1. Morphological and Physiological Traits⌂ Home

1.1. Gross Morphology: Spike characteristics⌂ Home

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.

Q

Q
Synonymk 1550; Ap2-5A
Chromosome5AL
Chromosome bin5AL-17 10541.
VarietiesCommon wheats. CS; Iranian spelts 140.
Type varietiesT. turgidum ssp. carthlicum, durum and polonicum 10457.
Marker associationsComplete linkage with cDNA clone PtAq22 0127.
cQ was cloned and shown to have similarity to AtAP2 (APETALA 2), the Q allele was more abundantly transcribed than the q allele transcription factors 10457. GenBank AY02956.1.
q
SynonymK 1550
VarietiesMacha wheats; European spelt wheats 10457; vavilovi wheats.
Sources / synonymsCS[*] 8/White Spring Spelt 5A 1048.
Type varietiesT. turgidum ssp. dicoccum, dicoccoides 10457.
Marker associationsCent – Xrsq805(Empb)-5A – 4.6 cM – Q – 4.3 cM – Xpsr370-5A 419; Q was physically mapped in 5AL, fraction length 0.87, bracketed by deletions 5AL-7 and 5AL-23 446; Q – 9.3 cM – Xpsr370-5A 9903. The speltoid phenotype of at least some spelts may be caused by genes at other loci 0140. Fine mapping of the 20 cM region possessing Q and delimited by deletions 5AL -7 and -23 is reported in 0324.
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. Pleiotropic features of the Q locus include effects on glume toughness, threshability, rachis fragility, spike length, flowering time, and plant height 11342. Final spike and reproductive morphology is affected by the Q/q sequence and its regulation by miR172 11344 along with direct or indirect interaction with the homoeologues 11344. Homoeologues of Q were described in 11192. Both have miRNA172 target sites close to the 3’ ends of the coding region. These genes were referred to as Ap2-5B , a transcriptionally active pseudogene, and Ap2-5D that encodes a functional protein that contributes to suppression of the speltoid phenotype 11342. Reduced height gene Rht23 , a mutationally derived allele in NAUH164, was caused by a SNP (G3147A, Ala416Thr) within the miR172 target site in 5DL that permitted up-regulation of Ap2-5D due to down-regulation of miR172 in leaves, stems and spikes 11345.

C

C
SynonymCd 47
Chromosome2DL
probably 2DL 10578. 2D 1192.
Cg
Chromosome2BL
Chromosome bin2BL-0.48-0.89, near breakpoint 0.69.
VarietiesAkage Gumbai 11114; Akage Gumbai 22 11114; Gumbai 22 11114; Kinoshita Komugi 11114; Nakote Gumbai 11114.
Marker associationsXhbg410/Xhbg440-2B – 18.1 cM – Cg – 15.3 cM – Xgwm47-2B 11114.
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 2A[m] 10578 Tetraploid wheat: A compact spike gene C[17648] in mutant line MA 17648 wad located in chromosome 5AL 10541. Xbarc319-5A – 9.7 cM – C[17648] – 24.8 cM – Xgwm179-5A 10541. C[17648] was distal to the Q locus 10541. QTL Courtot/Chinese Spring: Six QTL for spike compactness were detected 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.

S1

S-A1
Chromosome3A
VarietiesCS 0029.
S-A1a
VarietiesCS 0029; common wheats 0029.
S-A1b
SynonymS3 0056
VarietiesMS 1453 0056.
Marker associationsXgwm2-3A (S) – 5.1 cM – S-A1 – 6.6 cM – Xgwm720-3A (L) 0030.
S-B1
Chromosome3B
VarietiesCS 0029.
S-B1a
VarietiesCS 0029; common wheats 0029.
S-B1b
Synonyms[16219] 10541, S2 0030
VarietiesMSK 2452 0056; MSK 24540056.
Type varietiesMA 16219 10541.
Marker associationsXgwm685-3B (S) – 4.2 cM – S-B1 – 0.5 cM – Xgwm566/Xgwm845/ cent 0030.
S-D1
NoteTraesCSD01G137200 .
Chromosome3DL
Chromosome3D
Chromosome3DS
VarietiesCS 0029.
S-D1a
VarietiesCS 0029; common wheats 0029.
S-D1b
Synonyms1 , sp1 1286, Tasg-D1 11415
iS-615[*] 11/ T. sphaerococcum var. rotundatum 1500.
Sources / synonymsCS[*] 7/ T. sphaerococcum rubiginosum 3D 1304.
VarietiesNongda 4332 11415; Sphaerococcum wheats 0029; T. antiquorum K056397 & K56398 10234.
Marker associationsLocated between markers Xgwm341-3DS and Xgdm72-3DS 11415.
cThe sphaerococcum phenotype is caused by a gain of function mutation in serine/threonine kinase glycogen synthase kinase 3 (STKcGSK3) that in rice affects the brassinosteroid signaling pathway and grain shape 11415. There is a Lys286Glu substitution in the ninth exon of TraesCSD01G137200_ 11415.
S-D1c
SynonymS1 0056
VarietiesMS 3287 0056.
Marker associationsXgdm72-3D (S) – 8.0 cM – S-D1 – 2.9 cM – Xgwm456-3D/ cent 0030.

S2

s2
NotePartially dominant 1286.
Synonymsp2 1286
VarietiesSphaerococcoid wheats. " Sphaerococcum simulator " 1286.
Sphaerococcum-like tetraploid wheats were reported 122, 475, 1282, 1286, but comparisons between them, or with s2 , were not made. Whereas Schmidt & Johnson 1281 reported a single recessive controlling the sphaerococcum character in tetraploid wheat, Joppa 621 using the same stock found that two recessive genes were necessary to produce this phenotype.

1.2. Branched spike⌂ Home

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.

BH1

bh-A1
Synonymbh 665
Chromosome2AS
Type varietiesPI 349056 665.
bh-D1
Synonymmrs 10637
Chromosome2DS
Chromosome bin2DS5-0.47-1.0 10637.
VarietiesRa1 10637; Ruc163167-1-02 = Ra1 / ZGK242-81 10637; Ruc163167-1-02 = Alana /3/ Ra1 / ZGK242-82 // Ra1 10637.
Marker associationsXwmc453-2D/bh-D1 – 7.8 cM – Xgwm988-2D 10637; Xwwm484-2D – 3.3 cM – Xwmc453-2D/bh-D1 – 3 cM – Xwgm988-2D 10637.
Ra1 is a mutant stock maintained at the NI Vavilov Research Institute of Plant Industry, St Petersburg, Russia. A chromosome 2B gene of minor effect was identified 9907. In a monosomic analysis of the hexaploid line LYB with supernumerary spikelets, Peng et al. 9908 located recessive genes in chromosomes 2A and 4A that promote the development of supernumerary spikelets and a gene in chromosome 2D that prevents their expression.
bh-R1
Synonymmo 10637
Chromosome2R
alS. cereale D40 10637.
Marker associationsXrms056-2R – 15.7 cM – bh-R1 – 10.7 cM – Xcfe209-2R 10637.

1.3. Elongated glume⌂ Home

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.

P1

P1
SynonymP-A[pol] 1 0254, P 911, Eg 922, P-A[pet] 1 0254
Chromosome7AL
7A or 7B (based on linkage of 0.2 with a gene for red coleoptile) 922. 7AS {11606, 11587}.
P2
Chromosome7BL
itvLD222[*] 7/ T. ispahanicum 9990.
Type varietiesT. ispahanicum 9990. According to 0254 the loci of T. polonicum, T. petropavlovsky and T. isphanicum are allelic ('homoeoallelic') whereas other workers had claimed genes in the first two forms were not allelic. Wang et al. 0254 however concluded that loci bearing alleles for elongated glumes in T. turanicum and T. durum conv. falcatum were not part of the above series.

1.4. Ear length⌂ Home

QEl.ocs-5A.1
Chromosome5AL
VarietiesCS( T. spelta 5A)/CS(Cappelle-Desprez 5A) RI mapping population 9903.
Marker associationsAssociated with Xbcd9 -5A 0068.

1.5. Multi-gynoecium; Multi-ovary⌂ Home

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.

PIS1

Pis1
Chromosome2DL
Chromosome binC-2DL3-0.49 10636.
iCM28TP 11228.
VarietiesTP Mutant 10636.
Marker associationsXgwm539-2D – 17.6 cM – Pis1 – 19.5 cM – Xgwm349-2D 10636. KM69 132294739 – 3.5 cM – KM70136805221 – 3.0 cM – PIS1 – 1.1 cM – KM71140258883_ 11228.
A gene designated Mov-1 (multi-ovary) mapped to bin 2DL-9 in a CIMMYT line is assumed to involve the same locus as Pis1 . Mov-1 was mapped to a 3.5 cM interval (589.3 – 590.4 Mb) 11636.

1.7. Alkylresocinol content in grain⌂ Home

AR

Ar1
NoteHigh alkylresocinol content is dominant 0281.
Chromosome5AL
Type varietiesLangdon0281.
ar1
Type varietiesArdente 0281; this cultivar has a low content compared to all tested durum and common wheats 0281.

1.8. Aluminium tolerance⌂ Home

ALT1

Alt1
VarietiesET3 = Carazinho/4[*] Egret 234.
alt1
VarietiesES3 = Carazinho/4[*] Egret 234.

ALT2

Alt2
SynonymAltBH 1213
Chromosome4DL
suT. turgidum cv. Langdon 4D(4B) 848.
VarietiesBH1146{0115, 1213}; IAC-24 0115; IAC-60 0115; 13 induced mutants of Anahuac 0115.
Marker associationsAlt2 was mapped to a 4 cM interval flanked by Xpsr914-4D and Xpsr1051-4D 848; on a consensus 4B-4D map of T. aestivum; ALT2 – 1.1 cM – Xbcd1230-4D 1213; Alt2 cosegregated with Xbcd12304D and fell within the interval Xgdm125-4D – 4.8 cM – ALT21.1 cMXpsr914-4D 248. Malate transporter AlMT-D1 gene (GenBank AB081803) is completely linked to aluminium tolerance in chromosome arm 4DL between SSR markers Xwmc48b and Xwmc331 in a similar region to ALT2 10285. Almt1 transgenic expression in barley conferred and Al-activated efflux of malate with properties similar to those of Al -tolerant wheat 10286. Allelic variation at the promoter of Almt-D1 was 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.
Almt1 . TaALMT1 11242. 4DL 11242. 4DL 11242. QTL Atlas 66 / Century: A QTL in the region Xdgm125-4DLXwmc331-4DL 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 (R[2] = 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 ( Xbarc164-3BXbarc344-3B ) and Qalt.pser-2A ( Xgwm515-2AXgwm296-2A ) 10605. In D genome introgression lines of Chinese Spring a major QTL was located in the interval Xgwm125-4DXgwm976-4D , R[2] =0.31 10598, probably coinciding with Alt2 . A second QTL from CS, Qalt csipk-3B, R[2] =0.49, occurred in interval Xgwm1029-3BLXgwm1005-3BL in a CS / CS (Synthetic 3B) population 10598.

1.9. Anthocyanin pigmentation⌂ Home

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, PlbD1 ), 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.

PAN

Pan1
Chromosome7DS
VarietiesIlyitchevka 921; Mironovskaya 808 921; Novosibirskaya 67 921; Pyrothrix 28 921; Saratovskaya 210 921; Strela 921; Ukrainka 921.
Type varietiesT. polonicum 921.
Pan2
Chromosome7AS
Type varietiesT. turgidum ssp. dicoccoides acc. MG4343 9959.
Marker associationsPAN2 – 9.2 cM – RC1 – 12.2 cM – Xutv1267-7A (proximal) 9959.
An5
Chromosome5R
Pc/Pls/Plb
Chromosome7B
Type varietiesTRI 15744 (IPK GeneBank, Gatersleben) 10692.
Marker associationsXgwm951-7B – 6.7 cM – Pc/Pls/Plb – 8.2 cM – Pp1 – 8.9 cM – Xgwm753-7B 10692.
Ra1
SynonymRa 1645
1D Gulyeeva {474, 983}. 2D 1645.
Ra2
Chromosome4B
Ra3
Chromosome6B

RC1

RC-A1
SynonymRc
Chromosome7AS
dvPAU14087 10451.
Marker associationsXcfa2174-7AS – 11.1 cM – RCA1 – 4.3 cM – Xgwm573-7A/Xwmc17-7AL 10451.
Rc-A1a
SynonymRc1 , R 401
Chromosome7A
Chromosome7AS
Sources / synonymsCS[*] 6/Hope 7A 1293.
VarietiesHope Rc-B1a 1293.
Type varietiesT. turgidum ssp. dicoccoides acc. MG4343 9959.
Marker associationsPAN2 – 9.2 cM – RC-A1 – 12.2 cM – Xutv1267-7ª (proximal) 9959; RC-A1 (distal) – 11.9 cM – Xgwm913-7A 0250.

RC-B1

Rc-B1a
SynonymRc2 , R2 401
Chromosome7BS
Chromosome7B
Sources / synonymsCS[*] 6/Hope 7B 769.
VarietiesHope Rc-A1 .
Marker associationsXgwm263-7B – 26.1 cM – RC-B1 – 11.0 cM – Xgwm1184-7B 0250.

