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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}.
    2 Flour, semolina and pasta colour
      QTL: A QTL was detected on chromosome 7A {9936}. Cultivar Schomburgk contributed the yellow colour allele in a cross Schomburgk/Yarralinka {9936}. Markers Xcdo347-7A and Xwg232-7A accounted for 60% of the genetic variation {9936}. A Sequence Tagged Site PCR marker is available {0180}.
      A major QTL was detected in the distal region of chromosome 7BL in the cross Omrabi 5/ T. dicoccoides 600545. The QTL explained 53% of the variation and was completely linked to microsatellite marker Xgwm344-7B. Omrabi 5 contributed the allele for high yellow pigment level. Two additional small QTLs were detected on 7AL {0365}. Other references to flour colour are given under Flour Colour, Lr19, and Sr25.
      W9262-260D3 (low yellow colour)/Kofa (high colour): Four QTLs identified on chromosomes 2A (Xgwm425-2A), 4B (Xgwm495-4B), 6B (Xgwm193-6B) and Psy-B1 (chromosome 7BL) {10230}. See also Enzymes Phytoene synthase.
      Analysis of yellow flour pigment in a RIL population of PH82-2 (low)/Neixiang (high) revealed major QTL on chromosomes 7A co-segregating with marker YP7A (R2 = 0.2 - 0.28) (see Phytoene synthase 1), and 1B (R2 = 0.31 - 0.54) probably contributed by 1RS {10501}.
      Huapei 3 /Yumai 57: DH lines: 18 additive QTLs and 24 pairs of epistatic QTLs affected flour colour parameters; qa-1B, closely linked with Xbarc372-1B was associated with variation of a*, R2=0.256 {10625}.

      A further study confirmed major QTL on chromosome 1RS (R2=0.319) and 7A (R2=0.339), minor QTL occurred on 1A and 4A {10716}.
    3 Amylose content
      Amylose content has a significant effect on industrial quality; for example, reduced amylose wheats perform better in some types of noodles. The waxy protein genes have an important influence, but other genes are also involved.
    4 Milling yield
      QTL: A QTL was detected on chromosome 3A {0181}. Cultivar Schomburgk contributed an allele for the higher milling yield in cross Schomburgk/Yarralinka {0181}. RFLP markers Xbcd115-3A and Xpsr754-3A were associated with this QTL at LOD>3 {0181}.
      A QTL associated with Pinb on chromosome arm 5DS was detected in RILs from the cross NY6432-18/Clark's Cream {0241}. Cultivar Clarks Cream contributed the higher flour yield allele {0241}. This QTL coincided with QTL for hardness, hydration traits (dough water absorption, damaged starch and alkaline water retention capacity (AWRC) and baked product traits (cookie diameter and cookie top grain) {0241}.
    5 Alveograph dough strength W
      QTL: QTLs for W were detected on chromosome arms 5DS (associated with Xmta10-5D), 1AS (associated with Xfba92-1A), and 3B (associated with XksuE3-3B) in cross Courtot/Chinese Spring {0141}. The first two QTLs coincided with those for hardness.
      Ten QTL for W (39% of the variation), nine QTL for P (48% of the variation) and seven QTL for P:L (38% of the variation) were mapped in Forno/Oberkulmer spelt {0280}.
    6 Mixograph peak time
      QTL: A QTL associated with Glu-Dy1 on chromosome arm 1DL was detected in RILs from the cross NY6432-18/Clark's Cream {0241}. Clark's Cream contributed the higher mixograph peak time allele {0241}. This QTL coincided with a QTL for bread mixing time {0241}.
    7 Starch characteristics
      The Isoamylase-1 gene from Ae. tauschii (Iso-1) complements the deficient rice sugary-1 mutant line {10295}.
      QTL: QTLs for starch viscosity and swelling were associated with the Wx-B1 locus in Cranbrook (Wx-B1a)/Halberd (null Wx-B1b). An additional QTL for starch viscosity was found on 7BL between markers Xgwm344-7B and Xwg420-7B in the first cross. This QTL disappeared when amylase activity was inhibited indicating that it was determined by the late maturing a-amylase activity contributed by Cranbrook. A QTL for starch viscosity was associated with the Wx-A1 locus in the cross CD87/Katepwa {0362}.
    8 Loaf volume
      QTL: Loaf volume score consistent across three environments was scored in a RIL population Renan/Recital and revealed major QTL on chromosomes 3A (flanking markers Xfbb250-3A, Xgwm666-3A, positive effect from Renan) and 7A (flanking markers Xcfa2049-7A, Xbcd1930-7A, positive effect from Recital) {10536}.
      A total of 30 QTLs were located on 12 chromosomes, each of which explained between 5.85 and 44.69% of the phenotypic variation; the QTLs of largest effect were located on chromosomes 6B and 6D {10659}.
    9 Dough rheological properties
      QTL: In a Cranbrook/Halberd DH population, environmental factors were a major determinant of dough extensibility whereas additive effects of alleles at the hight and low molecular weight glutenin loci determined dough strength {10247}.
    10 Grain fructan content
      Fructans are non-digestible carbohydrates considered to have health benefits to consumers.
      QTL: Berkut (high fructan concentration) / Krickauff (low fructan concentration): QTL detected on chromosomes 2B, 3B, 5A, 6D, and 7A of which QGfc.aww-6D.2 (R2=0.17, nearest marker, Xbarc54-6D) and QGfc.aww-7A.1 (R2=0.27,Xgwm681-7A )had the largest effects {10631}.
    11 Water absorption
      Neixiang 188 / Yanshan 1 RIL population: XAbs.caas-5D.1 with positive effects from Yanshan 1, R2=0.3, Xcfd18-5DS - Xcfd189-5DS {10640}
    12 Chinese dry noodle quality
      Chuan 35050 / Shannong 483 RIL populations: 3 QTLs for noodle palate, elasticity and smoothness clustered near Glu-D1 with beneficial effects associated with subunits 5+10 coming from Chuan 35050. A very significant taste QTL, QStas.sdau-4A.1 and positive QTLs for stickiness and total score also on chromosome 4A came from Shannong 483 {10647}
33 Grain Traits
    Variation in grain traits based on gene homology with other species
34 Grain Weight
    QTL : Variation at locus QGw1.ccsu-1A, associated with Xwmc333-1A, accounted for 15% of the variation in a RIL population from RS111/CS {0165}.
    Rye Selection 111 (high GW)/CS (low GW) RIL: two definitive QTLs QGw.ccsu-2B.1 and QGw.ccsu-7A.1 and one tentative QTL, QGw.ccsu-1A.1, were detected by CIM analysis {10363}. The chromosome 7A QTL co-located with a QTL for early heading {10363}.
35 Grass-Clump Dwarfness/Grass Dwarfness
    Complementary dominant genes. Genotypes producing dwarfness: D1-D2-D3-, D1-D2D2, D1-D4-D3-, D1-D2-D4 and D1-D4D4.
36 Growth Rate and Early Vigour
    QTL analyses in Ae. tauschii: chromosomes 1D, 4D, and 7D carried QTLs for relative growth rate, biomass allocation, specific leaf area, leaf area ratio, and unit leaf rate. Chromosome 2D had QTLs for rate and duration of leaf elongation, cell production rate, and cell length. Chromosome 5D harbored QTLs for total leaf mass and area, number, and growth rate of leaves and tillers {10293}.
37 Hairy/Pubescent Auricles
38 Hairy Glume
39 Hairy Leaf
    A QTL analysis of the ITMI population identified loci determining hairiness of leaf margins and auricles in regions of chromosomes 4B and 4D orthologous to Hl1 {10516}.
40 Hairy Leaf Sheath
    Levy & Feldman {795} concluded that complementary genes determined hairy leaf sheath in T. dicoccoides.
41 Hairy Neck/Pubescent Peduncle
42 Hairy Node/Pubescent Node
    Inheritance of hairy (glabrous) node versus non-hairy node was attributed to a single, dominant gene difference {396,837,910,914} and the Hn/hn locus was shown to be linked with B1 (awn inhibitor). Observations on 5A trisomics and telosomics of Chinese Spring confirmed this location. Love & Craig {837} studied a cross involving Velvet Node CI 5877, and Gaines & Carstens {396} studied an offtype single plant designated Velvet Node Wash. No. 1981.
43 Heat tolerance
    QTL: QTLs contributing to grain-filling duration (GFD) under high temperatures were associated with Xgwm11-1BS (11% of variability) and Xgwm293-5AS (23% of variability) in Ventnor (tolerant) // Karl 92 (non-tolerant) {0327}.
44 Height
    Ht is the general symbol.
    2 Reduced Height : GA-sensitive
      Borner et al. {116} found no evidence of orthologous GA-sensitive genes in rye, but reviewed evidence for orthologous GA-insensitive genes. The close linkage of Rht8 and Xgwm261-2D permitted the use of the microsatellite as a marker for the detection of allelic variants at the Rht8 locus{9962}.
45 Herbicide Response
    2 2,4-D tolerance
      Randhawa et al. {1190} reported a single dominant gene in each of WL711, CPAN1874 and CPAN1922 controlling tolerance. HD2009 and PBW94 were described as susceptible.
    4 Imidazolinone resistance
      Resistance alleles found in mutagenized populations were incompletely dominant and additive in effect {10099}. Resistance is due to single base pair changes in acetohydroxyacid synthase.
46 Hybrid Weakness
    1 Hybrid necrosis
      [Progressive lethal necrosis {155}; Firing {971}].
      Complementary dominant genes. Descriptive alleles w (weak), m (medium) and s (strong) were allocated by Hermsen {532}. Phenotype is affected by modifying genes (and/or genetic background) and environment {566}. According to Dhaliwal et al. {257} progressive necrosis is suppressed at 28C.
    4 Apical lethality
      Apical lethality is caused by complementary recessive genes and is characterized by stunting and tiller death at the 4-5 leaf stage. The lethal genotype was designated apd1 apd1 apd2 apd2 {10492}.
47 Iron Deficiency
48 Lack of Ligules
    The liguleless character is controlled by complementary recessive genes in hexaploid wheat {077,738,942} and by a single recessive in tetraploid wheat {047,050,939,10133}. One gene at the tetraploid level is allelic with one of those in the hexaploid {939,10133}. Evidence for orthology of lg1 and lg2 with lg of rice {170}, lg1 of maize {004}, li of barley {1155} and al of rye was presented in {725}. al: Imperial rye chromosome 2R restored the liguled condition to a liguleless CS derivative {939}. Genotypes of selected tetraploid wheat {10133}
    Lg1Lg1 Lg3 Lg3: T. turgidum var. durum Ldn - dic DS 2A: T. turgidum var. dicoccum
    Khapli and Vernal; T. turgidum var. dicoccoides Israel A; MG4343
    Lg1Lg1 lg3 lg3: T. turgidum var. durum: Altaiskaya Niva; Castelpoziano; Langdon; Ldn-GB DS 2B; Golden Ball; Modoc; PI349056
    lg1lg1 Lg3Lg3: None identified.
49 Leaf Erectness
50 Leaf Tip Necrosis
51 Lesion Mimicry
    Lesion mimics that resemble the response of plants to infection by pathogens occur in many species ({10743} for examples).
52 Lodging
53 Male Sterility
54 Manganese Efficiency
    QTL: Variation associated with Xcdo583-4B explained 42% of the variation for Mn efficiency in the durum cross Stojocri 2 (Mn efficient)/Hazar (MN inefficient) {0320}.
55 Maturity time
56 Megasporogenesis
57 Meiotic Characters
58 Nitrate Reductase Activity
59 Nuclear-Cytoplasmic Compatability Enhancers
60 Nucleolus Organizer Regions
    1 18S - 5.8S - 26S rRNA genes
      NORs have been observed as secondary constrictions associated with nucleoli on satellited chromosomes {e.g.,221}, and by in situ hybridization to chromosome spreads {039,294,1014} of 18S-5.8S-26S ribosomal-DNA probes {038,433}. Allelic variation in gene number has been demonstrated at all wheat Nor sites and at Nor-R1 by filter {367} and in situ hybridization {1012}. Allelic variants of the Nor loci are detected by hybridization of rDNA probes to restriction endonuclease-treated DNA on Southern blots {037,288,917,1399}. Alleles Nor-B2a to Nor-B2f were identified using Taq1 digests of genomic DNAs hybridized to derivatives of the plasmid pTa250 {433} containing spacer-DNA fragments pTa250.4 {367,917} and pTa250.15 {288}.
      Other variants may have been isolated {1399} using BamH1/EcoR1 double digests and pTa71 {433}. The variants may or may not be equivalent to those described below. More detailed listings for allelic variation at Nor-B1 and Nor-B2 are given in {917,918}.
      Two sites designated temporarily as Nor-Ax and Nor-Ay were identified in T. monococcum ssp. boeoticum, but were absent in ssp. urartu.
61 Osmoregulation
    Osmoregulation is a specific form of solute accumulation regulating turgor pressure and hydration during periods of stress with positive effects on growth. Wheat lines selected for higher osmoregulation in the greenhouse have greater growth and seed yields under water limited conditions in the field.
62 Phenol Colour Reaction of Kernels
    Wheat genotypes vary in response when caryopses are treated with weak solutions of phenol, a dark colour response being indicative of a positive response. This response is believed to be related to the action of tyrosinase. There seems to a genetic relationship with polyphenol oxidase activity which causes a darkening of flour, pasta and noodle products (see also Polyphenol Oxidase (PPO) activity).
63 Pollen Killer
    Kato & Maeda {10164} reported both partial pollen and seed sterility in crosses involving certain landraces and Chinese Spring. They attributed sterility to recessive alleles of three complementary genes. The genes were designated Ki2, Ki3 and Ki4 {10164}, but the relationship of Ki3 to the earlier designated Ki was not established. Some genotypes:
    Ki2 Ki3 Ki4: v: Aka Kawa Aka {10165}; Hope {10165}; Marquis {10165}; Red Russian {10165}
    ki2 Ki3 Ki4: v: Akadaruma {10165}; Canthatch {10165}; Norin 61{10165}; Pakistani Landrace IL159 {10164}
    Ki2 ki3 Ki4: v: Gabo {10165}; Thatcher {10165}; Timstein {10165}; Zlatiborka {10165}
    Ki2 Ki3 ki4: v: Kagoshima {10165}; Komugi Jingoro {10165}; Sakobore {10165}
    ki2 ki3 Ki4: v: Finnish Landrace WAG4339 {10165}; Hungarian Landrace WAG4458 {10165}; Novosadska Jara {10165}
    ki2 Ki3 ki4: v: Chinese Spring {10165}; Eshima Shinriki {10165}; Ethiopian Landrace IL70 {10164}; Norin 26 {10165}
    Ki2 ki3 ki4: v: Cadet {10165}; Iraqi Landrace IL171 {10165}; Rex {10165}
64 Polyphenol Oxidase (PPO) Activity
    3,4 dihydroxyphenylalanine (L-DOPA) was used as a substrate in a non-destructive test of polyphenol oxidase activity in seeds. Chromosome 2D was shown to carry PPO gene(s) based on Langdon/Chinese Spring (2D) substitution lines and nullisomic-tetrasomic analysis {0342}. An orthologous series of genes affecting PPO activity in both common wheat and durum was proposed in {10149}. See also, Phenol Colour Reaction of Kernels
    Chara (mod high)/WW2449(low): DH population: PPO activity Associated with Xgwm294b-2A (R2=0.82), Xwmc170-2A, Xgwm312-2A and Xwmc178-2A (R2>0.7) {10410}.
    Chara (medium high PPO)/WW2449 (low PPO): one QTL was located on chromosome 2A. Two markers (one SNP, one CAPS) based on BQ161439 were polymorphic between the parents and showed linkage or allelism with PPO loci Xtc1 and XPPO-LDOPA. Xtc1 - 0.6 cM - XPPO-LDOPA/XPPO18/BQ161439 {10484}.
    A QTL on 2D, associated with Xfba314-2D was identified in an M6 / Opata 85 population using the L-DOPA assay. The high PPO activity was contributed by M6 {0344}. Markers significantly associated with PPO activity were also detected on chromosomes 2A, 2B, 3B, 3D and 6B in the population NY18 / Clark's Cream {0344}.
    A multiplex of markers PPO33 and PPO16 was reliable for selecting genotypes with low PPO activity {10418}.
    Tetraploid wheat
    Messopia/T. dicoccoides: RILs: Associated with RFLP Xutv1427-2A {10411}.
    Jennah Khetifa (high)/Cham 1 (low): Associated with Xgwm312-2AL {10411}. STS marker PPO18 based on a polyphenol oxidase (PPO) gene (GenBank AY596268) was closely linked to SSR markers Xgwm312 and Xgwm294 on chromosome arm 2AL. PPO18 explained 28-43% of the variation in PPO activity in the cross Zhongyou 9507/CA9632 {10290}.
65 Red Grain Colour
    Red colour is probably due to the polyphenol compounds phlobaphene or proanthocyanidin, synthesized through the flavanoid pathway. Himi & Noda {10107} provided evidence that the R genes were wheat forms of Myb-type transcription factors (Tamyb10-3A, Tamyb10-3B,Tamyb10-3D). Genetic evidence is provided in {10838}.
    Red colour is dominant to white. At each locus, the white allele is assigned a and the red allele, b. White-grained T. aestivum and amber-grained T. durum wheats carry recessive a alleles at each locus. White-grained CS*7/Kenya Farmer and CS*6/Timstein are considered near-isogenic to CS with R-D1b. Functional markers based on Tamyb10 sequences are given in {10838}.
66 Reaction to Black-Point of Grain
    Black-point, a common grain defect, is a dark discoloration of the embryo region of the kernels. Whereas black-point is often attributed to infection by a number of fungi, the presence of such fungi may be a consequence of saprophytic colonization of affected tissues rather than the cause (see {10148} for references). The condition may be triggered by high humidity {0845}.
    QTL: Sunco/Tasman DH populaion: QTL located in chromosomes 2B (15% of phenotypic variation), 3D, 4A (from Sunco) and 1D, 5A and 7AS (from Tasman {10148}. The 2B gene was associated with the presence of Sr36 {10148}.
    Markers Xgwm319-2B and Xgwm048-4AS were confirmed in a Batavia/Pelsart (resistant) DH population {10494}.
    Cascades/AUS1408 DH population: QTL from Cascades located in chromosomes 2D (5 cM from Xgwm484-2D, 18% of phenotypic variation), 2A (13%), and 7AS (12%) {10148}.
67 Response to Photoperiod
    One-gene {1169} and two-gene {638,1137,1170} differences were reported in inheritance studies. In Chinese Spring/Hope substitution lines for chromosomes 1A, 4B and 6B greater sensitivity to short photoperiod was found, whereas substitutions of 3B and 7D were less sensitive {487}.
    'a' alleles are dominant.
    There is an orthologous gene series on the short arms of homoeologous group 2. The "a" alleles confer the insensitive response {0063}, the contrasting allele may be referred to as "b". QTL: Trident (early)/Molineux (late): In addition to an effect associated with chromosome 2B, three QTLs were designated as follows: QPpd.agt-1AL (Xwmc304 - Xgwm497), QPpd.agt-7AS (Xbarc154 - Xbarc108) and QPpd.agt-7BS (Xgwm46 - Xgwm333) {10382}. The QTL in chromosome 1A is possibly orthologous to Ppd-H2 in barley.
68 Response to Salinity
    1 K+/Na+ discrimination
      Variation in K+/Na+ discrimination ratios correlate with salt tolerance, high ratios being indicative of higher tolerance.
    2 Salt tolerance
      QTL: Opata 85/W7984. 77 QTLs effective at different growth stages were mapped to 16 chromosome {10384}.
    3 Sodium exclusion
      QTL for Na+ exclusion and seedling biomass under salt stress were detected in the cross Berkut/Krichauff on chromosome 2A (Nax1 region) and 6A (Xcfd080-Xbarc171-6A) {10917}.
69 Response to Tissue Culture
70 Response to Vernalization
    The requirement for vernalization is particularly important for winter cereals to avoid cold injury of the sensitive floral organs during the winter. In wheat, vernalization requirement is controlled by four major genes designated Vrn-1, Vrn-2, Vrn-3, and Vrn-4. The first three genes were identified using map based cloning aproaches {10014,10299,10421}. The Vrn-1 gene encodes a MADS-box transcription factor closely related to the Arabidopsis AP1/FRUITFULL family, responsible for the transition of the shoot apical meristem from the vegetative to reproductive stage in wheat {10014}.
    Deletions in the promoter (Vrn-A1a, Vrn-A1b) {10198} or the first intron of this gene (Vrn-A1c, Vrn-B1a, Vrn-D1a) {10202} are the most common sources of spring growth habit among landraces and commercial cultivars of polyploid wheat worldwide {10617,10695,10709}.

