.0

1HABC307A102.1

1HcMWG706A112.0

1HABC257116.9

1Hipgd2117.6

1HcMWG733121.6

1HAtpbA127.6

1HABG702129.6

1HABC322B135.6

1HABC261137.9

1HMWG635C138.9

1HCab2140.9

1HAga7144.0

1HMWG912146.3

1HBCD340A147.0

1HABG055147.9

1HABG387A150.8


2HABG313A0.0

2HcMWG6821.6

2HABG0582.3

2HABG703B7.0

2HChs1B16.1

2HChs1A16.9

2HWG51618.0

2HABG00822.5

2HRbcS27.2

2HBCD351F29.9

2HABG31831.9

2HABC156A34.6

2HMWG85840.6

2HABG35843.3

2HCDO6446.0

2HABG45947.3

2HMWG520A48.6

2HABG00549.4

2HPox52.6

2HABC45453.9

2HMWG663-2B54.6

2HMWG95055.3

2HAdh856.0

2HB15C66.0

2HMWG55766.8

2HCDO53768.8


2HCDO474B70.3

2HABG01481.2

2HCDO58882.2

2HAdhIntC85.7

2HCDO474C87.1

2HHis3C88.2

2HABC152D90.2

2HRrn5S196.9

2HksuF15100.8

2HMWG503104.3

2HABG072123.0

2HCrg3A125.2

2HABC252140.3

2HGln2141.6

2HABC157147.6

2HABG317A158.4

2HABG317B161.7

2HABC153164.4

2HABC165165.1

2HABG316E171.1

2HABG316D175.1

2HPcr1176.4

2HcMWG720181.1

2HPrx2183.4

2HBG123A184.4

2HABA005185.7

2HbBE54C187.0


3HMWG571C0.0

3HABG316A2.1

3HABC17116.3

3HMWG798B25.3

3HMWG58429.3

3HABG46030.0

3HABG47132.7

3HABC32360.8

3HABG39673.0

3HABG39974.5

3HABG39875.3

3HDor4A77.0

3HBCD82877.8

3HMWG68080.5

3HABG703A83.6

3HMWG571B85.9

3HPSR15691.3

3HABC17692.7

3HABG37798.4

3HMWG555B104.1

3HABG315107.4

3HABG453109.4

3HMWG571A114.1

3HABR320115.4


3HABC307B118.6

3HCrg3B120.7

3HABG499121.4

3HCDO113B131.8

3HHis4B145.6

3HABG004157.4

3HWG110162.6

3HmPub164.8

3HABG389166.4

3HABC161168.8

3HABA302171.2

3HMWG902172.8

3HABG654176.2

3HBCD131177.9

3HGlb4180.1

3HGlb3180.8

3HiBgl183.1

3HABG495B186.0

3HMWG041190.1

3HABC166190.8

3HPrx1B192.1

3HABG319B196.2

3HABC172205.0


4HMWG6340.0

4HWG6221.4

4HABG313B12.0

4HMWG07714.1

4HCDO66917.2

4HB32E24.9

4HBCD402B33.7

4HBCD351D37.0

4HMWG635A41.0

4HBCD265B48.5

4HDhn653.7

4HABG00354.4

4HABC30355.1

4HWG1026B55.8

4HAdh458.9

4HABA00361.2

4HABG48463.2

4HPgk2A63.9

4HMWG05865.2


4HABG32167.9

4HABR31569.3

4HWG46476.6

4HbBE54A79.9

4HBCD453B90.7

4HABG47296.5

4HABG319A101.0

4HiAco2115.3

4HABG500B118.3

4HWG114121.4

4HcMWG652B124.0

4HABG054126.0

4HABG394127.3

4HABG366130.3

4HABG397134.0

4HABG319C147.7

4HBmy1166.7

4HksuH11168.4


5HMWG5020.0

5HABR3134.7

5HMWG920-1A7.4

5HABG316B9.4

5HABG70536.7

5HCDO669B41.2

5HDor542.8

5HAdh643.5

5HABR33644.2

5HABG49744.9

5HABG39545.6

5HCDO74946.9

5HRrn248.2

5HUbi250.2

5HB12DB51.5

5HLtp152.8

5HWG54157.5

5HWG53060.8

5HBG123B61.5

5HABC70662.3