RC-D1

Rc-D1a
SynonymRc 3
Chromosome7D
Chromosome7DS
VarietiesGaoyuan 115 11160; Mironovskaya 808 1444; Tetra Canthatch/ Ae. squarrosa var. strangulata RL 5271, RL 5404 1240; Tetra Canthatch/ Ae. squarrosa var. meyeri RL 5289, RL 5406 1240; Sears' T. dicoccoides /Ae. squarrosa = Sears' Synthetic 596.
Marker associationsRC-D1 (distal) – 3 cM – Xpsr108-7D 180; Xgwm44-7D – 6.4 cM – RCD1 – 13.7 cM – Xgwm111-7D 0250.
cTaMYB-D1 isolated from Gaoyuan 115 was proposed as the candidate gene 11160.
Tahir & Tsunewaki 1453 reported that T. spelta var. duhamelianum carries genes promoting pigmentation on chromosomes 7A and 7D and genes suppressing pigmentation on 2A, 2B, 2D, 3B and 6A. Sutka 1444 reported a fourth factor in chromosome 6B and suppressors in 2A, 2B, 2D, 4B and 6A. The Rc allele appears to encode a transcription activator of late biosynthesis genes involved in the lightregulation of anthocyanin systhesis (studies carried out on CS(Hope 7A) substitution line) 10317.

PC1

PC1
SynonymPc 743
Chromosome7BS
Chromosome7B
Sources / synonymsCS[*] 6/Hope 7B {743, 768}.
itvLD222[*] 11/CS (Hope 7B) 1546.
Marker associationsPc (proximal) – 5.7 cM – Xpsr490(Ss1)-7B .

PC2

Pc2
Chromosome7DS
VarietiesIlyitchevka 921; Mironovskaya 808 921; Novosibirskaya 67 921; Pyrothrix 28 921; Saratovskaya 210 921; Strela 921; Ukrainka 921.
Pc/Pls/Plb
Chromosome7B
Type varietiesTRI 15744 (IPK GeneBank, Gatersleben) 10692.
Marker associationsXgwm951-7B – 6.7 cM – PC/PLS/PLB – 8.2 cM – PP1 – 8.9 cM – Xgwm753-7B 10692.

PP1

Pp1
Chromosome7BL
Chromosome6A
iSaratovskaya 29[*] 8/Purple {Australia} Pp2 40.
VarietiesNovosibirsk 67 (this cultivar has white pericarp) 10392.
Varieties (alt.)Purple K49426 Pp3a 10392; Purple Feed Pp3b 10392.
Marker associationsXgwm983-7B – 15.2 cM – PP1 – 11.3 cM – Xgwm767-7B 10392.

PP2

Pp2
Chromosome7A
Type varietiesT. durum Desf. subsp. abyssinicum Vav 40. Piech and Evans 1138 located complementary genes on chromosomes 3A and 7B. Pp2 was renamed Pp3b .

PP3

Pp3
Note2A, not
Chromosome6A
Pp3a
Varieties (alt.)Purple K49426 Pp1 10392.
Marker associationsXgwm328-2AS – 2.7 cM – PP3a – 3.2 cM – Xgwm817-2AL 10392.
Pp3b
SynonymPp2
Varieties (alt.)Purple Feed 10392, 66.
Marker associationsXgwm328-2AS – 5.2 cM – PP3b/Xgwm817/Xgwm912-2A – 3.6 cM – Xgwm445-2A 10392.
pp1pp3
VarietiesSaratovskaya 29 (this cultivar has red pericarp) 10329.
A set of Saratovskaya 29 NILs is described in 11136. A homoeologous sysyem was later proposed:

PG

Pg
Chromosome2A
Type varietiesTRI 15744 (IPK GeneBank, Gatersleben) 10692.
Marker associationsXgwm328-2A – 19.2 cM – PG – 1.4 cM – PP3 – 5.1 cM – Xgwm817-2A 10692.

PLB

Plb
Chromosome7B
Type varietiesTRI 15744 (IPK GeneBank, Gatersleben) 10692.
Marker associationsXgwm951-7B – 6.7 cM – PC/PLS/PLB – 8.2 cM – PP1 – 8.9 cM – Xgwm753-7B 10692.
Transcription factor TaMYB3 on chromosome 4BL bin 0.62-0.95 isolated from purple grained cv. Gy115 appeared to be involved in purple pericarp color, but was not the candidate gene for purple grain color 11285. This may correspond to TaPPM2 located on chromosome 4BL, one of three PPM genes with no effect on purple pericarp 11313.

1.10. Awnedness⌂ Home

In cereals such as barley and rice awns are conferred by dominant genes. No homologous genes have been reported in wheat.

hd b1 b2
Bearded or fully awned genotype

HD

Hd
Chromosome4AS
iS-615[*] 11/CS 1500.
VarietiesChinese Spring B2 1293.
Marker associationsXcdo1387-4A – 8.2 cM – HD – 7.2 cM – Xpsr163-4A 0047 was mapped as a QTL with a peak on Xfba78-4A 0309.
hd
Sources / synonymsCS[*] 6/Hope 4A; CS[*] 5/Thatcher 4A; CS[*] 6/Timstein 4A.

B1

B1
NoteALI-1 11581.
Chromosome5AL
iS-615[*] 11/Jones Fife 1500.
VarietiesTimstein 741; Redman 160; WAWHT2046 10040.
Type varietiesLD222 10541. Glossy Huguenot 11570.
Marker associationsXgwm410.2-5A – 8.2 cM – B1 – 12.2 cM – YR34 10040; Terminally located 10189; Xgwm291-5A.3 – 5.3 cM – B1 10330.
matvXgwm291-5A – 8.0 cM – B1 10541.
cFunctionally confirmed by transforming awned Kennong 199 with a 2,017 bp fragment containing TraesCS5A02G542800 and UTRs from YMZ 11581.
B1 was mapped as a QTL with a peak on Xwmc182-6B 0309. Associated with increased number of spikelets per spike and decreased kernel size in a survey of global bread wheat germplasm 11571. Associated with lower grain length and 1000 grain weight 11581.
B1a
Sources / synonymsSaratovskaya 29[*] 8/Festiguay 5A 42.
B1b
Sources / synonymsSaratovskaya 29[*] 8/Aurora 5A 42.
B1c
Sources / synonymsSaratovskaya 29[*] 8/Mironskaya 808 5A 42.
b1 . ali-1 11581.

B2

B2
Chromosome6BL
iS-615[*] 11/CS 1500.
VarietiesChinese Spring Hd 1293.
b2
Sources / synonymsCS[*] 6/Hope 6B; CS[*] 5/Thatcher 6B; CS[*] 9/Timstein 6B.
A GWAS of 364 wheat accessions identified 26 loci associated with awn length 11581.

1.12. Blue Aleurone⌂ Home

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.

BA1

Ba1
NoteDerived from Elytrigia pontica (2n=70).
SynonymBa 643
Chromosome4B
trUC66049B (currently UC66049, PI 633834) 643.

BA2

Ba2
SynonymBa 10451
4A[m] L.

1.13. Brittle culm⌂ Home

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.

BRC1

brc1
Chromosome6AS
dvT. monococcum mutant brc3 11002.
Marker associationsXbarc37-6A – 1.9 cM – BRC1 – 10.3 cM – Xbarc113-6A 11002.

BRC2

brc2
Chromosome3AL
dvT. monococcum mutant brc2 11002.
Marker associationsXcfa2170-3A – 2.9 cM – BRC2 – 0.8 cM – Xcfd62-3A 11002.

BRC3

brc3
Chromosome1AL
dvT. monococcum mutant brc1 11002.
Marker associationsXwmc470-1A – 3.9 cM – Brc3 – 2.1 cM – Xgwm135-1A 11002.
A further recessive mutation in T. monococcum accession Pau 5088 was named brc5 (11505).

1.14. Brittle rachis⌂ Home

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 11080.

BR-A1

Br-A1
SynonymBr2 10061, Br-A2 10280
Chromosome3A
Chromosome3AS
sutvLDN(DIC 3A) 0130.
itvANW10A=LD222*7/LDN-DIC DS 3A 10242.
Marker associationsXgwm2-3A – 3 cM – BR-A1 – 8 cM – Xgwm666-3A.1/Xbarc356-3A/Xbarc19-3A/Xgwm674-3A/Xcfa2164-3A 10280.

BR-B1

Br-B1
SynonymBr3 0130, Br-A3 10280
Chromosome3BS
Chromosome3B
sutvLDN(DIC 3B) 0130.
itvANW10B=LD222*7/LDN-DIC DS 3B 10242.
Marker associationsXbarc218-3B – 22 cM – BR-B1 – 2 cM – Xwmc777-3B 10280.
Type varietiesSenatore Cappelli PI 342646 10242; Sammartinara 10242; others 10242.
The presence of Br-B1 in some durums apparently does not lead to significant shattering under conditions of Mediterranean agriculture 10242.

BR-D1

Br-D1
SynonymBr1 9970, Br[61] 10362
Chromosome3DS
VarietiesKU510, KU511, KU515 10061; R-61 10362; T. aestivum var. tibetanum 9970.
dvAe. tauschii KU2126 10227.
Marker associationsIn Ae. tauschii: Br[t] – 7 cM – Xgdm72-3D 10227.
Evidence for an orthologous series extending to many related species is discussed in 0130 and 10061.

BR-S1

Br-S1
Chromosome3SS
VarietiesIranian spelts 11080.
tv2Triticum timopheevii 11080.
dvAegilops tauschii 11080.
Marker associationsXpsr1196-3S – 32.3 cM – BR-S1 – 1.5 cM – Xabg471-3D 11080.

Br-D2

Br-D2
Chromosome3DL
VarietiesCommon wheat 11080; European spelts 11080.
dvAL8/78 (shattering) 11080; TA1604 (non-shattering) 11080.
Marker associationsXmwg2013-3D – 1.5 cM – BR-D2 – 2.9 cM – Xpsr170-3D 11080.

BR4

Br4
Chromosome2A
Type varietiesT. dicoccoides 10082.
Marker associations33 cM distal to Xgwm294-2A (LOD=6.3, R[2] =14.4%) 10082.

1.15. Boron tolerance⌂ Home

Genes controlling tolerance to high concentrations of soil boron act additively.

BO1

Bo1
SynonymBot-B5b 11432
Chromosome7BL
Chromosome7B
VarietiesCarnamah 10460; Frame 10460; Krichauff 10460; Yitpi 10460; Additional genotypes {10833, 10834}.
Varieties (alt.)Halberd Bo2Bo3 .
Type varietiesKalka 10834; Linzhi 10834; Niloticum 10834; Additional genotypes 10834.
Marker associationsBo1 co-segregated with several STS-PCR markers, including Xaww11-7BL , falling within a 1.8 cM interval 10460; The AWW5L7 ( Xaww11 ) PCR marker allele was a good predictor of boron tolerance 10460; Co-dominant PCR marker AWW5L7 co-segregated with Bo1 and was predictive of the response of 94 Australian wheat genotypes 10833; Xbarc32-7B – 2.4 cM – Xaww5L7 – 1.2 cM – Xbarc182-7B/BO1 – 1.2 cM – Xpsr680/Xmwg2062-7B 10833; Xbarc32-7B – 2.6 cM – Xaww5L7/BO1 10834.
cBoron transporter-like gene 11432. GenBank KF148625. Bo1 functions as a boron transporter 11432.

BO2

Bo2
Varieties (alt.)(W1[*] MMC)/Warigal Bo3 . Halberd Bo1 Bo3 .

BO3

Bo3
Chromosome4A
Varieties (alt.)Warigal Bo2 . Halberd Bo1 Bo2 .

BO4

Bo4
SynonymBOT(TP4A-B5C) 11432
Chromosome4AL
VarietiesG61450 11431.
cBo4 is a dispersed duplication of Bo1 11432.
A homoeologous nomenclature based on annotated boron transporters inferred from the IWGS CS sequence is provided in Extended Data Table 1 in 11432. Very sensitive genotype: Kenya Farmer bo1 bo2 bo3 . 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. QTL Cranbrook (moderately tolerant) / Halberd (tolerant): DH population; QTLs for tolerance were identified on chromosomes 7B and 7D 10832. For a review of boron tolerance in wheat, see 10835.

1.16. Cadmium Uptake⌂ Home

Low uptake is dominant.

CDU1

Cdu1
SynonymCdu 1128
Corrected to 5BL 10894. 5BL 10104.
cdu1
Synonymcdu 1 1128
itvKofa 10104.
Type varietiesKyle 1128; DT369 10894.

1.17. Chlorophyll abnormalities⌂ Home

V1

V1
Chromosome3BS
3B 122, 1294, 1311.
v1a
Synonymv 1294
iS-615[*] 11/Neatby's Virescent 1500.
Sources / synonymsCS[*] 9/Neatby's Virescent 1304.
VarietiesNeatby's Virescent 1055.
v1b
iCS[*] /Hermsen's Virescent v2b 1304.
VarietiesHermsen's Virescent v2b 1311.

V2

V2
Chromosome3A
VarietiesCS.
v2a
VarietiesViridis 508 1545.
v2b
NoteExpressed only when combined with v2b .
iCS[*] /Hermsen's Virescent v1a 1304.
VarietiesHermsen's Virescent v1a 1311.
v1b and v2b are expressed only when both are present. Corresponding normal alleles are designated V1 {3B} and V2 {3A} following Sears' 1295 demonstration of their effects on the expression of v1a .