    The Vrn-2 locus produces two linked and related proteins designated ZCCT1 and ZCCT2, characterized by the presence of a putative zinc finger and a CCT domain {10299}.
    Deletions and mutations involving both the ZCCT1 and ZCCT2 genes are frequent in diploid wheat and are associated with recessive alleles for spring growth habit {10299}.
    Among the cultivated tetraploid and hexaploid wheat species the Vrn-B2 gene is generally functional whereas the Vrn-A2 gene is not {10710}. At least one functional copy of Vrn-2 combined with homozygous recessive alleles at all three Vrn-1 loci is required to confer winter growth habit in hexaploid wheat.

    The Vrn-B3 locus (formerly known as Vrn-5 or Vrn-B4) is homologous to the Arabidopsis FT gene {10421}. This dominant allele, found in the variety Hope, is associated with the insertion of a transposable element in the Vrn-B3 promoter. Natural variation at the Vrn-A3 and Vrn-D3 loci has been also described in hexaploid wheat {10533}. Vrn-3 promotes the transcription of Vrn-1 and accelerates flowering {10421}.

    The Vrn-D4 allele for early flowering was originally identified in the Australian cultivar Gabo {671} and was backcrossed into Triple Dirk to develop the isogenic line TDF {1172}. This locus was mapped on the centromeric region of chromosome 5D between markers Xcfd78 and Xbarc205 {10711}. Natural variation for flowering time at the centrometric region of homoeologous group5 chromosomes has been found, so far, only in the D genome. Incorrect TDF seed stocks generated initial confusion about the existence of Vrn-D4 but molecular markers are now available to separate the incorrect stocks {10711}. Using genetic analyses, Iwaki et al. {10003} found the Vrn-D4 allele for spring growth habit occurred with a higher frequency in India and neighboring regions. The Vrn-D4 locus in TDF includes a duplication of -290kb region from chromosome arm 5AL inserted into the proximal region of chromosome arm 5DS. This translocated segment includes a functional copy of VRN-A1 that carries distinctive mutations in its coding and regulatory regions {11123}. Allelic variations at the Vrn-1 and Vrn-B3 loci in Chinese wheat cultivars are summarized in {10617}.

    Aneuploid and whole chromosome substitution experiments showed that all group 1 chromosomes of wheat carry genes affecting response to vernalization {773}.

    Stem-elongation in winter wheat: In regions where wheat is used as a dual purpose crop for grazing and grain production a relatively long vegetative phase is required to maximize the vegetative tissue and to delay the stem elongation phase. Variation in this attribute occurs among winter wheats such as Jagger (early stem elongation) and 2174 (late elongation).
    In a Jagger/2174 RIL population, QTL for stem elongation included Qste.0cs-5A (associated with the Vrn-A1 locus, Qste.ocs-1BL, Qste.ocs-2D (associated with the Ppd-D1 locus) and Qste.ocs-6A {1010}. In 2007 the respective R2 values were 0.289, 0.155, 0.067 and 0.058. Jagger alleles on chromosome 5A, 1B and 6A promoted stem elongation whereas the allele on chromosome 2D had a delaying effect {10722}.
71 Restorers for Cytoplasmic Male Sterility
    1 Restorers for T. timopheevi cytoplasm
      Minor restorer effects were associated with Xbarc330-5A in R18 and Xgdm130-7D in R9034 {10222}. The relationships of these QTL with previously located restorers in chromosomes 5A {860} and 7D (Rf2) are unknown.
72 Ribosomal RNA

    The 5S-Rrna-1 loci were physically mapped in 1AS, 1BS, and 1DS and the 5S-Rrna-2 loci were physically mapped in 5AS, 5BS and 5DS of Chinese Spring using deletion lines {1043}.
    Table 1 in {276} lists the chromosome or chromosome arm locations of rRNA loci in 12 Triticeae species.
    1 5S rRNA genes
      Within the Triticeae there are basically two sets of 5S rRNA loci. One set, identified by repetitive units 320-468 bp in length, is located on group 1 chromosomes. The other set, identified by repetitive units 469-500 bp in length, is on group 5 chromosomes. Within species the repetitive units at a locus are extremely uniform in size and sequence. They remain stable in foreign genetic backgrounds.
73 Seedling Leaf Chlorosis
74 Segregation Distortion
    See also, Gametocidal Genes.
75 Short Roots
    A 'very short root' phenotype was produced by heterozygous genotypes from selected crosses between Chinese Spring and certain synthetics. The Vsr1 locus was localized to a 3.8 cM interval on chromosome 5DL {11014}.
76 Soft Glumes
77 Sterol Esterification in Kernels - Synthesis of b-Sitosterol Esters
    Two sterol-ester phenotypes, p-L (palmitate + linoleate) and L (linoleate) are inherited as alleles at a single locus.
78 Stem solidness
    Solid stem, caused by increased pith in normally hollow stem regions, is associated with resistance to wheat stem sawfly, Cephus cinctus. Solid stem confers resistance to wheat stem sawfly. See also Reaction to Cephus spp.
79 Temperature-Sensitive Winter Variegation
    This phenotype involves reduced vigour and chlorotic patches on leaves of certain genotypes in Ae. umbellulata cytoplasm when grown at low temperatures {1596}.
80 Tenacious Glumes
    Soft Glume locus is not an orthologue of Tenacious Glumes {10769}.
    A QTL analysis of the relationship of glume tenacity (Gt) with threshability (Ft) and the size of the glume base scar (Gba) after glume detachment in two crosses, viz. the ITMI population and CS*/CS (Ae. tauschii 2D), was undertaken {10497}. In the first cross QFt.orst-2D.1 and QGt.orst-2D.1 were closely associated with Xgwm261-2D, and XFt.orst-2D.2 and XGt.orst-2D were associated with Xgwm455-2D, in the second population only the first pair along with Xba.orst-2D were detected; these appeared to correspond with Tg1 {10497}.
81 Tiller Inhibition
    A QTL of large effect on spike number per plant in a DH population of Fukuho-Komugi/Oligoculm mapping to the Hg - Xpsp2999(Glu3)-1A region {10218} probably corresponds to Tin1.
82 Uniculm Stunt
    Stunting is favoured by a combination of long days and low night temperatures {581}. Caused by duplicate recessive genes, us1 and us2, located in chromosomes 4A and 5B, respectively {200}.
    Genotypes: Normal v: Us1 us2: Alfa {581}; Jaral {581}.
    Normal v: us1 Us2: Mabruk {581}.
    Stunted v: us1 us2: Line 492 {581}.
83 Variegated Red Grain Colour
84 Yield and Yield Components
    1 Grain number per spike
      QTL: Three QTLs for kernel number per spike were assigned to chromosome 3A in RSLs from Cheyene*7/Wichita {0025}.
      2 1000-grain weight
        QTL: Two QTLs for 1,000-kernel weight were assigned to chromosome 3A in RSLs from Cheyenne*7/Wichita 3A {0025}. QTLs for grain size were identified on chromosome arms 1DS, 2DL and 6BL in a RIL population from RS111/CS {0236}. Eight QTLs for 1,000-kernel weight (54% of the variation) were mapped in Forno/ Oberkulmer spelt {0280}.
    5 Grain yield
      Non-glaucous (virescent) lines from a Shamrock/Shango DH population had higher yields than glaucous sibs {10543}; see Glaucousness, subsection Epistatic inhibitors of glaucousness. Grain yield under drought stress
      QTL: Dharwar Dry (drought tolerant)/Sitta: SSR locus Xwmc89-4AL was the marker most closely associated with QTL for grain yield, grain fill rate, spike density, grains/m2, biomass and drought susceptibility index covering a genetic distance of 7.7 cM {10488}.
    9 Spike number per square metre
      QTL: A QTL for spike number per square metre was assigned to chromosome 3A in RSLs from Cheyenne*7/Wichita 3A {0025}.
85 Yellow Berry Tolerance
    QTL : A QTL for yellow berry tolerance, contributed by RS111, was associated with Xgwm190-5D and Xgwm174-5D in a RIL population from RS111/CS {0237}. A tolerance QTL contributed by CS, the susceptible parent, was detected on 6B {0237}.
86 Proteins
    1 Grain protein content
      Thirteen QTLs for grain protein content were identified in a RI population from the cross WL711 (low protein content)/PH132 (high grain content) {10055}. The QTLs that were identified using more than one method or in more than one environment are listed below. Also listed is a QTL that was identified in the mean over the four environments and was therefore deemed important {10055}. QTLs for grain protein content were detected on chromosome arms 6AS (associated AFLP marker, XE38M60200) and 1BL (associated RFLP marker, Xcdo1188-1B) in Courtot/Chinese Spring {0141}.
      Nine QTLs (51% of the variation) were mapped in cross 'Forno'/ 'Oberkulmer' spelt {0280}.
      A QTL for grain and flour protein content, contributed by CS, was associated with XTri-1D/Centromere in a RSL population from the cross Cheyenne (high quality wheat)/CS (low quality wheat) {0251}.
      For QTLs conferring grain protein content detected in the cross Renan/Recital {10071}, only QTLs stable over at least 4 of 6 locations are presented. Renen contributed the four alleles for high grain protein content.
      Ning 7840/Clark: RILs: QTLs from Ning 7840 were detected on chromosomes 3AS (Xwmc749-3AS - Xgwm369-3AS; R2=0.09-0.11) and 4B (Xgwm368-4B - Xwmc617-4B, R2=0.08-0.11) {10702}. Durum: In 3BIL-85 (high protein introgressed from T. dicoccoides/Latino QTL were detected in chromosomes 2AS (associated with Xcfa2164-2A, R2=17%), 6AS (Xp39M37250-6A, R2=17%) and 7BL (Xgwm577-7B, R2=9%) {10338}.
      2 Alcohol dehydrogenase (Aliphatic)
        Three Adh genes were identified in Hordeum vulgare and H. spontaneum {144,490,493,520}. Two of these were tightly linked at the Adh-H1 locus {144}. The third gene was tentatively located in 5H {490}.
        A low-level of aliphatic alcohol dehydrogenase activity is commonly observed on zymograms in the absence of added substrate {513}; this may account for the observation of wheat lactate dehydrogenase that was reported in {1465}.
        The gene series formerly designated Adh-2 and Adh-3 appear under Aromatic Alcohol Dehydrogenase
      7 Esterase
        Genetic control of esterases [carboxylic ester hydrolases (E.C.] was the subject of a comparative study {814}. EST-2, EST-5 and EST-8 are controlled by genes on 3L and where a recombination test was possible between Est-D5 and Est-D8, no segregation was observed. The different gene symbols were retained because of the different tissue specificities and polymerisation profiles of the enzymes. The same arguments surround the EST-1 and EST-6 genes located in the 3S arms {814}.
        The Est-6 gene of rye was mapped {249}. The Est-6 genes of wheat were mapped comparatively in the proximal regions of chromosomes 2S {256}. The Est-2, Est-5 and Est-8 were mapped to the extreme distal regions in the 3L arms {247}.
    1 EST-1
      EST-1 is a dimeric enzyme that electrofocuses around pH4.0 and is expressed in all tissues except endosperm {814}.
    2 EST-2
      EST-2 is a coleoptile-specific monomeric enzyme that electrofocuses at low pI.
    3 EST-3
      EST-3 is a monomeric enzyme that is expressed in young seedlings (this enzyme was not observed in {814}.
    4 EST-4
      EST-4 is a monomeric, leaf-specific enzyme that electrofocuses around pH 4.5.
    5 EST-5
      EST-5 consists of 20 or more monomeric, grain-specific isozymes that electrofocus between pH 5.6 and 7.0.
    6 EST-6
      EST-6 is a dimeric enzyme that electrofocuses around pH 7.6 and is specific to endosperm. A group of leaf esterase isozymes controlled by the long arms of the homoeologous group 3 chromosomes were reported {919}. The relationship of these esterases to EST-2 and to the leaf esterase designed EST-6 reported in {629} has not been determined.
    7 EST-7
      EST-7 is a monomeric enzyme that electrofocuses in the same region as EST-6 but is specific to green tissues.
    8 EST-8
      EST-8 consists of about 10 isozymes that electrofocus between pH 4.5 and 6.5 and are expressed only in vegetative tissues. EST-8 is likely to be the enzyme previously described in {919} and {629}.
    9 EST-9
      EST-9 is a monomeric enzyme that electrofocuses around pH 5.0 and is expressed only in embryos.
      10 Hexokinase
        Allelic variation was observed in three of 55 hexaploid accessions {006}.
      11 Lipoxygenase
        The wheat Lpx-1 gene in wheat corresponds to barley LoxA (GenBank L35931). The Lpx-B1 locus is duplicated, with the Lpx-B1.1 and Lpx-B1.2 loci corresponding to GenBank sequences DQ474240 and DQ474241, respectively. The Lpx-B1b allele corresponds to a deletion associated with a 4.5-fold reduction in lipoxygenase activity. The Lpx-2 gene in wheat corresponds to the barley LoxC gene (GenBank L37358) whereas the Lpx-3 gene in wheat corresponds to the barley LoxB gene (GenBank L37359).
      13 Peroxidase
        Peroxidase (EC1.11.1.7) isozymes have high tissue specificity. Staining and electrophoretic systems are reviewed in {118}. PER-1, -2, -3, -4 and -5 are all reported in {816}.
    1 PER-1
      PER-1 is expressed in leaf {012} and coleoptile {816} tissues.
    2 PER-2
      PER-2 is expressed in young leaf {118}, coleoptile and root {816} tissues.
    1 Ndh-1
      Based on the correspondence of the electrophoretic patterns, isoelectric points (pIs) and chromosomal location, it was proposed that the Ndh1 (NADH dehydrogenase) and Dia3 (diaphorase) represent the same locus {0356}.
    2 Ndh-2
      Based on the correspondence of the electrophoretic patterns, isoelectric points (pIs) and chromosomal location, it was proposed that the Ndh-2 (NADH dehydrogenase) and Dia2 (diaphorase) represent the same locus {0356}.
    3 Ndh-3
      Based on the correspondence of the electrophoretic patterns, isoelectric points (pIs) and chromosomal location, it was proposed that Ndh-3 (NADH dehydrogenase), Dia1 (diaphorase) and Mnr1 (menadione reductase) represent the same locus {0356}.
      31 Benzoxinones
        The putative role of benzoxinones sets Bx-1 to Bx-5 is to catalyze the pathway Indole-3-glycerol phosphate to DIBOA. Primers designated from maize sequences were used to generate RT-PCR products utilised to screen a cDNA library from CS seedlings. Full-length cDNAs were heterologously expressed in yeast and the Bx gene products had enzymatic action. The Bx genes located by Southern analysis of CS deletion stocks occurred as clustered groups in homoeologous groups 4 (Bx-1, Bx-2) and 5 (Bx-3.1, .2, Bx-4, Bx-5) {10103}.
      33 Phytoene synthase
        Phytoene synthase, which condenses two molecules of geranyl geranyl diphosphate to produce phytoene, is the first of the specific enzyme necessary for carotene biosysthesis in plants.
    1 Phytoene synthase 1 (EC
      Homology with the same gene in rice (Psy1) {10230}.
      Phytoene synthase is involved in the carotenoid biosynthetic pathway and influences yellow pigment content in grain (See Flour colour and Grain quality parameters: Flour, semolina and pasta colour). The gene Psy-A1 was cloned and a functional marker developed from the sequence distinguishing Chinese common wheats with high and low pigment contents {10501}. Most hexaploid wheat cultivars have a 676-bp insertion in intron four that is absent in Australian cultivars Dundee, Raven, and Aroona with high yellow pigment. The Psy-B1b allele from tetraploid wheat Kofa is the result of a B-A intergenomic conversion event that probably occurred in Cappelli ph1c mutant 1 {10530}. An EMS mutation in the Psy-E1 gene is associated with whiter endosperm in lines carrying the Th. elongatum 7EL translocation.
      34 Polyphenol oxidase
        High PPO activity in kernels and flour leads to a time-dependent discolouration of end products such as noodles, pasta and breads.
        Primers different from those in {10386} were developed in {10504}, but their ability to distinguish phenotypic groupings (alleles) were similar. A null allele of Ppo-D1 was identified for this locus using primer pair WP3-2 {10504}.
      35 Protein disulfide isomerase (EC
        The genes for PDI and their promoters were sequenced in {10423}. A related sequence on 1BS was shown to be a partial, non-expressed copy in {10424}, but not detected in {10409}. PCR-RFLP markers for [TaPDI-4A] and [TaPDI-4B] were designated [Xvut(PDI)-4A] and [Xvut(PDI)-4B] in {10409}. These were also closely associated with Germin (oxalate oxidase {10441}) genes {10409}.
      39 Zeta-carotene desaturase
        Cv. Zhongyou 9507 has lower yellow flour pigment content, preferred for Chinese steamed bread and dry Chinese noodles. A QTL in the Zds-D1a region explained 18.4 % of the variation in yellow pigment content in Zhongyou 9507/CA 9632 {10906}.
      1 Glutenins
        These are heterogeneous mixtures of proteins comprising subunits linked by disulfide bonds. 'A' are high-molecular-weight (HMW) and 'B', 'C' and 'D' are low-molecular-weight (LMW) subunits.