5HWG88966.7

5HAle68.2

5HCDO348B69.5

5HABC31470.8

5HABC30278.2

5HCDO57B92.2

5HKsuA1B97.9


5HBCD351E99.7

5HABG069102.2

5HWG364103.8

5HABC717104.5

5HABG473105.2

5HMWG514B138.7

5HCDO504145.3

5HWG644146.8

5HABG712148.8

5HiEst9151.5

5HWG908153.5

5HMWG877157.5

5HABG495A160.4

5HABG496166.6

5HABC482173.0

5HABG391178.4

5HABG707180.2

5HABG390181.1

5HCDO484188.5

5HABG463191.2

5HMWG813A194.5

5HABC309196.5

5HABG314B197.2

5HABG314A197.9

5HMWG851C200.9

5HABA304203.7

5HMWG851B204.5


6HWG223B56.5

6HABG47460.4

6HBCD340E61.2

6HksuD1765.0

6HABG37967.3

6HABG38868.0

6HABC17569.3

6HksuA3D72.5

6HMWG82073.3

6HABC170B74.6

6HMWG684B75.4

6HNar776.0

6HAmy180.2

6HbBE54B82.9

6HMWG93487.6

6HMWG684A95.7

6HABC170A97.0

6HMWG798A139.7


6HLth0.0

6HPSR1671.5

6HMWG6206.6

6HABG4667.4

6HNar18.1

6HABG37813.3

6HCxp322.9

6HABC152A23.7

6HHis3D24.6

6HcMWG652A26.4

6HPSR10632.2

6HABG387B36.2

6HABG45847.8

6HABR33148.6

6HABC169B51.8

6HRrn153.1

6HB12DA53.8

6HksuA3B55.1

6HABR33555.8


7HABG7040.0

7HdRpg10.7

7HMWG036B3.4

7HMWG555A4.7

7HiPgd1A6.0

7HABR3036.7

7HBCD12910.0

7HABG32014.7

7HiEst518.0

7HGlx18.7

7Hprx1A19.6

7HWG789A24.1

7HCDO47525.9

7HMWG08926.8

7HABC169A27.5

7HABC167A29.5

7HdRcs130.2

7HABG38033.5

7HABC15840.2

7HKsuA1A44.2

7HABC154A46.2

7HMWG83648.2

7HBrz54.7

7HABC46560.2

7HABC25560.9

7HABC156D61.6


7HABG70175.0

7HABG022A77.4

7HABG01180.5

7HBCD340D82.4

7HABC30883.3

7HABC322A86.0

7HWG71986.7

7HABC45587.4

7HAdh789.2

7HCDO67391.5

7HABC154B93.3

7HYAtp57A94.0

7HAmy299.3

7HUbi1116.7

7HABC310B120.8

7HRisP103126.7

7HPSR129127.4

7HPgk2B128.7

7HbBE54E130.0

7HABC305132.0

7HABG461144.2

7HABG652148.6

7HWG420151.5

7HksuD14C152.2

7HMWG635B168.8

7HCat3170.1

7HDor4B170.9

Linkage group

No. of

Marker

Chromosome

length (cM)

Mean distance between 2 markers (cM)

1H54204.53.79

2H58187.03.22

3H47205.04.36

4H37168.44.55

5H53170.93.22

6H37139.73.77

7H41150.83.68

Total3271226.33.75

Markers distribution in the linkage groups map of Markers distribution in the linkage groups map of Steptoe × Morex Steptoe × Morex in 150 barley doubled haploid linesin 150 barley doubled haploid lines

RESULTSRESULTS

Mean squares (MS) for traits under studyMean squares (MS) for traits under study

SOVdfD

ays to flow

ering

Days to

spike emergence

Days to

maturity

Plant

height (cm)

Spike length (m

m)

Tiller per plant

Seed per spike

1000 seed w

eight (g)

Seed protein (%

)

