Published online November 21, 2006
Comparative Mapping of Growth Habit, Plant Height, and Flowering QTLs in Two
Interspecific Families of Leymus
Steven R. Larson,* Xiaolei Wu, Thomas A. Jones, Kevin B. Jensen, N. Jerry Chatterton, Blair L. Waldron,
Joseph G. Robins, B. Shaun Bushman, and Antonio J. Palazzo
Aggressive rhizomes and adaptation to poorly drained
alkaline sites, primarily within the western USA, characterize sod-forming L. triticoides (0.3–0.7 m). Conversely,
L. cinereus is a tall (up to 2 m) conspicuous bunchgrass
adapted to deep well-drained soils from Saskatchewan to
British Columbia, south to California, northern Arizona,
and New Mexico, and east to South Dakota and Minnesota. Most populations of L. cinereus and L. triticoides are
allotetraploids; however, octoploid forms of L. cinereus
are typical in the Pacific Northwest. Basin wildrye, and
octoploid giant wildrye [L. condensatus (J. Presl) Á.
Löve], are the largest cool-season bunch grasses native to
western North America. Artificial hybrids of L. cinereus,
L. triticoides, and other North American Leymus wildryes
display regular meiosis and stainable pollen (Stebbins
and Walters, 1949; Dewey, 1972; Hole et al., 1999). Both
L. cinereus and L. triticoides are highly self-sterile (Jensen
et al., 1990) and hybridize with each other in nature.
These species are naturally important forage and hay
grasses in the Great Basin and other regions of western
North America.
Growth habit is a highly variable and ecologically
important trait in perennial grasses. Aggressive rhizomes characterize quackgrass [Elymus repens (L.) Desv.
ex Nevski] and johnsongrass [Sorghum bicolor (L.)
Moench], which rank among the world’s worst perennial
grass weeds (Holm et al., 1977). In general, caespitose
(i.e., growing in bunches or tufts) and rhizomatous grasses
dominate semiarid and mesic grasslands, respectively
(Sims et al., 1978). Nutrient islands beneath caespitose
grasses may also contribute to clone fitness in this growth
form in both mesic and semiarid communities, whereas
the distribution of rhizomatous grasses may be restricted
to microsites characterized by higher soil organic carbon
and nitrogen concentrations (Derner and Briske, 2001).
We have observed L. cinereus and L. triticoides growing in
close proximity in mixed stands, at several disturbed sites,
with no apparent difference in microhabitat. Conversely,
we have observed L. triticoides in riparian or wet zones
and L. cinereus inhabiting dry adjacent uplands, restricted
to seemingly different natural microhabitats. In any case,
L. cinereus and L. triticoides display profound differences
in growth habit. Lateral branches of L. cinereus grow
strictly upward, often within the lower leaf sheaths as
tillers, whereas the lateral branches of L. triticoides frequently grow horizontally or downward under the soil
surface as rhizomes. In addition to obvious morphological
Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved.
ABSTRACT
Leymus cinereus (Scribn. & Merr.) Á. Löve and L. triticoides
(Buckley) Pilg. are tall caespitose and short rhizomatous perennial
Triticeae grasses, respectively. Circumference of rhizome spreading,
proportion of bolting culms, anthesis date, and plant height were evaluated in two mapping families derived from two interspecific hybrids of
L. cinereus Acc:636 and L. triticoides Acc:641 accessions, backcrossed
to one L. triticoides tester. Two circumference, two bolting, and two
height QTLs were homologous between families. Two circumference,
seven bolting, all five anthesis date, and five height QTLs were family
specific. Thus, substantial QTL variation was apparent within and between natural source populations of these species. Two of the four
circumference QTLs were detected in homoeologous regions of linkage
groups 3a and 3b in both families, indicating that one gene may control
much of the dramatic difference in growth habit between these species.
A major height QTL detected in both families may correspond with
dwarfing mutations on barley 2H and wheat 2A. The L. cinereus parent
contributed negative alleles for all four circumference QTLs, five of
nine bolting QTLs, two of five anthesis date QTLs, and one of seven
height QTLs. Coupling of synergistic QTL allele effects within parental
species was consistent with the divergent growth habit and plant height
of L. cinereus and L. triticoides. Conversely, antagonistic QTL alleles
evidently caused transgressive segregation in reproductive bolting and
flowering time.
L
wildryes are long-lived Triticeae grasses,
closely related to wheat (Triticum spp.) and barley
(Hordeum vulgare L.). The genus Leymus includes
about 30 species distributed throughout temperate regions of Europe, Asia, and the Americas (Dewey, 1984).
More than half of all Leymus species are allotetraploids
(2n 5 4x 5 28), but octoploid (2n 5 8x 5 56) and dodecaploid (2n 5 12x 5 84) variants may arise from interspecific hybrids (Anamthawat-Jónsson and Bödvarsdóttir,
2001) or autoduplication within species. These coolseason grasses display remarkable variation in growth
habit and stature with unusual adaptation to harsh
polar, desert, saline, and erosion-prone environments.
Leymus triticoides (creeping or beardless wildrye) and
L. cinereus (basin wildrye) are closely related but morphologically divergent North American range grasses.
EYMUS
S.R. Larson, T.A. Jones, K.B. Jensen, N.J. Chatterton, B.L. Waldron,
J.G. Robins, and B.S. Bushman, USDA-ARS, Forage and Range
Research Lab., Utah State Univ., Logan, UT 84322-6300; X. Wu,
Univ. of Missouri-Columbia, Columbia, MO 65211; A.J. Palazzo, U.S.
Army Corps of Engineers, Engineering Research and Development
Center-Cold Regions Research and Engineering Lab., Hanover, NH
03755-1290. Received 13 Dec. 2005. *Corresponding author (stlarson@
cc.usu.edu).
Abbreviations: AFLP, amplified fragment length polymorphism;
ANTH, anthesis date; BOLT, reproductive bolting; CIRC, plant circumference; HGHT, plant height; IM, interval mapping; MQM, multipleQTL model; QTL, quantitative trail locus (i); RFLP, restriction fragment
length polymorphism; rMQM, restricted MQM mapping; STS, sequencetagged site; SSR, simple-sequence repeat.
Published in Crop Sci. 46:2526–2539 (2006).
Genomics, Molecular Genetics & Biotechnology
doi:10.2135/cropsci2005.12.0472
ª Crop Science Society of America
677 S. Segoe Rd., Madison, WI 53711 USA
2526
2527
Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved.
LARSON ET AL.: LEYMUS WILDRYES
differences in growth habit and stature, L. cinereus and L.
triticoides display differences in salt tolerance, seed dormancy, seed color, seed shattering, mineral content, tillering, texture, and other characteristics.
The F1 hybrids of L. cinereus and L. triticoides are
robust and vigorous plants with relatively large biomass
potential compared to other range grasses of western
North America. Wu et al. (2003) recently constructed
high-density molecular genetic linkage maps for two
full-sib families, TTC1 and TTC2. The 164-sib TTC1 and
170-sib TTC2 families were derived from two different
L. triticoides 3 L. cinereus F1 hybrids, TC1 and TC2,
crossed to different clones of the same L. triticoides
tester genotype (T–tester). The TC1 and TC2 F1 hybrids
were derived from naturally heterogeneous L. triticoides
and L. cinereus seed accessions from Oregon and Alberta, respectively. Together, the TTC1 and TTC2 families capture a snippet of the divergence among and
heterogeneity within the L. triticoides and L. cinereus
source populations. These two experimental populations
provide unique system for gene discovery research and
development of breeding markers in perennial grasses.
A close relative of quackgrass, Leymus also provides a
useful system to study weedy traits such as rhizomes.
Our primary objective here was to identify, characterize, and compare QTLs controlling growth habit, plant
height, and flowering traits. Another major objective
was to compare the location of these Leymus QTLs with
genomic regions controlling related traits in other cereal
and grass species.
MATERIALS AND METHODS
Plant Materials and Genetic Maps
The full-sib TTC1 and TTC2 molecular genetic maps and
pedigrees were described by Wu et al. (2003). The 164-sib
TTC1 map included 1069 AFLP markers and 38 anchor loci in
14 linkage groups spanning 2001 cM. The 170-sib TTC2 map
contained 1002 AFLP markers and 36 anchor loci in 14 linkage
groups spanning 2066 cM. Some 488 homologous AFLP loci
and 24 anchor markers, detected in both families, showed similar map order among 14 homologous linkage groups of the
TTC1 and TTC2 families (Fig. 1). The 14 homologous linkage
groups of the allotetraploid TTC1 and TTC2 families were
tentatively numbered according to the seven homoeolgous
groups of the Triticeae grasses on the basis of synteny of two
or more anchor markers from each of the seven homoeologous
groups of wheat and barley. Moreover, genome-specific STS
markers were used to distinguish the Ns and Xm marker sequences for homoeologous groups four and five on the basis of
similarity to Psathyrostachys juncea (Fisch.) Nevski (genome
designation Ns), which is one of the diploid ancestors of allotetraploid Leymus (Wu et al., 2003). Otherwise, homoeologous chromosomes of allotetraploid Leymus were arbitrarily
distinguished by the letters “a” or “b.” For example, LG3a is
allegedly homoeologous with LG3b (Fig. 1), an assertion supported by synteny among 10 of the 11 anchor markers mapped
to these linkage groups (Wu et al., 2003). Additional SSR and
STS anchor markers described in the results below were
analyzed by methods described by Wu et al. (2003). Detailed
genetic maps containing all 1583 AFLP markers mapped in
the TTC1 and/or TTC2 families are shown in Supplementary
Data files online. For essential comparisons of homology
between the Leymus TTC1 and TTC2 families and comparisons of homoeology with other species, we showed all anchor
makers but only those AFLP markers that were detected (homologous) in both TTC1 and TTC2 families (Fig. 1).
Field Evaluations
Ramets from each of the two mapping families (TTC1 and
TTC2) were space planted in a randomized complete block
(RCB) design with two replicates (blocks) per family at the
Utah Agriculture Experiment Station Richmond Farm (Cache
Co., UT). Each block contained one entry of each of the 164
TTC1 siblings or one clone from each of the 170 TTC2 siblings
plus the parental clones (i.e., TC1, TC2 and T) and single-plant
representatives of the heterogeneous L. cinereus Acc:636 and
L. triticoides Acc:641 accessions. Individual ramets were
transplanted from soil containers (4-cm diameter) in the
spring of 2001 to field plots with 2-meter row spacing and 2meter spacing within rows (2-m centers). Plants were aligned
among rows such that each plant had four equidistant (2 m)
neighboring plants. Each row contained 34 centers (one plant
per center), thus blocks were restricted to six or seven rows.
The TTC1 rep1, TTC2 rep1, TTC1 rep2, and TTC2 rep2
blocks were arranged lengthwise, respectively, along the 66-m
rows to form one continuous array comprised of 24 3 34
centers (46 3 66 m). Quantitative traits, described below, were
measured in 2002, 2003, and 2004.
Rhizome proliferation was measured as plant circumference
(CIRC) in late spring or early summer. For caespitose plants,
this could be simply measured by stretching a tape ruler
around the tussock at the soil surface. The circumference of
irregular sods, characteristic of the more rhizomatous plants,
was approximated as the perimeter of a polygon where the
corners represent the outermost rhizome branches (i.e., the
shortest distance around the outermost rhizomes) at the soil
surface. Anthesis date (ANTH) was measured as the number
of days from 1 January until the first day of anther extrusion,
which is most apparent on warm dry mornings, beginning midJune. In practice, the first day of anther extrusion was interpolated between two or three observations per week. A
priori, one unexpected phenomenon in the TTC1 and TTC2
families was variation in reproductive bolting (BOLT), where
some genotypes flowered and some did not, first observed in
2002. Individual clones were retrospectively rated as 0 (not
flowered) or one (flowered), after seed harvest and mowing,
based on ANTH notes take in 2002. We subsequently rated
individual plants for the proportion of bolting culms from 0 (no
spikes produced) to 1 (virtually all major culms bolting), before seed harvests and mowing, in 2003 and 2004. Plant height
(HGHT)as measured after anthesis using a 2-m ruler. Plant
height (i.e., actual culm length from soil surface to the spike
terminal) was not measured on plants that did not flower.
Data Analysis
QTL analyses were based on trait means from two
replicants for each of the 164 TTC1 and 170 TTC2 segregates
in 2002, 2003, and 2004. QTL analyses were also performed on
averages over all 3 yr, using output from the LSMEANS procedure of SAS (Statistical Analysis Systems Institute Inc.,
Cary, NC). Coefficients of skewness and kurtosis, g1, and g2
were calculated as described by Snedecor and Cochran (1980).
Departures from normality were deemed significant if they
exceeded two standard errors of skewness (SES) and kurtosis (SEK) estimated by (6/N)22 and (24/N)22, respectively (Tabachnick and Fidell, 1996). Histograms of phenotypic values
Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved.
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CROP SCIENCE, VOL. 46, NOVEMBER–DECEMBER 2006
(clonal averages) from 2002, 2003, and 2004 were also prepared
for CIRC, BOLT, ANTH, and HGHT (Supplementary Data).
Broad-sense heritabilities within years were obtained using a SAS program for estimating heritability from lines
evaluated in an RCB design in a single environment (Holland
et al., 2003). Likewise, heritabilities among years were obtained using another SAS program for estimating heritability
from lines evaluated in RCB designs in multiple environments
(Holland et al., 2003), modified to account for repeated measurements on perennial plants. All class variables (i.e., rep,
clone, year) were treated as random effects. Genotypic and
phenotypic correlations between traits evaluated using a SAS
program for estimating correlations from RCB designs in
multiple environments (Holland et al., 2003), also modified to
account for repeated measurements. The original SAS
programs for estimating heritabilities, genotypic correlations,
and phenotypic correlations (Holland et al., 2003) are
available at http://www4.ncsu.edu/|jholland/homepage_files/
Page571.htm (verified 24 July 2006).