        Using proteomic analysis of 2D gels of seed storage proteins in 39 ditelocentric lines of cv. CS, 105 protein spots were resolved {03129}. Locations of structural genes controlling 26 spots were identified in 10 chromosomal arms (4 on 1BL, 5 on 1BS, 4 on 1DL, 4 on 1DS, 2 on 6AS, 3 on 6BS, 1 on 6DL, 1 on 6DS, 1 on 3BS and 1 on 3BL). Multiple regulators of the same protein located on various chromosome arms were observed. Two novel subunits, named 1Bz and 1Dz, were found to have very similar structures to HMW glutenin subunit 12 (encoded by Glu-D1-2a - see the relevant list below) and were located to chromosome arms 1BL and 1DL, respectively. PCR amplification of genomic DNA was used to isolate three LMW glutenin genes in cultivar Chinese Spring, named LMWG-MB1, LMWG-MB2 and LMWG-MB3 {01101}. The deduced amino-acid sequences showed a high similarity between these ORFs and with those of other LMW glutenin genes. The authors state that the study provided direct evidence that insertions and/or deletions provide a mechanistic explanation for the allelic variation, and hence the resultant evolution, of prolamin genes, and comment on relationships with gamma-secalins and beta-hordein families. Single-base substitutions at identical sites generate premature stop codons in both LMWG-MB2 and LMWG-MB3, indicating that these clones are pseudogenes.
    1 Glu-1
      The Glu-1 loci, all of which are compound, encode HMW glutenin subunits.
      Each Glu-1 locus in hexaploid wheat contains two genes, the products of which were described as 'x-type' and 'y-type' based on differences in molecular weight and isoelectric point {1118}.
      Other evidence has shown that these gene products differ in electrophoretic fingerprint pattern {1124} and cysteine content {1028}, and the genes themselves differ in nucleotide sequence {1470,1433,373}.
      Although early evidence suggested up to 6 genes in total at each locus {1471,373],it appears likely that only a single copy of each gene is present at the 1AL,1BL,and 1DL loci {495}.
      No 'y-type' protein from the Glu-A1 locus has been demonstrated in hexaploid wheat {1118}, although they are found in diploid wheats {1535,798}, and sequencing experiments have shown the presence of two stop codons in the transcribed portion of the gene {10088}. Definitive evidence that subunit 21* {602}, which has a mobility close to that of subunit 21, is a 'x-type' protein rather than a 'y-type' protein has not been obtained. The gene coding for 'x-type' proteins within Glu-A1 is also often silent {1118,420}.
      The symbols for the genes within the Glu-1 loci coding for 'x-type' and 'y-type' proteins will be Glu-1-1 and Glu-1-2, respectively, rather than Glu-1x and Glu-1y {1470}. The genes are closely linked but recombination has been observed between Glu-B1-1 and Glu-B1-2 with a frequency of 3 in 3,450 {1117}. The gene order, relative to the centromere, has not been ascertained.
      The subunit nomenclature used is that devised in {1116}; however, an alternative system based upon molecular weight was proposed in {1068}. A system of naming the Glu-A1-1, Glu-A1-2, Glu-B1-1 and Glu-B1-2 alleles in T. turgidum var. dicoccoides is given in {796}.
      In {00116}, a comparison between spelt wheats (T. spelta) and bread wheat was carried out for the glutenins using a nomenclature system described in {00117}.
      The Glu-1 loci may be recognised by the DNA probe pTag1290 {1471} and probe pwhe1(Dy10) {030}. Individual Glu-1-1 loci on 1A, 1B and 1D and the Glu-1-2 loci may be recognised by specific primers {263}.
      In {00105}, the evolution of the high molecular weight glutenin loci of the A, B, D and G genomes of wheat was explored; 30 partial allele sequences were compared, designated by Greek letters (alpha, beta, gamma, etc.) (5 of which were cited as Schlumbaum, pers. comm.; the remaining 25 were deposited as GenBank, accession nos. X98583-X98592, X98711-X98715 and Y12401-Y12410). These partial alleles derive from all six Glu-1-1 and Glu-1-2 loci in current-day samples taken from seven species of wheat, as well as from DNA extracted from charred grain of two samples from archaeological excavations, dated 3000 and 5000 years old, respectively.
      Following the first listing which considers the Glu-1 set for hexaploid wheat as a single locus, there is a provisional listing based on x- and y- type glutenins. These are not referenced.
      The importance of the HMW glutenin subunits for bread-making quality was first noted from observations in wheat cultivars of related pedigree on the effects of the presence of subunit 1 encoded by Glu-A1a {0197}, effects that have repeatedly been confirmed since (for example {0198,0199,01100}).

      A nomenclature system for prolamin banding patterns of triticale was proposed in {03139}. Extensive allelic variation in triticale at Glu-A1, Glu-B1, Glu-R1 and Gli-R2 loci was reported in {03121}.
    3 Glu-3

      The Glu-3 loci are defined as the cluster of LMW glutenin genes previously considered a component of the compound Gli-1 loci.
      More than 30 LMW glutenin complete genes, partial genes or pseudogenes have been sequenced from Triticum species (reviewed in {0245}).
      In T. aestivum, only Glu-B3 was shown to recombine with the gliadin genes (1.7 +/- 0.8) {1355,1358}. However, in T. durum, recombination was observed for both Glu-A3 and Glu-B3 with their respective Gli-1 loci: the map distance between Glu-A3 and Gli-A1 has been estimated as 1.3 +/- 0.4 cM {1242}, and that between Glu-B3 and Gli-B1 as 2.0 +/- 0.8 in {1144} and as 2.0 +/- 0.4 in {1242}. It appears that Glu-B3 is proximal to Gli-B1, and there is some evidence that Glu-A3 is proximal to Gli-A1 {1242}.
      Whereas hitherto it was widely thought that all LMW glutenin subunits were encoded by genes located on the chromosomes of homoeologous group 1, it has been demonstrated that, although the majority of the subunits are indeed controlled by genes on this group, some of the C subunits must be controlled by loci elsewhere in the genome {482}.
      A novel type of polymeric protein (Mr approx. 71,000) was reported in the Australian advanced breeding line DD118 {03125}. It participates in the polymeric structure of glutenin (possibly as a chain terminator), and has an Mr of approximately 71,000, could be considered as a D-subunit of LMW glutenin. However, N-terminal sequencing suggests it to be a Gli-B1 type omega-gliadin that has acquired a cysteine residue through mutation.

      In an electrophoretic survey of 51 primary tritordeums {03113}, 20 distinct whole banding patterns (a-t), each consisting of between one and three bands, were observed for D-zone prolamins exhibiting glutenin-like solubility characteristics.

      In 85 Japanese common wheat cultivars and 61 elite F6 breeding lines, 3 alleles were observed at each of Glu-A3 and Glu-B3, and 2 alleles at Glu-D3 were named according to their parental origins in three doubled haploid mapping populations {03135}.

      C-type LMW glutenin subunits in CS were assigned to chromosome groups 1 and 6, and shown to have sequences very similar to those of alpha- and gamma-gliadins {03134}. The authors suggest that they may be encoded by novel genes at loci tightly linked or present within the Gli-1 and Gli-2 loci, unlike other LMW glutenin subunits encoded by the Glu-3 loci.

      The HMW and LMW glutenin subunits carried by chromosome 1Am of T. monococcum accession G1777 were characterised electrophoretically and evaluated for quality characteristics using recombinant chromosome substitution lines with chromosome 1A of CS {03142}. The HMW subunits from G1777 are promising for bread-making quality, whereas its LMW subunits are promising for biscuit-making quality.

      The bread wheat cv. Salmone was shown to carry two DNA fragments designated as SF720 and SF750 located on the chromosome 1B satellite and associated with the presence of two LMW glutenin subunits {03143}. However, the authors suggest that they occur at a locus other than Glu-B3 due to their relatively high frequency of recombination with Gli-B3.

      A naming system in which Roman numerals are assigned to whole banding patterns for the LMW glutenin subunit is given in {03131} as an alternative to the LMW-1/-2 system described in {03136}. A further system naming whole banding patterns from LMW-1 to LMW-23 in emmer wheat is described in {03137}. In {00111}, in a study of common and durum wheats from Portugal, the authors used the nomenclature system described in {00113} for the LMW subunits in common wheat, and that described in {00114} for the LMW subunits in durum wheat. The latter system was updated according to {02110}, but has been changed herein to new alleles with the earlier durum designation {00114} given as synonyms.In {03116}, it was suggested that Glu-B3d (common wheat standard genetic stock) is equivalent to Glu-B3r (durum wheat standard genetic stock), and that (referring to article {03127}) LMWsubunits observed in some Portugese triticales could be of the durum type.
      A novel storage protein gene with chimeric structure was isolated from the old Hungarian cultivar Bankuti 1201, containing gamma-gliadin sequences in the 5' region, LMW-glutenin sequences in the 3' region and a frameshift mutation leading to a completely new polypeptide in the C-terminal region. A further seven recombinant prolamin genes were subsequently isolated. The eight genes, designated Ch1 to Ch8, seem to derive from four gamma-gliadin and three LMW-glutenin sequences and are probably the result of crossing over between the loci Gli-1 and Glu-3. However, the precise recombinational mechanism that gave rise to them has yet to be elucidated {10307}. Characterization of near isogenic lines for the different Glu3 alleles provides a useful quantification of their contribution to bread making quality {11129}.
    4 Glu-4
      The following loci, Glu-D4 and Glu-D5, encoding low molecular weight subunits of glutenin (30-32 kDa) were described in {02111}; the proteins encoded by them were first observed earlier {02114,02115}, and the former was later tentatively assigned the symbol Glu-4 {02116}, before its chromosomal location was established and the locus definitively named as Glu-D4 in {02111}. While this locus is located on chromosome 1D (in accordance with the position on the group 1 chromosomes of the remaining glutenin encoding loci found to date), the locus Glu-D5 is located on chromosome 7D. In SDS-PAGE, the proteins from both loci are detected only in the presence of 4-vinylpyridine added to the sample extract. Their amino acid compositions do not match those of the major prolamin groups; nonetheless, they classify as glutenins based upon solubility, immunological behaviour and N-terminal amino acid sequence (the latter suggesting an evolutionary link with the major (B and C) low molecular weight glutenin subunits).
    5 Glu-5
      A collection of 173 Ae. tauschii accessions were analysed for low molecular weight glutenin subunits by SDS-PAGE {02112}. Thirty three different patterns for B-subunits and 43 for C-subunits were identified, some of which were of identical electrophoretic mobility to those observed in common wheat. Also observed were subunits with the same mobilities as the D-subunits and as the subunits encoded by the Glu-D4 and Glu-D5 loci. This variation represents a source of novel germplasm of potential value for breeding programmes aimed at improving the D-genome of common wheat in the context of bread-making quality.
      2 Gliadins
        These are heterogeneous mixtures of alcohol-soluble polypeptides without quaternary structure. The Gli-1 loci are compound and are now considered to comprise the omega-gliadin and gamma gliadin {982,1415} multigene families {494}, which in some circumstances may be divided into Gli-1-1 and Gli-1-2, respectively. The LMW glutenin multigene families, which are closely linked to the Gli-1 loci {588}, are listed separately as the Glu-3 set {1358}; information on map distance and gene order in relation to Glu-3 and the centromere is given in the preamble for the Glu-3 loci. There is evidence that a few of the omega-gliadin genes are separated from the main omega-gliadin gene cluster {993}.
        Variation at the Gli-1 loci was described earlier {634,996,1126} and applied in mapping experiments {1243,1125,196,422,1120}. A rational system of naming the alleles was produced by Dr. E.V. Metakovsky{988}. This nomenclature is reproduced below. A considerable number of alleles were added to the original list given in {988}, and referenced here accordingly. A few alleles have been deleted, because, following much detailed comparison, there is now doubt that they can be reliably distinguished from existing alleles {9981}. The allelic letter in these cases has not been reused. To facilitate practical use of the list, the aim was to give at least three standard cultivars from a range of countries for each allele {9981}. This was achieved for the vast majority of entries and is a change from the original list compiled from {988}, where up to two standards were given. While the three or more standards described almost always include the original standards, some have been replaced for various reasons, such as international awareness of the cultivar, availability of seed, or the ease with which an allele can be identified in a particular genetic background {9981}. In the original list, where two cultivars were given as prototypes for an allele, the first named was from the USSR and the second from elsewhere; this is no longer the case, although care was taken to include a Russian cultivar where possible, to maintain a wide base of germplasm in which the alleles are available, as well as to acknowledge the research groups in the country where much of the pioneering work was carried out.
        For discussion of null alleles at the Gli-1 and Gli-2 loci, see {9984}.
        Recombination was observed within the gliadin multigene family at XGli-A1 {277}. These closely linked genes may correspond to Gli-A1 and Gli-A5, but they were temporarily designated XGli-A1.1 and XGli-A1.2 until orthology with Gli-A1 and/or Gli-A5 is established.