CN1

CN-A1
Chromosome7AL
1132.
cn-A1a
Synonymcn1a
iANK-32 10820; Chlorina-1 1311.
cn-A1b
Synonymcn1b
iCornell Wheat Selection 507aB-2B-21/6[*] CS 1133.
cn-A1c
Synonymcn2
iChlorina-448 (CS background) 1545.
cn-A1d
Type varietiesCDd6 {665, 666}.
itvANW5A-7A 10820.
Marker associationsHexaploid wheat: Xhbg234-7A – 8.0 cM – CN-A1 – 4.3 cM – Xgwm282/Xgwm332-7A 10820; Tetraploid wheat: Xbarc192-7A – 19.5 cM – CN-A1 – 11.4 cM – Xgwm63-7A 10820; Diploid wheat: Xgwm748-7A – 29.2 cM – CN-A1 – 33.3 cM – Xhbg412-7A 10820.
CN-B1
Chromosome7BL
VarietiesChinese Spring 1131.
cn-B1a
Type varietiesCDd1 {665, 666};
VarietiesCBC-CDd1 665.
cn-B1b
Type varietiesCDd2 {665, 666}.
CN-D1
SynonymCn3
Chromosome7D
Chromosome7DL
VarietiesChinese Spring 1131.
cn-D1a
Synonymcn-D1, cn3
iChlorina-214 1545.
VarietiesCD3 1583.
Two mutants in diploid wheat are reported in 10820.

1.18. Cleistogamous flowering⌂ Home

Cleisogamy in barley is controlled by the Chy1 allele that 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.

CL

clcl
Type varietiesHI8332 191; WH880 191. Chasmogamous genotypes
ClCl
Type varietiesIWP5308 191; PWB34 191; WH872 191.

1.19. Copper efficiency⌂ Home

Copper efficiency is a genetic attribute that enhances plant growth in copper deficient soil.

CE

Ce
Note4BL = T
Chromosome4BL
Chromosome5RL
VarietiesCornell Selection 82a1-2-4-7 462; Backcross derivatives of Cornell Selection to Oxley, Timgalen, Warigal 464; Hairy necked Viking 1276. 5BS = T5BS.5RL.
adCS+5R 463.
suCS 5R {5D} 463.
VarietiesSears' stock HN-2 464; Backcross derivatives to Warigal and Timgalen 464.

1.20. Corroded⌂ Home

CO1

co1
Synonymco 1297
wsl 11535. 6BS 1293.

CO2

co2
Chromosome6D
VarietiesKurrachee 1570; Shannong 33 Mutant I30 11534.
Marker associationsXcfd190-6DCO2 – 9.1 cM – 6DS-5 11534.
A gene(s) in chromosome 6A acted as an inhibitor of corroded {1039, 1570}. Corroded mutants are very frequent in EMS-treated populations 939.

1.21. Crossability with Rye and Hordeum and Aegilops Spp.⌂ Home

KR1

Kr1
Chromosome5B
Chromosome5BL
Marker associationsMapped to a 2.0 cM region flanked by Xw5145-5B and CA1500122/Xw9340-5B 10922.
A second gene in 5BL distal to the Ph1 locus and flanked by Oshypl and Os09g36440 , but including Xgwm371-5B , affected the temperature sensitivity of seed-set in Kr1 genotypes in wide crosses 10922.

KR2

kr2
Chromosome5A
Chromosome5AL
kr1 kr2
VarietiesChinese 446 790; Chinese Spring {1025, 1216, 762}; Martonvarsari 9[*] 4/CS 1016.
KR3
Chromosome5DL
Published paper not available.
kr3
A very weak effect.

KR4

kr4
Chromosome1A

KR5

Kr1 kr2
Sources / synonymsCS[*] 6/Hope 5B {1216, 762}.
VarietiesBlausamtiger Kolben 790.
kr1 Kr2
Sources / synonymsCS[*] 6/Hope 5A 1216.
Kr1 Kr2
VarietiesMarquis 790; Peragis 790.
kr1 kr2 kr3 kr4
VarietiesJ-11 1649.
Kr1 Kr2/Kr1 kr2
Note(heterogeneous).
VarietiesMartonvarsari 9 1016. Using the Chinese Spring/Cheyenne chromosome substitution series, Sasaki & Wada 1265 found significant differences in crossability for chromosome 5B, 7D, 1D and 4B. Differences between rye lines were also reported {1265, 1458}. Allelic variation in the potency of the dominant suppressor genes was reported {1385, 343}. Evidence for allelic variation in dominant supressors is reported in 1386. Lists of wheat/rye crossabilities:{1383, 1642, 850, 858}.
QTL 65% of the variability in a Courtot/CS population was associated with Xfba-367-5A (5AS), Xwg583-5B (5BL) and Xtam51-7A 0134. Only the second QTL appeared to coincide with known locations of Kr genes. The former was skr .

1.22. Dormancy (Seed)⌂ Home

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). For a review of genes involved in preharvest sprouting see 11569.

TaSDR-1

TaSdr-A1a
cGenBank KF021988 11119.
This allele is associated with lower germination index
TaSdr-A1b
cGenBank KF021989 11119.
This allele is associated with higher germination index TaSDR-B1 . [ TaSdr-B1 11119]. 2B11119.
TaSdr-B1a
VarietiesYangxiaomai 11119.
cGenBank KF021990 11119. This allele is associated with lower germination index.
TaSdr-B1b
VarietiesZhongyou 9507 11119.
cGenBank KF021991 11119. This allele was associated with higher germination index.

VP-1

VP-A1
Chromosome3AL
Vp-A1a
VarietiesNongda 311 10919.
c599 bp 10919. Higher germination index.
Vp-A1b
VarietiesWanxianbaimaizi 10919; Yannong 15 10919.
c596 bp 10919. Lower germination index.
Vp-A1c
VarietiesJing 411 10919.
c593 bp 10919. Higher germination index.
Vp-A1d
VarietiesXiaoyan 6 10919.
c590 bp 10919. Lower germination index.
Vp-A1e
VarietiesZhengzhou 6 10919; Bainong 64 10919.
c581 bp 10919. Higher germination index.
Vp-A1f
VarietiesYumai 34 10919.
c545 bp 10919. Higher germination index.
Vp-A1g
SynonymVp-1Ab 11047
VarietiesKalyansona 11047; Sonalika 11047; Yaqui 50 11047; Yecora Rojo 76 11047.
cGenBank GU385899 11047.
Vp-A1h
SynonymVp-1Ad 11047
VarietiesAttila 11047; Glenlea 11047; Tanori F71 11047.
cGenBank GU385901 11047.
Vp-A1i
SynonymVp-1Af 11047
VarietiesDebeira 11047; Kancahn 11047; Rayon F89 11047.
cGenBank GU385903 11047.
VP-B1
Chromosome3B
Vp-B1a
VarietiesCharger 10616; Zhongyou 9507 10615; 271 accessions 10616.
cGenBank AJ400713 10615.
Vp-B1b
VarietiesAltria 10616; Recital 10616; Yongchuanbaimai 10615; 2 accessions 10616.
c193 bp insertion in third intron relative to Vp-A1a .
Vp-B1c
VarietiesScipion 10616; Xinong 979 10615; 101 others 10616.
c83 bp deletion relative to Vp-B1a .
Vp-B1d
c25 bp deletion relative to Vp-A1a .
Vp-B1e
SynonymVp-1Be 10998
VarietiesFulingkemai 10999; Hongheshangtou 10621; Hongmangchum 10998; Wangshuibai 10999.
c83 bp deletion, 4 bp insertion and 2 SNPs relative to Vp-B1a 10621.
Vp-B1f
SynonymVp-1Bf 10998
VarietiesWanxanbaimaizi 10998.
Vp-B1g
SynonymVp-1Bg 11047
VarietiesHD2939 11047; Pavon 76 11047; Sonora 64 11047.
cGenBank GU385904 11047.
Vp-B1 allelic identifications for Chinese landraces, historical and current wheat cultivars are listed in 10621.
VP-D1
Chromosome3DL
cAJ400714 10919.
Vp-D1a
Chromosome3DL
Varieties81 Chinese wheat cultivars 10919.
c5 pairs of primers10919. There was a suggestion of a relationship between alleles and PHS response 10615.
Phs1
NoteSemi-dominant 9960. [ Phs 9960; Phs1-4AL ; Phs-A1 11546; TaMKK-A 11546.
Chromosome4AL
iHaruyokoi*6/Leader 10500; Haruyokoi*6/Os21-5 10500.
VarietiesChinese Spring 11737; Leader 10500; Os21-5 10500; Soleil 9960.
Marker associationsAssociated with Xpsr1327-4A 10346; Xhbe03-4AL – 0.5 cM – Phs1 – 2.1 cM – Xbarc170-4AL 10500.
phs1
VarietiesHaruyokoi 10500.
TaOsd1
CRISPR-Cas 9 knockout of TaQsd1 homoeologues in chromosomes 5A, 5B and 5D led to a 3-4-fold increase in dormancy in Fielder wheat 11738. Qsd1 encoding an alanine amino transferase controls dormancy in barley. Genotypes with and without favourable haplotypes are discussed in 11546. According to 11547 red grain colour increases the time to dormancy release and has a cumulative effect when combined with other dormancy genes not associated with grain colour. For a review of Preharvest Sprouting see 11595. 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 Xpsr9455A , Xpsr8(Cxp3)-6A and Xpsr563-6A , and Xpsr350-7B and Xbzh232(Tha)-7B 0169. AC Domain / Haruyutaka : one major QTL in chromosome 4AL and two lesser possibly homomeologous QTLs for dormancy in 4BL and 4DL 0226. 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, R[2] =0.44, was independent of seed colour 10626. Annong 0711 (res) / Henong 825 (sus) : RIL population. Differences in germination index were attributed to a 33 bp insertion in the promoter (possibly the AP2 binding site) of TaMFT-3A in Annong 0711 relative to Henong 825. TaMFt-3A encodes a phosphatidyl ethanolamine-binding protein 11410. 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. CN10955 (PHS resistant white seeded) / Annuello (PHS susceptible, white seeded) : F8 RIL population: QPhs.dpivic-4A.2 in the Xgwm637-4ASXgwm937/Xgwm894-4AL region and QPhs.dpivic4A.1 in the Xwmc48-4ASXgwm397-4AS region 10599. SPR8198 (red seeded, PHS tolerant) / HD2329 (white seeded, PHS susceptible) : Tolerance to preharvest sprouting (PHS) was associated with Xwmc104-6B and Xmst101-7D 0032. In a further study 7 QTL were located on chromosomes 2AL, 2DL, 3AL and 3BL, the most important, on 2AL and 3AL 10670. SPR 8198 (dormant) / HD2329 : QPhs.occsu-3A was located in the Xgwm155-3AXwmc153-3A region with R[2 ] = 75% across 6 environments 10261. Renan / Recital : 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 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. 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. Rio Blanco (white seeded, PHS resistant) / NW97S186 (white seeded, PHS susceptible): RIL population: QPhs.pseru-3AS , R[2] =0.41, Xgwm369-3AXbarc12-3A , and one minor QTL 10634. This major QTL was confirmed in a Blanco / NW98S079 RIL population, R[2] up to 0.58 10634. Qphs.pseru3A ( TaMFT- 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. Sun325B (dormant white seeded) / QT7475 (semi-dormant white seeded) : both parents with the chromosome 4A QTL: DH population: A QTL was located in the Xgwm77-3BXwmc527-3B interval (R[2 ] = 0.19) in the approximate region of the R-B1 locus 10669. Totoumai (res) / Siyong (sus) : RIL population: A QTL on chromosome 4AL was delimited to a 2.9 cM interval flanked by GBS109947 and GBS212432; nine and two SNP were associated with minor QTL on chromosomes 5A and 5B 11408, respectively. 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. 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. 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-4A.1 , may be the same as a QTL in AC Domain/Haruyutaka due to tight linkage with Xcdo7854A 10245. QPhs.ocs.4B.1 , a CS allele contributing to dormancy, was located in the region of Xgwm495-4B 10245. QPhs.sicau-3B.1 , distally located on chromosome 3B in T. spelta CSSR6 (res) / Lang (sus) ; nearest marker wPt-6157 ; transferred to durum cv. Bellaroi using SCAR markers 11246. 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. Association mapping of 198 winter wheat genotypes detected 8 QTLs on 7 chromosomes, viz. 1BS, 2BS, 2BL, 2DL, 4AL, 6DL, 7BS and 7DS 10959. A GWAS identified 12 QTL for PBS resistance among which those on chromosomes 3AS and 4AL were most commonly detected 11409. Diploid wheat QTL T. monococcum KT3-5 (non-dormant) / T. boeoticum KT1-1 (dormant) : RIL population: QTL on chromosome 5A[m] L, Xcdo1236c-5AXabc302-5A ), R[2 ] = 0.2-0.27. Weaker QTLs were found on 3A[m] ( TmAB18Xwmc102-3A and Xrz444-3ATmABF ) and 4A[m ] ( Xrz261-4AXrz141-4A ) 0892. The 3A QTL 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. For a review of Preharvest sprouting see 11595.