Seed yield (g)

year1457.89ns897.82ns263.64ns6885.25**2648.99**69.47ns7796.79**707.22ns201.13*878.18**

Block/year3468.36144.421530.65558.2866.6426.72438.08752.7320.66286569.90

Treatment (line)

7366.34**81.04**17.02**222.20**265.25**3.87*112.29**42.12*1.53**10257.66**

Treat x year7334.01**33.84**11.97**38.17ns120.72**3.75*65.24ns17.99**0.92**8815.93ns

covariate15.67120.082.06214.50208.4455.42445.341.880.40205083.6

Error29210.9717.407.3245.4333.162.7367.3413.280.426747.75

Mean179.4164.82.6.687.670.55.752.433.111.2228.5

(S.E.)1.92.41.63.93.30.94.72.10.447.4

C.V. %1.82.51.37.78.229.015.611.05.836.0

Add. Var.(VA)2.693.930.4215.3412.040.013.932.010.05120.14

Env. Var. (VE)10.9717.407.3245.4333.162.7367.3413.280.406747.75

Phenotypic correlation coeffietients between Phenotypic correlation coeffietients between quantitative traits in barley doubled haploid linesquantitative traits in barley doubled haploid lines

TraitX1X2X3X4X5X6X7X8X9X10

X1Day to flowering1

X2Days to spike emergence0.755**

1

X3Days to maturity0.421**

0.399**

1

X4Plant height (cm)-0.398*-0.267*-0.1621

X5Spike length (mm)0.1160.057-0.0670.569**

1

X6Tiller per plant0.286*0.283*0.259*

0.1750.1141

X7Seed per spike0.1640.1900.189-0.0550.1060.1191

X81000 seed weight (g)-0.227-0.1740.0220.113-0.2260.1720.1061

X9Seed protein (%)0.432**

0.438**

-0.0910.2100.008-0.151-0.0640.0411

X10Seed yield (g)-0.037-0.0020.1220.325**

0.0580.447**

0.324**

0.307**-0.2081

Diversity, genetic gain and heitability for Diversity, genetic gain and heitability for agronomic traits in 72 barley DHs and their agronomic traits in 72 barley DHs and their

parentsparentsItem

Da

ys

to

flow

erin

g

Da

ys

to s

pik

e

em

erg

en

ce

Da

ys

to m

atu

rity

Pla

nt

he

igh

t (cm

)

Sp

ike

len

gth

(mm

Tille

r p

er p

lan

t

Se

ed

p

er s

pik

e

10

00

se

ed

we

igh

t (g)

Se

ed

p

rote

in (%

)

Se

ed

yie

ld g

Steptoe (P1)174.57158.22206.2791.2777.494.4052.7531.0812.50202.06

Morex(P2)182.30169.78207.9393.6768.986.7153.9139.6011.21321.93

P1-P2-7.93*-11.561.66ns-0.24ns8.51*-2.31 *-1.16ns-8.52 *1.29 *-119.87 *

XP=(P1+P2)/2178.44164.00207.1092.4773.245.5753.3335.3411.58262.57

X DHs179.45164.82206.5887.4570.425.6752.4033.1511.24227.57

X DHs - XP1.01ns0.82ns0.52ns5.02ns-2.82ns0.1ns0.93ns-2.19ns-0.61ns-34.43ns

Best DH172.41157.52201.98103.37

84.547.4264.6138.8412.25348.60

GGC=BDH-Bpb-2.16ns0.7ns-4.29 *9.70 *7.05 *0.71ns10.70 *-0.76ns0.25ns26.67ns

10%BDH173.45158.96206.5899.0483.267.0759.9233.1512.03302.89

GGd-10%BDH-BP-1.12ns0.74ns0.31ns5.37ns5.77ns0.36ns6.01ns-6.45 *0.47ns-19.04ns

h2f77.8975.9970.1589.2959.1162.4271.9278.9874.2465.94

LSDe6.606.583.066.996.522.199.144.801.08106.25

ns=non-significant

A= mean DHs

B= best parent

C= genetic gain (compar. Best DH with best parent)

D= genetic gain (compare. 10% best DHs with best parent)

E= least sig. diff.