        Note: The catalogue entries reproduced here only refer to alleles in T. aestivum; there is, however, enormous variation in the gliadins in the close relatives of wheat; see, for example, {989} for studies in T. monococcum (more than 80 gliadin electrophoretic patterns observed in 109 accessions), {990} for studies in T. boeoticum (more than 50 electrophoretic patterns in 60 accessions), and {1076} studies in T. durum (19 electrophoretic patterns, referring only to variation in the omego-gliadins, in 243 accessions).
        In {00110}, variants for omega-gliadins were reported from study of twenty-four accessions of einkorn wheat (T. monococcum ssp. monococcum). In {00111}, in a study of common wheat and durum from Portugal, the authors used the nomenclature system described in {00112} for the omega-gliadins. In {00116}, a comparison between spelt and common wheat was carried out for the gliadins using a nomenclature system described in {00118}.
        The Gli-1loci may be recognised by probes pcP387 {372} and pTag1436 {065}, and by specific microsatellites primers {252}. Furthermore, it was shown that probe pTag1436 differentiates gliadin alleles rather well; using this probe, families of gliadin alleles and some of their relationships were described {9988}.
        Twenty eight gamma-gliadin gene sequences from GenBank were grouped into nine subgroups in {10063}. Primers were developed against some of the subgroups and the chromosomal locations of the gamma-gliadin genes were determined {10063}.
        Based upon morphological observation and RFLP analysis, it was proposed that the cultivar 'Chinese Spring' is a strain of the landrace 'Chengdu-guangtou' from the Chengdu Plain, Sichuan Province; this proposal is supported by the observation that CS and the landrace share the same alleles at all nine Gli-1, Gli-2 and Glu-1 loci {see 01102}.
        PCR primers GAG5 and GAG6 were applied to 35 cultivars of closely related spelt and hexaploid wheat, and to eight cultivars of durum, to yield products originating from two gamma-gliadin genes mapped to chromosomes 1B (termed GAG56B) and 1D (termed GAG56D) {01103}. Two alleles for GAG56D (differing in a 9 bp deletion/duplication and single nucleotide polymorphism) were found, one a new allele and the other previously published {01104}. Meanwhile two alleles found for GAG56B among the durum wheats correlated with the presence of gluten quality markers, gamma-gliadins 42 or 45.

        1B and 1D sulphur-poor omega-gliadins in cultivar Butte 86 were characterised by RP-HPLC, SDS-PAGE, two-dimensional PAGE, amino acid composition determination and sequencing, matrix assisted laser desorption ionisation-time of flight mass spectrometry and circular dichroism spectroscopy to reveal the detailed nature of the peptides belonging to the two groups, and showing that the complexity of mixtures of the peptides of the 1B group was greater than that of the 1D group {01105}. Although circular dichroism spectra were similar for the two groups of peptides, and suggested a mainly flexible random structure, there was evidence for a significant amount of left-handed polyproline II helical conformation in the case of the 1D components. The authors placed some of the results in the context of the possible ancestor of the B-genome and relationships with the barley C-hordeins and rye omega-secalins.
        Eleven new gliadin alleles were found in a collection of 52 Spanish landraces of common wheat {03141}.
        A new family of low-molecular-weight gliadin genes located on groups 4 and 7 were reported in {10117}. They appear to influence rheological properties and seem to be closely related to the 17kDa epsilon hordein, important in beer foam stability.
        A novel storage protein gene with chimeric structure was isolated from the old Hungarian cultivar Bankuti 1201, containing gamma-gliadin sequences in the 5' region, LMW-glutenin sequences in the 3' region and a frameshift mutation leading to a completely new polypeptide in the C-terminal region. A further seven recombinant prolamin genes were subsequently isolated. The eight genes, designated Ch1 to Ch8, seem to derive from four gamma-gliadin and three LMW-glutenin sequences and are probably the result of crossing over between the loci Gli-1 and Glu-3. However, the precise recombinational mechanism that gave rise to them has yet to be elucidated {10307}.
        Transcriptome analysis showed the presence of proteins called avenin-like a and b. The former contained a duplicated sequence of about 120 residues and corresponded to the LMW-gliadins. The latter was not previously characterized, but may form part of the glutenin fraction and hence influence quality. These avenin-like proteins showed higher expression levels in three Aegilops species (Ae. caudata, Ae. cylindrica and Ae. tauschii) than in common wheat {10321}. Four new classes of low molecular weight proteins related to gliadins, though not sufficiently similar to be classified as such, were reported in {02113}. One of the classes has no close association to previously described wheat endosperm proteins.
    1 Gli-1
      In barley, the B and C hordeins are controlled by the Hor2 and Hor1 loci, respectively, which are linked {1341} on chromosome 1HS {1063,1153}. The map distances and homology of the proteins indicate that Hor1, the locus closest to the centromere, is equivalent to the omega-gliadins (Gli-1-1) in Gli-1 {1338}.
      Three alleles at each of the Gli-1-1 (omega gliadin) loci were noted {1358}. The complexity of the Gli-1 compound loci is further emphasized by a report of individual genes being separable by recombination, where G1d-1A (a block of gamma and omega genes) is separable by 0.3% from Gld4-1A (omega gliadins) which is in turn, separable by 1.5% from Gld3-1A (omega gliadins) {1103}.
      Elsewhere, variation was described {634,996,1126} and applied in mapping experiments {107,196,422,1120,1125,1243}. Sixteen combinations of Gli-B1 and 4 combinations of Gli-D1 subunits are listed in {420}. Multiple alleles described in {996}, number 15 at Gli-A1, 18 at Gli-B1, and 8 at Gli-D1.
      The Gli-1 alleles present in 57 Yugoslav wheat varieties were reported in {994}.
    2 Gli-2
      Prior to the publication of {988}, allelic variation was demonstrated at all of the wheat Gli-2 loci, including 13 alleles at Gli-A2, 11 at Gli-B2, and 10 at Gli-D2, in a study of 39 cultivars {996}.
      The Gli-2 alleles present in 57 Yugoslav wheat varieties were determined {994}.
    3 Gli-3
      A Gli-3 set of loci coding for omega-type gliadins are located 22 to 31 cM proximal to Gli-1 on the short arms of group 1 chromosomes {422,1403,589}.
    4 Gli-4
      It is not clear how Gli-Sl4 and Gli-Sl5 relate to the Gli-4 and Gli-5 sets described below.
      A locus designated Gli-A4 controlling omega-gliadins in cv. Perzivan biotype 2 was mapped at 10 cM proximal to Gli-A1 on the short arm of chromosome 1A {1205}.
      However, Metakovsky et al. {9983} have since shown that this locus and Gli-A3 are, in fact, the same locus. Furthermore, Dubcovsky et al. {277} did not find evidence for the simultaneous presence of both Gli-A3 and Gli-A4 in five 1A or 1Am mapping populations and concluded that Gli-A4 should be considered to be Gli-A3 until conclusive evidence for the former is obtained. For these reasons, the locus Gli-A4 is deleted from the catalogue.
    5 Gli-5
      A locus designated Gli-5 controlling omega-gliadins was mapped to the short arms of chromosomes 1A and 1B, distal to Gli-1 {1147}. The map distance between Gli-B5 and Gli-B1 was estimated as 1.4 cM (recombination value of 1.4 +/- 0.4%), although there was significant variation in recombination values over crosses, ranging from 0 % to 5.9 % over the six crosses analysed. This variation was attributed to genotypic influence on the frequency of recombination.
    1 Triticin proteins
      The triticin proteins {1360} or [Triplet proteins {1357}] are storage globulins with homology to pea legumins and related proteins in oats, rice and several dicotyledonous species {1360}. Triticin gene segments including its hypervariable region were PCR-amplified, with preferential amplification of Tri-D1 for the only pair of primers giving consistent results {10322}.
      2 Inhibitors (dimeric) of heterologous alpha-amylases
        Chromosome 3BS has duplicated loci controlling two dimeric inhibitors of exogenous a-amylases, one known as 0.53 or Inh I {1260}, and the other as WDA I-3 {1260}. Chromosome 3DS has a homoeologous locus controlling a dimeric inhibitor of exogenous a-amylases, known as 0.19 or Inh III {1260,0124}, that is closely related to 0.53/Inh I. Intervarietal polymorphism for the WDA-3 protein was identified by isoelectric focussing of water-soluble endosperm proteins {0124}. This was interchromosomely mapped on 3BS using both a DH population of Cranbrook/Halberd, and a set of RILs of Opata 85/W-7984 (ITMI population) {0125}.
        Three genome allele specific primer sets were designed for the 3BS and 3DS alpha-amylase inhibitors in cv. Chinese Spring, based upon SNPs. Their validity was confirmed in 15 accessions of Triticum urartu, Triticum monococcum, Aegilops tauschii and Triticum dicoccoides. The results offered support that the 24kDa dimeric alpha-amylase inhibitors in cultivated wheat are encoded by a multigene family {10323}, previously proposed in {10324}, as the result of phylogenetic analysis of sequences characterized by cSNPs.
      3 Subtilisin inhibition
        Considerable genetic variation for Si-2 was noted in {701}. A chromosome location for Si-H2 on 1HL was inferred in {528} but questioned in {701}.
        Three subunits of the wheat tetrameric inhibitor of insect a-amylase, CM1, CM3 and CM16, with homology to the dimeric and monomeric a-amylase inhibitors and the trypsin inhibitors, were located by Southern analysis of cDNAs pCT1, pCT2, and pCT3 to 4A, 4B, 4D; 7A, 7B, 7D; and 4A, 4B, 4D, respectively {427}.
        Genes encoding proteins which inhibit the action of mammalian and insect, but not cereal, a-amylases, were located in chromosomes 3BS, 3DS and 6DS of Chinese Spring {1260}. Also, genes encoding inhibitors of insect a-amylases were located in H. chilense chromosomes 4Hch and 7Hch {1262}.
      2 Histone H1 Proteins
        The relationship of this gene series with a Hst-A1, Hst-B1, Hst-D1 series in group 5 chromosomes {0216} based on DNA hybridization studies was not established.
      6 Puroindolines and grain softness protein
        Puroindolines a and b are the major components of friabilin, a protein complex that is associated with grain texture (see 'Grain Hardness'). The name 'puroindoline' and the complete amino acid sequence of puroindoline a were given in {0382} from cv Camp Remy. Hard grain texture in hexaploid wheat results from unique changes in the puroindoline amino acid sequence or, currently, four null forms {0295} of the completely linked genes (max. map distance 4.3 cM) {452}. Tetraploid (AABB, AAGG) wheats lack puroindolines and are consequently very hard {03103}. A searchable database of wheat varieties and their puroindoline genotype is available at http://www.wsu.edu/~wwql/php/puroindoline.php. Grain softness protein-1 is a closely related gene which is closely located to the puroindoline genes {03111,1185}. 'GenBank' and 'dbEST' refer to sequence databases available at NCBI (also available throught EMBL and DDB).
        Recent reviews {10522,10523} provide comprehensive descriptions of the molecular genetics and regulation of puroindolines. Morris and Bhave {10524} reconciled the D-genome puroindoline alleles with DNA sequence data. Bonafede et al. {10525,10526} developed a CS line (PI 651012) carrying a 5AmS.5AS translocation from T. monococcum; the translocated chromatin carries A-genome Pina, Pinb and Gsp-1 alleles that confer softer kernel texture. Pinb-D1b, Pinb-D1c, Pinb-D1d, Pinb-D1e, Pinb-D1f, or Pinb-D1g are present in hard hexaploid wheats not carrying the Pina-D1b (null) mutation {452,1035,0082,0204}.

        Wheats with Pinb-D1b were slightly softer and a little superior to those with
        Pina-D1b in milling and bread-making characteristics although there was considerable overlap {0206}.
        Transgenic rice with the Pina-D1a and Pinb-D1a alleles possessed softer grain {0207}.

        Genotypes for a selection of North American wheats are given in {0204}.
        In T. monococcum the gene order was reported to be : tel - Gsp-1 - Pina - Pinb {0083,10122} whereas in Ae. squarrosa it was : tel - Gsp-1 - Pinb- Pina {10037}.
        The soft kernel trait was transferred to durum {10899}.
        Ikeda et al. {10305} reported a double-null with apparently no Pina-D1 or Pinb-D1 genes present in v: Bindokku, Cheyenne 'A', Chosen 68, Saiiku 18, Saiiku 44, and tentatively assigned it Pina-D1b/Pinb-D1h(t). How this deletion compares with the double null mutation reported by Tranquili et al. {10077} which was assigned Pina-D1k/Pinb-D1q is unknown.
        Lines possessing the alien-derived genes Lr57 and Yr40 lack puroindoline genes and therefore should be hard phenotypes {10770}.
      9 Serine protease inhibitors
        Serine proteinase inhibitors or serpins are salt soluble proteins (-43 kDa) representing about 4% of the total protein in wheat and barley endosperms. They may have a role in plant defense.
      10 Starch granule proteins
        The proteins, designated SGP-1, are starch synthases, encoded by SsII-A1, SsII-B1 and SsII-D1 {0042}. A triple null stock (SGP-1 null wheat) is reported in {0137}. Deletion mapping indicated that the gene order on the 7S arms is; centromere - Sgp-1 - Sgp-3 - Wx{1615}.
      13 Waxy proteins
        Waxy protein (granule-bound starch synthase = ADP glucose starch glycosyl transferase, EC 2.4 1.21 = GBSSI) is tightly bound within endosperm starch granules and is involved in the synthesis of amylose {1616}. Waxy variants, characterised by starch granules containing increased amylopectin and reduced amylose, are preferred for Japaness white salted or "udon" noodles {1650}. Similar waxy phenotypes are controlled by orthologous genes in barley, maize and rice but are not known to occur in rye {725}. All combinations of the null alleles were produced in Chinese Spring {0018}. Partial genomic clones of various diploid, tetraploid, and hexaploid wheats were sequenced {0278,0279}.
        A multiplex PCR assay for identifying waxy genotypes is described in {10032}. Lists of cultivars, lines and landraces of tetraploid and hexaploid wheats with different, mostly null, alleles at the Wx loci are given in {9910,9911,9912,1053,1054,9913,9915,9916,1650,9917}.
        The complete genomic sequence for Wx-D1a from CS {0073} and the cDNA sequence for the Wx-D1b allele from Bai Huo {0075} were determined.
        Isolation of a wheat cDNA encoding Wx-A1 and Wx-D1 was reported in {0123} and {0167}, respectively. Isolation of genomic sequences for the genes encoding granule-bound starch synthase (GBSSI or Wx) in T. monococcum, Ae. speltoides and Ae. tauschii was reported in {0168}. Cloning of a second set of GBSSI or waxy genes, GBSSII, which were shown to be located on chromosomes 2AL, 2B and 2D, was reported in {0167}.
        Various hard and soft wheats with the alleles Wx-A1b, Wx-B1b and Wx-D1b are listed in {0304}. Fifteen percent of Chinese wheats possessed Wx-B1 null alleles {10357}.
87 Abiotic Stress Responses:Dehydrin-response Element Binding Factors
    DREB proteins are a large family of transcription factors induced by abiotic stresses. Using genome-specific primers as probes for an orthologous Dreb1 gene series was placed on chromosomes 3A, 3B and 3D {10729}. SNPs in Dreb-B1 permitted mapping in chromosome 3BL in the ITMI (Opata 85 / W7984) mapping population.
88 Reaction to Barley Yellow Dwarf Virus
    Disease: Cereal yellow dwarf
89 Reaction to Bipolaris sorokiniana
    Diseases: Spot blotch and common root rot.
    Spot blotch
    Yangmai 6 (R)/Sonalika (S): RIL population: AUDPC was controlled by four QTLs derived from Yangmai 6, viz. QSb.bhu-2AL (Xbarc353-2A - Xgwm445-2A, R2=0.148), QSb.bhu-2BS (Xgwm148-3B - Xgwm375-2B, R2=0.205), QSb.bhu-5BL (Xgwm67-5BL - Xgwm371-5BL, R2=0.386) and QSb.bhu-6DL (Xbarc173-6D - Xgwm732-6DL, R2=0.225) {10719}.
90 Reaction to Blumeria graminis DC.
    Disease: Powdery Mildew.
    Resistance genes and their molecular associations are reviewed in {10141}.
    1 Designated genes for resistance
      Note: Chancellor, used as a susceptible genetic background, for some near-isogenic lines probably carries Pm10 and Pm15 {1479}.
      33 NILs, including 22 resistance genes and 3 genetic backgrounds are listed in {10389}. A further gene derived from T. monococcum PI 427772 was identified in BCBGT96A = PI 599036 = Saluda*3/PI 427772 {10479}.
      A single resistance gene was identified on chromosome 7AL in hexaploid germplasm NC96BGT4 (a T. monococcum derivative). This gene was proximal to Pm1 and considered to be different from Pm37, although possibly allelic {10274}. Genotype lists:Chinese wheats{1608,572}; Finnish wheats {0028}; French wheats{1629}; Hungarian wheats {02104}; Western Siberian wheats{1101}
      Complex genotypes:
      Drabent {heterogeneous} Pm2 Pm4bPm9/Pm1 Pm2 Pm4b Pm9 {1287};
      Nemares Pm1 Pm2Pm4b Pm6 Pm9 {1287};
      Planet, Sappo & Walter Pm1 Pm2 Pm4b Pm9 {096,097,540,1287,1428}

