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Gene Catalogue 2013
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Catalogue of Gene Symbols

Gene Symbol Class list

Class Count
1 Gross Morphology: Spike characteristics
    Major hexaploid wheat types are categorized into groups with respect to three major gene pairs; viz. Q, C and S1 {1038}.
    1. Common wheat Q c S1 v: vulgare group.
    2. Club wheat Q C S1 v: compactum group.
    3. Shot wheat Q c s1 v: sphaerococcum group.
    4. Spelt wheat q c S1 and q C S1 v: spelta group (including vavilovi).
    The majority of hexaploid wheat stocks are already, or can be readily, classified into these groups.
    Diploid wheat is assumed to be q. Durum and carthlicum groups have the genotype Q {1049}.
    1 Squarehead/spelt
      A nucleotid change in the microRNA172 binding site of the Q locus played a critical role in wheat domestication and the origin of free-threshing modern wheats {11192}.
    2 Club/Compact spike
      QTL:Six QTLs for spike compactness were detected in Courtot/Chinese Spring but only 4 on chromosome arms 1AL, 2BS, 2DS and 4AS were consistent for at least two years{0114} . Two additional QTLs for spike compactness were detected in Courtot/Chinese Spring {10080} on chromosome arms 5DL (QCp.icf-5D) and 6DL (QCp.icf-6D). Markers Xcfd26-5D and Xcfd38-6D explained 13.6% and 12.2% of the variance in spike compactness, respectively {10080}.
      Although gene C may be present in some forms of group macha {1447} and spelta {0623}, it is not universally present. Tsunewaki {1500} found that compact spike in one form was controlled by polygenes.
      C may be orthologous to gene Sog for soft glumes on chromosome 2Am {10578} Tetraploid wheat: A compact spike gene C17648 in mutant line MA 17648 wad located in chromosome 5AL {10541}. Xbarc319-5A - 9.7 cM - C17648 - 24.8 cM - Xgwm179-5A {10541}. C17648 was distal to the Q locus {10541}.
    3 Sphaerococcum
      The naturally-occurring sphaerococcum gene in chromosome 3D and various mutant alleles conferring a similar phenotype form a homoeologous series. The sphaerococcoid alleles are either recessive or incompletely dominant. All three mapped loci are closely linked to the respective centromeres {0030}. The "a" alleles are allocated to Chinese Spring or "normal" wheats.
    4 Branched spike
      Synonyms: branched spike, four-rowed spike, multi-rowed spike, supernumerary spikelet, tetrastichon spikelet.
      Branched spike and multi-rowed spike are phenotypes involving the presence of supernumerary spikelets, or the presence of additional spikelets at rachis nodes. A similar condition in rye is known as 'monstrosum ear' (reviewed in {10637}). Genetic studies of branched spike in tetraploid and hexaploid wheats indicate that the phenotype is recessive, involves one or more genes, and is strongly influenced by environmental effects. Comparative genetic studies suggest an orthologous gene series in homoeologous group 2 {10637}.
    5 Elongated glume
      Elongated glume is the phenotype associated with the polonicum group of tetraploid wheats. Expression in hexaploid wheat is much reduced compared with tetraploids. Matsumura {911} reported linkage of gene P and a gene for red coleoptiles implicating chromosomes 7A or 7B. A different gene was subsequently located in chromosome 7B {9990}.
    7 Multi-gynoecium
      Synonym: three pistils (TP).
      This trait describes a dominant phenotype consisting of 3 kernels within each wheat floret; that is, the flower consists of 3 separate ovaries, 3 anthers and 2 lodicules.
2 Accumulation of Abscisic Acid
    A QTL was mapped on 5AL between Xpsr575-5A {proximal} and Xpsr426-5A {distal} {1180}.
3 Alkylresocinols Content in Grain
4 Aluminium Tolerance
    Allelic variation at the promoter of Almt-D1 is associated with differences in Al tolerance. Molecular and pedigree analysis suggest that Al resistance in modern wheat germplasm is derived from several independent sources {10532}. QTL: Atlas 66/Century: A QTL in the region Xdgm125-4DL - Xwmc331-4DL accounted for nearly 50% of the phenotypic variation in root growth rate in hydroponic solution {10265}. An Al-activated malate transporter (LMT1) was earlier mapped to the same location {10266}.