A sequential and reiterative procedure of QTL detection
was performed using the MAPQTL five package (Van Ooijen,
2004). Genome-wide interval mapping (IM) (Lander and
Botstein, 1989; Van Ooijen, 1992) was performed in 1-cM increments to identify putative QTLs and possible cofactors
Fig. 1. Continued on next page.
used in a multiple-QTL model (MQM) (Jansen, 1993; Jansen,
1994; Jansen and Stam, 1994). Markers with the highest loglikelihood ratios (i.e., LOD test statistics) for each QTL (no
more than one per chromosome) were selected as the initial
set of possible MQM cofactors. A backward elimination procedure was applied to this initial set of cofactors using a
conservative significance level of 0.001 to ensure the independence of each cofactor. A reiterative process of restricted
MQM (rMQM) mapping, which excludes any syntenous cofactors (i.e., cofactors located on the same linkage group that
is being scanned), was used to refine the location of rMQM
cofactors (QTLs) and identify new rMQM cofactors (QTLs).
Moreover conventional MQM mapping was used to detect (or
exclude) syntenous QTLs, where that possibility was apparent.
Finally, all putative QTLs were also evaluated using the nonparametric Kruskal–Wallis rank sum test, equivalent to the
two-sided Wilcoxon rank sum test for two genotype classes
(this study), which means that no assumptions are being made
for the probability distributions of the quantitative traits
(Lehmann, 1975).
A threshold of 3.3 LOD was used throughout these QTL,
MQM, and rMQM detection procedures as a close approximation for a genome-wide 5% significance level as determined
from simulation tables based on genome size and family type
Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved.
LARSON ET AL.: LEYMUS WILDRYES
Fig. 1. Continued on next page.
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2530
Fig. 1. Continued on next page.
CROP SCIENCE, VOL. 46, NOVEMBER–DECEMBER 2006
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Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved.
LARSON ET AL.: LEYMUS WILDRYES
Fig. 1. Summary and comparison of plant height, growth habit, and flowering QTLs detected in the full-sib Leymus triticoides 3 (L. triticoides 3 L.
cinereus) TTC1 and TTC2 families based on 2002, 2003, 2004, and average measurements. The approximate location of QTLeffects (LOD 3.3),
corresponding to a genome-wide P # 0.05 signficance level, detected using a restricted multiple QTL model (rMQM) are indicated by black or
white bars for each trait 3 year. The updated molecular genetic linkage maps include 488 homologous AFLP markers and mapped in both
TTC1 and TTC2 families (Wu et al., 2003), 50 anchor markers (larger bold marker text) mapped in TTC1 and/or TTC2 families (Wu et al., 2003),
and 17 additional anchor markers (larger bold italic marker text) mapped in the TTC1 and/or TTC2 families. Annotation next to each anchor
marker indicate homoeologous groups of barley (H), wheat (ABD), cereal rye (R), and in parentheses oat chromosome designations.
Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved.
2532
CROP SCIENCE, VOL. 46, NOVEMBER–DECEMBER 2006
(Van Ooijen, 1999) and empirical thresholds based on permutation analyses with 1000 replications (Churchill and Doerge,
1994) specific to each trait. Both methods (Van Ooijen, 1999;
Churchill and Doerge, 1994) produced remarkably similar results for all four traits evaluated in this study.
Genome-wide IM and rMQM LOD scans based on 2002,
2003, 2004, and average phenotypic values were performed for
CIRC, BOLT, ANTH, and HGHT using MapChart version 2.1
(Voorrips, 2002) and the updated Leymus TTC1 and TTC2
genetic maps as described in the results section below. Detailed
graphs of each genome-wide LOD scan using the complete
high-density linkage maps are provided in the Supplementary
Data Online.
RESULTS
Molecular Map Update
A subset of 48 CSU or KSU wheat SSR primer pairs
described by Yu et al. (2004), the HVCABG SSR
primers described by Becker and Heun (1995), and the
HVM068 SSR primers described by Liu et al. (1996)
were tested for amplification and polymorphism among
the parental genotypes. Seven of these SSR primer pairs
(CNL45, KSU154, CNL39, HVCABG, HVN068,
KSU149, and KSU171) detected 11 loci in the Leymus
TTC1 and/or TTC2 families (Fig. 1). Likewise, another
subset of 48 published STS primer pair sequences (Mano
et al., 1999; Taylor et al., 2001; Lem and Lallemand,
2003), three sets of STS primers designed from the wheat
VRN2 (Yan et al., 2004), and one set of STS primers
designed from the BCD1117 barley cDNA clone (Heun
et al., 1991) were also tested for amplification and polymorphism among the parental genotypes. Seven of these
STS primer pairs (TRX1, TRX2, BCD1117, MWG2230,
VRN2, MWG2264, and ADP) detected 13 loci in the
Leymus TTC1 and/or TTC2 families (Fig. 1).
The thioredoxin h RFLP marker, Xbm2, maps near the
self-incompatibility gene locus, S, on homeologous group
1R of Secale (Korzun et al., 2001), and thioredoxin h STS
markers have been utilized in perennial ryegrass (Taylor
et al., 2001) and Kentucky bluegrass (Patterson et al.,
2005). The thioredoxin h STS primers used by Patterson
et al. (2005) amplified two distinct sets of sequences,
designated TRX1 (AY943821 and AY943822 from L. cinereus Acc:636; AY943823 and AY943824 from L. triticoides Acc:641) and TRX2 (AY943825-AY943827 from
L. cinereus Acc:636; AY943828-AY943830 from L. triticoides Acc:641). We designed TRX1 and TRX2 primers
that would amplify corresponding sequences from L. cinereus but not L. triticoides—thus providing informative polymorphisms for mapping in the TTC1 and TTC2 backcross
families. The TRX2 amplicons mapped to homologous positions of the LG6b linkage group, evident by the coliniearity with homologous TTC1 and TTC2 AFLP markers
(Fig. 1). The TRX1 primers amplified 289 bp sequences
that mapped to LG1a in TTC1 and LG2b in TTC2 (Fig. 1).
The TRX1 locus on LG1a is presumably homeologous to
the Xbm2 locus on Secale 1R. The role of thioredoxin
h genes in seed germination, self-incompatibility, grain
quality, and characteristics has been evaluated in cereals,
grasses, and other plants (Juttner et al., 2000; Langridge
et al., 1999; Li et al., 1994, 1997; Sahrawy et al., 1996).
Of special interest with regard to flowering traits, the
VRN1 and VRN2 genes are the two most potent genes
controlling differences in vernalization requirement between winter annual and spring annual barley, wheat,
and rye (Dubcovsky et al., 1998), and both of these genes
have been cloned (Yan et al., 2003, 2004). The VRN2
gene is present on homoeologous regions of 4H in winter
barleys (evidently absent in most spring barleys) and a
4A/5A translocation region of Triticum monococcum 5A
(Dubcovsky et al., 1998; Yan et al., 2004). Primers
designed from the wheat VRN2 gene amplified mixed
sequences from Leymus that were very similar in size
(supplemental data) and sequence (DQ486013) to the
wheat VRN2 gene. These putative Leymus VRN2 sequences map to a locus on the distal long arm of Leymus
5Ns (Fig. 1), which evidently corresponds to the VRN2
locus on Triticum monococcum 5A. Although we have
not yet mapped the Leymus VRN1 sequences, this gene
is closely associated with the CDO504 marker in other
Triticeae species and also maps to Leymus LG5Ns and
LG5Xm (Fig. 1) (Wu et al., 2003).
In summary, seven SSR primer pairs and seven STS
primer pairs (supplemental data) detected 15 additional
anchor loci in the TTC1 family and nine additional
anchor loci in the TTC2 family, not previously described
by Wu et al. (2003). The updated TTC1 map includes
1069 AFLP markers and 53 anchor loci in 14 linkage
groups spanning 2001 cM. The updated TTC2 map contained 1002 AFLP markers and 45 anchor loci in 14
linkage groups spanning 2066 cM. Some 488 homologous AFLP loci and 31 anchor markers have been
mapped in both families, showing similar map order.
Thus, 1583 AFLP markers and 67 different anchor loci
have been mapped into 14 linkage groups, which evidently correspond to the 14 chromosome pairs of allotetraploid Leymus (2n 5 4x 5 28). The map locations
of the 17 new anchor loci (24 total including seven
homologous pairs) (Fig. 1) were largely consistent with
the other anchor loci previously mapped in Leymus (Wu
et al., 2003).
Growth Habit
The L. cinereus Acc:636 and L. triticoides Acc:641
progenitor accessions displayed very different levels
of rhizomatous spreading, which became more pronounced each year (Table 1). Although CIRC measurements of the TC1 and TC2 F1 hybrids were significantly
greater than L. cinereus Acc:636, these means were wellbelow the midparent, TTC1, and TTC2 averages especially in 2004 (Table 1).
Circumference of plant (rhizome) spreading showed
relatively strong broad-sense heritabilities in the TTC1
family and moderate heritabilities in the more rhizomatous TTC2 family (Table 2). The CIRC trait also showed
especially strong heritabilities over years, in both TTC1
and TTC2 families; however, this is expected because
plant spreading is a cumulative trait. Not surprisingly,
the ratio of genotype 3 year to phenotypic variance was
negligible (0.02) for the CIRC trait. The CIRC trait was
weakly correlated with the BOLT and ANTH flowering
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LARSON ET AL.: LEYMUS WILDRYES
Table 1. Trait means 6 SD for circumference of plant spreading (CIRC), proportion of bolting culms (BOLT), anthesis date (ANTH), and
plantheight (HGHT) for Leymus cinereus Acc:636, L. triticoides Acc:641, L. triticoides T parental genotype, interspecific TC1 and TC2
parental hybrids, and full-sib L. triticoides 3 (L. triticoides 3 L. cinereus) TTC1 and TTC2 mapping families.
Trait
Year
Acc:636 (n 5 18)†
Acc:641 (n 5 15)†
t tester (n 5 13)‡
TC1 (n 5 14)‡
TC2 (n 5 14)‡
TTC1 (n 5 164)§
CIRC
2002
2003
2004
Avg.
2002
2003
2004
Avg.
2002
2003
2004
Avg.
2002
2003
2004
Avg.
23 6 10
51 6 26
76 6 21
57 6 23
0.28 6 0.46
0.63 6 0.43
0.85 6 0.33
0.60 6 0.58
176.8 6 2.1
171.1 6 5.1
178.3 6 3.9
175.9 6 2.4
99.0 6 39.8
135.6 6 23.0
182.9 6 18.5
142.2 6 26.0
354 6 186
594 6 326
816 6 416
702 6 208
0.67 6 0.48
0.56 6 0.32
0.78 6 0.35
0.74 6 0.24
173.3 6 2.6
169.3 6 1.4
172.6 6 1.6
172.2 6 1.1
94.5 6 12.6
98.5 6 8.5
123.3 6 12.7
107.3 6 8.7
487 6 61
853 6 120
1263 6 177
653 6 112
1.00 6 0.00
0.98 6 0.06
1.00 6 0.00
0.99 6 0.03
172.0 6 0.0
169.0 6 0.0
173.0 6 0.0
171.3 6 0.0
100.8 6 5.7
100.9 6 5.9
125.7 6 3.4
109.2 6 2.4
96 6 35
200 6 32
256 6 39
183 6 28
0.79 6 0.43
0.88 6 0.19
0.84 6 0.19
0.83 6 0.14
173.6 6 2.1
169.0 6 0.0
175.9 6 4.0
172.8 6 1.3
145.8 6 15.9
155.1 6 12.6
184.7 6 12.3
161.8 6 11.2
125 6 37
231 6 38
282 6 53
212 6 40
1.00 6 0.00
0.77 6 0.21
0.81 6 0.23
0.85 6 0.09
173.1 6 1.5
169.0 6 0.0
175.9 6 4.0
172.6 6 1.5
141.8 6 10.6
138.4 6 13.3
164.6 6 8.7
148.3 6 8.3
180 6 91
288 6 122
362 6 137
273 6 112
0.57 6 0.43
0.33 6 0.24
0.55 6 0.26
0.49 6 0.26
173.5 6 1.9
172.1 6 5.7
175.7 6 4.6
174.5 6 3.5
103.9 6 13.8
108.3 6 12.9
134.8 6 13.8
116.1 6 12.5
Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved.
BOLT
ANTH
HGHT
TTC2 (n 5 168)§
299
449
548
430
0.73
0.36
0.67
0.59
173.8
171.3
174.1
173.4
109.7
106.1
137.8
117.9
6 116
6 154
6 170
6 144
6 0.38
6 0.25
6 0.27
6 0.26
6 4.8
6 4.9
6 3.7
6 4.0
6 13.0
6 12.4
6 11.8
6 10.7
Measurements expressed in cm (CIRC and HGHT), proportion (BOLT), and days from January one (ANTH).
† Sample size based on individual plants.
‡ Sample size based on individual clones of each parental genotype.
§ Sample size based on means of two clones for each genotype (n).
traits, in the TTC1 and/or TTC2 families, but virtually
independent from plant height in both TTC1 and TTC2
families (Table 3).