1.23. Ear emergence⌂ Home

QEet.ocs-4A.1
Chromosome4AL
VarietiesCS/CS(Kanto107 4A) mapping population.
Marker associationsAssociated with Wx-B1 0047.
QEet.ocs-5A.1
Chromosome5AL
VarietiesCS( T. spelta 5A)/CS(Cappelle-Desprez 5A) RI mapping population 9903.
Marker associationsAssociated with Xcdo584-5A and morphological locus Q 0068.
QEet.ocs-5A.2
Chromosome5AL
Marker associationsXcdo 412-5AXbcd9-5A region 0026.
QEet.inra-2B
Chromosome2B
Marker associations2B linked to Xgwm148 (LOD=5.7, R[2 ] = 11.9%.
QEet.inra-2D
Chromosome2D
Marker associations2D linked to XksuE3 (LOD=2.7, R[2 ] = 6.5%).
QEet.inra-7D
Chromosome7D
Marker associations7D linked to Pch1 (LOD=3.9, R[2 ] = 7.3%).
QEet.ipk-2D
NoteQEet.ipk-2D coincides with a QTL for flowering time, QFlt.ipk-2D . Both QTLs may correspond to Ppd-D1 0255.
Chromosome2DS
VarietiesOpata/W-7984 (ITMI) RI mapping population0255; Lateness was contributed by W-79840255.
Marker associationsAssociated with Xfba400-2D and Xcdo1379-2D 0255.
QEet.ipk-5D
NoteQEet.ipk-5D coincides with a QTL for flowering time, QFlt.ipk-5D . Both QTLs probably correspond to Vrn-D1 0255.
Chromosome5DL
VarietiesOpata/W-7984 (ITMI) RI mapping population 0255; Lateness was contributed by W-7984 0255.
Marker associationsAssociated with Xbcd450-5D 0255.

1.24. Earliness per se⌂ Home

Genes for earliness per se 0023 affect aspects of developmental rate that are independent of responses to vernalization and photoperiod.

EPS-A1

Eps-A1a
Chromosome3AL
Chromosome3A
VarietiesChinese Spring 0024.
Eps-A1b
VarietiesTimstein 0024.
Eps-1A[m ] 0364. [ Eps-A[m] 1]. 1AL 0364.

EPS-B1

Eps-5BL.1
Chromosome5BL
Marker associationsQTL mapped on chromosome 5BL, linked to Xwmc73-5B (this QTL explained 8% of the variance in flowering time, P <0.03 10075.
Eps-5BL.2
Chromosome5BL
Marker associationsQTL mapped on chromosome 5BL, linked to Xgwm499-5B (this QTL explained 6% of the variance in flowering time) 10075.

EPS-D1

Eps-D1
Chromosome1DL
VarietiesEarliness allele: Cadenza and Spark 11193; Lateness allele: Avalon and Rialto 11193.
Marker associationsThe earliness allele was associated with a subtelomeric deletion containing three candidate genes one of which was TaELF-D1 11193.
A QTL for heading date co-segregated with TaELF3-1DL in a RIL population derived from Gaocheng 8901 / Zhoumai 16 11194; a deletion of the Eps-1D region was associated with earlier flowering.
epsCnn
VarietiesCheyenne0025.
Eps Wi
Chromosome3A
su: Cheyenne[*] 7 / Wichita 3A 0025. QTL CS / T. spelta var. duhamelianum KT19-1 : RIL population: Two QTL for narrow-sense earliness were detected on chromosome 2B in a 10057. These QTLs were associated with markers Xpsr135-2B and Xabc451-2B 10057. For both QTLs, earliness was conferred by the CS allele. Courtot / CS: 0132. Cutler / AC Barrie : Three QTLs were mapped on chromosomes 1B ( QEps.dms-1B.1 and QEps.dms1B.2 ) and 5B QEps.dms5B ) 11039. Grandin / BR34 : QEet.fcu.5AL identified in Xfcp359-5AXfcp231-5A interval (R[2 ] = 0.38), at or near the Q locus in 10256. Grandin was the earlier parent.

1.27. Flowering time⌂ Home

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 originated 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. The wheat TaHd1 and rice Hd1 genes were located in non-syntenic locations 10054. To date, no variation for flowering time has been identified on wheat group 6 chromosomes.

QFlt.ipk-3A
Chromosome3AL
VarietiesOpata/W-7984 (ITMI): RI mapping population0255; Lateness was contributed by W-7984 0255.
Marker associationsAssociated with Xbcd451-3A 255.
QFt.cri-3B.1
NoteNearest marker Xbarc164-3B ; identified in croses of substitution lines of Ceska Presivka and Zlatka or Sandra 10567. Arina (149 days) / Forno (150 days) : winter wheat cross: Six QTL were detected over six environments. The 3 most important, all from Arina, were in chromosomes 6DL (R[2] =16%), 3DL (R[2] =14%) and 7BL (R[2] =13%); 3 others in 2AL, 5BL and 6DL were from Forno 10172. CI 13227 / Suwon 92 : RIL population: Heading date: AFLP marker – 2.6 cM – QHd.pser-2DS – 121.1 cM – Xgwm261-2D 10269. This QTL could be Ppd-D1 10269. Ernie (early) / MO94-317 (late) : winter wheat cross: days to anthesis (dta): Qdta.umc-2D , linked to Xbarc95-2D , R[2] = 0.74 10456. Karl 92*2 / TA 4152-4 : F2:F4 population: Two QTLs, QHd.ksu-2D , associated with Xgwm261-2D (R[2] =0.17), and QHd.ksu-3D , associated with Xgwm161-2D 9 (R[2] ) 10273. Nanda 2419 / Wangshuibai : spring wheat cross: 7 QTL for flowering time identified with earlier alleles for five coming from Nanda 2419: QFlt.nau-1B (closest marker Xbarc80-1B , R[2] =11 %), QFlt.nau-1D ( Xbarc62-1D, Xgwm232-1D , R[2] =6.13 %), QFlt.nau-2B ( Xwmc35-2B , R[2] =10 %), XFlt.nau-2D ( Xwmc601-2D , R[2] =10 %), _XFlt.nau-
Chromosome4A
1 ( Xcfd2-4A, Xmag1353-4A , R[2] =10 %), XFlt.nau-4A.2 ( Xmag3386-4A, Xwmc161-4A , R[2] =18-19 %), XFlt.nau7B (Xmag2110-7B, Xmag1231-7B, Xgwm537-7B, Xwmc218-7B_ , R[2] =18 %) 10566.

1.28. Flour colour⌂ Home

Schomburgk/Yarralinka: RIL population: 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 9936. 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

LUTE

Lute
NoteHigh lutein ester.
Chromosome7DS
Chromosome bin7DS4-0.61-1.00.
VarietiesIndis 11189; Sunco*2/Indus Der. DM5685*B12 11189; Most bread wheat accessions.
Marker associationsXwmc438-7D – 15.1 cM – LUTE/XwPt-116/XwPt-3727 – 17.7 cM – Xbarc154-7 11189. Sunco is low lutein but high ester, whereas Haruhikari is low lutein and zero ester.
lute
NoteLow lutein ester.
VarietiesHaruhikari 11189.
Lutein esters were not detected in durum 11189.

1.29. Free-threshing habit⌂ Home

QFt.mgb-5A
Chromosome5AL
Type varietiesMessapia/ T. dicoccoides MG4343 mapping population 0046.
Marker associationsAssociated with XksuG44-5A 0046.
QFt.mgb-6A
Chromosome6A
Type varietiesMessapia/ T. dicoccoides MG4343 mapping population 0046.
Marker associationsAssociated with Xpsr312-6A 0046.

1.30. Frost resistance⌂ Home

FR-1

Fr1
Chromosome5AL
VarietiesHobbit1446.
Marker associationsMapped to the mid-region of 5AL, 2.1 cM distal from Xcdo504-5A and Xwg644-5A and proximal to Xpsr426-5A 419; Mapped 2 cM proximal to Xwg644-5A and VRN-A1 0291; and flanked by deletion points 0.67 and 0.68 0292.
Fr2
Chromosome5DL
Sources / synonymsCS[*] 7/Cheyenne 5D 0291.
Marker associationsFR2 mapped 10 cM proximal to VRND1 0291.
Studies using induced and natural mutants of VRN-1 suggested that differences in frost tolerance previously attributed to FR1 were pleiotropic effects of VRN-1 10708.

FR-2

Fr-A2
dvTriticum monococcum . Frost tolerant parent G3116, frost susceptible parent DV92.
Marker associationsThe QTL mapped on chromosome 5AL had a LOD score of 9 and explained 49% of the variation in frost tolerance. Closest markers: Xbcd508-5A and Xucw90(Cbf3)-5A. These markers are 30 cM proximal to Xwg644-5A , which is closely linked to frost tolerance locus FR-1 . Eleven different Cbf transcription factors were identified at the Fr-A2 locus 10302; QTLs for frost tolerance in the FR-A2 region were also identified in wheat chromosome 5B ( FR-B2 10079) and in barley chromosome 5H ( FR-H2 10083.
Fr-B2
SynonymFr-B1 10075
Marker associationsQTL mapped on chromosome 5BL, linked to Xgwm639-5B (this QTL explained 12-31% of the variance in frost tolerance) 10075. Xgwm639-5B mapped close to Xmwg9145B , and to Xbcd508-5B , a marker located at the peak of the Fr-A2 QTL 10075. This data suggests that this locus is more likely orthologous to FR-2 than to FR-1 .
QWin.ipk-6A
Chromosome6AS
VarietiesOpata/W-7984 (ITMI) RI mapping population 0255. Winter hardiness was contributed by W-7984 0255.
Marker associationsAssociated with Xfba85-6A and Xpsr10(Gli-2)-6A 0255.
Responses to cold exposure and their genetics are reviewed in {0020, 0274}. 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 (R[2] =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.

1.31. Gametocidal genes and segregation distortion⌂ Home

GC1-B1

Gc1-B1a
SynonymGc1a 1490, Gc1 1487
Chromosome2B
iCS[*] 8/ Aegilops speltoides subsp. aucheri 1487.
Gc1-B1b
SynonymGc1b 1490
Chromosome2B
i: C S[*] 8/ Ae. speltoides subsp. ligustica 1490.

GC1-C1

Gc1-C1
Chromosome2CL
adCS/2C 0189.
suCS2C(2A), CS2C(2B), CS2C(2D) 0189.
GC1-S[l]
Gc1-Sl1
SynonymGc-S[1] 3 1485
2S[1] 334.

GC2-S[l] 1

Gc2-S[l] 1a
SynonymGc-S[1] 1 1485
Chromosome4S
adCS/ Ae. longissima 866.
Gc2-S[l] 1b
SynonymGc-S[1] 2 1485
4S[1 ] 1013.

GC-C1

Gc3-C1
SynonymGc-C 1485
Chromosome3C
adCS/ Ae. triuncialis 338.
Gc1-B1a, Gc1-B1b and Gc1-S[1] , classified in the same functional group, are hypostatic to the genes Gc2S[1] 1a and Gc2-S[1] 1b . Gc3-C1 does not interact with the Gc genes in the other two groups. In addition to these genes, chromosomes carrying gametocidal genes occur in Ae. caudata 337, Ae. cylindrica 336 and other strains of Ae. longissima and Ae. sharonensis 335,1484. Gametocidal genes in chromosomes in the same homoeologous group have the same gametocidal action 0190. In monosomic additions of chromosomes with gametocidal effects, chromosome deletions and translocations are produced in gametes not having the gametocidal genes. This feature has been exploited to isolate genetic stocks suitable for physical mapping of wheat 0191 chromosomes, and of rye 0192 and barley {0193, 0194, 0195} chromosomes in a wheat background. Genes with gametocidal activity ( Sd1 1647 and Sd2 1161) in wheat are present in homoeologous group 7 chromosomes of Thinopyrum elongatum {653, 1647}. A segment earlier believed to be derived from Thin. distichum {889, 892} is probably the same as that from Thin. elongatum 1162. In the presence of both Sd1 and Sd2, Lr19 is transmitted preferentially in heterozygotes, the degree of distortion being determined by genetic background. In heterozygotes with the same background, and in the presence of only Sd2, Lr19 shows strong self-elimination. Based on these results, it seems likely that the Sears' translocation 7D-7Ag#7 does not carry Sd1 939. See also Pollen Killer.

IGC1

Igc1
NoteCauses suppression of the 3C chromosome gametocidal gene of Ae. triuncialis . This alien gametocidal factor also promotes chromosome breakage 1486.
Chromosome3B
VarietiesNorin 26 {1483, 1488}; Nineteen wheats listed in {1483, 1488}.
igc1
VarietiesChinese Spring1483,1488; Forty wheats are listed in 1483,1488.

SD1

Sd1
Chromosome7D
Varieties (alt.)Agatha Sd2 {1647, 1161}.
Marker associationsProximal to Lr19 and distal to Xpsr1657D 10255.

SD2

Sd2
Chromosome7BL
Varieties88M22-149 {1163, 1161}. Zhang et al. 10255 question the existence of this gene and alternatively suggested a duplication or deletion event influencing the transmission.
See also Pollen Killer

1.32. Gibberellic acid response (insensitivity)⌂ Home

GAI1

Gai1
SynonymGAI1 1246, 565
Chromosome4BS
Chromosome4B
iSee 408.
VarietiesNorin 10 Der. {407, 565}.
Marker associationsXpsr622-4B (distal) – 1.9 cM – GAI1 – 8.3 cM – Xbcd110-4B (proximal) 9959.
Type varietiesMessapia 9959.

GAI2

Gai2
SynonymGAI2 1246, 565
Chromosome4D
Chromosome4DS
iSee 408.
VarietiesMaris Hobbit 411; Norin 10 Der. 565; List in 407.