    Atlas 66 (insensitive)/Chisholm (sensitive) RILs: One QTL, located in chromosome 4DL, corresponded to ALMT1 and accounted for 50% of the phenotypic variation {10483}. A second QTL was located on 3BL(R2 = 0.11); nearest marker Xbarc164-3B {10483}. Both QTLs were verified in Atlas/Century {10483}.
    FSW (A1 tolerant) / ND35 (A1 sensitive): 3 QTLs for tolerance, Qalt.pser-4DL co-segregating with Xups4, a marker for the promoter of the ALMT1 gene, Qalt.pser-3BL (Xbarc164-3B - Xbarc344-3B) and Qalt.pser-2A (Xgwm515-2A - Xgwm296-2A) {10605}.
    In D genome introgression lines of Chinese Spring a major QTL was located in the interval Xgwm125-4D - Xgwm976-4D, R2=0.31 {10598}, probably coinciding with Alt2. A second QTL from CS, Qaltcsipk-3B, R2=0.49, occurred in interval Xgwm1029-3BL - Xgwm1005-3BL in a CS / CS (Synthetic 3B) population {10598}.
5 Anthocyanin Pigmentation
    The genetic determinants of anthocyanin pigmentation of various tissues are largely located in homoeologous regions in group 7, viz. 7BS (Rc-B1, Pc-B1, Plb-B1, Pls-B1) and 7DS (Rc-D1, Pc-d1, Plb-D1), and appear to be linked clusters rather than multiple alleles on each chromosome {10700}. Their relationship with genes for purple auricle and purple pericarp are still not clear.
    2 Purple/Red auricles. Purple leaf base/sheath
      For review see {1641}.
      Melz and Thiele {983} described a "purple leaf base" phenotype where anthocyanin pigmentation extended to the leaf base as well as auricles. Purple leaf base was expressed only when pigmentation occurred in the coleoptiles.
    3 Red/purple coleoptiles.
      There is an orthologous gene series on the short arms of homoeologous group 7. The 'a' alleles confer red coleoptiles. In chromosome substitution lines of wild emmer to common wheat both the 7AS and 7AL derivatives had red coleoptiles, placing Rc-A1 in the centromeric region {10974}. The Rc gene appears to encode a transcription activator of late biosynthesis genes involved in the light-regulation of anthocyanin systhesis (studies carried out on CS(Hope 7A) substitution line) {10317}.
    5 Purple grain/pericarp
      Genes for purple pericarp were transferred from tetraploid wheats to the hexaploid level {112,214,941,1138}. At the hexaploid level duplicate genes {112,941} and complementary genes {112,939,1138,438} were reported. At the tetraploid level, duplicate-gene {941} and single-gene {1327} inheritances were observed. Purple colour is dominant and may be affected by environment and genetic background. Complementary genes were located in chromosomes 3A and 7B {1138}. Possible pleiotropic relationships of genes affecting pigmentation of various tissues have not been studied in detail. Pc2 and Rc-B1a may be the same gene {769}. Also, complementary genes involved in determination of purple pericarp could be related to culm colour {112}.
      For review, see {1643}.
      Complementary dominant genes. A purple line PC was obtained from a cross of non-purple Line 821 (a 7S(7B) substitution from Ae. speltoides) and Line 102/00, a chromosome 2A introgression from T. timopheevii {10946}. Purple grained accessions are unknown in both Ae. speltoides and T. timopheevii. A set of Saratovskaya 29 NILs is described in {11136}.
6 Awnedness
      2 Tipped 1
        In a common genetic background, carriers of B1a have the shortest tip-awned phenotype; carriers of B1b and B1c have awns 2 to 3 times longer depending on environment. In F1 hybrids, differences between the substitution line combinations are significant. The postulation of B1 in both CS and Courtot {0309} based on the phenotype of a CS deletion stock is not supported by genetic observations
      4 Awnless
        Genotypes Hd B2 (e.g., Chinese Spring) and B1 B2 (e.g., Federation) are awnless. Presumably Hd B1 is awnless. Watkins & Ellerton {1551} noted the probability of a third allele "b1a" leading to a half-awned condition, and in discussion they consider the possibility of a similar third allele at the B2 locus. In view of more recent cytogenetic analyses, it seems that the half-awned condition could result from epistatic interactions between the alleles B1 and/or B2 and various promotor genes.
        Although hooded, half-awned, tip-awned and awnless variants occur among tetraploid wheats, these are relatively infrequent. It has not been established with certainty that the above inhibitors are involved.