At least three significant CIRC QTLs were detected
in each family; however, not more than two QTLs were
significant in any one year and population (Table 4,
Fig. 1). The strongest, most consistent CIRC QTLs in
both TTC1 and TTC2 families were located in homologous regions of LG3a (Table 4, Fig. 1). Likewise, homologous CIRC QTLs were also detected on LG3b in both
TTC1 and TTC2 families (Fig. 1). The LG3a and LG3b
CIRC QTLs, detected in both TTC1 and TTC2 families,
are all located near homoeologous copies of VP1 gene
markers (Fig. 1). The transcription factor Viviparous-1
(encoded by the Vp1 gene) induces and maintains seed
dormancy (McCarty et al., 1991) and has been mapped
to orthologous loci in wheat, maize, and rice (Bailey et al.,
1999).Inadditiontothehomologousandpossiblyhomoeologous QTLs detected on LG3a and LG3b, in both TTC1
and TTC2 families, a TTC1-specific CIRC QTL was detected on LG6a and a TTC2-specific CIRC QTL was detected on LG5Xm (Fig. 1). Thus, two CIRC QTLs were
common to both TTC1 and TTC2 families and two were
unique to only one family. The L. cinereus alleles had negative effects at all four (LG3a, LG3b, LG5Xm, and LG6a)
CIRC QTLs (Table 4, Fig. 1) consistent with divergent
phenotypes of the parental species.
Although significant skewness was detected in several
evaluations, all four CIRC QTLs (Table 4, Fig. 1) were
highly significant (P , 0.0001) on the basis of the nonparametric Kruskal–Wallis rank sum test.
Flowering Traits
The BOLT and ANTH traits showed relatively strong
negative genotypic correlations in both TTC1 and TTC2
families (Table 3), evidently controlled by several common
(probably pleiotropic) QTLs (Fig. 1). Thus, data from
these two traits are presented together in one section.
The L. cinereus Acc:636 and L. triticoides Acc:641 progenitor accessions; F1 interspecific hybrids (TC1 and
TC2); and TTC1 and TTC2 families displayed similar
BOLT and ANTH phenotypic means (Table 1), except
for the fact that L. cinereus showed relatively weak bolting in the first establishment year. Leymus cinereus has a
relatively large stature, requiring several years to reach
reproductive maturity. Yet, the TTC1 and TTC2 families
displayed transgressive BOLT and ANTH segregation
that persisted through 2002, 2003, and 2004 (see trait
distributions in Supplemental Data). Unlike the F1 hybrids or parental source populations, some transgressive
progeny failed to flower in 2002, 2003, 2004, and 2005.
The BOLT and ANTH traits both showed relatively
weak plot-mean heritabilities over years (Table 2). The
ratio of genotype 3 year interaction to phenotypic variance was substantially greater for ANTH (0.18) and
BOLT (0.21) compared with CIRC (0.02) or HGHT
(0.13), which explains the low plot-mean heritabilities
Table 2. Estimates of broad-sense heritability (H 2) 6 SE based on a plot basis and, in parentheses, genotypic mean basis for circumference
of plant spreading (CIRC), proportion of bolting culms (BOLT), anthesis date (ANTH), and plant height (HGHT) in the full-sib Leymus
triticoides 3 (L. triticoides 3 L. cinereus) TTC1 and TTC2 mapping families.
Trait
Family
2002
2003
2004
Over years
CIRC
TTC1
TTC2
TTC1
TTC2
TTC1
TTC2
TTC1
TTC2
0.72 6 0.04 (0.84 6 0.03)
0.54 6 0.06 (0.70 6 0.05)
0.44 6 0.06 (0.61 6 0.06)
0.43 6 0.06 (0.61 6 0.06)
0.50 6 0.07 (0.67 6 0.07)
0.73 6 0.04 (0.84 6 0.03)
0.69 6 0.05 (0.81 6 0.03)
0.66 6 0.05 (0.79 6 0.03)
0.77 6 0.03 (0.87 6 0.02)
0.55 6 0.06 (0.71 6 0.05)
0.63 6 0.05 (0.77 6 0.04)
0.64 6 0.05 (0.78 6 0.04)
0.42 6 0.08 (0.59 6 0.08)
0.43 6 0.08 (0.60 6 0.07)
0.51 6 0.07 (0.68 6 0.06)
0.53 6 0.06 (0.69 6 0.06)
0.73 6 0.04 (0.85 6 0.02)
0.57 6 0.05 (0.73 6 0.04)
0.64 6 0.05 (0.77 6 0.04)
0.69 6 0.04 (0.81 6 0.03)
0.51 6 0.06 (0.67 6 0.06)
0.46 6 0.07 (0.63 6 0.06)
0.73 6 0.04 (0.84 6 0.03)
0.59 6 0.05 (0.75 6 0.04)
0.64 6 0.04 (0.91 6 0.01)
0.50 6 0.05 (0.86 6 0.03)
0.38 6 0.04 (0.76 6 0.04)
0.41 6 0.04 (0.78 6 0.03)
0.31 6 0.05 (0.71 6 0.05)
0.41 6 0.04 (0.78 6 0.03)
0.54 6 0.04 (0.86 6 0.02)
0.48 6 0.04 (0.82 6 0.03)
BOLT
ANTH
HGHT
2534
CROP SCIENCE, VOL. 46, NOVEMBER–DECEMBER 2006
Table 3. Genotypic and in parentheses phenotypic trait correlations 6 SE for circumference of plant spreading (CIRC), proportion of
bolting culms (BOLT), anthesis date (ANTH), and plant height (HGHT) in the full-sib Leymus triticoides (L. triticoides 3 L. cinereus)
TTC1 (below diagonal) and TTC2 (above diagonal) families.
Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved.
CIRC
BOLT
ANTH
HGHT
CIRC
BOLT
ANTH
HGHT
—
0.22 6 0.08 (0.15 6 0.05)
20.33 6 0.09 (20.22 6 0.05)
0.01 6 0.09 (0.04 6 0.06)
0.24 6 0.09 (0.13 6 0.05)
—
20.81 6 0.06 (20.48 6 0.03)
0.32 6 0.09 (0.40 6 0.04)
20.22 6 0.09 (20.10 6 0.05)
20.83 6 0.04 (20.56 6 0.03)
—
0.07 6 0.11 (20.19 6 0.05)
0.12 6 0.09 (0.02 6 0.05)
0.32 6 0.09 (0.36 6 0.04)
20.26 6 0.09 (20.32 6 0.04)
—
over years for the ANTH and BOLT flowering traits.
Although, BOLT and ANTH traits show significant genotype 3 year interaction and the significance of QTLs
vary by year (Tables 4), several QTLs were detectable
over years (Fig. 1). One notable exception was a highly
significant TTC2 ANTH QTL on LG6a in 2002, which
was virtually nonexistent in subsequent years. Bolting
deficiencies were strongly correlated with late flowering
and weakly correlated with short caespitose characteristics (Table 3).
A total of nine BOLT QTLs were detected in the
TTC1 and/or TTC2 families (Table 4, Fig. 1). The TTC1
and TTC2 BOLT QTL peaks on the upper portion of
LG4Xm overlap well enough (Fig. 1) that we counted
these as one homologous QTL. Likewise, we believe that
there is insufficient separation of the TTC1 and TTC2
BOLT QTLs on LG6a to declare these as different (Fig. 1).
Thus, L. cinereus contributed five negative TTC1 and/or
TTC2 BOLT QTL alleles and four positive TTC1 and/or
TTC2 BOLT QTL alleles (Table 4, Fig. 1). Two of these
nine BOLT QTLs were significant in both TTC1 and
TTC2 families (i.e., homologous), whereas seven QTLs
were unique to TTC1 or TTC2. Four QTLs explained up
to 43.9% of the BOLT variation in the 2003 TTC2 data
set, which was the most explanatory QTL model in this
study (Table 4).
A total of five significant ANTH QTLs were detected
in the TTC1 or TTC2 families (Table 4, Fig. 1). All five
significant ANTH QTLs were unique to TTC1 or TTC2
(Table 4 and Fig. 1). However, the TTC1 family displayed
Table 4. Summary of QTL effects detected using multiple QTL model (MQM) scans of circumference of plant spreading (CIRC),
proportion of bolting culms (BOLT), anthesis date (ANTH), and plant height (HGHT) in the full-sib Leymus triticoides 3 (L. cinereus 3
L. triticoides) TTC1 and TTC2 backcross mapping families.
Trait
Family
Position (interval†)
CIRC
TTC1
3a (125–137)
3b (111–112)
6a (107–144)
Overall R2
3a (111–161)
3b (111–144)
5Xm (25–66)
Overall R2
3a (27–80)
4Xm (11–42)
6a (61–88)
6b (95–117)
7b (11–15)
Overall R2
1b (47–85)
4Ns (0–24)
4Ns (95–142)
4Xm (23–41)
4Xm (68–109)
6a (80–125)
Overall R2
2a (121–139)
6b (108–127)
Overall R2
4Ns (87–124)
4Xm (32–34)
6a (108–127)
Overall R2
2a (30–105)
2b (130–164)
5Ns (42–44)
5Xm (0–11)
6b (0–24)
2
Overall R
2a (37–103)
3a (20–34)
3a (77–89)
5Xm (6–82)
Overall R2
TTC2
BOLT
TTC1
TTC2
ANTH
TTC1
TTC2
HGHT
TTC1
TTC2
2
2
2
2002 LOD (R , effect‡)
2003 LOD (R , effect‡)
2004 LOD (R , effect‡)
NS
NS
NS
NS
4.36 (10.2%, 275)§
3.56 (8.4%, 268)§
NS
19.7%
NS
3.17 (7.4%, 20.24)§
3.57 (8.4%, 10.25)§
NS
3.20 (7.0%, 10.24)§
25.7%
NS
3.78 (8.7%, 20.25)§
4.36 (10.8%, 10.26)§
NS
NS
NS
19.9%
NS
NS
NS
3.54 (7.6%, 22.7)§
3.70 (7.5%, 12.7)§
5.85 (15.6%, 23.9)§
30.9%
7.66 (23.5%, 113.3)§
NS
NS
NS
NS
20.6%
8.56 (17.5%, 111.1)§
4.10 (8.7%, 17.74)¶
NS
5.64 (11.1%, 28.8)§
34.1%
3.88 (10.7%, 279)§
3.47 (8.5%, 273)¶
NS
16.9%
5.15 (12.0%, 2107)§
NS
3.84 (8.7%, 294)§
21.0%
NS
4.12 (10.9%, 20.16)§
NS
NS
NS
10.9%
5.38 (8.9%, 20.15)§
NS
4.53 (9.0%, 10.16)§
4.05 (7.2%, 20.13)§
4.61 (8.1%, 20.14)¶
7.33 (15.4%, 10.20)§
43.9%
NS
NS
NS
5.21 (15.9%, 24.5)§
NS
NS
13.7%
6.74 (15.2%, 110.1)§
NS
3.44 (7.4%, 17.0)¶
NS
3.86 (11.7% 19.1)§
31.4%
4.20 (9.3%, 17.6)§
4.15 (9.2%, 17.5)¶
3.42 (7.2%, 16.7)§
NS
28.8%
3.16 (9.3%, 283)§
NS
4.08 (9.9%, 287)§
18.4%
3.9 (9.0%, 2102)§
NS
4.76 (12.1%, 2121)§
20.3%
6.12 (13.3%, 10.19)§
5.12 (11.4%, 20.18)§
NS
3.47 (9.3%, 20.16)¶
NS
28.9%
NS
NS
5.65 (15.3%, 10.21)§
NS
NS
NS
15.3%
NS
NS
NS
NS
NS
NS
NS
10.8 (24.5%, 113.6)§
5.16 (10.7%, 19.0)§
NS
NS
NS
33.8%
3.50 (8.8%, 17.0)§
NS
4.96 (13.2%, 18.6)§
NS
20.3%
2
Average LOD (R , effect‡)
3.41 (10.0%, 269)¶
NS
3.33 (8.2%, 64.4)§
16.4%
5.44 (12.8%, 2104)§
NS
4.49 (10.4%, 296)§
20.4%
4.00 (7.6%, 10.14)¶
6.33 (13.6%, 20.19)§
4.01 (8.1%, 10.15)§
4.11 (10.0%, 20.16)§
NS
35.9%
NS
NS
5.64 (11.3%, 10.18)§
4.37 (9.2%, 20.16)§
NS
4.89 (12.4%, 10.18)§
32.6%
4.59 (11.4%, 12.5)§
4.00 (9.9%, 12.2)¶
19.2%
5.42 (14.7%, 23.12)§
NS
NS
14.7%
10.36 (21.2%, 111.6)§
5.07 (9.6%, 17.8)§
NS
e
3.77 (7.0%, 26.8)
NS
38.8%
6.40 (14.2%, 18.1)§
3.98 (7.9%, 16.0)§
NS
4.65 (9.1%, 26.9)§
34.5%
† Approximate positions (cM) where LOD scan exceeded 3.3 threshold as shown in Fig. 1.
‡ Mean of heterozygous genotype (L. cinereus and L. triticoides alleles)– mean of homozygous genotypes (two L. triticoides alleles) expressed in centimeters
(CIRC and HGHT), proportions (BOLT), or days (ANTH).
§ Cofactors selected by interval mapping (LOD . 3.3) and backward elimination (significant at the 0.001 level).
¶ Additional cofactors identified by restricted multiple QTL model mapping (LOD . 3.3) and backward elimination.
Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved.
LARSON ET AL.: LEYMUS WILDRYES
ANTH LOD values that were nearly significant on a
chromosome region homologous to the TTC2 ANTH
QTL on LG4Xm (see ANTH LOD scan in Supplemental Data). Similarly, both TTC1 and TTC2 displayed
nearly significant ANTH LOD values on homologous
regions of LG3b. In any case, L. cinereus contributed
three positive and two negative ANTH QTL alleles at
the five significant QTLs (Table 4, Fig. 1).