      Scandinavian wheats {10681}.
    2 Suppressors of Pm
      Some wheats which, on the basis of cytological and rust tests carry 1RS from Petkus rye, do not express resistance due to presence of a suppressor {385}. Zeller & Hsam {1625} located a suppressor of Pm8 and Pm17 in chromosome 7D of Caribo. Mildew resistance was suppressed in Florida, Heinrich, Ikarus, Olymp and Sabina, which are derivatives of Caribo with 1BL.1RS. According to Ren et al. {1209}, SuPm8 does not suppress Pm17. Hanusova et al. {492} listed 16 wheats that carry a suppressor of Pm8; 111 wheats did not carry the suppressor. In contrast, a high frequency of suppression occurred in CIMMYT wheats {108,1208}. Further genotypes are identified in {491,11025}. Although Line 81-7241 carries Pm8 as well as Pm23, evidence was presented to indicate that Pm8 was suppressed in Line 81-7241{1618} and , by inference, indicated that Chinese Spring possessed SuPm8. Pm8 was suppressed when locus Pm3 is transcribed (including Chinese Spring and Thatcher which have no currently detectable Pm3 resistance alleles) {10828}.
    4 QTLs for resistance to Blumeria graminis
      QTL: Several QTLs were detected in two RE714/Hardi populations when tested at two growth stages and with different cultures over three years. The most persistent and effective QTL was located in the vicinity of Xgwm174-5D {0272}. Three QTLs, QPm.vt-1B, QPm.vt-2A and QPm.vt-2B, with additive gene action, accounted for 50% of the variation in a population developed from Becker/Massey{0284}. These QTLs were confirmed by the addition of extra markers to the Becker/Massey map and in a separate analysis of USG 3209 (A Massey derivative)/Jaypee (susceptible) {10505}. USG 3209 possessed Pm8 (1BL.1RS) and an unknown specific resistance factor and their combination had a positive effect on APR even though neither was effective against the races used to identify the QTL {10505}. QTLs on chromosomes 1A, 2A, 2B, 3A, 5D, 6A and 7B were detected in a RE714/Festin population in multiple locations and over multiple years. The QTL on chromosome 5D was detected in all environments and all years and was associated with markers Xgwm639-5D and Xgwm174-5D. Resistance was contributed by RE714. A QTL coinciding with MlRE on 6A was also detected in all environments. The QTL on chromosome 5D and 6A accounted for 45% to 61% of the phenotypic variation {0354}.
      Avocet R(S)/Saar (R) F6 RILs: QTL located on chromosomes 1BL (close to Xwmc44-1B) (Pm39), 7DS (Xgwm1220-7D) (Pm38) and 4BL (XwPt-6209) (resistance allele from Avocet R {10481}.

      AGS 2000 (Pm3a + Pm8 / Pioneer 26R61 (Pm8). QSuSuPm.uga-1AS(SuPm8) with an inhibitory effect on powdery mildew response was located at or near Pm3a.QPm.uga-7AL from Pioneer 26R61 flanked by Xcfa2257-7A and Xwmc525-7A was in the region of the Pm1 locus, even though the test culture was virulent for known Pm1 alleles {11025}.

      SHA3/CBRD(S)/Naxos(R): RIL population: A major QTL on chromosome 1AS accounted for 35% of the phenotypic variation; other QTL from Naxos were on 2DL, 2BL and 7AL. Although SHA3/CBRD possessed a Pm3 allele suppressed Pm8 which appeared to be effective in Norway {10934}. Bainong 64(R)/Jingshuan 16(S). DH lines: Four QTL from Bainong 64: Qpm.caas.1A.
      Xbarc148-1A - Xgwmc550-1A interval. R2=0.074-0.099; QPm.caas-4DL proximal to Xwmc331-4D. R2=0.15-0.23; QPm.caas-6BS, proximal to Xbarc79-6BS, R2=0.09-0.13; and QPm.caas-7AL, proximal to Xbarc174-7AL, R2=0.067-0.071{10680}. CI 13227(S)/Suwon 92(R), SSD population; APR (field resistance) was closely associated with Hg. Xpsp2999-1A and Xpm3B.1 and Xpm3B.2 designed from the Pm3b sequence.
      Fukuho-Komugi/Oligoculm, DH population. QTL for adult plant resistance located on 1AS (R2=22%, Pm3 region, Xgdm33 - Xpsp2999), 2BL (R2=8%, Xwmc877.1-Xwmc435.1) and 7DS (R2=10%) derived from Fukuho-komugi, and 4BL (R2=6% at one of two sites, Xgwm373-Xgwm251) from Oligoculm {10335}. The QTL on 7DS, flanked by Xgwm295.1-7D and Ltn, is likely to be Lr34/Yr18.
      Lumai 21(R)/Jingshuang 16(S), F3 lines: Three QTLs from Lumai 21: QPm.caas-2BS, Xbarc98-2BS - Xbarc1147-2BS interval, R2=0.106-0.206; QPm.caas-2BL, Xbarc1139-2BL - Xgwm47-2BL interval, R2=0.052-0.101; and QPm.caas-2DL, Xwmc18-2DL - Xcfd233-2DL interval, R2=0.057-0.116 {10707}.
      RE9001(R)/Courtot(S) RIL population:QPm.inra-2B (R2=10.3-36.6%), in the vicinity of Pm6, was consistent over environments {10360}. Eleven QTL, detected in at least one environment were identified by CIM {10360}.

91 Reaction to Cephalosporium gramineum
    Disease: Cephalosporium stripe

    QTL: Coda (more resistant)/Brundage (less resistant): RIL population: 7 QTLs identified based on whiteheads; three from Coda - QCs.org-2D.1 (nearest marker C, R2=0.11), QCs.orp-2B (nearest marker Xwmc453-2B, R2=0.08), and QCs.orp-5B (nearest marker Xgwm639-5A, R2=0.12) and four from Brundage (QCs.orp-2D.2 (nearest marker Xbarc206-2D, R2=0.04), QCs.orp-48 (nearest marker wpt-3908, R2=0.05), QCs.orp-5A.1 (nearest marker wpt-3563, R2=0.08), QCs.orp-5A.2 (nearest marker B1, R2=0.05) {10836}.
92 Reaction to Cephus spp.
    Pest: Wheat stem sawfly. North American species C. cinctus; European species C. pygmeus. Resistance to wheat stem sawfly is associated with solid stem (see also: Stem solidness).
    Tetraploid wheat
93 Reaction to Cochliobolus sativus Ito & Kurib.
    Disease: Cochliobolus root rot.
94 Reaction to Colletotrichum cereale
    Susceptibility to this non-pathogen of common wheat is rare, with only one susceptible genotype being documented. A few susceptible tetraploid genotypes were identified {10939}.
95 Reaction to Diuraphis noxia (Mordvilko)
    Insect pest: Russian aphid, Russian wheat aphid. QTL: QTLs for antixenosis were associated with Xpsr687-7D (7DS) and Xgwm437-7D (7DL) in CS/CS (Synthetic 7D) {10136}. Separate antibiotic effects were demonstrated for the same chromosome {10136}.
    A QTL, QDn.unlp.6A, for antixenosis was associated with Xgwm1393-6AL and Xgwm1150-6AL in a CS/CS(Synthetic 6A) DH population {10216}.
96 Reaction to Fusarium spp.
    1 Disease: Fusarium head scab, scab
      Type II resistance. Whereas much of the recent genetic work involved FHB caused by F. graminearum, according to {10514}, F. culmorum is more damaging than F. graminearum in terms of FHB severity, kernel damage, yield reduction and DON/NIV contamination. Patterson (mod sus)/Fundulea 201R RILS: QTLs accounting for 19% and 13% of phenotypic variation were found on chromosomes 1BL (Xbarc8-1BS - Xgwm131-1BL region) and 3AS (Xgwm674-3A/Xbarc67-3A region) {10114}. Two weak QTLs were possibly associated with chromosomes 3D (Patterson allele) and 5AS {10114}.
      Alve(S)/Line 685 R: DH population: QTL on chromosomes 4D (Rht-D1), 3BS, 5A and 2BL {10972}. Two resistance QTL were needed to counteract the negative effect of the Rht-D1b semi-dwarfing allele {10972}. Arina(R)/Forno(S): Three QTLs, QFhs.fal-6DL (R2=22%), QFhs.fal-5BL.1 (in Forno, R2=14%) and QFhs.fal.4AL (R2=10%) and 5 minor QTLs in 2AL, 3AL, 3BL, 3DS and 5DL were detected {10172}.
      Arina/Riband DH lines: QTL affecting ADUPC were identified in 1BL(2), 2B, 4DS, 6BL and 7AL (Arina), and 7AL and 7BL (Riband). The most effective was the 4DS QTL that appeared to be an effect of Rht-D1a rather than height per se {10464}.
      Baishanyuehuang(R)/Jagger(S): RIL population: Four genes/QTLs derived from the resistant parent included Fhd1(R2=0.16), Qfhb.hwwg-3BSc(R2=0.09), Qfhb.hwwg-3A(R2=0.05-0.08) and Qfhd.hwwg-5A(R2=0.05 in one trial) {10950}. Cansas (moderately resistant)/Ritmo (susceptible): Map based analysis across environments revealed seven QTL, QFhs.whs-1BS (1RS), QFhs.whs-3B (not Fhb1), QFhs.whs-3DL, QFhs.whs-5BL, QFhs.whs-7AL and QFhs.whs-7BL (cumultatively, R2 = 0.56). The chromosome 1D gene was primarily involved in resistance to fungal penetration and the others in resistance to spread {10503}. There were significant correlations of FHB response with height and heading date {10503}. Cansas/Ritmo: After QFHS.whs-5BL '(renamed Qfhs.lfl-1BL in {10768})'. Qfhs.lfl-1BL was verified in F4:7 lines and detected in Biscay, History and Pirat {10768}.
      More detailed mapping led to the relocation of the 5B QTL to chromosome 1BL. The renamed Qfhs.lfl-1BL reduced FHB severity by 42% relative to line lacking it {10698}. This gene was also present in Biscay, History and Pirat {10698}.
      Three RGA sequences putatively assigned to chromosome 1A explained 3.37-12.73% of the phenotypic variation in FHB response among F7 and F10 populations {10364}. STS marker FHBSTS1A-160 was developed from one of the RGA.
      CS/CS(Sumai 3 7A): QFhb7AC, nearest marker Xwmc17-7A, explained 22% of phenotypic variance for Type II and 24% of phenotypic variance for Type III resistance {10798}.
      Dream(R)/Lynx(S) RIL population. Following inoculation with F. culmorum 4 QTL for AUDPC were identified on chromosomes 6AL (R2=19%), 1B (12%), 2BL (11%) and 7BS (21%). The resistance allele in 1B came from Lynx and was associated with T1BL.1RS {10260}.
      Dream*4/Lynx lines were developed by selection of QTL on chromosomes 6AL, 7BS and 2BL. Lines carrying QFhs.lfl-6AL and QFhs.lfl-7BS were more resistant than lines lacking them; the 2BL QTL effect was not verified {10470}.
      Frontana(R)/Remus(S): Major QTLs in chromosomes 3AL (Xgwm270-3AL - Xdupw227-3A region) and 5A (Xgwm129-5A - Xbarc-5A region) accounted for 16% and 9% of the phenotypic variation (mainly type 1 resistance) over 3 years {10174}.
      Frontana(MR)/Seri82(S), F3 and F3:5 populations: QTLs were located in chromosomes 1BL (R2=7.9%), flanked by AFLP markers, 3AL (R2=7.7%), flanked by Xgwm720-3A and Xgwm121-3A, 7AS (R2=7.6%), flanked by an AFLP and Xgwm233-7A {10349}.
      Soissons (relative resistant)/Orvantis (susceptible): Increased susceptibility associated with the Rht-D1b allele was further confirmed in crosses of semi-dwarf cultivars Apachi, History and Romanus {10793}.
      Sumai3(R)/Y1193-6(S):RIL population: Three resistance QTL on chromosomes 3BS, 6BL and 2DS with R2 values of 0.26, 0.11 and 0.19, respectively; the last was derived from Y1193-6 {11001}. Treho(S)/Heyne(MR): RIL population: Three QTLs from Heyne, viz Qfhb.hwwg-3AS (R2, up to 0.18), Qfhb.hwwg-4DL (R2=0.14-0.23) and Qfhb.hwwg-4AL (R2, up to 0.18) {11005}. VA00W-38(mod. R)/Pioneer26R46(S):RIL population:Consistent QTL from VA00W-38 detected on chromosomes 1BL, 2A, 2DL, 5B, 6A and 7A explained 6.5-21.3% of the phenotypic variation; one QTL from 24R46 was identified on chromosome 7A {11022}. Major QTL on 2DL, 6A and 5B decreased FHB index, Fusarium damaged kernels, and DON, respectively {11022}. Veery (S) / CJ9306 (R): Four QTLs, XQFhs.ndsu-3BS (Xgwm533b - Xgwm493), QFhs.nau-2DL (Xgwm157 - Xwmc-041), QFhs.nau-1AS (Xwmc024 - Xbarc148) and QFhs.nau-7BS (Xgwm400 - Xgwm573) accounted for 31, 16, 10 and 7%, respectively, of the average phenotypic variation over three years {10490}
      Type I resistance and DON accumulation: Hobbit Sib/T. macha 4A DH population: Both traits were assigned to a small region distal to Xgwm601-4A and cosegregating with Xgwm165-4A {10254}.
      DH181(R)(Sumai 3/HY 386 Seln.): QTL identified in 2DS, 3AS, 3BS, 3B Cent. region, 4DL, 5AS, 6BS {10213}.
      Field resistance: Wuhan-1/Maringa, QTLs were located on chromosomes 2DS, 3BS (Proximal) and 4B {10020}.
      Resistance to DON accumulation: Wuhan-1/Maringa, QTLs were located on chromosomes 2DL and 5DS {10020}.
      Veery/CJ 9306 (R): Four QTLs contributed to resistance; QFhs.ndsu-3BS nearest marker Xgwm533b (R2 = 0.23), QFhs.nau-2DL, Xgwm539 (R2 = 0.2), QFhs.nau-1AS, Xbarc148 (R2 = 0.05) and QFhs.nau-5AS, Xgwm425 (R2 = 0.05) {10496}.
      Haplotype diversity among a large number of FHB resistant and susceptible (mainly Canadian) germplasms indicated similarities in Asian, Brazilian and other materials {10173}. Brazilian cv. Maringa was more similar to Asian than to other Brazilian lines {10173}.
      For review see{0283}.
      Mesterhazy et al.{0006} reported a strong genetic correlation in resistance to different species of Fusarium.
      Bobwhite plants transformed with AtNPR1, an Arabidopsis thaliana gene that regulates activities of SAR, displayed a heritable type II response equal to that of Sumai 3 {10237}.
      In cross Patterson (open)/Goldfield (closed) RILs, narrow flower opening was correlated with FHB resistance. The major QTL effect associated with narrow flower opening and low FHB incidence occured in map interval Xbarc200 - Xgwm210 (29% of variation in FHB incidence); these genes were probably located in chromosome 2BS {10243}.
      Wangshuibai/Annong8455:RIL population: CIM analysis over 2 years detected QTL for FHB response on chromosome 3B (R2=0.17) and 2A (R2=0.12) and for DON levels in 5A (R2=0.13), 2A (R2=0.85) and 3B (R2=0.06) {10447}. The regions involved were Xgwm533-3B - Xbarc133-3B, Xgwm425-2A, and Xgwm186-5A - Xgwm156-5A {10447}.
      In a reciprocal backcross analysis of Chris monosomics/Frontana, Frontana chromosomes 3A, 6A and 4D reduced visibly diseased kernels, kernel weight and DON content, whereas Frontana chromosomes 2A, 2B, 4B and 7A increased the same traits {10398}.
      Further study of the 3A, 6A and 4D reciprocal substitution lines indicated that chromosome 3A of Frontana had the largest effect on incidence, severity, spread and kernel damage, 4D less so and 6A possibly not at all {10900}.
      Soissons (relatively resistant)/Orvantis (susceptible): Soissons carried QFhs.jic-4D (R2=0.106-0.161) associated with Rht-D1a (tall allele) {10718}. FHB susceptibility tended to be associated with the Rht-D1b allele {10718}. Supporting studies with NILs indicated that the presence of Rht-B1b led to reduced type 2 resistance relative to presence of Rht-D1b or the tallness alleles at both loci {10718}.
      Ernie(I)/MO94-317(S):RIL population: 3 QTLs on chromosomes 3BSc, 4BL and 5AS accounted for 31 and 42% of the total phenotypic variances for DON and Fusarium damaged kernels (FDK), respectively. A minor QTL (R2=0.04) for FDK was on chromosome 2B {10831}.
      Grandin(S)/PI277012(I): DH population: Two QTLs, Qfhb.rwg-5A.1 on 5AS (R20.06-0.2) and Qfhb.rwg-5A.2 on 5AL (R2=0.12-0.2) conferred type I and II resistance and reduced DON content {0147}. The new QTL on 5AL was closely but not completely linked with gene q which is present in PI277012 {10860}.