        The inhibitor alleles have a pleiotropic effect on glume-beak shape {1348}. Acuminate beak is associated with full beardedness and occurs only in b1 b2 types. B2 reduces beak length producing an acute beak shape. B1 reduces beak length producing an obtuse beak shape. In this effect B1 is epistatic to B2.
    2 Promotors
      The effects of (recessive) awn-promoting genes were documented in a number of studies, mainly through monosomic and disomic F1 comparisons, and in tetraploids, whereas Heyne & Livers {549} provided genetic evidence of their effects. A series of "a" genes was documented, but the evidence for the existence of at least some of these was not well supported. Hence symbols for this gene series are not recognized.
    3 Smooth awns
      Smooth-awned tetraploid wheats were reported {016,045,690,1259} and genetic analyses {016,045,690} suggested a single recessive factor, with modifiers in most instances, relative to rough awns. The phenotype has not been reported in hexaploid wheats. No gene symbol is applied.
7 Basal Sterility in Speltoids
    The presence of gene Q ensures the fertility of the first and subsequent florets in wheat spikelets {378}. In speltoids lacking Q, fertility of the second and subsequent florets is ensured by the dominant allele Bs (designated A in {378}) located on chromosome 5D {377}. In the presence of Bs the fertility of the first floret is under polygenic control.
    In bs bs speltoids floret development is under polygenic control, and stocks with varying levels of basal fertility were isolated.
    All group vulgare genotypes so far studied carry Bs.
    The following stocks were described {378}:
    GenotypeApprox. sterile-base score
    Group vulgare----QQ Bs Bs0.00
    Speltoids StFFqq Bs Bs0.00
    StFqq Bs Bs0.08
    St1Aqq Bs Bs0.39
    St1qq Bs Bs0.96
    St2qq bs bs1.41
8 Blue Aleurone
    The Ba allele in T. monococcum spp. aegilopoides acc. G3116 determines a half-blue seed phenotype and is different from the allele present in Elytrigia pontica that determines a solid blue phenotype {282}. They are treated as different genes.
    For review see {1643}.
9 Brittle Culm
    Three independent mutants with brittle tissues were obtained as EMS-induced mutants in T. monococcum accessions PAU 14087 {11002}. The mutations likely affected cellulose synthesis and involved all tissues {11002}.
10 Brittle Rachis
    Brittle rachis in T. durum was defined as a spike that disarticulated when the tip was bent by 45 degrees relative to the peduncle {10242}. In chromosome substitution lines of wild emmer to common wheat, the 3AS derivative was more brittle than the 3BS derivative {10974}.
    Wedge(W) type disarticulation is associated with the Br-1 gene set whereas barrel(B) type disarticulation is caused by a different gene and is limited to species with the D genome {15033}.
11 Boron Tolerance
    Genes controlling tolerance to high concentrations of soil boron act additively. Boron efficiency In contrast to tolerance, boron efficiency was studied in {10135}. Monogenic segregation occured in Bonza (B inefficient)/SW41 (moderately B inefficient) and SW41/Fang60 (B efficient). Two genes, designated Bod1 and Bod2 segregated in Bonza/Fang60.
12 Cadmium Uptake
    Low uptake is dominant.
13 Chlorophyll Abnormalities
    3 Striato-virescens
      A mutant of this type was described {376} but has been lost.
14 Cleistogamous Flowering
    Cleisogamy in barley is controlled by the Chy1 gene, which encodes an AP2 protein. The Cly and cly1 alleles differ by a single nucleotide within the miR172 binding site. Three wheat homologues of Cly1, viz, TaAP-2A, TaAp-2B and TaAp-2D were located in the terminal bins of chromosomes 2AL, 2BL and 2DL, respectively in Chinese Spring and Shinchunaga {11013}. Cleistogamous flowering in durums Cleistogamy, a rare flowering habit in durum wheats, is controlled by a single recessive gene relative to chasmogamy {191}.
    Cleistogamous genotypes clcl. tv: HI8332 {191}; WH880 {191}.
    Chasmogamous genotypes ClCl . tv: IWP5308 {191}; PWB34 {191}; WH872 {191}.
15 Copper Efficiency
    Copper efficiency is a genetic attribute that enhances plant growth in copper deficient soil.