GAI3

Gai3
SynonymGAI3 1246, 565
Chromosome4B
Chromosome4BS
iSee 408.
VarietiesMinister Dwarf 413; Selection D6899 359; Tom Thumb 405; Tom Thumb Der. 565, 567. In wheats with Gai3 , the aleurone layer failed to respond to applied GA 405
Two studies involving crosses between Tom Thumb derivatives and tall parents suggested that gibberellic acid insensitivity and reduced height were controlled by one gene, i.e., Gai3 359, 413. In a third study involving a Tom Thumb derivative, recombinants were isolated, indicating separate but linked genes, i.e., Gai3 and Rht-B1c 565, 567. Further evidence was obtained for linkage between genes for gibberellic acid insensitivity and Norin 10 genes for reduced height in hexaploid 568 and durum 720 wheats. Hu & Konzak 567 reported 27% recombination between Gai1 and Rht-B1b and 10% recombination between Gai2 and Rht-D1b in hexaploid wheats involving Norin 10 and Suwon 92 derivatives. In durum derived from crosses involving Norin 10, 15% recombination was obtained between one of the genes for reduced height and gibberellic acid insensitivity {1246, 1247}. Gale & Law 403 considered Gai1 and Rht-B1b, Gai2 and Rht-D1b, Gai3/ and Rht-B1c to be pleiotropic genes.

1.33. Glaucousness (Waxiness/Glossiness)⌂ Home

The W loci are complexes of closely linked genes involved in beta-diketone synthesis.

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.

W1
iChinese Spring mono-2D/S615//10*wS615 10001.
VarietiesBethlehem 11458; Chinese Spring 1493; P86 11247.
itvLD222[*] 11/ T. turgidum var. pyramidale recognitum 1546.
Varieties (alt.)S615 W2 10001.
Type varietiesKofa+Lr19 11458; AUS2499 11458.
cW1 is a highly duplicated, variable gene
cluster containing type III polyploid synthase, hydrolase and cytochrome P450 genes and is homologous to the Cer-cqu cluster in barley 11458.
w1
NoteRecessive allele for reduced glaucousness.
Chromosome2BS
suBethlehem CASL*/ T. dicoccoides TTD140 2BS(2B) 11458.
VarietiesCS mono-4B mutant 1064; J87 11247; Mentana 1432; Salmon 1493.
Type varietiesAUS2499 11458.

W2

W2
iChinese Spring mono-2B/S615//11*w-S615 10001.
VarietiesT. compactum cv. No 44 10001.
Varieties (alt.)S615 W1 10001.
W2a
dvGlaucous forms of Ae. tauschii .
W2b
VarietiesChinese Spring - weak hypomorph recognized at increased dosage 1432.
A non-glaucous spike phenotype in line L-592, a 7S(7A) substitution line, is described in 0113.
w1w2
iw-S615 = S615*11/Salmon 10001.
VarietiesSalmon 10001; Mentana 1432; CS mono-4B mutant 1064.

W3

W3
Chromosome3DL
dvAegilops tauschii KU-2126 11457.
Marker associationsXgwm645-3DL – 8.0 cM – W3 – 8.9 cM – Xbarc42-3DL 11457.
w4.

IW

Iw1
SynonymW1[I ] 1493, I1-W 1493
iS615/Cornell 5075//10*S615 10001.
VarietiesWE74 11094; Shamrock 11090. tvsu : LDNDIC521-2B 11245. tv T. dicoccoides PI 481521 11245.
Marker associationsJIC007 – 1.47 cM – IW1 – 0.18 cM – JIC010/JIC011 11090; Co-segregation with BF474014, CJ876545 and CD927782 and flanked by BE498358 and CA499581 within a 0.96 cM interval 11094.
cIw1 encodes a long non-coding RNA (LncRNA) that putatively arose from an inverted repeat of a carboxylesterase gene (80% homology) in the W1 cluster that consists of WI-COE (non-annotated
carboxylesterase), WI-PKS ( Traes2BS9E10D26DB , polykedide synthase) and WI-CYP ( Traes2BS163390FC4, cytochrome P450-type hydroxylase) 11459. GenBank C-DNA sequence, KX823910. The IR region has >94% identity to an IR region in Ae. tauschii chromosome 2 that also produces MiRNA and a marker-based location similar to that of Iw2 11459. Xgwm614-2BIW1/Xbarc35-2B/CD893659/CD927782/BQ788707/CD938589Be498111 11245.

IW2

Iw2
SynonymIw3672 10510, I2-W 1493, IwT 11207
Chromosome2DS
Chromosome bin2DS5-0.47-1.00 10578; 2DS5-0.84-1.00.
iS615/Golden Ball Synthetic//10*S615 10001.
VarietiesGolden Ball Synthetic 10001; Synthetic hexaploid line 3672 10510; TA4152-60 11094; Vernal Synthetic 10001.
Type varietiesT. dicoccoides PI 481521 11245.
dvNon-glaucous forms of Ae. tauschii 1493.
Marker associationsIn Ae. tauschii: IW2 – 30.1 cM – Xgdm35-2DS 10227; Xbarc124-2D – 0.9 cM – IW2 – 1.4 cM – Xwe6(AL731727) 10510; Xcfd56-2D – 6 cM – IW2 – 10 cM – Xcfd51-2D 10578; Co-segregation with BF474014 and CJ876545 and flanked by CJ886319 and CJ519831 within a 4.4 cM interval 11094. Xgwm614-2BIW1/Xbarc35-2B/CD893659/CD927782/BQ788707/CD938589Be498111 11245.

IW3

Iw3
SynonymIW3 277, I3-W 277
Chromosome1BS
sutvLangdon*/ T. dicoccoides 1B 11455.
Type varietiesT. turgidum var. dicoccoides 277.
Marker associationsXWL1967/Xfcp168-1B – 0.15 cM – XWL3096 – 0.015 cM – IW3 – 0.12 cM – Xpsp3000-1B 11455.
A non-glaucous spike phenotype in line L-592. A 7S(7A) substitution line, is described in 0113. 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 glaucous sibs 10543. Although maps constructed from three tetraploid crosses suggested that w1 , W1 and Iw1[DIC] = Vir remain unresolved 10815.

WS

Ws
Chromosome1AS
Chromosome bin1AS1-0.47-1.00 10666.
VarietiesSvenno 10666.
Marker associationsBJ23702a – 3.5 cM – Tc95235 – 4.8 cM – Bla 10666.
ws
VarietiesCiccio 10666.

1.34. Glume colour and awn colour⌂ Home

RG-A1

RG-A1
SynonymRg3 923, 924, 562
1AS 923, 9906, 924, 562.
Rg-A1a
VarietiesTRI 542 10378; White glumed genotypes.
dvDV92 282; G2528 10378.
Rg-A1b
SynonymRg3
iSaratovskaya 29*3//F2 CS mono 1/Strela 924.
VarietiesCS/Strela Seln 9906; Iskra 9906; L'goskaya-47 1405; Zhnitsa {9906, 10378}.
Varieties (alt.)Milturum 553 Rg-B1b 9906; Milturum 321 Rg-B1b 9906; Strela Rg-B1b 9906, 924; Sobko & Sozinov {1405, 1406}; reported a further group of 30 international wheats which, by inference from their Gli-A1 alleles, probably carry Rg-A1b .
Marker associationsA linkage order of Rg-A1bHg – cent – Glu-A1 1405; Xgwm1223-1A / Rg-A1 / Hg – 2.2 cM – Xgwm136-1A – 4.2 cM – Xgwm33-1A 10635.
Rg-A1c
SynonymBg {1304, 282}, Bg(a)
1A 1304, 282.
Rg-A1d
SynonymBg(b)
dvG3116 282.
At the diploid level Rg-A1c (Bga) and Rg-A1d (Bgb) were determinant and caused a solid black glume and a black line at the margins of the glume, respectively 282. A single factor for black glumes was reported in diploid, tetraploid and hexaploid wheats 1347. Linkage with Hg was demonstrated at all levels of ploidy, indicating a common gene on chromosome 1A; Bg is epistatic to Rg . 1
RG-B1
SynonymRg1 , Rg
Chromosome1B
Chromosome1BS
TraesCS1B02G005200.
Rg-B1a
VarietiesTRI 542 10378; White glumed genotypes, including Chinese Spring.
dvT. turgidum ssp. dicoccoides acc. MG4343 9959.
Rg-B1b
SynonymRg1
Sources / synonymsCS*5/Red Egyptian 1B 1304.
VarietiesDiamant I 9906; Federation 41 1517; Golubka 10635; Highbury 1121; Jagger 11538; Norin 60 11538; Red Egyptian 1304; Red glume spelts 11538; T. petrapavlovsky 9906.
Varieties (alt.)Milturum 321 Rg-A1b 9906; Milturum 553 Rg-A1b 9906; Strela Rg-A1b 9906.
Type varietiesMessapia 9959; Ward 792. ma Xutv1518-1B (distal) – 7.7 cM – RG-B1 – 0.8 cM – Gli-B1 (proximal) 9959; Xgwm1078-1B – 1.5 cM – RG-B1 – 3.1 cM – Xgwm0550-B1 10378; Xutv1518-1B – (distal) – 7.7 cM – RG-B1 – 0.8 cM – GLI-B1 (proximal); Xgwm1078-1B – 4.6 cM – RG-B1 – 2.0 cM – MW1B002(Gli-B1) – 4.1 cM – Xgwm550-1B 10635.
cEncodes an R2R3-MYB transcription factor 11538. TraesJAG1B01G000800 and TraesNOR1B01G001100 in red glume Jagger and Norin 40, respectively, carried the same Rg-B1bh1_ sequence; haplotype comparisons revealed that a specific group of MYB alleles was conserved in red glume genotypes 11538.
RG-D1
SynonymRg2
Chromosome1DS
1DL 769, 1241.
Rg-D1a
VarietiesNovosibirskaya 67 10378; L301 10378; White glumed genotypes.
Rg-D1b
NoteDerived from Ae. tauschii
SynonymRg2
iSaratovskaya 29*5// T. timopheevii ssp. timoppheevii/T. tauschii 9906.
VarietiesITMI Synthetic W7984 10635; Synthetic Hexaploid-11 10218; ( Triticum turgidum ssp. dicoccoides/Ae. tauschii ) 769; ( Tetra Canthatch/Ae. tauschii var. strangulata RL 5271); RL5404 1240; ( Tetra Canthatch/Ae. tauschii var. meyeri RL5289); RL5406 {648, 1240}.
Varieties (alt.)Sears Synthetic Rg-A1c 10638.
dvAegilops squarrosa accessions.
Rg-D1c
NoteBrown or smokey-grey phenotype 729.
SynonymBrg 729
iANK-23 = Novosibirskaya 67*10/K-28535 729.
VarietiesGolubka 10378; K-28535 729; K-40579 729; T. aestivum botanical varieties cinereum, columbina and albiglaucum 10378.
Marker associationsXgwm1223-1D – 1.5 cM – RG-D1 – 13.1 cM – Xbarc152-1D 10378; Xbarc149-1D – 6.3 cM – RG-D1 – 26.5 cM – Xbarc152-1D 10378.
QTL Opata / W-7984 (ITMI) mapping population : QRg.ipk-1D was mapped in the 0255; Linkage with GLI-D1 implied RG-D1 ( Rg2 ). This QTL coincided with a QTL for awn colour, QRaw.ipk-1D 0255. ma: Xpsp2000-1D – 9.3 cM – RG-D1 – 21.2 cM – Xgwm106-1D 10128; Xgwm1223-1D – 6.6 cM – RG-D1 / Xksud14-1D – 13.9 cM – Xgwm33-D1 10635; RG-A1 – 3.9 cM – Xgwm1223-1D 10638.

PBC

Pbc
Note3B 742
Chromosome3BS
Sources / synonymsCS[*] 6/Hope 3B 742; CS[*] 6/Ciano 5B 939.

CC

Cc
Chromosome7B
Chromosome7BS
Type varietiesLangdon mutant 719; PI 349056 665.
dvCBC-CDd1 665.
The chocolate chaff phenotype was suppressed by a gene(s) in chromosome 7D 719.

BLA1

Bla1
Chromosome1AS
Chromosome bin1AS1-0-0.47 10666.
VarietiesSvenno 10666.
Marker associationsTC95235 – 4.8 cM – BLA1 10666.
QRaw.ipk-1A
Chromosome1AS
VarietiesOpata/W-7984 (ITMI) RI mapping population 0255; Awn colour was contributed by W-7984 0255.
Marker associationsAssociated with Gli-A1 0255.
QRaw.ipk-1D
Chromosome1DS
VarietiesOpata/W-7984 (ITMI) RI mapping population 0255; Awn colour was contributed by W-7984 0255.
Marker associationsAssociated with GLI-D1 0255.

1.35. Grain Hardness/Endosperm Texture⌂ Home

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.