The genome-wide LOD scans were similar but not
identical for the BOLT and ANTH flowering traits (Fig. 1
and Supplemental Data). The TTC1 BOLT QTL on
LG3a had no apparent effects on ANTH. The TTC2
BOLT QTLs on LG4Ns were associated with significant
or nearly significant ANTH QTLs. Likewise, the BOLT
QTLs on LG4Xm were associated with significant or
nearly significant ANTH QTLs in the TTC1 and TTC2
families. The TTC1 ANTH QTL on LG 5Ns was associated with elevated BOLT LOD, albeit less than the
3.3 LOD threshold (Supplemental Data). Interestingly,
the TTC1 and TTC2 BOLT QTLs on LG6a were associated with significant ANTH QTLs only in the TTC2
family. The TTC1 BOLT QTL on LG6a was not associated with elevated ANTH LOD values. The latter
observation raises some doubt about the putative homology of the TTC1 and TTC2 BOLT QTLs on LG6a
(Fig. 1). Finally the TTC1 BOLT QTLs on LG6b and
LG7b were closely associated with a significant or nearly
significant ANTH QTLs, respectively.
Significant skewness and kurtosis was detected for
BOLT and ANTH; however, all QTLs (Table 4, Fig. 1)
were highly significant (P , 0.0001) on the basis of the
nonparametric Kruskal–Wallis rank sum test.
Plant Height
The L. cinereus Acc:636 plants were substantially
taller than the L. triticoides Acc:641 plants in 2003 and
2004 (Table 1). However, the L. cinereus plants require
several years to reach full-size; thus, interspecific differences in plant height were not apparent in 2002.
Interestingly, the TC1 and TC2 F1 hybrid genotypes
displayed greater plant height than the L. cinereus Acc:
636 or L. triticoides Acc:641 reference individuals, especially in 2002 and 2003 (Table 1). The 2004 HGHT means
were substantially greater in 2004, which may be attributable to better rainfall (i.e., 2002 and 2003 were severe
drought years) and/or longer plant establishment.
Heritabilities over years (Table 2) for HGHT were
intermediate between CIRC and the two flowering traits
(ANTH and BOLT). Likewise, the ratio of genotype 3
year interaction to phenotypic variance for HGHT (0.13)
was intermediate between CIRC (0.02) and the flowering traits, ANTH (0.18) and BOLT (0.21). Weak positive genotypic correlations between HGHT and BOLT
were detected in TTC1 and TTC2 families, but no
other trait correlations with HGHT were detected in
both families.
We detected a total of seven HGHT QTLs in the
TTC1 and/or TTC2 families (Table 4, Fig. 1). The TTC2
HGHT QTL on LG3a was split into two QTLs using a
combination of rMQM and unrestricted MQM mapping
2535
(Table 4, Fig. 1). The single largest HGHT QTL was
located in homologous regions of LG2a in both TTC1
and TTC2 families. Likewise, overlapping TTC1 and
TTC2 HGHT QTLs on LG5Xm were too close to separate (Fig. 1), which we count as one homologous QTL.
Moreover, LG5Xm was the only chromosome that contributed negative HGHT QTL alleles from the taller L.
cinereus species, which also seems to support our interpretation that these are homologous QTLs in the TTC1
and TTC2 families. Thus, L. cinereus contributed six
positive and one negative HGHT QTL alleles in the
TTC1 and/or TTC2 families, which is consistent with divergent phenotypes of the parental species.
DISCUSSION
Leymus Molecular Genetic Maps
Seventeen additional anchor markers described in this
study support previous linkage group identifications, tentatively numbered according to the seven homoeologous
groups of the wheat, barley, and rye Triticeae cereals
(Wu et al., 2003). Among the well-characterized Triticeae
genomes, researchers have detected one rearrangement
in Aegilops longissima Schweinf. & Muschl. (SI genome), 11 in Ae. umbellulata Zhuk. (U genome), 0 in
Ae. speltoides Tausch (S genome), seven in Secale cereale
L. cultivated ryes (R genome), and two paracentric
inversions in H. vulgare cultivated barleys (H genome)
relative to Ae. tauschii Coss., the donor of the hexaploid
wheat D genome (Devos and Gale, 2000). The A, B, and
D genomes of allohexaploid wheat are evidently colinear
except for several large reciprocal translocations involving chromosome arms 2BS and 6BS and chromosomes
4A, 5A, and 7B (Devos and Gale, 2000). One of these
rearrangements involves a reciprocal translocation between 4AL and 5AL, which includes the VRN2 gene in
T. monococcum L. (Dubcovsky et al., 1998; Yan et al.,
2004). Interestingly, the location of the VRN2 gene on
Leymus LG5Ns evidently corresponds with the location
of VRN2 in T. monococcum (Devos et al., 1995; Dubcovsky et al., 1996). Thus, the Leymus Ns and T. monococcum genomes evidently share the same 4AL/5AL
translocation arrangement. Although we detected seven
instances of marker synteny in Leymus that were not
syntenous in other Triticeae species, we have no firm
evidence of rearrangements other than the putative 4Ns/
5Ns translocation that has already been documented in
wheat. Incongruent RFLP marker locations can often
be attributed to paralogous duplications or, in the case
of PCR, priming annealing at nonhomologous loci. Although there is substantial evidence of colinearity between
the Leymus Ns, Leymus Xm, wheat A, wheat B, wheat D,
and barley H genomes, we cannot exclude the possibility of
a few yet undetected rearrangements in Leymus.
These are the first QTL analyses conducted using the
high-density linkage maps published by Wu et al. (2003).
Throughout much of the linkage map, relatively small
irregularities in the LOD scans (Supplemental Data) resulted from imperfect genotyping, missing data, and ambiguous marker orders. Nevertheless, these QTL maps
2536
CROP SCIENCE, VOL. 46, NOVEMBER–DECEMBER 2006
provide a useful starting point for high-resolution QTL
mapping experiments using six advanced backcross
populations that are currently being genotyped and evaluated in clonally replicated field trials.
Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved.
Comparison of TTC1 and TTC2 QTLs
Two circumference, two bolting, and two height QTLs
were evidently homologous in both TTC1 and TTC2
families. Conversely, two circumference, seven bolting,
all five anthesis date, and five height QTLs were unique
to TTC1 or TTC2 families. Differences between the
TTC1 and TTC2 families can be attributed to differences
between the TC1 and TC2 hybrids, which in turn can only
be attributed to genetic variation within the L. cinereus
Acc:636 and L. triticoides Acc:641 natural germplasm
sources. Thus, functionally important QTL variation was
apparent within and between natural source populations
of these experimental families and species.
Growth Habit
The coincidence of QTL effects near the VP1 loci on
LG3a and LG3b, in both TTC1 and TTC2, suggest that
these QTLs may be controlled by homoeologous copies
of the one gene. If so, this gene evidently controls much
of the dramatic difference in growth habit between L.
cinereus, L. triticoides, and perhaps other grasses. The
only other CIRC QTLs, on LG5Xm and LG6a, were
unique to the TTC1 and TTC2 families, respectively. A
preponderance of mapped barley mutations (four of six
loci) that affect vegetative axillary development have
been localized to chromosome 3HL (minus arm), including absent lower laterals (als), low number of tillers1
(int1), and semi brachytic (usu), which produce fewer
tillers, in addition to granum-a (gra-a), which produces
significantly more tillers (Babb and Muehlbauer, 2003;
Franckowiak, 1996). A recessive gravitropic lazy dwarf
gene (lzd) was also mapped to the short arm of barley
chromosome 3 (Franckowiak, 1996; Takahashi et al.,
1975), but this mutation seems to be more proximal to the
centromere than the Leymus LG3a and LG3b growth
habit QTLs. Chromosome 3 is highly conserved within
the Triticeae (Devos and Gale, 2000). Colinear from end
to end with rice chromosome 1, Triticeae group three is
also the most conserved of all chromosome groups when
compared with rice (La Rota and Sorrells, 2004). Rice
chromosome 1 contains putative transmembrane auxin
efflux carrier and DNAJ-like genes (International Rice
Genome Sequencing Project, 2005), near the Vp1 locus,
originally identified in Arabidopsis pin-forming (Gälweiler et al., 1998) and arg1 (altered response to gravity)
(Sedbrook et al., 1999) mutants, respectively. However,
pin1- and arg1-like sequences are also present in other
regions the rice genome. Rhizome (Hu et al., 2003) and
tiller angle (Li et al., 1999) QTLs also map to rice chromosome 1, but these rice loci were not located near the
rice Vp1 gene.
The TTC2 CIRC rMQM QTL on LG5Xm mapped
near the TCP-like sequence (TCP2) amplified from L.
triticoides rhizome cDNA using primers designed from
the teosinte branched one gene (Doebley et al., 1997),
BCD1130, BCD1707, and PSR128 loci. The PSR128
marker also maps in the centromere region of maize
chromosome 4 near the recessive lazy1 (la1) gene (Lawrence et al., 2004). Lazy maize mutants, allelic to la1,
actively grow downward (prostrate) under light (Firn
et al., 2000). We found BCD1707 and BCD1130 sequences on rice chromosome 11, using a TBLASTX
search (Ware et al., 2002) of the rice genome (International Rice Genome Project, 2005), near another lazy (la)
gene (Miura et al., 2003). The centromere and short arm
regions of Triticeae group 5 are homoeologous to rice
chromosomes 9 and 12, respectively (Van Deynze et al.,
1995; La Rota and Sorrells 2004). Rice chromosome 9
contains a major QTL (Ta) for tiller angle (Li et al., 1999).
The TTC1 CIRC QTL peak on LG6a is located approximately (minus) 40 cM from the MWG2264 locus.
Interestingly, the uniculm (cul2) mutation and MWG2264
loci are both located (minus) 8.8 and 3 cM relative to the
cMWG679 locus on barley 6H (Babb and Muehlbauer,
2003; Graner et al., 1991; Graner, 2004). Compared with
other barley mutants, uniculm2 (cul2) is unique in that it
inhibits formation of axillary meristems and does not
produce tillers (Babb and Muehlbauer, 2003).
We were not able to identify rhizome QTLs in Leymus that were homeologous to rhizome QTLs of Sorghum and Oryza. Paterson et al. (1995) detected about
12 rhizome QTLs including a conspicuous cluster of
QTLs affecting rhizome number, rhizome distance, and
seedling tillers on Sorghum linkage group C, which
corresponds to wheat group 4 (Draye et al., 2001). The
latter Sorghum QTL also corresponds with sucker and
stalk number QTL in Saccharum (Jordan et al., 2004), in
addition to the Rhz2 gene and 11 other QTLs controlling rhizome proliferation differences between O.
sativa L. and O. longistaminata A. Chev. & Roehr. (Hu
et al., 2003). We did not detect any plant circumference
QTLs on Leymus LG 4Ns or LG4Xm, which include the
PRC140 rhizome QTL marker (Draye et al., 2001)
derived from a Sorghum LG C and sequences orthologous to the maize teosinte branched one gene (Doebley
et al., 1997). The Rhz3 gene and rhizome QTL on Oryza
chromosome 4 correspond to rhizome QTLs on Sorghum LG D (Hu et al., 2003) and Saccharum (Jordan
et al., 2004). This region of Oryza chromosome 4 and
Sorghum LG D evidently corresponds to Triticeae group
2 (Paterson et al., 1995; Van Deynze et al., 1995). We did
not detect any significant rhizome QTL effects on Triticeae group 2.
Bolting and Anthesis Date
The genetic control of flowering time is complex,
involving three major groups of genes on all seven
Triticeae chromosome groups (Snape et al., 2001). Two
of these groups interact with the environment, namely
those controlling vernalization (Vrn) and photoperiod
(Ppd). The third set of genes controls developmental rate
independent of vernalization and photoperiod, so-called
“earliness per se” (Eps) genes. A relatively large number
of Eps genes have been mapped to all seven homoeologous
groups of wheat and/or barley (Laurie et al., 1995; Snape
2537
Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved.
LARSON ET AL.: LEYMUS WILDRYES
et al., 2001), several of which may correspond to BOLT
and/or ANTH QTLs detected in Leymus.
The Vrn genes map to Triticeae groups 1, 4, 5, and 7
(Dubcovsky et al., 1998; Snape et al., 2001). Moreover,
the two most potent genes, on groups 4 and 5, have been
cloned and characterized (Yan et al., 2003; Yan et al.,
2004). Surprisingly, no significant bolting or anthesis
date QTLs were associated with the Leymus CDO504
markers on LG5Ns and LG5Xm, which are presumably
closely linked with genes that are orthologous to VRN1.
We have amplified VRN1 sequences from Leymus but
have not yet found polymorphisms needed for mapping.
Likewise, no significant bolting or anthesis date QTLs
were associated with the VRN2 gene on LG5Ns. A second yet undetected VRN2 gene may or may not be present on the Xm genome. The most conspicuous wheat and
barley Ppd genes are located on homoeologous regions
of the 2A, 2B, 2D, and 2H short (plus) chromosome arms.
Other Ppd genes have also been reported on barley 1H
(Laurie et al., 1995) and 6H (Strake and Börner, 1998).
We did not detect ANTH or BOLT QTLs on short (minus) arm of Leymus group two chromosomes.
The TTC1 and TTC2 families displayed substantial
genetic variation for anthesis date and bolting, including
genotypes that have not flowered in the field, which was
not apparent among the parental species accessions.
This transgressive segregation can be explained by a
mixture of antagonistic (i.e., positive and negative) QTL
alleles from each parental species. Leymus cinereus contributed five positive and four negative BOLT QTLs
and three positive and two negative ANTH QTLs. Thus,
correspondence of bolting and anthesis date QTLs (and
relatively strong negative genotypic correlations) suggest that outbreeding depression (i.e., transgressive genotypes that to flower) was associated with flowering
(i.e., lateness) genes under balancing selection. Although
population means and standard deviations may not
demonstrate significant bolting depression, the fact that
a substantial number of progeny failed to flower year
after year indicates that some outbreeding depression
is occurring. We have not observed this phenomenon
within the L. cinereus Acc:636 or L. triticoides Acc:641
natural source populations.