16 Corroded
17 Crossability with Rye and Hordeum and Aegilops spp.
    1 Common wheat
      High crossability of some wheats, particularly those of Chinese origin, viz. Chinese 446 {790}, Chinese Spring {1216}, and TH 3929 {939}, with cereal rye, weed rye (S. segetale L.) {1646}, and other species, e.g., Aegilops squarrosa {691}, Hordeum bulbosum {1387,1397,1469} and H. vulgare {349,693],is determined by additive recessive genes. The kr genes influence crossability with H. vulgare. Allele kr1 is more potent in suppressing crossability than Kr2 which is stronger in effect than Kr3 {1387}. According to Zheng et al. {1649}, the effect of Kr4 falls between Kr1 and Kr2.
    2 Tetraploid wheat
      The Chinese tetraploid, Ailanmai, possesses recessive crossability genes on chromosomes 1A, 6A and 7A with the 6A gene being the least effective {0017}.
18 Dormancy (Seed)
    Seed dormancy in wheat has several components, including factors associated with vivipary and red grain colour. Dormancy is an important component of resistance/tolerance to pre-harvest sprouting (PHS).
    2 Vivipary
      Orthologues of maize viviparous 1 (Vp-1) are located in chromosomes 3AL, 3BL and 3DL {9961} approximately 30 cM distal to the R loci. Variability at one or more of these loci may be related to germination index and hence to PHS {10468}.
      Alleles of Vp-A1 were recognized using STS marker A17-19 {10919}.
      Three sequence variants of Vp-B1 identified in {10468} were used to develop STS marker VpiB3 whose amplified products showed a significant, but not complete, association with germination index used as one measure of PHS.
      Alleles of Vp-B1 were recognised using STS marker Vp1B3 {10615,10621}. There was a suggestion of a relationship between alleles and PHS response {10615}. Vp-B1 allelic identifications for Chinese landraces, historical and current wheat cultivars are listed in {10621}.
    3 Pre-harvest sprouting
      QTL: Several QTL for falling number and alpha-amylase activity, two indicators for pre-harvest sprouting resistance, were identified in {0169}. The most significant were associated with Xglk699-2A and Xsfr4(NBS)-2A, Xglk80-3A and Xpsr1054-3A, Xpsr1194-5A and Xpsr918-5A, Xpsr644-5A and Xpsr945-5A, Xpsr8(Cxp3)-6A and Xpsr563-6A, and Xpsr350-7B and Xbzh232(Tha)-7B {0169}.
      In cross AC Domain/Haruyutaka, one major QTL in chromosome 4AL and two lesser possibly homomeologous QTLs for dormancy in 4BL and 4DL {0226} were found. Tolerance to preharvest sprouting (PHS) in the cross SPR8198/HD2329 was associated with Xwmc104-6B and Xmst101-7D {0032}. In AC Domain (red seeded, PHS resistant) / RL4137 (white seeded, PHS moderately resistant) most measures of PHS occurred as clusters at the R loci. However, QSi.crc-5D for sprouting index, R2=0.44, was independent of seed colour {10626}.
      QTL for preharverst sprouting were identified on chromosomes 3A (associated with Xfbb293-3A at P=0.01), 3B (associated with Xgwm403-3B and Xbcd131-3B at P=0.001), 3D (associated with Xgwm3-3D at P=0.001) and 5A (associated with Xbcd1871-5A at P=0.001) in the population Renan/Recital {0347}. The resistant alleles on the group 3 chromosomes and on 5A were contributed by Renan and Recital, respectively. All QTL for preharvest sprouting co-located with QTL for grain colour {0347}. Zenkoujikomugi/CS: Qphs.ocs-3A.1 on chromosome 3AS was associated with Xbcd1380-3A and Xfbb370-3A accounting for 38% of the phenotypic variation {10195}. Zenkoujikomugi/Spica: White seeded wheats with the dormancy-related QTL, QPhs-3AS from Zenkoujikomugi were more resistant to PHS than counterparts with the contrasting allele from Spica {10377}. White seeded wheats with contrasting alleles of QPhs-4AL were not different {10377}.
      QPhs.ocs.3A-1 was localized to a 4.6 cM interval flanked by Xbarc310-3A and Xbcd907-3A {10245}. A weaker QTL, Qphs.ocs-3A.2 in 3AL, was not associated with TaVp1 {10195}, the wheat orthologue of the maize transcription factor Viviparous-1.
      Qphs.ocs-4A.1 may be the same as a QTL in AC Domain/Haruyutaka due to tight linkage with Xcdo785-4A {10245}.
