qtl analysis for yield and other agronomic traits in barley
TRANSCRIPT
QTL ANALYSIS FOR YIELD AND QTL ANALYSIS FOR YIELD AND
OTHER AGRONOMIC TRAITS IN OTHER AGRONOMIC TRAITS IN
BARLEYBARLEY
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.
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
acid detergent fibre contentHan et al. 2003
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
resistance to net blotch and spot blotchSteffenson et al. 1996
Chromosometrait references
3H
resistance to leaf stripeArru et al. 2003
head shattering, plant height & grain yieldKandemir et al. 2000
grain yieldRomagosa et al. 1999
seed dormancyLarson et al. 1996
genetic variation in barley of crossability with wheatTaketa et al. 1998
partial resistance to bacterial leaf streakEl-Attari et al.1998
resistance to net blotch and spot blotchSteffenson et al. 1996
4H
acid detergent fibre contentHan et al. 2003
malting quality Han et al. 1997
salt tolerane at germination and the seedling stageMano et al. 1997
powdery mildew resistance Iwasa et al, 1997
resistance to net blotch and spot blotchSteffenson et al. 1996
green malt beta-glucanse activityHan et al. 1995
Chromosometrait references
5H
alpha-amylase activity of maltAyoub et al. 2003
resistance to leaf stripeArru 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
powdery mildew resistance Iwasa et al, 1997
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
genetic variation in barley of crossability with wheatTaketa et al. 1995
partial resistance to bacterial leaf streakEl-Attari et al.1998
Chromosometraitreferences
6H
salt tolerance at germination and the seedling stageMano et al. 1997
resistance to net blotch and spot blotchSteffenson et al. 1996
7H
genetic variation in barley of crossability with wheatTaketa et al. 1995
powdery mildew resistance Iwasa et al, 1997
green malt beta-glucanseHan et al.1995
containing malting quality Han et al. 1997
polymorphic microsatellite in the limit dextrinase gene
resistance to net blotch and spot blotchSteffenson et al. 1996
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
ChromosomeLocusMap Position
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
ChromosomeLocusMap Position
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
ChromosomeLocusMap Position
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
ChromosomeLocusMap Position
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
ChromosomeLocusMap Position
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
ChromosomeLocusMap Position
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
ChromosomeLocusMap Position
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
ChromosomeLocusMap Position
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
ChromosomeLocusMap Position
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
ChromosomeLocusMap Position
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
ChromosomeLocusMap Position
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
ChromosomeLocusMap Position
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
ChromosomeLocusMap Position
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
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
QTL on chromosome 1H
Date o
f flow
ering
(16.3
%)
1000 seed
s weig
ht (23.8
%)
88.7
169.2 Year 2
62.0
169.2
QTL on chromosome 1H
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
QTL on chromosome 1H
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
QTL on chromosome 1H
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
QTL on chromosome 2H
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
QTL on chromosome 2H
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
%)
QTL on chromosome 2H
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
%)
QTL on chromosome 2H
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
QTL on chromosome 2H
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
QTL on chromosome 3H
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
QTL on chromosome 3H
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
QTL on chromosome 3H
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
QTL on chromosome 3H
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
QTL on chromosome 3H
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
QTL on chromosome 4H
Pro
tein p
ercentag
e (18.6
%)
13.9
112.4
Pla
nt h
eig
ht (1
5.3
%)
71.4
GxE
187.3
QTL on chromosome 4H
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%
)
QTL on chromosome 5H
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
%)
QTL on chromosome 5H
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
%)
QTL on chromosome 5H
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
QTL on chromosome 5H
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%
)