      Nanda 2419 /Wangshuibai:Backcross-derived NILs with Qfh.nau-2B, Qfhs.nau-3B, Qfhi.nau-4B (syn. Fhb4), and Qfhi.nau-5A were developed with Mianyang 99-323 as the recurrent parent {10884}.
      Wheaton(I)/Haiyanzhong: RIL population: Four QTLs, Qfhb.uhgl-7D [syn. Qhb.hyz-7D], nearest marker Xwmc121-7D, R2=0.16-0.2), Qfhb.uhgl-6B.1 [Qhb.hyz-6B.1], R2=0.4), Qfhb.uhgl.6B.2 [Qhb.hyz-6B.2], R2=0.07), Qfhb.uhgl-5A [Qhb.hyz-5A], R2=0.04-0.07) were from Haiyanzhong, and Qfhb.uhgl-1A [Qhb.hyz-1A], R2=0.05) was from Wheaton {10837}. Tetraploid wheat T. dicoccum line Td161 crossed to three durum parents: small effect QTL were detected on chromosomes 3B, 4B, 6A, 6B and 7B; all except the 6A QTL were located at previously known positions {10993}.
      T. dicoccoides Mt. Gerizim#36/*2 T. durum Helidur F6 lines: two QTL for type2 resistance located on chromosome 3A (Xbarc45-3A - Xbarc67-3A and 6B (Xs13m22-2 - Xgwm626-6B {11088}.
    2 Disease: Crown rot caused by Fusarium pseudograminearum, F. culmorum and other Fusarium species
      QTL: Kukri(R)/Janz(S) DH population. Simple interval mapping in the region Pst1 ACG.Mse1 CAC - Xgwm251-4B accounted for 48% of the variation in crown rot response {10034}.
      Lang (S)/CSCR6(R):RIL population: tested under controlled conditions with F. pseudograminearum and F. graminearum: Qcrs.cpi-3BL from CSCR6, R2=0.49 and Qcrs.cpi-4B from Lang R2=0.23 {10703}. Six of 9 NIL pairs made by MAS for Xgwm01081-3B earlier located near the 3BL QTL {10703} in CSCR6 showed significant differences (P < 0.01) in crown rot response {10891}.
      Lang/T. spelta CSCR6: Qcrs.cpi-3BL from CSCR6 was flanked by wPt8438 and wPt9495; R2 up to 0.49, validated in other crosses {10723}. Qcrs.cpi-4B from Lang; R2 up to 0.23 {10723}.
      Sunco/2-49: DH population: Three QTLs for seedling resistance, viz. QCr.usq-1D.1 and QCr.usq-4B.1 (R2=0.19) from 2-49 and QCr.usq-2B.1 from Sunco {10883}. 2-49/W21MMT70: DH lines: Three QTLs fo seedling resistance, viz. QCr.usq-1D.1, and a weaker QTL on chromosome 7A from 2-49 and QCr.usq-3B.1 (R2=0.41) from W22MMT70 {10883}.
      2-49 (partially resistant) / Janz(susceptible) DH population: Analysis of partial seedling resistance indicated major QTL in chromosomes 1D (R2=0.21) and 1A (R2=0.09) and minor QTL in 2A, 2B (from Janz), 4B and 7B {10132}. W21NMT70/Mendos: DH population: three consistent QTLs for seedling resistance were identified with CIM; these were located in chromosome 5D and 2D (resistance alleles from W21NMT70) and 2B (resistance allele from Mendos) {10358}. Sunco/Macon: RIL population: QTLs were located in chromosomes 2B, 3B, 4B and 4D. Qcrs.wsu-3BL from Macon and flanked by Xgwm299-3B was the most effective {10932}. Sunco/Otis: RIL population: QTLs were located in chromosomes 2B, 3B, 4B and 7A. Qcrs.wsu-3BL from Ots was the most effective {10932}.
97 Reaction to Heterodera avenae Woll.., H. filipjeva (Madzhidov) Stelter
    Cereal root eelworm; cereal cyst nematode. QTL: Qcre.src-1B was located to the Xwmc719-1B (R2=12%) - Xgwm140-1B (R2=12%) region in Trident/Molineux {10343}.
98 Reaction to Magnaporthe spp.
    1 Reaction to Magnaporthe grisea (Herbert) Barr: Syn. Pyricularia oryzae
      M. grisea is a pathogen of blast on many graminaceous species, the best known of which is rice. In Brazil it has become a pathogen of wheat. The wheat pathotype(s) is different from those attacking other species such as rice, oat, millets and weeping lovegrass. A second gene designated Rwt3 {0302} was present in CS and Norin 4. Genes Rwt3 and Rwt4 were detected using hybrids of Triticum-virulent and Avena-virulent pathogen isolates.
    2 Reaction to Magnaporthe oryzae
      Wheat cultivars carrying the 2NS translocation from Aegilops ventricosa had 50.4 to 72.3% less head blast than those without 2NS when inoculated with an older isolate (MoT) of Magnaporthe oryzae (Triticum pathotype) under growth chamber conditions. When inoculated with recently collected isolates from wheat, cultivars with 2NS had 64.0 to 80.5% less head blast {11127}.
99 Reaction to Mayetiola destructor (Say) (Phytophaga destructor) (Say)
    Insect pest: Hessian fly. A recombination value of 12.0% between leaf-rust reaction {possibly Lr10} and Hessian-fly reaction in Selection 5240 was reported {018}. temporary designations: Qhf-hwwg-1A {11018}. 1AS{11018}. v: Clark H34 {11018}. ma: Closely linked to Xwgm33-1A {11018}; Located within a 6 cM region flanked by Xwgm33-1A and Xsnp5150-6B, R2=0.1 {11018}.
    Halotype analysis was used to postulate Ae. tauschii-derived genes H13, H22, H23, H26 and H32 in a set of synthetic wheat lines {10983}.
100 Reaction to Meloidogyne spp.
    Root rot nematode, root knot eelworm
101 Reaction to Mycosphaerella graminicola (Fuckel) Schroeter, Zymoseptoria tritici
    Disease: Septoria tritici blotch QTL: Four QTLs for resistance to Mycosphaerella graminicola were identified in replicated field experiments in a double haploid population from Savannah (susceptible)/Senat(resistant). Senat contributed all the alleles providing resistance {10067}
    QStb.riso-2B was mapped on chromosome arm 2BL linked to SSR marker Xwmc175-2B (LOD>5, R2>17%) {10067}.
    QStb.riso-3A.2 was mapped on chromosome arm 3AS linked to SSR markers Xwmc489-3A, Xwmc388-3A and Xwmc505-3A (LOD>4, R2>18%). Also detected at the seedling stage {10067}. Xgwm369-3A is present on chromosome arm 3AS {0187}. A resistance gene from Senat located at or near the Stb6 locus was mapped 5 cM from microsatellite Xgwm369-3A on chromosome arm 3AS {10067}.
    QStb.riso-6B was mapped on the centromeric region between SSR markers Xwmc494-6B and Xwmc341-6B (LOD>16, R2>68%). Also detected at the seedling stage {10067}.
    QStb.riso-7B was mapped on chromosome 7B close to SSR marker Xwmc517-7B (LOD>4, R2>11%) {10067}.
    ITMI Population: Three QTL, QStb.ipk-1DS, QStb.ipk-2DS and QStb.ipk-6DS conferred seedling-stage resistance to 2 isolates, whereas 2 QTL QStb.ipk-3DL and QStb.ipk-7BL conferred separate adult-stage resistances to each isolate {10151}.
    A weak QTL, QStb.psr-7D.1, giving partial resistance to Portuguese isolate IPO92006, was detected in the Xcdo475b-7B - Xswm5-7B region in chromosome 7DS {10341}.

    Apache/Balance: Analyses with a panel of M. graminicola cultures identified QTLs on chromosomes 1BS (Apache, considered to be Stb11), 3AS (Balance, considered to be Stb6), 6DS (Balance, named as Stb18), 7DS (Apache, considered to be Stb4) and 7DL (Apache) {10827}.
    Florett/Biscay(S): RIL population: two QTLs for APR were located on chromosomes 3B and 6D {10901}.
    Tuareg/Biscay(S): RIL population: two QTLs for APR were located on chromosomes 4B and 6B {10901}. Solitar(R)/Mazurka(S): DH population: Resistance under field conditions was associated with QTL on chromosomes 5A, 6D and 7D which accounted for 20% of the genotypic variation; all three were derived from Solitar, but there was no evidence that Stb6 and Stb11, also present in Solitar, were involved {10984}. Steele-ND(R)/ND735(S): RIL population: A consistent QTL (R2=0.1) for seedling resistance flanked by DArT markers XwPt-7101 and X377410 was mapped to chromosome 5BL in the region of Stb1 {10992}. Two other QTLs on chromosomes 1D and 7A were detected in single experiments {10992}.
102 Reaction to Phaeosphaeria nodorum (E. Muller) Hedjaroude (anamorph: Stagonospora nodorum (Berk.) Castellani & E.G. Germano); Parastagonospora nodorum
    Disease: Septoria nodorum blotch, Stagonospra nodorum blotch.
    1 Genes for resistance
      A QTL analysis of SNB response in the ITMI population found significant effects associated with chromosome 1B (probably Snn1) and 4BL, with an interactive effect involving the 1BS region and a marker on chromosome 2B {10009}. An additional QTL on 7BL was effective at a later stage of disease development {10009}.
      Four QTLs, on chromosomes 2B (proximal part of long arm), 3B (distal part of short arm), 5B and 5D, were mapped in a Liwilla / Begra doubled haploid population. Longer incubation period and lower disease intensity were contributed by Liwilla {10045}.
      Two QTLs for glume blotch resistance under natural infection were identified on chromosomes 3BS and 4BL in Arina / Forno RILs {10065}. The 3BL QTL, designated QSng.sfr-3BS, was associated with marker Xgwm389-3B and explained 31.2% of the variation. The resistance was contributed by Arina {10065}. The 4BL QTL, QSng.sfr-4BL, was associated with Xgwm251-4B and explained 19.1% of the variation. Resistance was contributed by Forno {10065}. A QTL on 5BL, QSng.sfr-5BL, overlapped with QTLs for plant height and heading time {10065}. QSng.sfr-3BS peaked 0.6 cm proximal to Xsun2-3B {10465}. Association mapping involving 44 modern European cultivars indicated that the association was retained in a significant proportion of genotypes {10465}.
      A QTL, QSnl.ihar-6AL, identified in DH lines of Alba (R) / Begra (S) accounted for 36% of the phenotypic variance in disease severity and 14% of the variance in incubation period {10143}.

      Br34 / Grandin: Three QTLs with resistance effects from BR34; Qsnb.fcu-5BL.1 (Tsn1), R2 = 0.63, Qsnb.fcu5BL.2, R2 = 0.06, and Qsnb.fcu-1BS (vicinity of Snn1), R2 = 0.10 {10458}. QTL analysis of the RIL population with Culture Sn6 revealed four QTLs, Qsnb.fcu-2DS (R2 = 0.3 - 0.49) associated with Snn2, Qsnb.fcu-5BL (R2 = 0.14 - 0.2) associated with Tsn1, Qsnb.fcu-5AL (R2 = 0 - 0.13) associated with Xfcp13-5A, and Qsnb.fcu-1BS (R2 = 0 - 0.11) associated with Xgdm125-1BS {10507}.
      Forno (S) / Oberkulmer spelt (R). Among 204 RILs leaf and glume response were genetically different but correlated (R2=0.52). Ten QTLs for glume blotch (SNG) resistance were detected, 6 from Forno. A major QTL (R2=35.8%) was associated with q. Eleven QTLs (4 from Forno) affected leaf blotch; 3 of these (chromosome 3D, 4B and 7B) with R2>13% were considered potential candidates for MAS {10250}.
      ITMI population: A major QTL, coinciding with Snn1, was located in chromosome 1BS (R2 = 0.58, 5 days after inoculation), minor QTL were found in 3AS, 3DL, 4AL, 4BL, 5DL, 6AL and 7BL {10009}.
      P91193D1 (partially resistant) / P92201D5 (partially resistant) RIL populations were tested in Indiana and Western Australia for glume resistance. Two QTL were identified: Qng.pur-2DL.1 from P91193D1 (R2 = 12.3 in Indiana and 38.1% in WA, respectively; Xgwm526.1-2D - Xcfd50.2-2D) and QSng.pur-2DL.2 from P99201D5 (R2 = 6.9% and 11.2%, respectively; Xcfd50.3-2D - wPT9848) {10471}.
      HRWSN125(R)/WAWHT2074(S): Constant detection of QSnl.daw-2DL for flag leaf resistance, and QSng.daw-4BL for glume resistance over two years {10584}.
      Tetraploid wheat
      Langdon/Langdon (T. turgidum ssp. dicoccoides Israel-A 5B): QSnb.ndsu-5B located 8.3 cM proximal to tsn1 for tan spot resistance; R2 = 0.38 {10597}.
      A summary of QTL analyses is provided in {10726}.
      Salamouni/Katepwa: RIL population: Two QTLs. QSnb.fcu-1A (Snn4) (R2=0.24) and QSnb.fcu-7A (R2=0.16) were associated with SNB response to isolate Sn99CH 1A7a {10867}.
    2 Sensitivity to SNB toxins (necrotrophic effectors)
      A discussion on the origin and role of host-specific toxins is provided in {10726}.
      Australian cultivars with Tsn1 and tsn1 are listed in {10540}.
      Tetraploid wheat
      In a reevaluation of this work Faris and Friesen {10688} attributed all of the variation in SNB response to the presence or absence of SnTox1.
      QTL: ITMI population: A major QTL, coinciding with Snn1, was located in chromosome 1BS (R2=0.58, 5 days after inoculation), minor QTLs were found in 3AS, 3DL, 4AL, 4BL, 5DL, 6AL and 7BL {10009}.
      P91193D1 / P92201D5 RIL population: tested in USA and Australia: QSng.pur-2DL.1 from P91103D1, R2=0.123 (Indiana) and 0.381 (South Perth); and QSng.pur-2DL.2 from P92201D5, R2=0.069 (Indiana) and 0.112 (South Perth) {10776}.
      Host sensitivity genes in US southern winter wheats are listed in {1241}.
103 Reaction to Pratylenchus spp.
    Root lesion nematode; prats
    2 Reaction to Pratylenchus thornei
      QTLs were located on chromosomes 2BS and 6DS {0122}.
104 Reaction to Puccinia coronata var. hordei.
105 Reaction to Puccinia graminis Pers.
    Disease: Black rust; black stem rust; stem rust.
    Note: Some near-isogenic lines are based on Marquis. The genes present in the Marquis background are not listed for those NILs. Additional temporary designations are listed in {1230}.
    Genotype lists: {323,970,10270,10511,10697}.