      QPhs.ocs.4B.1, a CS allele contributing to dormancy, was located in the region of Xgwm495-4B {10245}.
      In cross SPR 8198 (dormant)/HD2329, QPhs.occsu-3A was located in the Xgwm155-3A - Xwmc153-3A region with R2=75% across 6 environments {10261}.
      QTL analyses in several crosses {10275} indicated a common region in chromosome 4A associated with dormancy, dormant genotypes included AUS1408, SW95-50213 and Halberd. The location was consistent with Japanese and U.K. work even though different flanking markers were involved.
      CN10955 (PHS resistant white seeded) / Annuello (PHS susceptible, white seeded) F8 RIL population: QPhs.dpivic-4A.2 in the Xgwm637-4AS - Xgwm937/Xgwm894-4AL region and QPhs.dpivic-4A.1 in the Xwmc48-4AS - Xgwm397-4AS region {10599}.

      Rio Blanco (white seeded, PHS resistant) / NW97S186 (white seeded, PHS susceptible) RIL population: QPhs.pseru-3AS, R2=0.41, Xgwm369-3A - Xbarc12-3A, and one minor QTL {10634}. This major QTL was confirmed in a Blanco / NW98S079 RIL population, R2 up to 0.58 {10634}. Qphs.psweru-3A was fine mapped to a 1.4 cM region flanked by two AFLP markers and was tightly linked to Xbarc57-3A and seven other AFLP markers {10893}.

      RL4452 (red seeded, low PHS tolerance)/AC Domain (red seeded, high PHS tolerance): DH lines: Genes associated with falling number, germination index and sprouting index contributing to PHS were locatged on chromosomes 3A, 4A (locus-2) and 4B in AC Domain and 3D, 4A (locus-1) and 7D in RL4452 {10671}.

      SPR8198 (red seeded, PHS tolerance)/HD2329 (white seeded, PHS susceptible): RIL population: 7 QTL located on chromosomes 2AL, 2DL, 3AL and 3BL, the most important on 2AL and 3AL {10670}.

      Sun325B (dormant white seeded)/QT7475 (semi-dormant white seeded), both parents with the chromosome 4A QTL: DH population: A QTL was located in the Xgwm77-3B - Xwmc527-3B interval (R2=0.19) in the approximate region of the R-B1 locus {10669}.

      Diploid wheat
      QTL:T. monococcum KT3-5 (non-dormant)/T. boeoticum KT1-1 (dormant): RIL population: QTL on chromosome 5AmL, Xcdo1236c-5A - Xabc302-5A), R2=0.2-0.27. Weaker QTLs were found on 3Am(TmAB18 - Xwmc102-3A and Xrz444-3A - TmABF) and 4Am(Xrz261-4A - Xrz141-4A) {0892}. The 3AmQTL co-located with TmABF and TmAB18 {10417}, derived from orthologous ABA signaling genes in Arabidopsis. The 5A QTL may be orthologous to the barley dormancy gene SD1 {10417}.
      Argent (non-dormant, white seeded) / W98616 (dormant, white seeded): 90 DH lines: Strong QTLs on chromosomes 1A, 3A, 4A and 7A and weaker QTLs on 2B, 5B, and 6B, all from W98616 {10740}.
      Association mapping of 198 winter wheat genotypes detected 8 QTLs on 7 chromosomes, viz. 1BS, 2BS, 2BL, 2DL, 4AL, 6DL, 7BS and 7DS {10959}.
19 Ear Emergence
20 Earliness Per Se
    Genes for earliness per se {0023} affect aspects of developmental rate that are independent of responses to vernalization and photoperiod. QEet.fcu.5AL identified in Xfcp359-5A - Xfcp231-5A interval (R2=0.38), at or near the Q locus in Grandin/BR34 {10256}. Grandin was the earlier parent.
    Cutler/AC Barrie: Three QTLs were mapped on chromosomes 1B (QEps.dms-1B.1 and QEps.dms-1B.2) and 5B QEps.dms5B) {11039}.