F= heritability (from mean of two years using 2 GXE

T-values for comparisons between traits in T-values for comparisons between traits in irrigated(I) and non-irrogated(NI) replicationsirrigated(I) and non-irrogated(NI) replications

Comparison

Da

ys

to flo

we

ring

Da

ys

to s

pik

e e

me

rge

nc

e

Da

ys

to m

atu

rity

Pla

nt h

eig

ht (c

m)

Sp

ike

len

gth

(mm

)

Tille

r pe

r pla

nt

Se

ed

pe

r sp

ike

10

00

se

ed

we

igh

t (g)

Se

ed

pro

tein

( %)

Se

ed

yie

ld (g

)

I vs. NI (1st year)1.86ns1.55 ns11.96**2.87**0.10 ns4.60**2.26*2.39*3.52**2.42*

I vs. NI (2nd year)8.90**5.68**27.41**1.83 ns0.65 ns0.09 ns1.81 ns17.56**2.99 ns8.25**

I vs. NI (mean 2 years)9.35**5.29**32.85**3.22**0.48 ns4.06**0.46 ns12.26**4.62**7.87**

ANOVA results for traits under study with respect to ANOVA results for traits under study with respect to year and stress(drought)year and stress(drought)

ns,*,**,*** non- significant, significant at 0.05, 0.01 and 0.001, respectively

SOVdf

Days to

flow

ering

Days to

spike em

ergen

ce

Days to

matu

rity

Plan

t h

eigh

t (cm)

Sp

ike len

gth

(mm

)

Tiller

per p

lant

Seed

p

er spike

1000 seed

weig

ht (g

)

Seed

p

rotein

( %)

Seed

yield

(g)

Year (α)11722.3ns1691.1ns1458.3ns29965.5**4834.9***822.0*13933.6*1504.4ns686.1**662307.8 ns

Stress/ year (β)

2502.1***319.2**2705.7***233.3*4.1 ns35.2***175.0*1541.5***3.9***306719.4***

Error29221.025.46.662.783.92.851.717.40.66918.8

Number of QTLs recognizedNumber of QTLs recognized

Chromosome

1H2H3H4H5H7H

1st year(16 items)

LOD=2.03Sig.=0.039

45313-

2nd year(13 items)

LOD=2.04Sig.=0.024

24313-

Mean of 2 years(12 items)

LOD=2.59Sig.=0.035

34311-

I Year interaction(23 items)

LOD=2.23Sig.=0.032

574232

I Stress interaction(21 items)

LOD=2.79Sig.=0.027

364341

Locaions and effects of QTLs found for agronomic traits Locaions and effects of QTLs found for agronomic traits in 72 barley DH lines and their parents(Mean of 2 in 72 barley DH lines and their parents(Mean of 2

years)years)

+ According to Kosambi mapping (cM)

++ Proportion explained variance (r2)

TraitQTLLinkage group

Chromosome Location +

LOD% P.E.V.++Additive effects

Day To floweringflo-1H-11H1512.3918.72.6

flo-2H-12H806.2035.03.82

Days to spike emergencespi-2H-12H804.7229.63.72

Days to maturitymat-3H-13H613.7834.71.92

Plant height (cm)hei-3H-13H716.3637.3-7.25

Spike length (mm)spl-3H-13H7810.87

53.2-10.3

Tiller per plantspp-1H-11H1022.1216.90.57

spp-5H-15H575.7418.82.54

Seed per spikesps-2H-12H1045.2537.75.33

1000 seed weight (g)tsw-1H-11H1273.7226.1-2.55

Seed protein (%)prp-2H-12H673.3226.8-0.48

prp-4H-14H1482.4821.4-0.43

Locaions and effects of QTLs found for 10 agronomic traits in 72 Locaions and effects of QTLs found for 10 agronomic traits in 72 barley DH lines and their parents (in 2 years)barley DH lines and their parents (in 2 years)

+ According to Cosambi mapping (cM)

++ Proportion explained variance (r2)