    Complex genotypes:

    AC Taber: Sr2, Sr9b, Sr11, Sr12{9905}.
    Centurk: Sr5 {979}, Sr6 {979}, Sr8a, Sr9a {979}, Sr17 {979}.
    Chris: Sr5 {679,1371}, Sr7a {1371}, Sr9g {1371}, Sr12 {1371}.
    Egret: Sr5 {939}, Sr8a {939}, Sr9b {939}, Sr12 {939}.
    FKN: Sr2, Sr6, Sr7a, Sr8a {791}, Sr9b {791}.
    H-44: Sr2, Sr7b {677}, Sr9d {677}, Sr17.
    Hartog: Sr2 {127}, Sr8a, Sr9g, Sr12 {939}.
    Hope: Sr2 {677}, Sr7b {677}, Sr9d {677}, Sr17.
    Kenya Plume: Sr2 {1370}, Sr5 {1370}, Sr6 {1370}, Sr7a {1370}, Sr9b {1370}, Sr12 {1370} Sr17 {1370}.
    Khapstein: Sr2, Sr7a, Sr13 {674}, Sr14 {674}.
    Lawrence: Sr2, Sr7b {939}, Sr9d, Sr17.
    Lerma Rojo 64: Sr2, Sr6, Sr7b {979}, Sr9a {979}.
    Madden: Sr2, Sr9b, Sr11, Sr13 {842}.
    Manitou: Sr5 {679}, Sr6 {679}, Sr7a, Sr9g {965}, Sr12 {939}.
    Mendos: Sr7a {939}, Sr11 {879}, Sr17, Sr36.
    Pasqua: Sr5, Sr6, Sr7a, Sr9b, Sr12. Gene Lr34 acted as an enhancer of APR{9905}.
    PI 60599: Sr7a {689}, Sr8a, Sr9b, Sr10.
    Selkirk: Sr2 {499}, Sr6 {468}, Sr7b {499}, Sr17, Sr23 {950}.
    Redman: Sr2, Sr7b {939}, Sr9d {939}, Sr17.
    Reliance: Sr5 {1308}, Sr16 {1238}, Sr18, Sr20.
    Renown: Sr2, Sr7b {939}, Sr9d {939}, Sr17.
    Roblin: Sr5, Sr7a? Sr11, Sr12.
    Timgalen: Sr5 (heterogeneous) {1555}, Sr6 {1555}, Sr8a, Sr36.
    Thatcher: Sr5 {1308}, Sr9g {965}, Sr12 {939}, Sr16 {1308}.
    WW15 = Anza = Karamu = T4: Sr5 {939}, Sr8a {939}, Sr9b {939}, Sr12 {939}.
    Arina / Forno:Qsr.sun-5BL {10565}; resistance contributed by Arina, associated with Xglk356-5B, R2 = 11-12% {10565}. Qsr.sun-7DS {10565}; resistance contributed by Forno, associated with markers XcsLV34 and Xswm10 diagnostic for Lr34/Yr18 {0828}.
    HD2009/WL711 RILs: Three of several QTLs gave consistent effects across environments, viz. QSr.sun-3BS, R2 = 0.09-0.15, probably Sr2, QSr.sun-5DL, R2 = 0.2-0.44, probably Sr30, and QSr.sun-7A, R2 = 0.07-0.13, nearest marker wPT-4515 {10632}.

    RL6071/RL6058(R): RIL population: RL6058, a Tc backcross line with Lr34/Sr57 is more resistant than Tc. Enhancement of resistance in both Kenya and North America was attributed to a QTL in the region wPt5044 - Xgwm-2B in chromosome 2BL {10902}. Avocet S/Pavon 76: RIL population of lines lacking Sr26:Five QTLs, QSr.cim-3B(Sr2), QSr.cim-1B(Lr46/Yr29/Pm39 region) and QSr.cim-3D (R2=0.2) from Pavon 76; QSr.cim-4B and QSr.cim-5A from Avocet S {10975}. Carberry (Resistant in Canada) / AC Cadillac (Resistant in Canada and Kenya): DH population: QTLs effective in Kenya were located in chromosomes 2B, 5B, 7B and 7D, those effective in Canada were on 3B(Sr2), 5A and 5B; those effective in Kenya and Canada were on 4B and 6D(Sr2); both parents had Lr34/Sr51 {11040}. PBW343(S)/Muu(I): RIL population:4 consistent QTLs were identified, QSr.cim-2BS, QSr.cim-3BS(Sr2) and Sr.cim-7AS from Muu, and QSr.cim-5BL from PBW343 {11019}.
106 Reaction to Puccinia striiformis Westend.
    Disease: Stripe rust, yellow rust.
    2 Temporarily designated genes for resistance to stripe rust
      North American workers {181,186,184} allocated a number of temporary designations to uncatalogued genes detected with North American P. striiformis accessions. Druchamp, Yamhill and Stephens were reported to carry 'Yr3a or Yr4a" because these genes could not be distinguished with the cultures that were used.
    3 Stripe rust QTLs
      Two QTLs in Camp Remy/Michigan Amber were located on chromosome 2BL (QYR1, LOD score 12) and 2AL (QYR2, LOD 2.2) {0287}. Four QTLs were scored in the ITMI population. The most effective (QYR3, LOD 7.4) on chromosome 2BS was probably Yr27, the others were located in 7DS (QYR4, LOD 3.4), 5A (QYR5, LOD 2.8), 3D (QYR6, LOD 2.8) and 6DL(QYR7, LOD 2.4) {0287}.
      Camp Remy/Recital: 217 RILs. Six QTLs for APR were detected over 4 years. QYr.inra-2BL (R2=0.42-0.61) corresponded largely to seedling resistance gene Rsp and possibly Yr7. The other genes were Qyr.inra-2AL, QYr.inra-2BL, QYr.inra-2DS (perhaps Yr16), QYr.inra-5BL.1 and QYr.inra-5BL.2 {10279}. Differential reactions of RILs possessing different QTL occurred between old and new P. striiformis races {11144}.

      Coker 9835 (S) / VA96W-270 RIL population: Adult plant resistance was conferred by QYr.ar-3BS (nearest markers Xbarc147, ger9-3p, IWA6092) and QYr.ar-4BL (nearest markers Xbarc163, Xcfd39 and several IWA markers {11175}. Cultivar Pat had the same haplotype {11175}.

      USG 3555 / Neuse: Three QTL on chromosomes 1AS, 4BL and 7D (not Yr18) were derived from USG 3555 and one QTL on chromosome 3A was from Neuse {11142}. Seven QTLs were identified for stripe rust severity in a joint analysis of five datasets from a Fukuhokomugi/Oligoculm doubled haploid population {10060}. Their location, associated marker, percentage variation explained, and genotype contributing to enhanced resistance at that locus, are listed below.
      3BS; Xgwm389-3B; 0.2-4.9%; Oligoculm {10060}. 4BL; Xgwm538-4B; 1.8-12.3%; Oligoculm {10060}. 4DL; Xwmc399-4D; 2.5-8.0%; Oligoculm {10060.} 5BL; Xwmc415-5B; 2.4-16.1%; Oligoculm {10060}. 6BS(centromeric); Xgwm935-6B; 0.5-3.8%; Oligoculm {10060}. 7BS; Xgwm935-7B; 1-5.2%; Oligoculm {10060}. 7DS; Xgwm295-7D; 10.7-23.7%; Fukuho {10060}; the 7DS QTL was probably Yr18 {10060}. Four QTLs were identified for stripe rust infection in a joint analysis of three datasets from a Fukuhokomugi/Oligoculm doubled haploid population {10060}. Their location, associated marker, percentage variation explained and parent contributing to enhanced resistance at that locus are listed below.
      2DL; Xgwm349-2D; 6.5-9.6%; Fukuho {10060}. 3BS; Xgwm389-3B; 15.1-24.5%; Oligoculm{10060}. The 3BS QTL may be Yr30 {10060}. 5BL; Xwmc415-5B; 6.4-12.7%; Oligoculm {10060}. 7BL; Xwmc166-7B; 2.5-9%; Oligoculm {10060}. Otane (R)/Tiritea (S) DH population: QTL in 7DS (probably Yr18), 5DL (from Otane) and 7BL (Tiritea) {10150}. Interval mapping of 7DS indicated that the presumed Yr18 was 7 cM from Xgwm44-7D {10150}.
      Kariega/Avocet S DH population: Two QTLs QYr.sgi-7D (probably Yr18) and QYr.sgi.2B.1 accounted for 29 and 30%, respectively, of the phenotypic variation for stripe rust response. The nearest marker to the latter was Xgwm148-2B {10184}.
      Four QTLs were detected in a multiple cross analysis {10283}: Chromosome 2AL (probably Yr32 in Deben, Kris and Soloist), 2AS (probably Yr17 in Kris), 2BL (Xwmc149-2B - Xwmc317a-2B in Deben) and 6BL (Xwmc397-6B - Xwmc105b-6B in Soloist and Kris).
      Avocet S/Pavon 76: QTL identified in 1BL (Xgwm259), 3BS (PstAATMseCAC2), 4BL (Xgwm495), 6AL (Xgwm617), 6BL (PstAAGGMseCGA1) {10443}.
      T. monococcum PAU14087 (resistant)/T. boeoticum PAU5088 (resistant): RIL population: One adult plant resistance QTL identified in each parent and named QYrtm.pau-2A (in a 3.6 cM interval between Xwmc407-2A and Xwmc170-2A; R2 = 0.14) and QYrtb.pau-5A (in a 8.9 cM interval between Xbarc151-5A and Xcfd12-5A; R2 = 0.24) {10518}.
      Luke (R) /Aquileja (R): Two QTLs for high-temperature adult plan resistance, QYRlu.cau-2BS.1 (distal, flanked by Xwmc154-2B and Xgwm148-2B, R2 = 0.366) and QYrl.cau-2BS.2 (proximal, flanked by Xgwm148-2B and Xbarc167-2B, R2 = 0.415) from Luke, and QYraq.cau-2BL (flanked by Xwmc175-2B and Xwmc332-2B, R2 = 0.615) in Aquileja for stripe number {10582}.
      Avocet S / Attila: QTLs were located on chromosomes 2BS (probably Yr27), 2BL (a race-specific effect) and 7BL (XP32/M59 - Xgwm344-7B {10586}.
      Guardian / Avocet S: F3 lines. One major QTL, QPst.jic-1BL (Xgwm818-1 - Xgwm259-1B, R2 up to 0.45), and two minor resistance QTLs on chromosomes 2D and 4B originating from Guardian {10589}. The major QTL was in the region of Yr29.
      Stephens / Michigan Amber: Two QTLs for high temperature APR were located in chromosome 6BS; QYrst.wgp-6BS.1 located in a 3.9 cM region flanked by Xbarc101-6B and Xbarc136-6B and QYrst.wgp-6BS.2 located in a 17.5 cM region flanked by Xgwm132-6B and Xgdm113-6B {10602}.
      Pingyuan 50 (R) / Mingxian 169 (S): DH population: APR: QYrcaas-2BS (Xbarc13-2BS) - Xbarc230-2BS, R2=0.05-0.09), QYr.caas-5AL(Xwmc410-5AL - Xbarc261-5AL, R2=0.05-0.2), QYrcaas-6BS(Xgwm361-6BS - Xbarc136-6BS, R2=0.05-0.08) {10693}.

      Renan (R) / Recital (S) RIL population: Tested for AUDPC in 1995/6 and 2005/6 with pathogen isolates avirulent and virulent, respectively, for Yr17:QYr.inra-2AS.2, (=Yr17), R2=0.45, 1995/6; QYr.inra-2AS.1, R2=0.9, 2005/6; QYr.inra-2BS, R2=0.11 & 0.13, QYr.inra-3Bcent, R2=0.06 in 2005/6; QYr.inra-6B, R2=0.04 & 0.06; from Renan; and QYr.inra-2AS.1, R2=0.09; QYr.inra-3DS, R2=0.08 & 0.12 from Recital. Other QTLs were effective only at certain growth stages {10689}.

      Express / Avocet S: RIL population: Relative AUDPC for high temperature APR was controlled by QYrex.wgp-6AS, R2=0.326, interval Xgwm334-6A - Xgwp56-6A; QYrex.wgp-3BS, R2=0.274, interval Xgwm299-3B - Xwgp66-3B, QYrex.wgp.1BL, R2=0.094, interval Xwmc631-1B - Xwgp78-1B {10672}. When rust phenotyping was based on infection type only the 6S and 3BL QTLs were evident {10672}. Alcedo (R) / Brigadier (S): DH population: Two major QTLs QPst.jic-2DL (R2 up to 0.36) and QPst.jic-4BL (R2 up to 0.29) for percent infection contributed be Alcedo {10785}. A seedling-expressed QTL was located at the same position in 2DL {10774}.
      Flinor (R) / Mingxian 169 (S): Two independent QTLs for high temperature (24/18C) seedling resistance located in chromosome 5BL, designated QYr-tem-5B.1 (Xbarc89 - Xgwm67) and QYr-tem-5B.2 (Xbarc140n - Xwmc235) and R2 = 0.37 and 0.33, respectively {10797}.
      Kukri (MR) / Janz (MR): DH population: Tested with pre- and post-2003 Australian Pst races in several environments. QYr.sun-7B (Kukri) and Qyr.sun-7D(=Yr18)) (Janz) were consistent over environments; QYr.sun-1B, -5B and -6B were detected in most environments and QYr.sun-3B was identified in only one season. Two genes, QYR.sun-1A from Janz and QYr.sun-2A from Kukri, were detected only with pre- and post-2003 races, respectively, and likely contributed to differential responses of these cultivars to the two groups of races {10751}.

      Pioneer 26R61(R)/AGS2000(S): RIL population: Two QTLs, QYr.uga-2AS (R2=0.56) flanked by Xbarc124-2A and Xgwm359-2A (also named YrR61) and QYr.uga-6AS (R2=0.06) {10914}. Minor QTLs were also on other chromosomes.

      UC1110(MR)/PI 610750(MR): RIL population: QYr.ucw-3BS ex UC1110, R2=0.22, associated with Xgwm522-3B.1. This marker differs from Xgwm533-3B.2 that is associated with Yr30 {10705}; QYr.ucw-5AL, R2=0.1, ex PI 61075 - syn. Yr48 {10705}; QYr.ucw-2BS, R2=0.045, ex UC1110, located in the centromeric region near Xwmc474-2BS {10705}, Qyr.ucw-2AS, R2=0.023, ex PI 61725, near wPt-5839 {10705}.