21 Embryo Lethality
    1 Embryo lethality in wheat x rye hybrids
      The Chinese Spring (Omperial rye) addition lines 6R and 6RL crossed with different inbred rye lines (R2, R6, R7) produced hybrid seeds with different proportions of differentiated embryos. R2 with (Eml-R1a) gave only undifferentiated embryos; R6 and R7 (with Eml-R1b) gave 74-100% differentiated embryos {10748}. Cross of R2 with the CS nulli-tetrasomics gave differentiated embryos only with N6AT6B and N6AT6D, indicating the presence of a complementary factor Eml-A1 chromosome 6A {10748}.
22 Flag Leaf Width
    Two NILs in backgrounds of Mianyang 99-323 and PH691 possessing Fhb5 in a Xbarc303-5A - Xbarc100-5A interval from Wangshuibai spanning the centromere had a narrow leaf phenotype. QFlw.nau-5A, re-designated as TaFLW1, was mapped to a 0.2 cM region, Xwmc492-5A - Xwmc752-5A: bin 5AL12-0.37-0.57, and was separated from Fhb5: bin 5AS3-C-0.75 {10934}.
23 Flowering Time
    The isolation of wheat genes orthologous to the Arabidopsis Co and rice Hd1 genes was reported in {10054}. The genomic clones TaHd1-1, TaHd1-2 and TaHd1-3 originate from the long arms of chromosomes 6A, 6B and 6D, respectively. The orthology of the TadHd1 genes with Co/Hd1 was demonstrated by complementation of a rice line deficient in Hd1 function with the TaHd1-1 genomic clone. It should be noted that the wheat TaHd1 and rice Hd1 genes are located in non-syntenic locations {10054}. To date, no variation for flowering time has been identified on the wheat group 6 chromosomes.
    Winter wheat cross, Arina (149 days)/Forno (150 days): Six QTL were detected over six environments. The 3 most important, all from Arina, were in chromosomes 6DL (R2=16%), 3DL (R2=14%) and 7BL (R2=13%); 3 others in 2AL, 5BL and 6DL were from Forno {10172}.
    Winter wheat cross Ernie (early)/MO94-317 (late), days to anthesis (dta):
    Qdta.umc-2D, linked to Xbarc95-2D, R2 = 0.74 {10456}.
    Spring wheat cross: Nanda 2419 / Wangshuibai: Seven QTLs for flowering time identified with earlier alleles for five coming from Nanda 2419: QFlt.nau-1B (closest marker Xbarc80-1B, R2=11 %), QFlt.nau-1D (Xbarc62-1D, Xgwm232-1D, R2=6.13 %), QFlt.nau-2B (Xwmc35-2B, R2=10 %), XFlt.nau-2D (Xwmc601-2D, R2=10 %), XFlt.nau-4A.1 (Xcfd2-4A, Xmag1353-4A, R2=10 %), XFlt.nau-4A.2 (Xmag3386-4A, Xwmc161-4A, R2=18-19 %), XFlt.nau7B (Xmag2110-7B, Xmag1231-7B, Xgwm537-7B, Xwmc218-7B, R2=18 %) {10566}. Heading date QTL: CI 13227/Suwon 92 RIL population: AFLP marker - 2.6 cM - QHd.pser-2DS - 121.1 cM - Xgwm261-2D {10269}. This QTL could be Ppd-D1 {10269}.
    Karl 92*2/TA 4152-4 F2:F4 population: Two QTLs, QHd.ksu-2D, associated with Xgwm261-2D (R2=0.17), and QHd.ksu-3D, associated with Xgwm161-2D 9 (R2) {10273}.
24 Flour Colour
    Loci controlling flour colour were identified and mapped in a recombinant inbred population derived from Schomburgk/Yarralinka {9936}. Regions in 3A and 7A accounted for 13% and 60% of the genetic variation, respectively, and Xbcd828-3A, Xcdo347-7A and Xwg232-7A.1 were significantly associated with flour colour. The association was highly significant in all three replicates only for the 7A QTL. Symbols were not assigned to the flour colour loci. See also 29.2. Flour, semolina and pasta colour. Lutein is one of the carotenoids contributing to flour colour. Esterification of lutein contributes to its stability during storage. A locus controlling esterification was located in chromosome 7D.
    Lutein esterification
25 Free-threshing Habit
26 Frost Resistance
    QTL:Norstar(tolerant)/Winter Manitou(non-tolerant): DH population: Norstar possessed major and minor QTL for tolerance on chromosomes 5A and 1D. The 5A QTL was 46 cM proximal to the vrn-A1 locus (R2=0.4); its peak co-incided with Xwmc206-5A and Xcfd2-5A, and expression of C-repeat Binding Factor genes with strong homology to Cfb14 and Cfb15 located at the Fr-2 locus in T. monococcum {10414}.