HA

Ha
NoteSoft phenotype.
Chromosome5DS
iFalcon/7*Heron, Heron/7*Falcon3109; Paha*2//Early Blackhull/5*Paha 203, 298; Early Blackhull Derivative/5*Nugaines 203, 298.
VarietiesChinese Spring 3106, 777; Cappelle Desprez 470; Heron {470, 1452}; Paha, Nugaines203, 298; NY6432-18 241.
ha
NoteHard phenotype
iFalcon/7*Heron, Heron/7*Falcon 3109; Paha*2//Early Blackhull/5*Paha 203, 298; Early Blackhull Derivative/5*Nugaines 203, 298.
Sources / synonymsCS*6/Cheyenne 5D 915; CS*6/Hope 5D 777; Capelle Desprez*7/Besostaya 5D 470.
VarietiesFalcon {470, 1452}; Holdfast 470; Early Blackhull, Early Blackhull Derivative {203, 298}; Cheyenne 3106; Clark's Cream 241.
Marker associationsHa was closely linked to Xmta9(Pui1)-5D 1414.
Single factor effects on hardness were found for chromosomes 2A, 2D, 5B and 6D, and interactive effects were found for chromosomes 5A, 6D and 7A 1414. The addition of King II rye chromosome 5R converted Holdfast wheat from hard to soft 470. A 14.5 kD rye analogue was also isolated from 6x triticales which have soft texture 470. All ryes have soft texture. Two genes for grain hardness were reported in 55. Hard and soft NILs are listed in 0298. QTL Courtot / CS: DH population: a major locus in chromosome 5DS coincided with Ha ; minor QTLs mapped in chromosomes 1A (associated with Xfba92-1A ) and 6D (associated with Xgwm55-6D ) 0141. Forno / Oberkulmer: Ten QTLs for kernel hardness (54% of the variation) were mapped in spelt 0280. Karl*2 / TA 4152-4: QHa.ksu-3B , associated with Xksum9-3B (R[2] =0.09, and QHa.ksu-5D (Ha) , associated with Xcfd-5D (R[2] =0.3), were identified 10273. Neixiang 188 (hard) / Yanshan 1 (medium hard): RIL population: QGh.caas-1B.1 with hardness allele from Yanshan 1, R[2 ] = 0.28, Xwms153-1BLXbarc81-1BL 10640. Opata 85 / W-7984 Synthetic (ITMI population): RIL population: Two QTLs were detected 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. Using proteomic analysis of 2D-protein gels applied to 101 lines of the 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 comprised 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 among 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.

1.36. Grain quality parameters⌂ Home

Qsev.mgb-6A
Chromosome6AL
Type varietiesNessapia/ T. dicoccoides MG4343 mapping population 9920.
Marker associationsAssociated with Xrsq805-6A 9920.
Qsev.mgb-7A
Chromosome7BS
Type varietiesMessapia/ T. dicoccoides MG4343 mapping population 9920.
Marker associationsAssociated with Xpsr103-7A 9920.
Cheyenne (high quality) / CS (low quality): RIL population: QTL were associated with Glu-1 on chromosome arms 1AL and 1DL and Gli-1/Glu-3 on 1BS 0251. Cultivar Cheyenne contributed the higher SDS sedimentation values 0251. The QTL on 1AL coincided with a QTL for bread loaf volume 0251. The QTL on 1DL and 1BS coincided with QTL for bread mixing time 0251.
TaPod-A1a
Chromosome3AL
VarietiesDoumai 11233.
TaPod-A1b
VarietiesShi 4185 11233.
There was no apparent relationship to the Per- series identified by isozyme analyses and listed in the Protein section.
QAmc.ocs-4A.1
Chromosome4AS
VarietiesCS/CS(Kanto107 4A) mapping population 0047.
Marker associationsAssociated with Xbcd1738-4A and Xcdo1387-4A 0047.

LVL

Lvl1
SynonymLvl 1 10312
Chromosome3A
Sources / synonymsCappelle Desprez*7/Bezostaya 1 3A 10312.
Marker associationsXgwm720-3ALvl1 appeared to be located in the Xgwm2-3AXgwm720-3A region 10312.
QTL Renan / Recital: RIL population: Loaf volume score was consistent across three environments 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. Thirty 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.

TaBAS1

Tabas1-B1
Chromosome2BL
Marker associationsXbarc167-2B – 10.38 cM – Tabas1 – 5.23 cM – Xcfa2278-2B 11198.
cBAS1 is a type of 2-Cys peroxiredoxin in a large peroxidase family.
Tabas1-B1a
VarietiesJing 41111198. Associated with higher TGW.
Tabas1-B1b
VarietiesHongmanchun 2111198. Associated with lower TGW

TaGASR

TaGASR7-A1
NoteSnakin/GASA gene family.
Chromosome7AL
Marker associationsXwmc301-7A – 17.9 cM – TaGASR7 – 10.6 cM – Xwmc9-7A 11115.
cGenBank KJ000052 11115. Hap1c in Lumai 14 and Xiaoyan 81 conferred higher grain length and grain weight than Hap1g in Hanxuan 10 and Xinmai 10 11115.

TaGS1

TaGS-D1
Chromosome7DS
Marker associationsTaGs-D1 – 8.0 cM – Xbarc184 11116.
TaGs-D1a
VarietiesDoumai 11116; Jingdong 8 11116.
cKF687956 11116. Associated with higher TGW and grain length 11116.
TaGs-D1b
VarietiesShi4185 11116; Yumai 21 11116.
cKF687957 11116. Associated with lower TGW and grain length 11116.

TaGW

TaGW-A2
Chromosome6A
Orthologous to the rice RING-type E3 ubiquitin ligase OsGW2 that functions as a negative regulator of grain weight.

TaSAP1

TaSAP1-A1
NoteStress association protein gene family.
Chromosome7A
Marker associationsXwmc530-7A – 2.1 cM – TaSAP1-A1 – 13.9 cM – Xbarc174-7A 11117.
cGenBank KC193579 11117. Variation at this locus was associated with 1,000-grain weight, number of grains per spike, spike length, penuncle length and total number of spikelets per spike, but different haplotypes had different effects various traints 11117.

TaTGW-7A

TaTGW-7Aa
VarietiesJing 411 11197. Associated with higher TGW.
TaTGW-7Ab
VarietiesHongmanchun 21 11197. Associated with lower grain weight.

TaTGW-A1

TaTGW-A1a
VarietiesDoumai 11196; Zhou 8425B 11196. Associated with higher TKW.
TaTGW-A1b
VarietiesChinese Spring 11196. Associated with lower TKW.

TaTGW6

TaTGW6-A1
Chromosome3AL
Marker associationsGene-366561 – 2 cM – TaTGW-A1 – 18 cM – BobWhitec4730456_ 11196.
cTGW6 in rice encodes an indole-3-acetic acid-glucose hydrolase 11196.
TaTGW6-B1
Chromosome3BL
cTGW6 in rice encodes an indole-3-acetic acid-glucose hydrolase 11196.
TaTGW6-D1
Chromosome3DL
cTGW6 in rice encodes an indole-3-acetic acid-glucose hydrolase 11196.

1.38. Grass-clump dwarfness/Grass dwarfness⌂ Home

Complementary dominant genes. Genotypes producing dwarfness: D1-D2-D3-, D1-D2D2, D1-D4-D3-, D1-D2-D4 and D1-D4D4 .

D1

D1
SynonymG 972
2D 939, 534, 1595. 2DS 942.

D2

D2
SynonymB I 972
Chromosome2BL
2B 574, 536.

D3

D3
SynonymA 972
Chromosome4AL
4A 534, 1595.

D4

D4
Chromosome2D
Chromosome2DL
Sources / synonymsCS[*] 7/Cheyenne 2D 1000.
Varieties (alt.)Cappelle-Desprez D2 1595; Cheyenne D2 1000; Brevor D2 1000.
d1d2d3d4
VarietiesChinese Spring {1000, 534}.
Genotype lists in can be found in 358, 534, 972. The effects of multiple allelism at D2 , and possibly at D1 , and modifying genes were demonstrated 1595. Knott 683 described a lethal dwarf condition controlled by a dominant gene closely linked with Sr30 (chromosome 5D) in Webster and a complementary recessive gene in LMPG. Phenotypes resembling grass clump dwarfs in hybrids carrying a 2BL.2RS translocation were reported in 916. The complementary gene{s} in wheat was not D1, D2 or D3 . The effect was suppressed at high temperature.

1.40. Hairiness/Pubescence traits⌂ Home

PA

pa
VarietiesGabo 886; Saratovskaya 29 886. This phenotype was expressed in Diamant ditelo 4BL886.

HG1

hg1
VarietiesUlyanovka 1405; Pionerskaya {1405, 715}.
Evidence for multiple alleles in T. monococcum is given in 744. The likelihood of three alleles, hg (hairless), Hg1 (weakly hairy) and Hg (very hairy), with hg1 being recessive to Hg and causing a short (weak) hairy phenotype, was mentioned in 1405.

HG2

Hg2
Chromosome2BS
VarietiesCIGM86.944 [syn. Croc1 / Ae. tauschii_ 518] 11508.
Type varietiesCroc_1 11508.
Marker associationsXicsH020 – 1.18 cM – HG2 – 0.84 cM – XicsHS358, corresponding to physical interval 740.0-741.1 Mb in cv. Svevo 11508.

HL1

Hl1
NoteWeakly hairy.
SynonymHl 884
Chromosome4BL
Chromosome4B
VarietiesArtemovka 925; Caesium 111 925; Lutescens 53/12 925; Lutescens 62 925; Milturum 321 884; Poltavka 925; Pyrothrix 28 925; Saratov 321 884; Saratovskaya 29 760, 884; Sarrubra 925.
Marker associationsXgwm375-4B – 12.1 cM – Hl1 – 2.1 cM 10516.

HL2

Hl2
Chromosome7BS
VarietiesHong-mang-mai 316. The hairy leaf gene ( Hl[Aesp] ) in Ae. speltoides introgression line 102/00[I] was allelic with Hl2 10516.
hl1 hl2
VarietiesChinese Spring 884.
Kuspira et al . 744 provided evidence for at least three alleles at an Hl locus in T. monococcum . 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. Trichome number on leaf margins in Ae. tauschii was mapped to a 530 kb region in chromosome arm 4DL 11612.

HS

Hs
SynonymHls 761
VarietiesCertain hexaploid derivatives of G25 produced in Israel 939.
Type varietiesT. dicoccoides G25 761.
hs
VarietiesMost hexaploid wheats 939.
Type varietiesT. dicoccoides G7 761.
Levy & Feldman 795 concluded that complementary genes determined hairy leaf sheath in T. dicoccoides .

HP

Hp
NoteDerived from Secale cereale 4BL {T
Chromosome4B
5R} 274, 275.

HN

Hn
Chromosome5AL
VarietiesAurore 722; Fylgia 722; Extra-Kolben II 722; Marquis 910; Tammi 765; T. vulgare erythrospermum 910.
Type varietiesT. polonicum vestitum 910.
hn
VarietiesGarnet 722; Kimno 722; Pika 722; Timantii 722.
Multiple alleles were reported in T. monococcum 744.

1.42. Reduced height⌂ Home

RHT-A1

Rht-A1a
Chromosome4A
Chromosome4AL
VarietiesChinese Spring 0019.
Marker associationsXwmc48-4AS – 2 cM – Xgwm610-4A – 1 cM – Rht-A1 – 2 cM – Xgpw4545-4AL 11017.
cGenBank KC767924. All common wheats are assumed to be monomorphic. A functional Rht-A1a allele is expressed at a similar level to its orthologues 10923.
Haplotypes named as Rht-A1b to Rht-A1g are described in 11620.

RHT-B1

Rht-B1a
VarietiesTall wheats116; e.g. Chinese Spring0019.
cGenBank KC767925.
Rht-B1b
NotePartially recessive 024, recessive 357, semi-dominant 408.
SynonymRht1 15, Sd1 15
Chromosome4B
Chromosome4BS
iSee 414, 2102, 408.
VarietiesFrontier 1597; Guardian 1597; Selection 14-53/Burt, 5 15; Siete Cerros 407; Wren 1174; WW15 407.
Varieties (alt.)Norin 10-Brevor, 14 Rht-D1b 15; Oleson Rht-D1b 357; Selection D6301 Rht-D1b 357; Shortim Rht-D1b 243; See1062, 407, 1386, 415.
Type varietiesCocorit 71 109, 416; Creso 109, 451,
416; Malavika 1442; Mida 450; Sansone 109; Valgerado 109, 416; Valnova 450; Valselva 450. ma,tv: Gai1/Rht-B1b – 1.8 cM – Xpsr622-4B 110; Co-located with Xbarc10-4B 10189. The development of allele-specific primers for Rht-B1b was reported in 0378. An EcoTILLING study of >1,500 Chinese wheat accessions identified 7 sequence variations in RHT-A1 , 8 new variants in RHT-B1 and 4 new variants in RHT-D1 11697. QTL QTL for reduced plant height, peduncle length and coleoptile length contributed by Cranbrook were associated with XcsMe1-4B (up to 49% of variability for plant height and peduncle length and 27-45% of variability for coleoptile length) in the cross Cranbrook (semidwarf) / Halberd (tall) . The dwarfing effect underlying the QTL was caused by the Rht-B1b allele 0379.
Rht-B1c
NoteSemi-dominant 1040.
SynonymSd3 565, Rht3 565
iTom Thumb/7[*] Kharkov//Lancer 1040; See 408.
VarietiesMinister Dwarf 404; Selection D6899 (Tom Thumb-Sonora 64/Tacuari) 357; Tom Thumb 405; Tom Pouce Blanc 1634, 407; Tom Pouce Barba Rouge1634, 407; Topo; Tordo.
Marker associationsXmwg634-4B (distal) – 30.6 cM – Rht-B1c – 11.9 cM – Xpsr144-4B (proximal) 117; Allele-specific markers were designed from the gene sequence 10923.
cThe Rht-B1c transcript carries a 90 bp in-frame insertion within the region encoding the conserved N-terminal DELLA domain plus two SNPs upstream of the insertion. A much larger insertion occurs in the gDNA 10923.
Rht-B1d
NoteSemi-dominant 1599, 116.
SynonymRht1S 1599
VarietiesSaitama 27 1599; Occurs frequently in Italian and Yugoslavian wheats 1599; Argelato, Centauro, Chiarano, Etruria, Farnesse, Gallo, Gemini, Lario, Pandas, Produttore, Orlandi, Orso, Salvia, Sprint, Strampelli.
cHas the same point mutations as in Rht-B1b - there is likely to be another mutation outside the coding region 10923. Rht-B1c carries a 2,026 bp insertion of a terminal repeat transposons in miniature (TRIM) insertion at position 147 bp relative to Rht-B1a ; this leads to an additional 30 amino acids in the DELLA domain affecting affinity between GID1 and Della 11390. Genbank JN857970 (gDNA), JN859791 (cDNA) 11390.
Rht-B1e
SynonymRht11 718, Rht1 ( B-dw ) 1600, RhtKrasnodari1 452
VarietiesKarlik 1 PI 504549 10924; Krasnodari 1 (a spontaneous GA-insensitive offtype of Bezostaya 1) 1600; Polukarlikovaya 49 and 11 derivatives 10924.
Marker associationsA PCR marker distinguishes this allele from Rht-B1a and RhtB1b 10923.
cA stop codon occurs three codons upstream of the Rht-B1b mutation 10923.
Rht-B1f
NoteSemi-dominant 116.
SynonymRhtT. Aethiopicum 116
Type varietiesT. aethiopicum accessions W6824D 116; W6807C 116.
Rht-B1g
VarietiesHighbury mutants M3 103-3 and M3 103-9 0019.
Allele Rht-B1g is a fast neutron-induced mutation of Rht-B1b and produces a tall gibberellin responsive phenotype 0019. Haplotypes named Rht-B1h to Rht-B1o are described in 11620.
Rht-B1p
NoteRht17 .
VarietiesChris Mutant CI 17241 1129.
cContains a C-to-T substitution at position 178 leading to a stop codon 11621. GenBank KT013263.
Rht-B1[IC2196 ] 10144.