Plant Height
The HGHT QTLs on LG2a displayed remarkably
strong effects and similar map locations in both TTC1
and TTC2 families, suggesting that these QTLs are homologous. The LG2a HGHT QTL was the only major
positive HGHT effect associated with chromatin of the
much taller L. cinereus species and detected in both
TTC1 and TTC2 families. We interpret these results to
mean that this is one of the key QTLs controlling relatively large differences in plant height between L.
cinereus and L. triticoides. Börner et al. (1999) mapped
gibberellin-sensitive gal (GA-less) and gibberellininsensitive gai (GA-ins) mutations about 55 cM apart
on barley 2H. The Leymus LG2a HGHT QTL spans a
40 cM interval from the XANTHA locus down below
the CNL045 locus (Fig. 1). The XANTHA–CNL045
LG2b interval also includes the cWMG763 locus (Fig. 1),
which was mapped to the short (plus) arm of barley 2H
about 36 cm distal from the MWG2287 marker in the
barley IGRI 3 FRANKA population (Graner et al.,
1991; Graner, 2004). The MWG2287 locus is closely
linked to the barley gai mutation in the centromeric region of barley chromosome 2H (Börner et al., 1999). We
speculate that the gai (sdw3) locus, described by Börner
et al. (1999) and Gottwald et al. (2004), may correspond
with a major plant height QTL associated with the cluster of DNA markers near the CNL045 locus of LG2a.
The CNL045 loci are associated with high-density clusters of markers on Leymus LG2a and LG2b linkage
groups, probably caused by reduced recombination near
the centromere. Yang et al. (1995) also described a partially dominant GA-ins dwarfing gene on the short arm
of chromosome 2A of wheat (Rht21), which could be
homeologous to the gai mutation mapped by Börner
et al. (1999). In any case, the barley, wheat, and Leymus
plant height genes on homeologous group two are evidently different from the so-called “green revolution”
GA-ins dwarfing genes of wheat, barley, and rice encode
gibberellin response modulators that map to Triticeae
group four (Peng et al., 1999).
ACKNOWLEDGMENTS
This work was supported by joint contributions of the USDA
Agriculture Research Service, Utah Agriculture Experiment
Station, Department of Defense Strategic Environmental
Research and Development Program CS1103 project, and US
Army BT25-EC-B09 project (Genetic Characterization of
Native Plants in Cold Regions). Trade names are included for
the benefit of the reader, and imply no endorsement or preferential treatment of the products listed by the USDA. The authors gratefully acknowledge the technical assistance of Mauro
Cardona, Linnea Johnson, Neil Rasmussen, and Ron Reed.
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LG1a--TTC1
Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved.
120
125
130
4
115
P33M61.137
E38M60.108
P35M61.177
E38M49.399
P33M61.233
P33M47.234
P44M62.119
P44M62.368
E37M47.327
E41M49.134
P35M61.174
E38M60.311
E38M60.294
P33M47.273
E36M48.168
E36M49.306
E36M59.250
E38M60.177
E36M48.225
E36M48.069
E41M49.278
E41M59.271
E38M47.222a
E41M48.322
P44M62.164a
E37M47.270
P42M62.153
E37M61.182
E41M60.361
E36M59.217
E41M47.148
E38M60.162
BCD1150
E41M49.278
E37M62.244
P33M60.290
E36M48.257b
E37M62.270
E41M47.119
E38M47.222a
85
110
3
E41M47.148
E38M60.107a
P44M49.388
E38M59.101
E38M49.146
90
E41M59.163
E38M60.159
P42M61.114
E37M61.344
95 1H
HVA.094
E41M61.122
E37M47.196
P42M61.218
100
P33M62.354
E41M61.398
P35M59.188
P35M62.180
105
E41M59.310
2
E38M49.066
E36M50.397
E36M61.291
P33M60.147
E41M59.335
E41M61.367
P33M60.360
P35M61.361
P33M47.161
E38M60.181
E36M59.167
E36M61.231
E41M61.271
P42M62.293b
P33M61.331
E41M48.388
E38M49.396
E37M61.173
E36M49.287
E38M47.293
E37M49.146
E41M59.124
E38M59.215
P33M62.346
E36M61.286
E41M59.193
E36M48.163
E41M47.113
E41M60.123a
E41M49.217
E38M49.266
E37M49.361
E37M49.321
E37M60.153
E38M60.074
E38M59.330
E41M59.082
E41M49.072
E37M60.146
E36M50.254
P33M50.281
P33M59.158
E36M61.336
P33M60.290
75
80
P33M50.162
P33M62.220
P42M61.072
E36M61.377
70
E38M49.140
P33M47.280b
P33M47.280b
P33M50.162
P35M59.213
E38M47.185
E37M61.306
1
E36M61.336
0
E41M49.220
E41M49.072
P35M50.213a
4
65
E37M49.321
3
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LG1b--TTC2
2
55
1
50
0
45
P33M47.258
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2
30
1
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LG1b--TTC1
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E41M48.389
E36M62.240
P44M62.293
E41M60.123a
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LG1a--TTC2
2
5
1
0
0
1B(A)
E38M60.107a
E36M59.239
P35M59.211
E41M59.163
P44M62.164b
HVA.094
1H
E38M59.101
BCD1562
1BD(A)
E41M47.110
P33M62.354
P42M61.218
E37M47.196
E38M60.291
E41M61.398
E36M59.296
E41M60.195
P35M61.177
GDM126.146 1D
E38M60.108
E41M47.163
P44M62.368
P33M47.234
P44M62.124
P33M62.138
E41M49.134
E37M47.327
E38M60.294
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P35M50.127
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E38M49.283b
P35M62.350
E37M60.097
E36M61.240
E36M62.140
E37M60.103
E37M47.319
E37M62.388b
P35M61.396
E41M62.173
E38M60.213
E36M62.346
E36M61.323
E41M61.385
E41M48.245
P33M61.198
E37M62.232
E41M48.131
P35M59.154
P44M62.105
P33M61.367
P42M61.117
P33M62.319
E38M47.229
E41M47.355
P33M50.154
E38M59.317
E37M61.095
P33M47.255
E37M62.201b
E37M61.290
E37M47.151
E37M47.150
E41M61.252
E37M49.092
E36M61.192
E41M49.171
E36M61.267
E36M61.191
E38M49.283b
1AD WMC336.388
P35M62.350
E41M61.265
E37M47.126
E36M62.140
E37M60.103
P44M62.378
P35M61.394
P44M49.200
P33M61.367
P35M49.334
P35M49.311
P33M61.198
P33M47.328
P35M49.139
P42M61.117
E36M61.323
E41M48.245
E38M60.213
E36M62.346
E41M62.189
E41M61.385
E37M49.051
E37M47.319
E37M47.320
E41M47.355
E41M60.360
E37M49.079
E37M61.073
E36M59.317
E41M61.141
E38M49.269
E37M60.097
E37M61.095
E38M59.204
E41M60.336a
E41M47.237
E37M49.092
E36M61.267
P44M62.209
P33M50.246b
E38M49.214
P33M62.271
E41M48.053
E37M60.284
P33M59.118a
P44M62.351
P33M50.246b
E38M49.241
E38M59.153
E38M49.386
E36M62.057
E41M48.147
P44M49.133
P35M61.141
E36M62.192
E41M47.078
E36M62.150
P33M50.180b
P44M49.250
E37M60.259
E38M49.241
E38M49.256
P35M60.119
P44M49.133
P42M62.312
P42M61.157
E36M62.192
E41M47.078
E41M62.337
135
E38M49.063a
CIRC_IM
BOLT_IM
ANTH_IM
HGHT_IM
CIRC_rMQM
BOLT_rMQM
ANTH_rMQM
HGHT_rMQM
Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved.
P44M62.176
E38M60.056
30
E38M60.153
P44M62.228
35
P33M62.117
E41M59.094
P35M62.365
E37M62.072
P42M62.303
P42M62.298
E36M59.221
40
E37M60.172b
E41M61.111a
E38M59.282
2H XANTHA.395
P44M62.228
45
E36M50.224
E41M61.352
50
60
65
70
75
2A
85
90
P33M48.212
E37M60.192
E41M60.139
2H XANTHA.270
55
80
E36M61.152a
E36M50.224
E38M47.180
P33M48.212
P33M50.249
E37M60.192
E41M47.249
E36M59.234
E36M61.361
P42M61.367
E36M49.271
E37M60.342
E41M62.194
E37M47.145
E41M61.188
E37M62.228
E36M61.297
E41M49.092
E36M61.253
P33M61.216
CNL045.154
E37M62.182
E36M48.098
P35M50.385
P33M48.215
E41M48.070
E36M61.361
E36M61.297
E41M61.188
E41M61.254
E41M62.194
E41M48.085a
E37M60.342
E37M62.228
E38M60.087
E38M60.069
E41M49.092
P42M61.367
E41M62.319
P33M61.216
P33M60.143
E41M48.062a
P33M48.154
P33M61.203
E41M62.152
E41M50.202
E38M59.329
E37M61.232
E37M62.252
E37M62.252
E36M48.248
P33M48.224
95
100
E41M60.123b
E41M60.238
P35M50.279
E37M49.102
P33M50.180c
E36M48.202
P35M50.279
E36M59.222
E37M49.102
E41M61.099
E41M47.139
E36M49.130
E36M48.356
E36M59.222
105
110
E41M59.318
P33M59.318
P33M62.231
E36M48.356
P33M50.246a
E41M59.154
E41M47.356
P42M61.185
E36M49.339
P42M61.122
P42M61.196
135
P33M48.126
P35M50.307
2ABD GWM382.095
P33M62.291
140
E38M49.152
P35M62.111
P35M49.140
P44M62.088a
145
E36M49.348
E36M48.075
E36M49.366
E38M49.067
150
E41M48.112
E36M59.182
E41M47.285
E38M59.074
155
130
160
165
E37M60.188
P35M62.320
E37M61.110
P33M60.130
E37M62.131
E37M60.297
E41M50.159
E37M62.272
E37M60.257
E38M60.280
E41M60.172
E41M49.215
E37M49.304
P35M59.253
P33M48.079
P33M60.249
P35M61.364
P33M61.285
E37M62.183
E36M59.171
E41M48.226
E38M60.234
E37M62.261
E38M60.130
E37M62.136
E38M47.105
E36M49.313
P33M59.241
P42M62.115
E37M61.141
P44M49.223
E36M61.157
E37M49.147
2A CNL045.172
P33M60.180
E41M60.293a
E37M47.370
E37M60.294
P33M59.278
P35M60.227
E37M60.257
E36M61.157
E37M62.272
P33M59.241
E37M60.188
P35M60.248b
E41M59.257
P33M61.285
P33M60.130
E41M48.251
TRX1
1R
E37M62.183
P35M61.364
P33M48.079
P35M59.253
P33M60.250
E37M62.260
E36M59.171
E37M62.212
E36M62.145
E41M49.247
E41M61.350
E37M61.281
E37M61.141
E37M49.147
E36M62.097
P33M60.180
E37M47.143b
E36M49.179
P33M59.278
P35M60.280
E41M61.119
P33M59.172
E37M62.269
E41M61.359
E38M47.076a
125
2H MWG763.277
E41M61.099
E38M49.074
120
P42M61.142
E41M61.119
P33M59.172
P33M59.242
115
P33M47.345
P35M60.227
CIRC_IM
CIRC_rMQM
P35M62.377
E38M47.076a
P33M62.231
E41M60.088
E36M62.339
E41M47.357
P33M47.243
P44M49.149
P33M62.296
E41M48.060
P33M62.333
P33M48.126
P35M49.242
E37M49.160
GWM382.095 2ABD
P35M62.399
P35M60.144
P33M62.291
E36M49.366
E41M47.132
E41M48.112
E41M47.131
P44M49.369
P42M62.146
E41M47.214
P44M49.323
P33M48.397
P35M50.258
E41M48.188
P33M50.180a
P33M50.180a
E36M49.266
E41M47.359
E36M61.091
P35M60.272
P42M62.067
P35M60.272
E41M59.323
E37M49.165
E37M49.165
E37M62.269
P35M59.372
P33M60.268
P33M48.350
E37M60.186
P33M48.348
E37M47.198
P33M61.071b
E41M61.156
P42M61.081
E38M59.220
E36M49.092
P35M61.243
P33M60.217
P35M49.090
2ABD GWM382.126
P33M62.394
E41M47.155
E38M59.095
E41M60.144
E37M47.356
E38M59.179
P42M61.140
E41M59.126a
E38M47.278
P42M61.081
E38M59.220
E36M49.109
E37M47.056
P35M61.243
P44M49.286
P33M60.217
E41M47.155
E38M59.095
E41M60.144
E37M47.356
E38M59.179
P33M50.245
E41M59.126a
GWM382.126 2ABD
P33M48.067
E38M47.278
P33M48.067
BOLT_IM
ANTH_IM
HGHT_IM
BOLT_rMQM
ANTH_rMQM
HGHT_rMQM
P42M61.140
4
P35M50.208
P33M48.392
3
E36M59.293
E41M47.204
E36M50.311
P44M49.095b
P33M50.227
P33M50.152
P35M62.194
P35M62.163
E41M62.160
E37M49.270
P44M49.258
E37M61.102
2
E41M49.131
E36M61.248
1
0
E41M48.241a
E36M59.293
E41M49.131
E36M49.311
E36M59.392
E38M47.072
P35M62.194
P33M60.125
P33M50.152
P33M48.328
P35M50.208
P33M48.394
P44M62.299
5
E38M60.153
LG2b--TTC2
4
E36M61.248
3
25
2
20
LG2b--TTC1
1
15
P33M61.175
E38M47.364
E36M48.172
E37M61.246
E41M48.255
E38M60.263
E37M61.155
P33M61.312
P42M61.145
P33M48.139
P35M49.202
P33M61.212
P33M59.256
E38M59.217
P44M62.223
P35M62.178
P44M49.246
E41M59.184
E36M61.221
E38M60.056
P33M62.364
P35M49.297
P35M62.365
E36M48.305
0
10
LG2a--TTC2
6
5
4
3
2
1
0
5
10
9
8
7
6
5
4
3
2
1
0
LG2a--TTC1
P33M62.285
E41M48.255
P33M48.139
P33M61.175
P33M61.312
P33M47.078
P35M62.187
P35M49.202
P33M61.213
P35M61.225
E36M48.172
E38M60.263
E37M61.155
E41M59.077
E38M47.364
E36M61.138
E41M59.345
P44M62.223
E36M61.175
P44M49.246
P33M59.256
E36M50.247
0
LG3a--TTC1
Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved.