      Stephens I/Platte(S): RIL population: 13 QTLs were identified across several environments; significant QTL x environment interactions suggested that plant stage specificity, pathogen genotype and temperature as well as host genotype were important in determining rust response {10890}. Avocet(S)/Chapio(I): F6RIL population: In Mexico QTLs were located in chromosomes 2BS (Yr31), 3BS(Yr30) and 7DS(Yr18); only the last two were effective in 2009. In China QTLs were located in chromosomes 3BS, 5BL and 7DS. A 3DS QTL was effective in Mexico in 2009 and in China in 2013 {11020}. Avocet(S)/Pastor(I): RIL population: QTL mapped on 1BL (Yr29), 2BS(Yr31), 5A,6B and 7AL plus minor QTLs on 1AL, 1B, 3A, 3B, 4D, 6A, 7AS and 7AL {10928}. Claire/Lemhi: DH population: 4 QTLs for APR:Qyr.niab-2D.1 (at or near Yr16, R2=0.1-0.25, Qyr.niab2DL.2 (R2=0.14-0.32), Qyr.niab-2BL, and Qyr.niab-7B (R2=0.11-0.13) {10962}. An unknown seedling resistance gene was located in chromosome 3BL {10962}. Jagger(MR)/2174(MS): RIL population: Qyr.osu-2A(Yr17) and Qyr.osu-5A (in Xgwm156-5A - centromere region) from Jagger and Yr18 from 2174 (but only in tests in China) {10973}. Yr16DH70 (Cappelle Desprez/2*Palmiet Selection)/Palmiet: DH population: One major effect QTL, Qyr.ufs-2A, and three less effective QTLs in 2D (possibly Yr16), 5B and 6D were from Yr16DH70, and a minor effect QTL on 4B was from Palmiet {10933}. UC1110(MR)/PI 61070(MR): RIL population: Four QTLs for APR:two, Qyr.ucw-3BS, peaking at Xgwm533.1, R2=0.22, and Qyr.ucw-2BS, R2=0.05 from UC1110, and Yr48 and Qyr.ucw-2AS, R2=0.02, from PI 61070 {10705}.

      A summary of published QTL locations is provided in {11089}; 49 chromosome regions on 20 of the 21 wheat chromosomes were proposed.

107 Reaction to Puccinia triticina
    Disease: Brown rust, leaf rust.
    1 Genes for resistance
      A series of temporary designations for seedling and adult plant resistance genes in six durums is given in {1648}.
      A potentially novel resistance gene was located in chromosome 5BS of Iranian landrace PI 289824. Xgwm234-5B - 8.9 cM - Lr - 2.3 cM - STS Xtxw200 {10253}. Complex genotypes:
      AC Domain: Lr10 Lr16 Lr34 {820}.
      AC Splendor: Lr1 Lr16 Lr34 {10179}
      AC Teal: Lr1 Lr13 Lr16 {821}
      Alsen: Lr2a Lr19 Lr13 Lr23 Lr34 {10152}
      Alsen: Lr2a Lr10 Lr13 Lr23 Lr34 {10223}.
      Benito: Lr1 Lr2a Lr12 Lr13 {1256}.
      Buck Manantial: Lr3 Lr13 Lr16 Lr17 Lr34? {300}.
      Coker 9663 Lr9 Lr10 Lr14a {10742}.
      Estanzuela Benteveo: Lr13 Lr26 Lr34 {10980}.
      Estanzuela Pelon: Lr1 Lr17a Lr26 Lr34 {10980}.
      Estanzuela Tarariras: Lr3bg Lr13 Lr34 {10980}.
      Era: Lr10 Lr13 Lr34 {342}.
      Grandin: Lr2a Lr3 Lr10 Lr13 Lr34 {821}.
      INIA Boyero: Lr13 Lr26 Lr34 {10980}.
      INIA Churrinche: Lr10 Lr24 {10980}.
      INIA Tero: Lr17a Lr24 {10980}.
      Mango: Lr1 Lr13 Lr26 Lr34 {1374}.
      MN7529: Lr1 Lr2a Lr10 Lr16 {976}.
      Norm: Lr1 Lr10 Lr13 Lr16 Lr23 Lr34 {10152,10223}
      Opata 85: Lr10 Lr27+Lr31 Lr34 {1058}.
      Pasqua: Lr11 Lr13 Lr14b Lr30 Lr34 {304}.
      Pioneer 26R61 Lr13 Lr14b Lr26 {10742}.
      Prospect: Lr1 Lr2a Lr10 Lr13 {197}.
      Roblin: Lr1 Lr10 Lr13 Lr34 {303,713}.
      Trap: Lr1 Lr3 Lr10 Lr13 Lr34 {1374}.
      Genotype lists:Australian cultivars {0288}; Chinese cultivars {0013,10682}; Combinations with Lr34{1361}; Cultivars from the former USSR {1380}; Czechoslovakian cultivars{855,0102}; European cultivars {0229,0260,0288,0337,10345,10794}; Indian cultivars {1365,1345}; Indian Subcontinent{1365}; Mexican cultivars{1373}; U.S.A. cultivars {1219,978,0334,10111,10146,10152}, French cultivars {10792}, Croatian cultivars {11135}. Kazakhstan cultivars {11161}, see also {970}.

      See {11178} for review and analysis of leaf rust resistance genes in six durum wheats.
    3 QTLs for reaction to P. triticina
      Two QTLs, located distally on chromosome arm 1BL and on chromosome 7DS, were mapped for leaf rust severity in a Fukuho-komugi/Oligoculm doubled haploid population {10060}. The resistance on 1BL was contributed by Oligoculm and explained 15% of the variation. The 1BL QTL may correspond to Lr46 and was associated with marker Xwmc44-1B {0460}. The resistance on 7DS was contributed by Fukuho-komugi and explained 41% of the variation. The 7DS QTL corresponds to Lr34 and was associated with marker Xgwm295-7D {10060}.
      Two major QTL, located on chromosomes 7D and 1BS, for leaf rust resistance were mapped in an Arina/Forno RIL population {10066}. The resistance on 7D was contributed by Forno and explained 32% of the variation. This QTL most likely corresponds to Lr34 {10066}. The resistance on 1BS (QLr.sfr-1BS) was associated with Xgwm604-1B and was contributed by Forno {10066}. Additional minor QTLs were identified on chromosome arms 2DL, 3DL, 4BS and 5AL {10066}.
      QTLs for leaf rust resistance were identified in {0050} and were named by the catalogue curators as follows:

      QTLs: Two QTLs for slow leaf rusting, located on chromosomes 2B and 7BL, were mapped for final severity, area under disease progress curve, and infection rate in a CI 13227 (resistant)/Suwon (susceptible) SSD population {10211}. QLr.osu-2B was associated with microsatellite markers Xbarc18-2B and Xbarc167-2B (R2 = 9-18%). QLr.osu-7BL was associated with microsatellite marker Xbarc182-7B (R2 = 12-15%) {10211}. CI 13227 constributed the resistant alleles for both QTLs. QLrid.ocu-2D, linked to Xgwm261-2D, affected the duration of infection {10211}.
      However, Thatcher backcross derivatives of CI 13227 appeared to have Lr3c and Lr46 {11021}. Avocet/Pastor RIL population: QTLs mapped on 1BL (Lr46, 2BS, 5A, 6B and 7BL plus minor QTLs on 1B, 2A and 2D {10928}. Avocet S/Pavon 76: QTL identified included: 1BL (PstAFAMseCAC1&2), 4BL (Xgwm368), 6AL (Xgwm617), 6BL (PstAGGMseCGA1) {10443}. Avocet S/Attila: At least two additive genes for slow rusting {10586}. In addition to Lr46 there were small effects on chromosomes 2BS, 2BL and 7BL {10586}.
      Beaver/Soissons DH population: QTL for resistance to Australian pathotypes were located on 4-6 chromosomes over 3 years; the most consistent being 1B(1BL.1RS), 4BS (proximal to Xbarc20-4B) and 5AS (QTLBvr5AS, proximal to Xbarc10-5A) and in the vicinity of wPt-8756 and wPt-1931 {10687}.
      TA 4152-60 (MR) / ND495 (MR): DH population: Five QTLs for APR were identified in the field, viz QLr.fcu-3AL (R2 = 0.18), QLr.fcu-3BL (R2 = 0.19), QLr.fcu-5BL (R = 0.07), and QLr.fcu-6BL (R2 = 0.12) from TA 4152-60 and QLr.fcu-4DL (R2 = 0.13) from ND495 {10757}. The 3AL gene also conferred seedling resistance to some races and the 3BL gene conferred resistance to race MFPS {10757}.
      Tetraploid wheat
      Colosseo/Lloyd: A major QTL, QLr.ubo-7B.2, for seedling and adult plant resistance from Colosseo, was located between Xgwm344.2-7B and DART 378059, bin 7BL 10-0.78-1.00 {10600}.

      TA4152-60/ND 495 DH population: Four QTLs for APR, QLr.fcu-3AL(Xcfa2183-3AL - Xgwm666-3AL, R2=0.18), Qlr.fcu-3BL(Xbarc164-3BL - Xfcp544-3BL, R2=0.19), Qlr.fcu5BL, and Qlr.fcu-6BL(Xbarc5-6BL - Xgwm469.2-6BL, R2=0.12) were from TA4152-60 and Xlr.fcu-4DL(Xgdm61-4DL - Xcfa2173-4DL, R2=0.13) was from ND495 {10717}. The 3AL QTL conferred seedling resistance to all 3 races, and the 3BL gene gave race-specific seedling resistance to one race. Xlr.fcu-3BL was effective only in the presence of an allele associated with Xgwm359-5DS {10717}.
      Association mapping in durum wheat indicated genomic regions affecting leaf rust response in chromosomes 1A, 1B, 2A, 2B (Lr13, Lr23 region), 3B, 5A, 5B, 6B, 7A, and 7B (see Lr14 {10736}.
108 Reaction to Pyrenophora tritici-repentis (anomorph: Drechlera tritici-repentis)
    Disease: Tan spot, yellow leaf spot. Virulence in the pathogen is mediated by host-specific toxins and host resistance is characterized at least in part by insensitivity to those toxins. Three toxins, Ptr ToxA, Ptr ToxB and Ptr ToxC have been identified (see {10153}). Toxin sensitivity determined by use of toxins extracted from pathogen strains and resistance determined by infection experiments are treated as different traits, although common genes may be involved.
    A review is provided in {10690}. Grandin(S)/BR34(R) RILs: QTL in 1BS, QTs.fcu-1BS, (13-29% of variation depending on race) and 3BL, (13-41%) were involved in resistance to 4 races. Five other QTL showed race specific responses {10248}.
    Introgressions of genes for insensitivity to Ptr ToxA and Ptr ToxB are outlined in {10153}.
    Batavia (S) / Ernie (R): DH population tested over three years. Four (1A(Ernie), 7A, 2BS, 3BS(Batavia)), five (2BS, 5BL(E), 3D, 6A, 7D(B)) and four (2BS, 5BL(E), 1A, 6A(B),) QTL accounted for most of the variation in each year. The greatest effect across years was the QTL on chromosome 2BS (R2=0.382, 0.298 and 0.362, respectively). This QTL was validated in four additional populations {10782}.
    TA4152-60(R)/ND495(S)DH population. Five QTLs for resistance, all from TA4152-60 {10580}, viz., QTs.fcu-2AS and QTs.fcu-5BL.1 conferring resistance to all races used, QTs.fcu-5AL conferring resistance to races 1, 2 and 5, QTs.fcu-5B.2 conferring resistance to races 1 and 2, and QTs.fcu-4AL conferring resistance to race 3.
    WH542(R)/HD29(S) RIL population: SIM indicated QTL on chromosomes 1B, 3AS, 3BL, 5B and 6BS, but only two were confirmed by CIM, Qts.ksu-3AS flanked by Xbarc45-3A and Xbarc86-3A (LOD 5,4, R2 = 0.23) and Qts.ksu-5BL (probably Tsn1) flanked by Xgwm499-5B and Xest.stsbe968-5B (LOD 6,5, R2 = 0.27) {10552}.
    Wangshuibai / Ning 7840: RIL population: Race 1: QTs.ksu-1AS, R2=0.39 (nearest marker Xcfa2153-1A and QTs.ksu-2BS, R2=0.04) (nearest marker Xbarc2-2B {10753}.
    2 Insensitivity to tan spot toxin (chlorosis)
      QTLs: 'ITMI population': In addition to tsc2 which accounted for 69% of the phenotypic variation in response to race 5, a QTL in chromosome 4AL (Xksu916(Oxo)-4AS, W-7948) accounted for 20% of the phenotypic variation {10015}.

      Salamouni/Katepwa: RIL population: variation at the Tsc2 locus explained 54% of the variation in response to race DW5 {10871}.
109 Reaction to Rhizoctonia spp.
    Cause of Rhizoctonia root rot.
110 Reaction to Sitobion avenae
    English grain aphid.
111 Reaction to Sitodiplosis mosellana (Gehin)
    Insect pest: Orange blossum wheat midge, Wheat midge. This pest should not be confused with Contarinia tritici, the yellow blossom wheat midge. QTL:
    Reeder I/Conan: RIL population: QSm.mst-1A, flanked by Xwmc59-1A and Xbarc1022-1A was the most effective and constant QTL for reduced larval infection over two years (R2=0.17 and 0.34) {10841}. RILs with this QTL in three genetic backgrounds had reduced infestations of 42% {10841}.
112 Reaction to Schizaphis graminum Rond. (Toxoptera graminum Rond.)
    Insect pest: Greenbug QTL: Antibiosis was associated with several markers, including Rc3 (7DS) in chromosome 7D {10167}. QGb.unlp.6A, for antixenosis was associated with Xgwm1009-6A and Xgwm1185-6A in a CS/CS(Synthetic 6A) DH population {10216}.
113 Reaction to Soil-Borne Cereal Mosaic Virus
    Syn.: Soilborne wheat mosaic. Vectored to the roots by the fungus, Polymyxa graminis A major QTL, QSbv.ksu-5D, (R2=0.38) was found in Karl 92*2/TA4152-4 {10273}; the resistance was contributed by Karl 92.
114 Reaction to Tapesia yallundae. (Anomorph: Pseudocerosporella herpotrichoides (Fron) Deighton)
    Disease: eyespot, strawbreaker footrot.
115 Reaction to Tilletia caries (D.C.)Tul., T. foetida (Wallr.) Liro, T. controversa
    Disease: Bunt, dwarf smut, stinking smut. QTL: Blizard (R) / 8405-JC3C (S): DH population. Resistance and markers Xgwm374-1BS, Xgwm364-1BS and Xbarc128-1BS were within a 3.9 cM interval {10783}.
    Trintella/Piko: DH population: One major gene in the chromosome 1BS centromere region, nearest marker Xgwm273-1B {11003}. Smaller QTL effects were detected on chromosomes 7A, 7B and 5B in different years.
    Idaho 444 (R) / Rio Blanco S: RIL population: Three QTL for dwarf bunt resistance: QDB.ui-7DS (R2 = 0.3 ? 0.6), QDB.ui-1A (r2 = 0.11 ? 0.15) and QDB.ui-2B (R2 = 0.06). Two PCR-based markers were developed for the wPt-2565 sequence on chromosome 7DS {11182}.
116 Reaction to Tilletia indica Mitra
    Disease: Karnal bunt.
117 Reaction to Ustilago tritici (Pers.) Rostrup
    Disease: Loose smut.
118 Reaction to Wheat Spindle Streak Mosaic Bymovirus (WSSMV)
    QTL : 79% of the variation between Geneva (resistant) and Augusta (susceptible) was associated with markers Xbcd1095-2D and Xcdo373-2D located 12.4 cM apart in chromosome 2DL {0131}. WSSMV is soil-borne and vectored by the fungus Polymxa graminis. This virus has some sequences similarity to Wheat Yellow Mosaic {10285}.
119 Reaction to Wheat Streak Mosaic Virus
    Vectored by wheat curl mites, Eriophyes tulipae and E. tosichella. See: Resistance to colonization by Eriophyes tulipae. According to {10226} WSMV may also be see-borne. At least some sources of resistance to WSMV are also effective against Triticum mosaic virus.
120 Reaction to Xanthomonas campestris pv. undulosa
    Disease: Bacterial leaf streak
121 Resistance to Colonization by Eriophyes tulipae (Aceria tulipae)
    Mite pest: Wheat curl mite.
    Eriophyes tulipae is the vector of wheat streak mosaic virus (WSMV) and the wheat spot mosaic agent (WSpM).
122 Reaction to Wheat Yellow Mosaic Virus
    WYMV is soil-borne and vectored by the fungus Polymxa graminis. This virus has some sequence similarity to Wheat Spindle Streak Mosaic {10258}. another bymovirus. QTL:
    RIL population: Xifeng(R)/Zhen 9523(S): Three QTLs, Qym.njuy5A.1 (R2 = 0.26-0.54), Ym.njau-3B.1 (R2 = 0.03-0.01) and QYm.njau-7B.1 (R2 = 0.03-0.05 in some trials). The chromosome 5A gene was closely associated with Xwmc415.1, CINAU152 and CINAU153 and was phenotyped as a single Mendelian gene {11073}.