27 Gametocidal Genes
28 Gibberellic Acid Response (insensitivity)
29 Glaucousness (Waxiness/Glossiness)
    Glaucousness refers to the whitish,wax-like deposits that occur on the stem and leaf-sheath surfaces of many graminaceous species. The expression of glaucousness depends on the arrangement of wax deposits rather than the amount of wax {603}. Non-glaucous variants also occur and genetic studies indicate that non-glaucousness can be either recessive or dominant. Recessive forms of non-glaucousness are apparently mutants of the genes that produce the wax-like deposits. Dominant non-glaucous phenotypes (as assessed visually) appear to be due to mutations that affect the molecular structure, and reflectance, of the wax-like substances {10001}. The genes involved in wax production and the "inhibitors" are duplicated in chromosomes 2B and 2D. There appear to be independant genes for wax production and "inhibitors" {912,1493,10001}. In earlier issues of the gene catalogue the two kinds of genes were treated as multiple alleles {1432}. All forms of wild and cultivated einkorn are non-glaucous {10001}. Orthologous loci occur in barley chromosome 2HS (gs1, gs6, gs8) {467}, rye chromosome 7RL (wa1) {725} and maize (gl2) {211}. A gene for spike glaucousness, Ws, was mapped distally on chromosome 1BS in the cross T. durum cv. Langdon / T. dicoccoides acc. Hermon H52 {0171}.
    2 Epistatic inhibitors of glaucousness
      Each inhibitor inhibits all genes for glaucousness. A dominant gene (Vir) for non-glaucousness was located in chromosome 2BL of cv. Shamrock, a derivative of T. dicoccoides {10543}. This gene mapped 2 cM distal to Xgwm614-2B {10543} whereas the W1/Iw1 locus was placed distal to Xgwm614-2B in {10189}. Lines with Vir had delayed senescence ('staygreen') and an average yield advantage over their glaudous sibs {10543}. Although maps constructed from three tetraploid crosses suggested that w1, W1 and Iw1DIC = Vir remain unresolved {10815}. QTL
      Leaf glaucousness
      RAC875 (glaucous) / Kukri (non-glaucous). Several QTL affected leaf glaucousness, the strongest of which was QW.aww-3A; QTL of lesser effect QW.aww-3B and QW.aww-3D were detected at homoeoogous regions on chromosomes 3B and 3D {11131}.
30 Glume Colour and Awn Colour
    Black glumes are now included in the following homoeologous series with red/brown/bronze glumes.
    1 Red (brown/bronze/black) glumes
      The majority of studies report a single dominant gene for red glume colour. A few papers report two factors {1009,1477,1520}. Red glume colour in Swedish land cultivars is apparently associated with hairy glumes {1277} suggesting, because Hg is located in chromosome 1A, that a red glume factor different from Rg1 is involved in the Swedish stocks. Nothing was known of the possible association of such a gene with Bg, another glume colour gene on chromosome 1A. See {1640} for review. A chromosome 1A gene, Rg3, was eventually identified by linkage with Gli-A1 {1405} and shown to cosegregate with Hg {624}.
    2 Pseudo-black chaff
      This is a blackening condition transferred from Yaroslav emmer to Hope wheat by McFadden at the same time as stem-rust resistance was transferred. The association of this condition with mature-plant stem-rust reaction (Sr2) has been noted in a number of papers. According to {742}, the condition is recessive. Pan {1102} considered linkage with stem-rust reaction could be broken, but this seems unlikely.
    3 Black-striped glumes
      This phenotype was reported in group dicoccon. v: E4225 {1417}.
    4 Inhibitor of glume pigment
      An inhibitor of glume pigment was reported on chromosome 3A {106}.
    6 Awn colour
      The literature on awn colour is not clear. In general, awn colour is associated with glume colour {045}. Occasionally, however, awn colour and glume colour may be different. According to Panin & Netsvetaev {1103}, black awns were determined by three complementary genes designated Bla1, Bla2, Bla3. Bla1 was located in chromosome 1A and linked with Gld 1A (= Gli-A1) and Hg.