E41M48.133
P33M50.328
P33M61.149a
E41M61.290
E37M49.132
65
70
P35M59.284
P42M61.316
E37M61.350
E37M60.356a
75
80
P44M49.331
E37M62.274
P35M62.385
E41M61.234a
E41M61.235
P33M60.305
E41M60.080b
E41M62.354
E41M62.350
E38M49.084
BCD1532
3AB(C)
E41M60.111
P33M62.144
3H
110
115
120
125
E37M47.386
P44M49.110
P35M60.074b
E38M47.374
E37M49.329
P33M47.167
P33M62.190
E36M62.173
E37M61.199
E37M60.339
E36M61.263
P33M61.153
E36M59.253
P44M49.329
E36M61.097
E37M62.082
E37M60.203
E36M61.141
E37M49.303
E41M61.131
E36M49.351
E36M50.351
E38M60.242
E41M59.113
P35M49.242
E36M61.234
E38M60.316
E37M47.124
E38M47.140
E41M49.163
E38M60.305
E38M47.192
E41M47.225
E38M47.076b
P35M62.199
E37M47.302
P33M50.328
P33M61.149a
E41M61.290
E36M61.303
E41M61.127
E41M48.133
P35M59.284
130
135
E37M47.224
140
E41M62.069
145
P44M49.139
E37M60.302
150
E37M49.349
P35M50.264
P33M47.097
E37M60.082
E38M60.278
P44M49.324
VP1i5.406
E41M61.220
E37M60.083
P44M62.204
E36M49.248
E37M47.225
E36M59.219
P33M50.146
E36M62.090
E38M47.185
E37M49.141
E36M48.167
P33M61.174
E41M48.248
E37M49.118
P35M49.224
E41M62.341
E37M49.201
E41M61.063
P35M61.089
P35M62.147
E37M60.303
E41M49.232
E37M47.224
E37M49.355
E41M62.069
P35M59.174
170
E38M47.172
E41M60.286
P33M60.207
E41M47.134
P35M62.242
E36M62.227
E41M61.077
P33M50.345
E37M62.162
3ABD
3ABD
E41M49.304
P33M60.227
E36M62.227
P33M47.099
VP1i2.394
E37M62.162
E37M47.326
E38M59.249
E36M48.204
E37M47.263
E41M59.206
E41M62.204
P42M62.249
P33M48.155
E36M59.262
E41M47.173
E37M47.326
E36M48.284
E41M62.204
E41M62.260
P42M62.249
P33M59.245
E37M49.179
E41M59.222
E41M59.325b
E37M49.358
4B(F)
CDO1401
E41M49.285
E41M59.210
E36M50.237a
E37M49.358
E38M59.065
E37M62.235
P33M62.090
E37M62.075
P33M60.393
P33M61.129
P44M62.093
160
165
E38M49.064
P33M60.207
E41M49.302
E41M48.102
P33M60.192
P44M49.139
P33M59.118b
155
E36M61.243
P44M62.360
P35M49.270
P33M47.111
P35M61.089
P35M62.147
E38M47.359
E37M61.191
E36M50.170
E36M48.308
P44M62.093
P33M61.129
P33M47.072
P33M47.073
P33M47.167
E37M47.386
P44M49.110
E38M47.374
E37M61.199
E37M60.339
P33M62.190
E37M49.329
P33M61.153
E36M62.173
P33M50.116
E36M49.351
E36M59.253
E36M61.097
E36M61.141
E37M60.203
P44M49.329
E37M60.200b
E37M62.082
E36M61.263
E41M61.131
E37M49.303
E36M61.234
E38M47.140
E38M60.242
E37M47.242
E36M48.066
E36M61.206
E38M60.316
P33M47.073
E37M47.225
E36M59.219
E41M59.365
E41M60.296
P33M48.378
P33M50.149
P33M48.119
P33M50.285
P44M49.324
P44M49.197
E41M60.166
P33M47.072
E36M62.203
P44M62.360
105
3H
E38M49.057
E36M62.338
E41M60.166
E37M61.349
100
SIP1.413
E41M62.154
SIP1.413
E36M62.338
P33M60.192
E41M61.220
P44M62.204
VP1i5.406
E37M60.083
P35M60.135
P35M50.120
P42M61.197
E38M49.057
E38M47.165
3ABD
E41M49.135
E41M47.167
E41M62.223
E36M48.164
P42M61.232
E36M61.311
GWM005.080 3A
P35M62.225
E36M62.302
P35M50.154
E41M49.122b
E36M48.101
E37M47.226
E38M60.359
E41M47.071
E36M59.287
E36M61.330
90
95
E41M49.135
E41M47.167
P42M61.197
E41M60.343
E41M48.102
P33M47.097
85
P35M50.120
3H
SIP1.291
E41M61.095
P35M60.074a
P33M59.118b
E41M61.107
CIRC_IM
ANTH_IM
CIRC_rMQM
ANTH_rMQM
BOLT_IM
HGHT_IM
BOLT_rMQM
HGHT_rMQM
4
60
3
E41M47.071
E41M49.090
E36M61.330
E38M47.076b
E41M47.225
2
E38M60.359
55
E41M61.394
P33M62.393
P33M47.179
P33M47.279
P44M62.334
1
E41M62.223
50
E36M61.339a
P42M62.177
E36M59.261
P33M62.343
P33M47.310
P33M47.179
P33M47.280a
E41M60.156
P33M62.393
CDO460
3ABR(C)
E36M49.206
E36M61.092
E41M48.085b
E41M61.313a
P35M49.082
0
E41M49.122b
P33M47.228
4
45
E38M47.228
E41M59.148
LG3b--TTC2
3
3A GWM005.080
P35M49.082
2
3H
E41M48.390
1
40
E38M47.214
E36M48.164
P42M61.232
P35M50.314
P35M62.225
SIP1.324
0
35
5
30
4
25
3
20
2
P35M62.385
P44M62.197a
P35M61.082
E36M62.130
P44M49.253
E37M62.257
E38M49.084
E37M47.128
E41M59.268
E37M47.305
P33M62.144
E37M62.096
P33M60.308
1
15
LG3b--TTC1
0
10
4
E38M49.363
E38M47.228
P33M47.131
P33M48.194
P35M59.195
P33M60.104
P42M61.294
P42M62.178
E38M59.298a
P35M61.158
3
5
LG3a--TTC2
2
P33M50.250
1
0
0
LG4Ns--TTC1
Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved.
P42M62.164
E38M59.346
P35M61.277
E41M60.069
P35M59.112
40
4ABD GWM165.202
50
E38M60.090
E41M61.373
E41M60.069
P33M59.133
E38M59.346
P35M61.277
E41M47.205b
E41M60.233
P35M61.370
GWM165.202 4ABD
E38M59.387
E41M62.393
E38M47.063
55
60
65
70
75
80
E41M60.132a
E36M61.342
E41M49.143
E41M49.142
E41M50.282
E36M49.353
E38M47.103
E36M61.287
E41M61.084b
E41M60.196
E41M60.243
E38M60.259
E41M59.342
E41M62.335
E36M62.319
3A CNL039.229
E36M50.376
E41M61.109
P42M61.317
E36M62.249
E36M59.374
E41M48.185a
85
P33M59.302
90
P33M62.070
95
4ABD
(F)
4H
E41M62.393
E36M49.187
E37M49.316
P33M59.091
E37M49.313
E41M59.152
P33M47.094
E41M49.143
E37M61.186
E41M61.109
E41M48.067
E38M60.259
E37M60.262
E36M61.287
E38M47.103
E36M62.319
E41M62.336
E36M49.376
P33M50.264
E41M61.084b
E36M50.376
E41M60.247
E38M60.122
E37M49.174
E36M50.353
E41M60.196
E37M60.325
E36M49.353
P35M62.370
P33M62.070
E36M62.249
E41M59.342
E37M60.183
E37M61.332
E37M62.286
4D
P33M62.351
P33M59.134
E41M61.186
E41M49.198b
P33M48.399
E41M60.326
P35M50.371
P35M50.372
P35M60.392
E37M47.304
E38M60.325
E41M62.235
E41M62.082
P35M49.187
E41M62.239
P33M48.359
E36M61.378
E41M61.381
E41M47.150
E41M47.301
E41M61.381
E36M62.087
E41M47.150
GWM165.172
E41M47.301
E41M62.091
E41M49.343
E41M62.246
E36M48.272
BCD250
E36M61.332
P33M47.142
PRC140.409
P42M61.182
P33M60.110
E38M60.146
E41M50.330
E36M61.306
E38M47.284
E38M47.363
E41M61.317
E41M59.230
E36M59.113
E36M49.238
E37M60.135
E38M47.068
E37M49.273
KSU149.240
E38M49.170
E38M49.291
E37M47.108
E36M50.171
E36M62.341
E36M48.118
E36M61.099
E41M62.369
E36M62.087
P35M50.262
E41M62.246
E36M61.332
E41M61.317
P35M62.123
P33M60.110
E38M60.146
E36M61.306
E38M47.284
E41M59.230
E36M59.113
E36M49.238
E41M61.340
PRC140.409
KSU149.243
E37M47.108
E38M49.291
E36M48.118
4H
4D
E36M48.277
E41M47.171
P33M61.340
E41M47.172
E38M60.201
KNOX.239
4H
E38M59.100a
E41M47.060
E38M60.396
4H TCP1Xm.459
TCP1Xm.459 4H
E38M60.196
4H
KNOX.239
E41M61.269
E36M49.076
E38M59.100a
E41M47.060
P33M62.206
E41M61.269
E36M49.076
E37M60.190
E37M60.090
P44M49.326
P35M49.205
P35M60.313
P35M62.276
P35M62.274
100
105
E36M50.064
P35M60.392
P35M50.371
P35M50.372
P42M61.285
E37M62.368
E41M62.082
E41M62.233
P44M49.141
P33M48.359
E41M62.200
P42M62.362
E37M49.360
P44M62.317
4
E38M47.398
E41M49.223
E41M49.283
E41M61.172
P35M62.140
P42M61.073
E41M49.277
E36M59.152
P42M62.263
E41M62.219b
3
E36M61.368
2
P33M61.316
E36M61.289b
35
45
E38M59.239
1
30
P44M62.317
P44M62.331a
E38M60.091
4H
MipsA.284
4B KSU154.137
E41M62.396
4B KSU154.137
E36M62.210
P33M59.260
0
25
E38M60.090
P33M62.351
E36M50.064
E37M49.207
E37M49.206
P33M59.134
4BH GWM006.170
P33M48.399
E41M49.194
E36M48.109
E37M62.129
E37M62.128
P35M61.153
4H MLOH1A.352
P35M59.230
E41M47.169
LG4Xm--TTC2
6
5
4
3
2
1
0
20
LG4Xm--TTC1
6
5
4
3
2
1
0
15
4
10
3
5
LG4Ns--TTC2
2
P33M59.260
P35M62.140
E41M61.172
E41M49.283
E41M49.223
E36M62.349
P42M61.073
P44M49.095a
E41M49.277
E37M47.143a
1
0
E38M59.239
0
P33M47.125
E38M47.198
E41M48.093
P35M49.201
P33M61.273
P35M62.276
P33M47.126
P35M62.274
110
E36M62.211
115
120
P35M62.127
TCP1Ns.430
125
E36M62.211
130
4H
E38M49.390
E38M60.223
E37M60.252
E41M61.380
E41M47.384
E36M59.247
TCP1Ns.430
E37M60.375
P35M62.127
135
140
145
P33M61.096
E41M60.301
E36M49.279
E41M48.195
E38M49.390
E36M59.247
E41M61.212
E41M61.272
4H
CIRC_IM
BOLT_IM
CIRC_rMQM
BOLT_rMQM
ANTH_IM
HGHT_IM
ANTH_rMQM
HGHT_rMQM
LG5Ns--TTC1
LG5Ns--TTC2
Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved.