TraitQTLLinkagegroup

ChromosomeLocation+LOD

++P.E.V. (%)Additive effects

2002200120022001

Day to flowering

flo-1H-11H1413.3214.015.33.371.54

flo-2H-12H808.4930.720.55.571.93

flo-3H-13H1093.7118.114.14.101.42

Days to spike emergence

spi-1H-11H1003.0814.914.33.831.52

spi-2H-12H756.97-26.318.35.431.85

spi-5H-15H505.15-15.519.13.671.91

spi-7H-17H1052.83-12.112.43.311.25

Days to maturity

mat-1H-11H1443.645.622.20.481.44

mat-3H-13H614.2726.912.02.650.95

Plant height (cm)

hei-2H-12H1863.9612.816.1-2.89-3.99

hei-3H-13H7010.6234.230.9-6.75-7.75

hei-4H-14H593.7215.311.93.953.94

Spike length (mm)

spl-2H-12H804.7920.814.12.876.49

spl-3H-13H7715.820.861.2-3.26-17.4

Tiller per plant

spp-1H-11H1182.705.120.2-0.041.12

spp-5H-15H506.0212.529.90.421.48

Seed per spike

sps-1H-11H756.2224.724.95.485.00

1000 seed weight (g)

tsw-1H-11H1185.3920.922.4-2.02-2.98

tsw-5H-15H665.7419.620.2-1.90-3.02

tsw-7H-17H1462.8614.613.91.552.43

Seed protein (%)

prp-2H-12H654.8010.226.0-0.26-0.63

prp-4H-14H1342.238.818.6-0.25-0.53

Seed yeild (g)

syi-2H-12H723.66.921.919.241.7

QTL on chromosome 1H

Date o

f sp

ike in

itiation

(17%

)

1000 seed

s weig

ht (22.4

%)

70.4

142.9

155.3

Year 1

31.1

Date o

f ma

turity(2

2.3%

)

41.6

158.8

Tiller/p

lant (2

0%

)67.3

151.6


Date o

f flow

ering

(16.3

%)

1000 seed

s weig

ht (23.8

%)

88.7

169.2 Year 2

62.0

169.2


Tiller/p

lant (2

0.2%

)

Date o

f flow

ering

(15.3%

)

1000 seed

s weig

ht (21

%)

148.6

75.1

169.2

63.6

154.8

85.6

Date o

f sp

ike in

itiation

(14.9%

)

48.8

167.1

39.1

158.8

Date o

f ma

turity(2

2.2%

)

GxE


Tillers

/pla

nt (16.9

%)

Date o

f flow

ering

(18.7

%)

1000 seed

s weig

ht (26.1

%)

83.7

160.0

86.1

169.2

54.4

164.6Mean


Date o

f Flo

werin

g (1

6.5%

)

1000 seed

s weig

ht (24.4

%)

169.2

76.1

149.9

Date o

f sp

ike in

itiation

(15.7%

)

46.8

Drought tolerance

100.7

169.2


Date o

f flow

ering

(22.3

%)

Date o

fs

pike

init.(1

9.5%

)

Seed

/spik

e(27.8

%)

35.0

101.4

125.5

Year 1

52.6

23.8

Seed

yield (2

1.9%

)121.5

29.2

121.8

Pro

tein p

ercentag

e(28.9

%)

42.2

87.0


46.5 Date o

f flow

ering

(31.5

%)

Date o

fs

pike

init.(2

8.1%

)

Seed

/spik

e(2

7.7%

)

42.6

94.5

129.6

111.9

Year 2

56.2

16.7

151.0

Sp

ike leng

th (21.2

%)


Date o

f flow

ering

(30.7

%)

Date o

fs

pike

init

(26.3

%)

Seed

/spik

e(2

4.9%

)

59.6

87.0

59.5

108.1

17.5

108.4

Pro

teinp

ercentag

e(2

6%

)

Seed

yield(2

1.5%

)

17.3

193.0

42.7

Plan

t heig

ht (16.1

%)

GxE

61.7

91.9

28.3

129.5

141.9

Sp

ike leng

th(20.8

%)


58.8

Date o

f flow

ering

(35%

)