RHT-D1

Rht-D1
4DS 1266 980, 116. 4D 583, 1544, 411. **bin:
Rht-D1a
VarietiesTall wheats 116; e.g. Chinese Spring.
cKC767927.
Rht-D1b
NotePartially recessive 024, recessive 357, semi-dominant 408.
SynonymSd2 15, Rht2 15
Chromosome4D
Chromosome4DS
iCommon wheat and durum NIL pairs are listed in 2102. See 414, 2102, 408.
VarietiesBiscay 10574; Combe 567; Era 407; Gaines Sib 2 15; Jaral 407; Kite 1174; Maris Hobbit 411; Pirat 10574; Pitic 62 567; Rubens 10574; Songlen 243.
Varieties (alt.)Oleson Rht-B1b 357; Norin 10-Brevor, 14 Rht-B1b 15; Selection D6301 Rht-B1b 357; List in 1386.
Marker associationsXpsr1871(Pki)-4D – 4 cM – Rht-D1 – 6 cM – Xubc821(PhyA)-4D 410; Rht-D1 – 2.8 cM – Xglk578-4D 9966; Xpsr1871 – 1 cM – Rht-D1b – 4 cM – Xpsr821 (PhyA) 0019. The development of allele-specific primers for Rht-D1b was reported in 378.
Rht-D1c
NoteDominant 114.
SynonymRht10 1266
VarietiesAi-bian {1266, 1544}.
Marker associationsXpsr921-4D (4DS) – 0.8 cM – Rht-D1c - 28 cM – Xgwm165-4D (4DL) 117.
Rht-D1d
NoteSemi-dominant 116.
SynonymRhtAi-bian 1a 115
VarietiesAi-bian 1a (spontaneous mutant of Ai-bian 1) 115.
Haplotypes named as Rht-D1e to Rht-D1h are described in 11620. Line XN004, earlier considered to have Rht21 0230, was shown to carry an allele at the Rht-D1 locus 0231. Various common wheat and durum N1Ls differing at the RHT-B1 and RHT-D1 loci are listed in 02102. Genotype lists in 402, 1382, 1612, 1613. Rht-D1b, Rht-D1c and Rht-D1d are identical across the coding region, but Rht-D1c has a fourfold increase in copy number relative to Rht-D1b ; Rht-D1d has a reduced copy number relative to Rht-D1c {10923, 11016}.

RHT4

Rht4
NoteRecessive.
Chromosome2BL
VarietiesBurt ert 937, CI 15076 {717, 566}.
Marker associationsAssociated with Xwmc317-2B 10249.
RHT5
Traes3B02G025600 (predicted).
Rht5
Chromosome3BS
VarietiesMarfed ert 1, M1, CI 13988 1593, 717, 718.
Marker associationsApproximately 10 cM from Xbarc102-3B 10249. Located to an ~1 Mb interval flanked by Kasp25 and Kasp23 in the 0-30 Mb region 11625. The predicted gene in Marfed M has a 30 bp deletion in the first intron 11625.

RHT6

Rht6
NoteRecessive.
VarietiesBrevor 569; Burt 718, 569.
Varieties (alt.)Norin 10-Brevor, 14 Rht-B1b RhtD1b 569.

RHT7

Rht7
Chromosome2A
VarietiesBersee Mutant A 1602; Bersee Mutant C 1602.
**RHT8.** TraesCS6A02G221900 ; TraesCSU03G0022100 (CS RefSeq v2.
Rht8
Note2D 1601, 1598, 772.
Chromosome2DS
Sources / synonymsCappelle-Desprez[*] /Mara 2D 1601.
VarietiesChuan Mai 18 10249; Novasadska Rana 1 1604; Sava {414, 1601}; Yumai 8679 11624.
Varieties (alt.)Akakomugi Rht9 1191; Mara Rht9 1191.
Marker associationsXgwm484-2D (proximal) – 19.9 cM – Rht8 – 0.6 cM – Xgwm261-2D (distal) 727; Close linkage with Xgwm261-2D 10249; A survey of Chinese cultivars showed 13 alleles of Xgwm261-2D 10284.
cRHT8 contains two ORF with near-identical sequences. Encodes an unknown 808 aa protein containing a zinc finger BED-type motif and predicted Ribonuclease H-like domain {11624, 11642}. CRISPR-Cas9 editing of the RNHL-A1 ( Traes2A02G059900 ) and RNHL-B1 ( Traes2B02G073600 ) homoeologues caused reduced height 11624.
rht8
RNHL-D1 11624. Close linkage of Rht8 and Xgwm261-2D permitted the use of the microsatellite as a marker for the detection of putative allelic variants at the Rht8 locus 9962. Allele sizes for Xgwm261 in U.S. eastern and central wheat cultivars are given in 10868. A series of ‘alleles’ was based on variation in the closely linked marker Xgwm261-2D
Rht8a
NoteAssociated with a 165-bp fragment of WMS 261 9962.
VarietiesAutonomia 9962; Bobwhite 9962; Brevor 9962; Chaimite 9962; Ciano 67 9962; Chris 9962; Dugoklasa 9964; Federation 9962; Frontana 9962; Glennson 81 9962; Hope 0243; Jupateco 73 9962; Kenya 9962; Klein 32 9962; Lerma Rojo 9962; Lusitano 9962; Maringa 9962; Marquis 0243;
Mentana 9962; Michigan Amber 0243; Nainari 60 9962; Newthatch 9962; Opata 85 9962; Othello 9962; Penjamo 62 9962; Quaderna 9962; Rex 9962; Riete 9962; Saitama 27 9962; Spica 9962; Veery S 9962; Victo 9962.
Rht8b
NoteAssociated with a 174-bp fragment of WMS 261 9962.
Sources / synonymsCappelle Desprez[*] /Mara 2D 1601.
VarietiesArthur 0243; Balkan 9962; Bunyip 9962; Cappelle-Desprez 9962; Carstens 0243; Diakovchanka 0243; Eureka 9962; Festival 9962; Fronteira 9962; Fultz 9962; Gabo 9962; Heine VII 9962; Inallettabile 95 9962; Jena 9962; Klein Rendidor 9962; Leonardo 9962; Lutescens 17 9962; Mironovskaya 808 9962; Norin 10 9962; Norin 10/Brevor 14 9962; Oasis 243; Odom 0243; Podunavka 9962; Purdue Abe 0243; Record 9962; Red Coat 9962; Salzmunder Bartweizen 14/44 0243; Soissons 9962; Talent 9962; Tevere 9962; Timstein 9962; Tp114/65 0243; Wilhelmina 9962; Wiskonsin 245 C/11226 0243.
Rht8c
NoteAssociated with a 192 bp fragment of WMS 261 9962.
VarietiesAkakomugi 1191; Alfa 9962; Aquila 9962; Ardito 9962; Argelato 9962; Avrora 9962; Banija 9964; Baranjka 9964; Beauchamps 9962; Bezostaya 9962; Biserka 9962; Campodoro 9962; Centauro 9962; Chikushi-Komugi (Norin 121) 9962; Chuanmai 18 10512; Damiano 9962; Djerdanka 9964; Dneprovskaya 9962; Duga 9964; Etoile-de-Choisy 9962; Etruria 9962; Fakuho-Komugi (Norin 124) 9962; Farnese 9962; Favorite 9962; Fedorovka 0243; Fiorello 9962; Fortunato 9962; Funo 9962; Gala 9962; Haya Komugi 9962; Impeto 9962; Irnerio 9962; Jarka 9964; Jugoslavia 9962; Kavkas 9962; Kalyan 0243; Khar'kovskaya 50 0243; Khar'kovskaya 93 0243; Khersonskaya 86 0243; Kolubara 9964; Kosava 9964; Libellula 9962; Lonja 9964; Lovrin 32 9962; Macvanka-2 9964; Mara {9962, 119}; Marzotto 9962; Mv 03-89 0243; Mv 06-88 0243; Mv 17 0243; Neretva 9962; Nizija 9962; Novasadska Rana 1 1604; N.S. Rana 1 9962; N.S. Rana 2 9962; N.S. 649 9962; N.S. 3014 9962; Obrii 243; Odesskaya 51 0243; Odesskaya 117 0243; Odesskaya 132 0243; Odesskaya Krasnokolosaya 0243; Odesskaya Polukarlikovaya0243; Orlandi9962; Osjecanka 9964; OSK 5 5/15 9964; OSK 4 57/8 9964; OSK 3 68/2; Partizanka 9962; Partizanka Niska 9962; Poljarka 9964; Posavka 1 9964; Posavka 2 9962; Pomoravka 9962; Produttore 9962; Radusa 9962; Roazon 0243; Salto 9962; Sanja 9962; San Pastore 9962; Sava {414, 1601, 9962}; Siete Cerros 9962; Sinvalocho 9962; Simvol Odesskii 0243; Sivka 0243; Strumok 0243; Skopjanka 9962; Skorospelka 3B 9962; Slavonija 9964; Somorka9964; Sremica 9964; Superzlatna 9962; Svezda 9962; Tira 0243; Tisa 9964; Transilvania 9962; Ukrainka Odesskaya 0243; Una 9962; Villa Glori 9962; Vympel 0243; Yubileinaya 75 0243; Zagrebcanka 9964; Zelengora 9964; ZG 6103/84 9964; ZG 7865/83 9964; Zitarka 9964; Zitnica 9962; Zlatna Dolina 9964; Zlatoklasa 9964; Zolotava 0243. Although CS carries a 192 bp fragment, sequencing showed it was a different allele than other genotypes with Rht8c 02103.
Although the 'diagnostic' association of Rht8c and Xgwm261 192 applied in many Strampelli derivatives and European wheats, there was no association between reduced height and this allele in Norin 10 and its derivatives 10512. The pedigrees of a number of Chinese wheats postulated to have Rht8c on the basis of the marker traced to Italian sources 10515.
Rht8d
NoteAssociated with a 201-bp fragment of WMS261 9962.
VarietiesPliska 9962; Courtot 9962.
Rht8e
NoteAssociated with a 210-bp fragment of WMS261 9962.
VarietiesChino 9962; Klein Esterello 9962; Klein 157 9962.
Rht8f
NoteAssociated with a 215-bp fragment of WMS261 9962.
VarietiesKlein 49 9962.
Rht8g
NoteAssociated with a 196-bp fragment of WMS261
Synonym0243
VarietiesMirleben 0243
Rht8h
NoteAssociated with a 206-bp fragment of WMS261
Synonym0243
VarietiesWeihenstephan M1 0243.
Genotypes of Indian semi-dwarf wheats based on the Ellis et al. 0378 markers are listed in 10404 and those for U.S. eastern and central and winter wheat cultivars are listed in 10868.

RHT9

Rht9
Chromosome5AL
7BS 1601, 772.
RHT10
See Rht-D1c .

RHT11

Rht11
NoteSee Rht-B1e.
VarietiesKarlik 1 718.
RHT12
TraesCS5A02G543100
Rht12
NoteDominant.
Chromosome5A
Chromosome bin5AL-23, based on co-segregation with B1 1606.
iN98-2105, Yangmai 5*5 / Karcagi 522M7K 11428.
VarietiesKarcagi 522M7K 721; Mercia, Vigour 18 and Halberd derivatives 11622.
Marker associationsRht12 is located distally on 5AL cosegregating with B1 and closely linked to b-AmyA1 1606; Xgwm291-5A – 5.4 cM – Rht12 726. The Rht12 phenotype was due to deletion of a 10.73 Mb terminal deletion of chromosome 5AL {11428). Reduced plant height might be due to activation of TaGA2ox-A14 11428.
cEncodes a GA2oxA13 enzyme 11622 (previously predicted as GA2oxA14 11428. Rht12 delayed ear emergence by 6 days 1606.