45
5B
50
55
E36M62.340
P33M62.259
60
E36M59.111
65
E36M48.254
P33M50.232
70
E36M62.189
75
80
85
E41M62.107
E37M60.105
E41M61.201
E41M61.256
E36M61.139
E37M47.322
5ABD
E37M61.149
E41M60.285
E41M61.136a
E37M47.133
E41M60.293b
E36M61.383
E41M61.091b
E36M48.180
E38M49.222
E37M62.136
E38M49.369
E37M47.378
P33M60.366
KSU171.279
P33M50.340
P33M50.158
E38M59.268
E41M49.140
E37M62.273
E36M62.311
E38M60.107b
E41M47.281
E36M48.296
E38M60.169
P33M50.233
E36M48.308
E36M62.189
E36M48.254
E41M62.107
E41M59.267
E41M61.201
E41M61.256
5B
5ABDHR
PSR128.412
P35M62.264
E36M61.173
E41M48.173
E41M48.103
E36M48.088
E41M59.130
95
100
E37M60.307
E37M61.327
E37M60.092
E41M48.103
5ABDHR CDO504Xm.167
E37M47.096
E37M47.136
E37M47.324
E38M59.199
E37M60.105
E37M49.183
E36M49.188
110
115
E36M50.333
E36M62.180
P33M59.264
P35M61.295
P33M62.323
E41M61.194
E38M47.084
E41M60.244
E38M59.248
E41M50.064
P33M62.172
P35M60.115
P33M62.178
E41M60.140
E41M59.311
125
E41M59.312
E36M59.140
E41M60.099
E36M59.265
130
E37M47.153
E41M48.056b
E37M47.244
P42M62.112
135
E37M60.235
P33M60.077
P42M61.143
P33M48.283
140
P33M48.258
P35M49.344
4H-5A
VRN2.199
P33M50.292
145
E36M49.319
E36M50.319
E36M48.207
E41M61.279
150
120
CDO504Xm.167 5ABDHR
E36M62.220
P35M59.137
E41M60.229
E37M49.077
P35M59.102
E36M48.200
P44M62.088b
E41M49.213
P44M62.089
P35M60.240
E37M61.215
E41M62.122
P33M59.262
P35M62.167
P35M60.100
P33M47.216
E36M62.252
E41M60.147
P33M48.361
E36M62.159
E36M50.333
E36M49.333
E37M61.190
E41M59.303
E37M49.100
E37M47.063
E37M49.063
E36M61.125
P35M59.137
E41M49.213
E41M62.122
P44M62.088b
P44M62.089
E36M62.334a
E41M60.058
P35M60.240
P35M60.239
P33M48.153
CDO504Ns.086 5ABDHR
P35M62.167
E38M47.084
E37M60.200a
E36M49.138
P44M62.296
P35M60.100
P33M47.216
E41M48.075
E37M62.155
E37M49.100
P35M49.065b
P33M62.335
E41M60.244
E41M60.079
E38M59.248
P33M62.172
E36M59.265
P35M60.115
E37M49.352
E41M48.056a
E38M59.149
E41M48.056b
P42M62.112
E36M48.158
155
P33M62.208
E37M61.190
E36M59.312
P35M49.251
105
5ABD
E41M48.119a
E36M62.180
P33M59.264
P42M62.191
E38M49.230
E41M47.149
P33M47.211
E38M47.338
E36M59.291
P33M59.104
E37M62.120
5H MWG2230.285
P35M59.376
P33M48.340
5ABD(F)
(F)
7B
E38M59.199
E41M59.199
90
5B
5H
E37M47.244
P33M60.077
CIRC_IM
ANTH_IM
CIRC_rMQM
ANTH_rMQM
BOLT_IM
HGHT_IM
BOLT_rMQM
HGHT_rMQM
160
165
P42M61.139
E37M60.235
P35M49.344
P33M50.292
VRN2.193
P44M49.296
4H-5A
4
40
E38M60.094
5AD
3
35
E37M47.133
E38M60.327
E41M61.091b
E36M61.339b
E36M48.180
E36M59.225
P33M60.366
E37M60.287
E36M61.383
E36M62.311
E38M59.268
E37M61.204
E37M60.398
KSU171.279
P33M50.158
E41M48.280
E37M47.378
E38M60.107b
E41M47.281
2
E37M61.149
E41M60.357
30
1
E38M60.094
25
E41M59.333
E36M49.272
E41M48.244a
E36M59.308
P33M61.089
E37M61.269
E36M61.218
E37M49.073
E41M47.364
E37M60.156
E41M49.094
P35M61.131
E41M59.260
E41M49.120
P33M59.200
GWM205.142
E37M49.250
P33M59.169
GDM101.197
P33M59.170
P35M61.102
TCP2Xm.252
E41M49.333
E38M60.204
E38M47.124
E38M47.147
E38M49.245
E38M47.313
E41M60.205
E37M60.172a
E36M59.133
BCD1130
BCD1707
E36M50.284
KSU220.308
E36M62.330
E36M49.284
E41M48.059
P35M50.363
E41M60.163
P33M61.250
E38M59.139
P44M49.085
P42M61.257
E37M47.307
E36M62.325
P33M50.244
E38M49.191
E41M49.192
GDM068.116
E41M60.267
P33M48.298
E38M49.207b
0
P33M50.274
E38M60.248
E37M60.228
E37M62.163
20
4
P42M62.381
GDM101.196
E38M59.137
E41M59.333
E41M48.244a
E36M59.308
P42M61.162
P33M61.123
P33M59.169
E41M49.094
E37M60.156
E41M61.224
E37M49.251
E41M49.120
E41M59.114a
E38M49.136
E36M50.284
E41M48.336
E41M47.258
E38M47.147
P35M50.363
E36M49.284
E38M60.204
E38M47.313
E41M60.205
E36M61.082
E37M60.172a
E38M49.245
E41M60.164
GDM068.114
P33M61.250
E37M61.291
E38M49.191
P44M49.085
P33M50.244
E37M47.341
E41M60.276
E38M59.139
E41M49.191
E38M60.178
P42M61.100
P33M48.298
E36M61.352
E41M48.062b
P35M49.199
P33M50.197
3
5H
P35M50.281
P33M62.133
P35M62.390
E36M48.290
LG5Xm--TTC2
2
5B
1
0
4
15
3
5H
2
10
P33M60.202
P42M62.081
P35M62.389
TCP2Ns.117
P33M62.093
P33M62.097
E36M59.169
E37M62.172
1
P33M62.175
E41M48.077
E37M62.172
TCP2Ns.117
5
LG5Xm--TTC1
0
4
3
2
1
0
P33M62.133
0
LG6a--TTC1
LG6a--TTC2
E38M47.183
P33M48.087
P35M61.198
P44M62.219
P33M61.168
E37M47.093
E41M61.134
E41M48.235
E37M49.233
P33M61.280
E36M62.114
E38M59.319
P33M48.296
P44M62.200
E36M59.260
P33M48.087
P35M61.198
P44M62.219
P33M61.168
E41M61.134
E37M47.093
E37M60.158
E36M62.114
E41M59.185
E38M59.321
E36M62.374
P44M62.200
E41M47.397
P44M49.151
P33M60.356
E37M62.189
P35M49.269
P33M61.149b
P33M48.131
E37M47.083
E38M60.144
P35M49.164
80
E36M62.292
E36M62.177
E41M47.385
P42M62.269
E41M62.076
E38M49.336
E36M49.310
E38M59.160
P33M48.386
P35M62.209
P42M62.293a
P42M61.099
E41M61.399
E36M59.245
E38M59.291
85
90
95
E36M61.170
E38M47.157
P35M50.389
E41M47.205a
E41M61.259
E41M59.353
P33M60.252
P35M62.313
P35M62.344
E37M61.282
E36M48.329
E41M48.079a
E41M62.150
E41M48.205
P35M62.205
P33M47.170
P33M47.206
E41M47.385
60
P35M60.093
E36M61.112
P35M62.312
E36M48.329
P35M60.106
P44M62.196
55
E41M60.287
E41M61.276
P35M60.093
P33M48.131
E36M61.344
75
4
6B
E41M60.122
E41M47.350
70
3
P35M62.066
P33M60.154
P44M49.121
P33M62.219
P33M47.398
P42M62.206
E37M61.063
6H MWG652.264
P35M49.196
P35M49.194
6B
GIRi5.351
P44M49.065
P33M50.323
6H
45
65
2
P35M50.213b
P33M50.323
P44M49.345
50
1
E38M49.197
E38M49.103
E41M49.122a
E37M49.135
E36M48.257a
P44M49.121
P42M62.205
MWG652.264
P33M62.219
P44M49.205
GIRi7.394
P35M59.091
E37M61.063
E37M60.089
E41M62.334
E41M49.246
E41M62.279
P33M60.203
E41M47.350
40
0
4
3
2
Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved.
LG6b--TTC2
P35M50.213b
30
35
1
0
6
5
4
3
25
2
1
20
0
15
LG6b--TTC1
P35M62.193
P33M60.214
5
10
4
P35M62.193
3
2
1
0
0
P35M62.344
P33M47.133
P35M62.183
E36M62.178a
P35M62.209
E38M49.336
MWG2264.403 6H
E36M49.310
E41M62.113
E41M62.076
E38M47.252
E38M59.161
E36M62.292
E37M47.346
P35M62.251
P33M47.206
P33M47.132
P42M61.099
P33M50.278
E38M59.291
E36M61.170
E38M47.157
E41M61.259
P33M60.252
E41M60.363
E38M60.205
1R
E41M60.098b
E41M47.058
P35M62.183
E36M61.255
E36M48.196a
E41M60.098a
P42M62.163
E37M62.190
E38M59.264
E41M61.105
E36M61.252
E38M49.315
P33M60.310
E41M61.098
E41M62.313
E36M48.327
E41M60.098a
E36M61.255
E37M49.363
TRX2
E36M62.118
E41M62.157
E36M49.274
E37M62.190
P35M60.149
E36M61.252
E37M49.363
TRX2
E38M49.315
P35M60.149
E41M60.363
100
1R
E37M61.133
E41M47.099
7D GDM150.365
E41M60.232
P33M50.258
P42M62.284
P35M50.236
E37M60.290
E41M61.192
P33M60.390
P44M49.312
P44M62.284
P35M61.301
E37M47.366
P35M59.243
P42M61.319
P35M49.381
P35M49.378
P33M62.108
E41M59.252
E37M62.268
E37M60.328
E36M62.219
P35M62.223
105
110
P35M62.224
P35M61.164
P33M62.217
115
P35M60.169
120
P33M62.213
E41M62.185
125
130
135
140
145
E41M62.185
E41M59.349
E37M62.371
P44M62.103
E38M59.286
E38M49.278
E41M59.367
P33M59.267
P35M59.214
P35M50.093
P35M61.205
P35M59.093
E38M47.329
E36M48.152
E37M47.102
E41M59.348
E41M59.368b
P44M62.103
E38M49.278
E38M47.353
E36M48.196b
P33M59.273
E37M61.133
E41M47.099
P42M62.284
P33M50.258
GDM150.365 7D
E41M60.232
P35M50.236
E36M49.295
E41M61.192
P35M49.378
E37M62.268
P35M59.243
P35M49.381
E37M47.366
E37M60.356b
E41M59.291
E41M59.313
P44M62.284
P44M49.070
P33M62.313
P35M62.100
E38M60.334
E36M62.219
P35M59.214
P35M50.093
P35M61.205
E41M60.071
E41M60.071
E36M61.367
P35M60.286
CIRC_IM
BOLT_IM
ANTH_IM
HGHT_IM
CIRC_rMQM
BOLT_rMQM
ANTH_rMQM
HGHT_rMQM
LG7a--TTC1
E41M59.278
E41M60.070
40
P44M62.197b
P42M62.326
E38M60.140
45
E37M62.297
50
55
7H
60
7H(D)
65
6ABDHR
5A,5B,5D
5A,5B,5D
70
75
80
85
90
E41M49.357
E41M49.170
E41M47.162
E36M62.306
MWG741.287
E41M62.351
BCD421
E36M62.225
E37M62.323
MWG669.492
ADP.350
P42M62.289
ADP.190
E38M49.297
E41M48.294
E36M59.391
E36M62.298
E36M49.380
E38M60.092
E41M61.112
E41M50.267
P33M62.126
P33M47.242
E36M59.279
E41M60.338
E37M61.340
E41M62.342
E41M62.103
E38M59.259
E41M47.066
E41M61.313b
E41M47.162
E38M59.342
E36M61.067
E36M59.263
E41M60.336b
MWG741.287 7H
E38M49.136
E36M61.122A
E36M62.225
E37M62.323
E41M61.111b
E41M62.116
P44M62.226
E38M49.297
E41M60.239
E38M60.092
E41M61.267
E41M48.294
E41M61.112
E37M61.340
E41M47.220
E36M49.380
E41M60.338
E41M59.114b
P44M62.288b
MWG669.492 6ABDHR
E37M47.259
E41M60.378
P44M62.063
P44M49.129b
E36M50.314
E38M59.300
P33M48.319b
P42M61.322
P44M49.130
E36M50.202
95
4
E37M49.142
E37M49.143
E38M60.140
3
35
2
E36M61.152b
E37M49.198
E41M49.193
P33M61.326
1
30
0
P33M48.240
4
E38M60.148
3
P44M62.197b
E37M49.382
E41M48.254
LG7b--TTC2
2
E41M62.212
P35M50.219
1
7H
P33M50.143
P33M48.159
E41M48.215
E37M49.340
E41M48.153
MWG703.194 7H
E36M62.088
E36M48.336
P44M62.397
E41M62.212
E41M49.259
E41M48.162a
P35M50.101
E41M59.215
E36M59.095
E36M49.090
E41M61.084a
E37M62.291
E37M62.071
P33M61.387
7H
6-SFT.579
E41M59.072
P44M49.179
P33M50.284
P35M59.166
P33M48.135
P35M61.154
E38M47.385
P33M62.071
E41M60.130
P33M62.352
E38M47.222b
0
6-SFT.101
4
P33M47.119
3
P44M62.094
2
1
Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved.