31 Grain Hardness/Endosperm Texture
    Grain hardness or endosperm texture significantly influences flour milling, flour properties and end-use. The difference in particle size index between a hard wheat (Falcon) and a soft wheat (Heron) was reported by Symes {1452} to be due to a single major gene. Symes {1452} also found evidence for "different major genes or alleles" which explained differences amongst the hard wheats Falcon, Gabo and Spica. Using Cheyenne (CNN) substitution lines in CS and a Brabender laboratory mill, Mattern et al. {915} showed that the hard wheat milling and flour properties of Cheyenne were associated with 5D. Using Hope 5D substitution line in CS [CS(Hope 5D)] crossed to CS, and CS(Hope 5D) crossed to CS ditelosomic 5DL, Law et al. {777} showed that grain hardness was controlled by alleles at a single locus on 5DS. The dominant allele, Ha, controlling softness was present in Chinese Spring and the allele for hardness, ha, was present in the others. A similar study using CS (CNN5D)/CS recombinant inbred lines was reported by Morris et al. {03106}.
    A pleiotropic result of hardness is the decreased level of a 15 kD starch granule protein, friabilin, on the surface of water-isolated starch {470}. In endosperm, soft and hard wheats have similar amounts of friabilin, consequently the distinction between the two textural types depends upon the manner in which the friabilin co-purifies with starch. Friabilin is also referred to by the name 'Grain Softness Protein' (GSP) {0384}, and was later shown to be comprised primarily of puroindoline a and puroindoline b {0295}. Grain hardness of reciprocal soft x hard F1 kernels was well correlated with friabilin occurrence on starch in triploid endosperm {0381}. See IV, Proteins: 5.8 Puroindoline. GSP-1 genes, which are closely related to puroindolines, are also listed in the Protein section. Two QTLs were detected for grain hardness in RILs of the ITMI population (Synthetic / Opata 85) {10051}. The QTL on the short arm of chromosome 5D was associated with Xmta10-5D, and increased hardness was contributed by Opata {10051}. The locus located proximally on the long arm of 5D was associated with Xbcd450-5D and increased hardness was contributed by the Synthetic allele {10051}.
    Two QTLs, QHa.ksu-3B, associated with Xksum9-3B (R2=0.09, and QHa.ksu-5D(Ha), associated with Xcfd-5D (R2=0.3), were identified in Karl*2/TA 4152-4 {10273}.
    Using proteomic analysis of 2D-protein gels applied to 101 lines of the Opata/W-7984 (ITMI) RI mapping population, and after a preliminary study of a sub-group of these lines {10086}, 446 amphiphilic protein spots were resolved, 170 specific to either of the two parents and 276 common to both {10087}. An important category of these proteins comprises the puroindolines. Seventy-two loci encoding amphiphilic proteins were conclusively assigned to 15 chromosomes. At least one Protein Quantity Locus (PQL) was associated with each of 96 spots out of the 170 spots segregating; these PQL were distributed throughout the genome. The majority of the amphiphilic proteins were shown to be associated with plant membranes and/or play a role in plant defence against external invasions. Not only the puroindolines were associated with kernel hardness - a number of other amphiphilic proteins were also found to influence this trait.
    Neixiang 188 (hard)/Yanshan 1 (medium hard) RIL population: QGh.caas-1B.1 with hardness allele from Yanshan 1, R2=0.28, Xwms153-1BL - Xbarc81-1BL{10640}
32 Grain Quality Parameters
    In the comprehensive study of 46 quality-related traits in a RL4452/AC Domain RIL population, 99 QTLs involving 41 traits were located in 18 chromosomes {10361}; 14 QTLs clustered in the Glu-1B region (50cM), 20 QTLs occurred in the Xwmc617-4D - Xwmc48-4D region (30cM), 10 QTLs mapped to the Xgwm130-7D - Xwmc405-7D region (14cM) and 66 QTLs were dispersed {10361}. In a large study of 11 seed quality traits in a AC Karma/87E03-S2B1 DH population, 26 QTLs were detected in 7 chromosomes {10434}; 6 were clustered in the Glu-D1 region and 5 were clustered in the Rht-D1 region.
    QTL analyses of 10 milling and baking quality traits (grain hardness, flour yield, grain and flour protein, alkaline water retention capacity (AWRC), sedimentation properties, cookie properties, lactic acid retention, dough strength, extensibility and mixograph properties) in the ITMI population grown in Mexico, France and USA (California) are reported in {10436}.
    Neixing 188 / Yanshan 1 RIL population: 75 QTLs for 5 quality-related traits are reported in {10640}.