Date o

fs

pike

init. (2

9.6%

)

Seed

/spik

e(3

7.7%

)

60.3

94.6

68.9

115.8

18.7

108.5

Pro

teinp

ercentag

e(2

6.8%

)

104.6

Mean

5


Date o

f flow

ering

(30.7

%)

Date o

fs

pike

initiatio

n

Seed

/spik

e(3

2.6%

)

59.6

87.0

108.6

Pro

teinp

ercentag

e(1

7.3%

)

Seed

yield(1

6.2%

)

209.5

Plan

t heig

ht (14.9

%)

Drought tolerance

62.9

91.7

124.9

19.9

85.4

26.5

90.1

59.1


86.1

50.5

Sp

ike leng

th (60.5

%)

Year 1

Plan

t heig

ht (31.7

%)

Date o

f flow

ering

(17.7%

)

126.1

59.4

74.6

43.0


126.1

98.9

41.1

Sp

ike leng

th (23.6

%)

Year 2

132.7

0.0

Plan

t heig

ht (36.3

%)

0.0

Date o

f ma

turity (2

7.9%

)

126.1


Plan

t heig

ht (34.2

%)

Sp

ike leng

th(61.1

%)

62.8

46.2

0.5

Date o

f flow

ering

(18.1%

)

Date o

f ma

turity(2

6.9%

)

161.9

34.3

50.3

85.2

GxE

113.0


Date o

f ma

turity

101.8

Plan

t heig

ht (37.3

%)

87.7

48.2

Sp

ike leng

th (53.2

%)

77.0

53.9

7.5

Mean

Date o

f ma

turity (3

4.7%

)

101.8


Plan

t heig

ht (36.4

%)

Sp

ike leng

th (56.6

%)

47.1

0.0

Date o

f flow

ering

(17.4

%)

Date o

f ma

turity (2

7.9%

)

188.8

50.8

87.8

16.8

64.3

Drought tolerance

113.5


Year 1

77.6

192.4

Pro

tein p

ercentag

e (61.5

%)


Year 2

3.4

128.0

Plan

t heig

ht (16.7

%)


Pro

tein p

ercentag

e (18.6

%)

13.9

112.4

Pla

nt h

eig

ht (1

5.3

%)

71.4

GxE

187.3


Pro

tein p

ercentag

e (21.4

%)

202.8

84.1

Mean


Pro

tein p

ercentag

e (23.7

%)

23.6

118.1

Pla

nt h

eig

ht (1

5.3

%)

75.9

Drought tolerance

206.8

93.0

171.2

Date o

f flow

ering

(16.6%

)


210.1

42.4

Year 1

114.8D

ate of s

pike

initiatio

n (19.9

%)

0.0

18.2

125.3

1000 seed

s weig

ht (22.1

%)

0.0

142.2T

iller/plan

t (30.5

%)


Tiller/p

lant (1

8.6%

)

210.1

42.4

Year 2

132.9

0.0

159.2

0.0

Date o

f sp

ike in

itiation

(16.9%

0)

10.2

116.3

1000 seed

s weig

ht (21

%)


Tiller/p

lant (2

9.9%

)

210.1

42.4

0.0

102.5

Date o

f sp

ike in

itiation

(26.4%

)

97.3

1.3

36.7

94.5

1000 seed

we

igh

t (20.5

%)

GxE


Tiller/p

lant (1

8.8%

)

210.1

42.4

42.4

Mean

210.1


Tiller/p

lant 27

.6%

)

210.1

42.4

0.0

132.8

Date o

f sp

ike in

itiation

(16.5%

)

113.9

0.0

39.0

91.4

1000 seed

we

igh

t (22.6

%)

Drought tolerance

0.0

122.1

Date o

f flow

ering

(16%

)


22.1

155.9

28.0

146.4

Date o

f sp

ike in

itiation

(12.4%

)

1000 seed

we

igh

t (14.6

%)

GxE

qtl analysis for yield and other agronomic traits in barley

Documents

different traits

quantitative traits

agronomic traits

mapping of quantitative

yield kghaproduction

genes qtls

genetic map

determination of qtls