LG7b--TTC1
0
25
4
20
3
15
LG7a--TTC2
2
10
1
5
E36M49.124
P33M50.361
E38M49.207a
E38M60.308
P33M59.082
P44M62.094
7H
6-SFT.101
P33M50.267
E36M62.193
E41M48.215
E41M48.152
7H MWG703.194
P35M49.204
E41M49.204
E36M61.095
E36M49.061
P33M61.386
0
0
E41M59.215
E41M60.235
6-SFT.579
7H
E36M59.100
E36M62.261
E36M61.316
E37M62.291
E41M61.084a
E38M59.323
P44M49.228b
P33M50.284
E37M61.205
E41M59.072
P35M62.184
P35M59.165
P33M62.071
E38M47.385
E41M60.130
P44M62.117
E36M48.130
E37M61.193
E37M47.078
E41M61.091a
P35M60.246
E36M59.371
E38M59.304
P35M60.247
LIMDEX.227 7H
P35M59.116
P42M62.386
P33M50.349
E38M59.298b
E37M61.193
E41M47.266
E38M49.375
E41M61.090b
E41M61.091a
P35M60.247
E36M59.134
E38M59.304
E41M59.234
P35M60.189
P35M59.116
E37M62.180
E36M49.190
E38M47.096
P44M49.261
P33M48.384
E37M62.180
E41M60.190
E38M47.096
P35M62.109
P33M62.325
E41M47.170
P33M62.325
P35M62.254
P33M48.319a
E36M61.093
E38M59.082
E41M48.321
E41M48.111
E36M61.289a
E36M49.327
E38M60.165
E38M47.171
E37M60.144
E37M47.082
P33M61.368
P33M62.327
P44M62.290
E37M47.222
E41M47.134
E37M60.148
E38M59.082
E41M48.111
E41M59.095
E37M49.380
E37M62.157
E37M60.144
E41M59.203
E41M48.355
E36M62.334b
P44M62.290
E41M59.203
P35M49.347
P35M49.347
P35M61.311
E36M61.320
E38M47.384
E41M48.338
P35M61.382
7A WMC488.105
P33M62.122
E36M61.122
WMC488.105 7A
E41M48.240
E41M59.273
100
E41M60.378
105
110
P44M49.129b
E37M60.246
E36M49.258
E36M50.202
115
E41M62.362
P44M49.130
120
125
130
135
140
145
E36M62.325
E36M59.237
E37M60.107
E41M47.310
E36M61.142
E37M61.374
P33M48.294
P33M50.156
P33M50.142
P35M62.166
E37M47.169
E36M49.270
E38M47.230
P33M48.174
P35M62.280
P33M60.318
P33M48.178
E41M47.257
E37M60.106
E37M60.092
E38M59.327
P35M50.209
P35M62.166
E37M61.374
E36M62.300
E41M62.287
E41M59.252
E37M47.294
P33M60.318
P33M48.174
E41M61.090a
E36M49.269
P35M62.280
P33M48.178
E41M47.257
E41M49.313
E41M60.185
E41M47.313
E38M47.261
E41M49.065
E37M62.230
E36M50.212
P33M47.108
E41M61.262
E41M61.240
E37M61.225
E37M61.283
E41M59.220
P33M48.122
P33M47.230
E41M62.229
P33M60.378
P33M50.332
P35M61.183
E36M49.368
E37M49.356
E37M60.323
E41M47.399
E38M59.151
E36M62.383
E36M50.325
E36M62.172
P33M62.181
E36M48.313
P35M50.285
P35M61.381
E37M49.108
P35M59.237
P33M59.186
E38M49.305
P35M61.311
E36M62.172
P33M48.122
P44M62.156
E41M62.229
E41M48.377
E36M49.101
E36M50.398
E37M49.113
E41M47.255
P33M60.378
E36M48.293
E36M50.325
E36M50.269
P35M62.095
E41M48.384
P33M61.074
E38M49.305
E36M48.313
P35M61.143
P35M61.381
P33M60.225
E41M60.185
150
155
P33M61.074
CIRC_IM
BOLT_IM
ANTH_IM
HGHT_IM
CIRC_rMQM
BOLT_rMQM
ANTH_rMQM
HGHT_rMQM
Histograms for circumference (cm) of plant spreading in the full-sib Leymus TTC1 (n=164)
and TTC2 (n=170) mapping families, based on means of two clones, compared to the
heterogeneous L. cinereus Acc:636 (Lc) and L. triticoides Acc:641 (Lt) progenitors,
interspecific hybrid parents (TC1 and TC2), and recurrent parent (T).
60
55
2002
ID
Lc
Lt
T
TC1
TC2
TTC1
TTC2
Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved.
50
45
40
35
30
AV (SD)
23 (10)
354 (186)
487 (61)
96 (35)
125 (37)
180 (91)
299 (116)
TTC1
TTC2
25
20
15
10
5
0
70
Lc
140
210
280
350
TC1 TC2 TTC1 TTC2
420
Lt
490
560
630
700
770
840
910
980 1050 More
T
50
45
2003
ID
Lc
Lt
T
TC1
TC2
TTC1
TTC2
40
35
30
25
AV (SD)
51 (26)
594 (326)
853 (120)
200 (32)
231 (38)
288 (122)
449 (154)
TTC1
TTC2
20
15
10
5
0
70
140
Lc
210
280
350
420
TC1 TC2 TTC1
490
560
TTC2
630
700
770
840
Lt
980 1050 More
T
40
35
910
ID
Lc
Lt
T
TC1
TC2
TTC1
TTC2
2004
30
25
20
AV (SD)
76 (21)
816 (416)
1263 (177)
256 (39)
282 (53)
362 (137)
548 (170)
TTC1
TTC2
15
10
5
0
70
140
Lc
210
280
350
420
TC1 TC2 TTC1
490
560
630
TTC2
700
770
840
Lt
910
980 1050 More
T
Histograms for proportion of bolting culms in the full-sib Leymus TTC1 (n=164) and TTC2
(n=170) mapping populations, based on means of two clones, compared to the
heterogeneous L. cinereus Acc:636 (Lc) and L. triticoides Acc:641 (Lt) progenitors,
interspecific hybrid parents (TC1 and TC2), and recurrent parent (T).
110
Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved.
100
ID
Lc
Lt
T
TC1
TC2
TTC1
TTC2
2002
90
80
70
60
AV (SD)
0.28 (0.46)
0.67 (0.48)
1.00 (0.00)
0.79 (0.43)
1.00 (0.00)
0.57 (0.43)
0.72 (0.39)
TTC1
50
TTC2
40
30
20
10
0
0.1
0.2
0.3
0.4
0.5
Lc
0.6
0.7
TTC1
Lt
0.8
0.9
1
T
TC2
TTC2 TC1
40
35
2003
ID
Lc
Lt
T
TC1
TC2
TTC1
TTC2
30
25
20
AV (SD)
0.63 (0.43)
0.56 (0.32)
0.98 (0.06)
0.88 (0.19)
0.77 (0.21)
0.33 (0.24)
0.55 (0.26)
TTC1
TTC2
15
10
5
0
0.1
0.2
0.3
0.4
0.5
TTC1
0.6
TTC2 Lt
0.7
Lc
0.8
TC2
0.9
1
TC1
T
40
35
2004
30
25
20
ID
Lc
Lt
T
TC1
TC2
TTC1
TTC2
AV (SD)
0.85 (0.33)
0.78 (0.35)
1.00 (0.00)
0.84 (0.19)
0.81 (0.23)
0.55 (0.26)
0.67 (0.27)
TTC1
TTC2
15
10
5
0
0.1
0.2
0.3
0.4
0.5
0.6
TTC1
0.7
TTC2
0.8
0.9
Lt TC2 TC1 Lc
1
T
Histograms for anthesis date (days from January 1) in the full-sib Leymus TTC1 (n=164)
and TTC2 (n=170) mapping populations, based on means of two clones, compared to the
heterogeneous L. cinereus Acc:636 (Lc) and L. triticoides Acc:641 (Lt) progenitors,
interspecific hybrid parents (TC1 and TC2), and recurrent parent (T).
100
90
Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved.
80
ID
Lc
Lt
T
TC1
TC2
TTC1
TTC2
2002
70
60
50
AV (SD)
176.8 (2.1)
173.3 (2.6)
172.0 (0.0)
173.6 (2.1)
173.1 (1.5)
174.8 (1.9)
173.8 (4.9)
TTC1
TTC2
40
30
20
10
0
166
169
175
178
181
184
187
190
193
T-tester Lt TTC2
Lc
TC2 TC1 TTC1
55
50
172
ID
Lc
Lt
T
TC1
TC2
TTC1
TTC2
2003
45
40
35
30
AV (SD)
171.1 (5.1)
169.3 (1.4)
169.0 (0.0)
169.0 (0.0)
169.0 (0.0)
172.1 (5.7)
172.1 (5.5)
25
TTC1
TTC2
20
15
10
5
0
166
169
172
175
178
181
184
187
190
193
T-tester Lt Lc TTC1
TC1 TC2
TTC2
70
65
60
55
50
45
40
35
30
25
20
15
10
5
0
ID
Lc
Lt
T
TC1
TC2
TTC1
TTC2
2004
AV (SD)
178.3 (3.9)
172.6 (1.6)
173.0 (0.0)
175.9 (4.0)
175.9 (4.0)
175.7 (4.6)
174.2 (4.0)
TTC1
TTC2
166
169
172
175
178
Lt TTC2 TC1
T-tester
TC2
181
Lc
184
187
190
193
Histograms for plant height (cm) in the full-sib Leymus TTC1 (n=164) and TTC2 (n=170)
mapping populations, based on means of two clones, compared to the heterogeneous
Leymus cinereus Acc:636 (Lc) and L. triticoides Acc:641 (Lt) progenitors, interspecific
hybrid parents (TC1 and TC2), and recurrent parent (T).
50
Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved.
2002
ID
Lc
Lt
T
TC1
TC2
TTC1
TTC2
40
30
AV (SD)
99.0 (39.8)
94.5 (12.6)
100.8 (5.7)
145.8 (15.9)
141.8 (10.6)
103.9 (13.8)
109.7 (13.0)
TTC1
TTC2
20
10
0
60
70
80
90
100
110
120
130
140
Lt Lc T TTC1 TTC2
150
160
170
180
TC2 TC1
70
ID
Lc
Lt
T
TC1
TC2
TTC1
TTC2
2003
60
50
40
AV (SD)
135.6 (23.0)
98.5 (8.5)
100.9 (5.9)
155.1 (12.6)
138.4 (13.3)
108.3 (12.9)
109.7 (13.0)
TTC1
TTC2
30
20
10
0
60
70
80
90
100
110
Lt T
120
130
TTC1 TTC2
140
150
Lc TC2
160
170
180
TC1
60
ID
Lc
Lt
T
TC1
TC2
TTC1
TTC2
2004
50
40
30
AV (SD)
182.9 (18.5)
123.3 (12.7)
125.7 (3.4)
184.7 (12.3)
164.6 (8.7)
134.8 (13.8)
137.8 (11.8)
TTC1
TTC2
20
10
0
60
70
80
90
100
110
120
130
140
150
Lt T TTC1 TTC2
160
170
TC2
180
Lc TC1
Description of additional SSR (CNL and KSU) and STS markers added to the Leymus TTC1 and TTC2 maps originally published by WU
et al. (2003)
Loci(groups)
Primers
Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved.
TRX1 (LG1a, LG2b) GGATAACATGACCCTAAAAACT†
GTTGTGCATCACAGGGTTATA
TRX2 (LG6B)
GGATAACATGACCCTAAAAAG†
TTGTTCATCACAGGGTTATC
CNL45
GAGAGAGCTTCGTCCCACTC
CACCACTCGTTTCTTTCACTTT
(LG2A, LG2B)
KSU154 (LG4Ns)
CNL39 (LG4Ns)
BCD1117
(LG4Ns, LG4Xm)
HVCABG (LG4Ns)
HVM068 (LG4Xm)
KSU149 (LG4Xm)
KSU171 (LG5Ns)
MWG2230 (LG5Ns)
VRN2 (LG5Ns)
MWG2264 (LG6a)
ADP (LG7a)
GGAGACTCTGGTCATCTCGC
ATACTGGAGTGAAGGCACGG
TACCTGTGCGGCGATGAAT
CAGGAGCAGGAGAACGTGAA
TCAGTTCTCAATAGAAGTGCTGTG
TCCTGAATAAGGTCTTCATACCAA
ACACCTTCCCAGGACAATCC
CAGAGCACCGAAAAAGTCTGTA
AGGACCGGATGTTCATAACG
CAAATCTTCCAGCGAGGCT
GAGCCACCAGAGCAGAAATC
CGAGCTCCCCTTCTTCTTCT
TCTTGCTTGCATTGTAACCG
TCATGTCTGGGAGCATGGTA
AATGATGTTGCTTTCCTGTTTGCTC
ACAGATGATGATGGCGTGCAGCTTT
AGTACCAGTTCTTCRCCCAAGG
CTGCASYAGGTGAGCCAT
AGGTAGAAGTCAAACTGTGTGGGAT
GTATTACTTTACGAGTTAGATGCTA
CCTCCGTGAACAATTTCCTG
TCCAATACGAGCATTCTTGT
Gene or motif
Anchor loci
Expected
amplicon
Mapped
amplicons§
AY204511 thioredoxin, Xbm2,
1R
287‡ Leymus 289
AY204511 thioredoxin, Xbm2
1R
287‡ Leymus 288
(ga)9, putative RNA binding protein
2A
167‡ wheat
154, 161, 170, 172
(cac)7
4B
146‡ wheat
137
AF251264 (atgc)5 rubisco activase B
3A
220‡ wheat
229
BCD117 barley cDNA clone
4HD
209
152, 211
Rubisco (AT)29
4H
182 barley
152
(GA)22
4H
204 barley
199
(acc)9
4D
228‡ wheat
240, 243
(agt)7
5B
243‡ wheat
279
MWG2230 barley genomic DNA
5H
320 barley
285
Wheat vrn2 (AY485644)
4H, 4A/5A
203 wheat
193,199
MWG2264 barley genomic DNA
6H
400-450
403
1003‡ rice
350 TaqI
190 TaqI
M31616 ADP glucose phosphorylase 5ABD
† FAM 5’ labeled primer.
‡ Amplicon size based on sequence data.
§ Amplicon size based on comparisons with internal size standards in capillary electrophoresis.