RHT13

Rht13b
NoteRht13 718.
Chromosome7BS
VarietiesMagnif 41M1 CI 17689 718.
Marker associationsAssociated with Xwms577 ( gwm577 ) -7B 10249.
cA serine-240-phenylalanine mutation caused autoactivation leading to transcriptional upregulation of pathogenesis-related genes including class III peroxidases associated with cell wall modelling 11626. The same sequence was identified in TILLING line Cadenza0453 11626.

RHT14

Rht14
NoteAllelic with XRht16, Rht18 and Rht24 {10767, 10818}.
Chromosome6AS
VarietiesCp B 132 123 = Castelporziano PI 347331718.
Marker associationsRht14 – 11.7 cM – Xbarc3-6A 10767. Mapped to genomic region 383-422 Mb flanked by GA20xA9 and Xwmc753-6A 11372. GA2oxA9 expression was higher in Castelporziano than in its tall parent Capelli – see Rht18 11301.
cSee RHT24 .

RHT15

Rht15
Type varietiesDurox 718.

RHT16

Rht16
NoteAllelic with Rht14 and Rht18 {10767, 10818}.
Chromosome6AS
VarietiesEdmore M1 718.
Marker associationsRht16 – 28.0 cM – Xbarc3-6A 10767.

RHT17

Rht17
NoteRht-B1p {M23014}.
VarietiesChris Mutant CI 17241 1129.

RHT18

Rht18
NoteAllelic with Rht14 , Rht16 and Rht24 { 10767, 10818}.
Chromosome6AS
Type varietiesIcaro718.
Marker associationsRht18 – 25.1 cM – Xbarc3-6A 10767. Hexaploid derivatives in the backgrounds of Fengchan 3, Jinmai 47 Rht8 , and Xifeng 20 are reported in 11096.
matvXbarc118-6A – 1.4 cM – RHT18/TdGA2Ox-A9/S470865SSR4/Xbarc37-6A – 0.4 cM – IWA4371 – 0.4 cM –Xgwm82.1-6A 11295.
cSee Rht24 .
Independent ‘overgrowth’ mutants isolated from Icaris contained changes in the GA2oxA9 coding region; this gene is predicted to encode GA2-oxidase that metabolizes GA biosynthetic intermediates into inactive products thus reducing bioactive GA1 11301.

RHT19

Rht19
Type varietiesVic M1 718.

RHT20

Rht20
VarietiesBurt M860 718.

RHT21

Rht21
The existence of this gene was not confirmed 0231.

RHT22

Rht22
Chromosome7AS
Type varietiesAiganfanmai 10857. Ailanmai 11587.
Marker associationsXgwm471-7A – 29.5 cM – Rht22 – 20.1 cM – Xgwm350-7A 10857.

RHT23

Rht23
Chromosome5DL
VarietiesNAUH164 11077.
Marker associationsXgdm63-5D – 4.7 cM – Rht23 – 11.1 cM – Xbarc110-5D 11077. NAUH164 is an EMS-derived mutant of Sumai 3 11077.
RHT24 . c: Encodes TaGA2ox-A9 which is more highly expressed by the Rht24b allele 11623. Same locus as Rht14 , Rht16 , and Rht18 . Rht24a . Rht24 11185. [ QTLheight6A1 11183, QPH.caas-6A 11184, Rht24b_ {11293, 11294}.]. 6AL 11185.
Rht25b
NoteQHt.ucw-6AS 11300.
Chromosome6AS
VarietiesPatwin-515HP 11659; U C1110 Rht-D1b 11300.
Marker associationsQHt.ucw-6AS was located in a 0.2 cM interval flanked by 6A13699/6A13791/6A14397 and 6A14825 11300.
RHT26 Rht26a
Varieties (alt.)Lunxuan 987 RhtB1b {11739.
Rht26b . 3DL 11739.
Rht27
dvT. urartu G1812 11740.
Marker associationsFlanked by 3T-387 and 3T-306 11740. Delimited to a 1.55 Mb region (517.5 – 528.6 Mb, CS RefSeq v1.0) containing 20 genes, 2 of which, or homoeologs of which, coded genes affecting plant height 11740.
rht27
dvReduced Height 27 11740. The height of the mutant was 27% of the wild type 11740.
Rht_NM9
Chromosome2A
VarietiesInduced mutant NM9 11273.
Marker associationsXgwm122-2A – 1.7 cM – SNP34 – 1.9 cM – RhtNM9 – 1.9 cM – SNP41 – 14 .0 cM – Xwmc261-2A_ 11273.
QHt.fcu-4BL
Marker associationsAssociated with Xbarc125-4B (R[2 ] = 0.57) 10256. Reduced height allele in Grandin 10256.
QHt.fcu-6AS
Marker associationsAssociated with Xbarc23-6A - Xcp201-6A (R[2 ] = 0.07) 10256. Reduced height allele in BR34 10256.
In RL4452 / AC Domain:
QHt.crc-2D
Chromosome2D
Marker associationsLinked to BE497718-260 (LOD 4.2).
QHt.crc-4B
Chromosome4B
Marker associationsLinked to Rht-B1 (LOD 7.7) 10287. Associated with QTLs for lodging and 1000-grain weight.
QHt.crc-4D
Chromosome4D
Marker associationsLinked to Rht-D1 (LOD 30.9) 10287. Associated with QTLs for lodging 1000-grain weight, yield, height, and test weight.
QHt.crc-5B
Chromosome5B
Marker associationsLinked to Xwmc640-5B (LOD 6.1) 10287.
QHt.crc-7A
Chromosome7A
Marker associationsLinked to Xwmc139-7A (LOD 3.3) 10287.
QHt.crc-7B
Chromosome7B
Marker associationsLinked to Xgwm333-7B (LOD 3.3) 10287. In Courtot / CS:
QHt.fra-1A
Marker associationsLinkage with Xfba393-1A 9957.
QHt.fra-1B
Marker associationsLinkage with Xcdo1188-1B.2 9957.
QHt.fra-4B
Marker associationsLinkage with Xglk556-4B 9957.
QHt.fra-7A
Marker associationsLinkage with Xglk478-7A 9957.
QHt.fra-7B
Marker associationsLinkage with XksuD2-7B 9957.

In Renan / Recital:

QHt.inra-2B
Marker associationsAssociated with Xgwm249-2B (LOD=5.8, R[2] =15.4%) 10069.
QHt.inra-4A
Marker associationsAssociated with Xfba243-4A (LOD=6.5, R[2] =15.0%) 10069.
QHt.inra-5A
Marker associationsAssociated with Xgwm639b-5A (LOD=5.7, R[2] =10.8% 10069.
QHt.inra-6D
Marker associationsAssociated with Xcfd76-6D (LOD=3.7, R[2] =8.1% 10069.
QHt.inra-7A
Marker associationsAssociated with Xcdo545-7A (LOD=3.2, R[2] =7.7%) 10069.
In Opata 85 / W-7984 (ITMI) RIL mapping population:
QHt.ipk-4A
Chromosome4AL
Tall allele contributed by Opata 85 0255.
QHt.ipk-6A
Chromosome6A
Tall allele contributed by W-7984 0255.
QHt.ocs-4A.1
Chromosome4AL
Marker associationsAssociated with Xpsr119-4A and Wx-B1 0047.
QHt.ocs-4A.2
Chromosome4AS
Marker associationsAssociated with Xbcd1738-4A and Hd 0047.
In CS( T. spelta 5A) / CS(Cappelle-Desprez 5A) RI mapping population:
QHt.ocs-5A.1
SynonymQt.ocs-5A.1 0068
Chromosome5AL
Marker associationsAssociated with the interval Xcdo1088-5AXbcd9-5A 0068.
This QTL coincided with a QTL for culm length, QCl.ocs-5A.1 0068. In: Sevannah / Senat DH population:
QHt.riso-3A
Marker associationsMapped on the centromeric region between SSR markers Xwmc505-3A and Xwmc264-3A (LOD >6) 10067.
QHt.nau-2D
NoteRecessive. Gibberellin-sensitive.
Chromosome2DS
Chromosome bin2DS-0.47-1.00.
VarietiesEMSInduced Dwarf Wangshuibai 11463.
Marker associationsXbarc-2D – 2.6 cM- 2DQHt.nau-2D – 0.3 cM- 2DXgpw361-2D 11463.
Two QTLs for plant height were assigned to chromosome 3A in RSLs from Cheyenne[*] 7/ Wichita 3A substitution line 0025. Seven QTLs on chromosomes 1A, 1D, 2B, 2D and 4B affected plant height among RILs of CS /
T. spelta duhamelianum
Effects linked with the CS alleles of Xbcd1160-1A , Xksu127-1D and XksuF11-2D increased height whereas those CS alleles associated with Xpsr131-2B , Xpsr125-2B , Xpsr934-2D and Xcs22.2-4B reduced it 0196. For review and identification of 65 QTL-rich clusters associated with plant height see 11741.

1.43. Herbicide Response⌂ Home

DFQ1

Dfq1
NoteInsensitive.
Chromosome2B
Chromosome2BL
VarietiesCS1396.
dfq1
NoteSensitive.
Sources / synonymsCS[*] 6/Ciano 67 2B 1396; CS[*] 7/Marquis 2B 789; CS[*] /Sicco 2B 1396.
VarietiesCiano 67 1396; Sicco 1396.
Busch et al . 153 reported a single dominant gene for tolerance of Era and Marshall compared with the susceptibility of Eureka and Waldron, but its relationship to Dfq1 is unknown.

SU1

Su1
NoteInsensitive.
Chromosome6BS
Chromosome6B
VarietiesCappelle-Desprez 1402.
Type varietiesB-35 735.
su1
NoteSensitive.
VarietiesChinese Spring 1402; Poros 1402.
Type varietiesB-7 735.
Marker associationsXpsr312-6B – 5.3 cM – Su1 – 6.8 cM – Xpsr477(Pgk2)-6B 736. ma,tv: Nor2 (6BS) – 2.7 cM – Su1 1401; Su1 – 5.2 cM – Xpsr371-6B (6BL) 735.
Su1 also controls insensitivity to metoxuron 1402. A single dominant gene for tolerance to isoproturon was found in tetraploid wheats derived from a tolerant T. monococcum source 1044. This gene is presumably different from Su1 .

IMI1

Imi1
SynonymFs-4 10100, AhasL-D1 10101
Chromosome6DL
VarietiesBW755 = Grandin*3/Fidel-FS-4 10099; CDS Teal IMI 1A 10099; CDC Teal IMI 9A 10099; CDC Teal IMI 10A = Fidel-FS-2 10099; Clearfield WHS Janz = Janz*4/Fidel-FS-2; Clearfield WHS Stiletto = Stiletto*3//Spear/FidelFS-3; Fidel-FS-2 = ATCC40997 10100.
Varieties (alt.)CDC Teal IMI 15A = PTA 3955 Imi3 10099.

IMI2

Imi2
SynonymAhasL-B1 10101
Chromosome6BL
VarietiesCDC Teal IMI 11A = PTA 3953 10099.

IMI3

Imi3
SynonymAhasL-A1 10101
Chromosome6AL
Varieties (alt.)CDC Teal IMI 15A Imi3 10099.
dvT. monococcum mutant EM2 (mutant of susceptible line TM23 10102).
Mutant EM2 has a serine to asparagine substitution near the carboxyl end of the enzyme. The same change led to imidazolinone resistance in hexaploid wheat, rice and Arabidopsis 10102.

1.44. Hybrid Weakness⌂ Home

NE1

Ne1
SynonymLe 550, 155, F 971, Le1 1491
Chromosome5B
Chromosome5BL
Marker associationsXbarc216-5B – 8.3 cM – Ne1 – 2 cM – Xbarc74-5B 10334. Ne1 – 11 cM – Xgwm639-5B 11343. Mapped to a 4.06 Mb region (383.03 – 3.87.10 Mb) that was deleted in all tested non- Ne1 carriers 11517. Co-segregation with the null allele of indel marker 5B-InDel385 11517; Mapped to a 4.45 Mb interval represented by Xwgrc3074-5B – 0.07 cM – NE1/5markers – 0.12 cM – Xwgrc3009-5B 11518. Xbarc216-5B – 3.8 cM – Xwgrc3030 – 0.3 cM – NE1/Xwgrc1426/ 3009 – 4.8 cM – Xbarc74 - 5B 11537; Xgwrc3030 – 1.4 cM – Ne1/Xwgrc3146/3147/3150/Xmag1426 – 0.12 cM – Xwgrc3150 11537; markers Xwgrc3146, Xwgrc3147 and Xwgrc3150 were dominant 11537. Nwu5B-4137 – 0.2 cM – NE1 – 0.3 cM – Nwu5B4114_ at 383.30 – 388.01 Mb in CS RefSeq 1.0 11594.
Ne1s
VarietiesBig Club 550, 532, 155; C306 1475; Felix 531; Gaza 141 PI 220429 532; Luteseens 1163 1264; Marquillo 550, 115, 532; Ponca 532; Spica 939; Synthetics TA4152-19, TA4152-37, TA4152-44, TA4152-60 10334.
Type varietiesGaza 1E PI 133460; Gaza PI 189262 532; Iumillo 532; Kubanka 532; PI 94587 {532, 155}; Quanah 532.
Ne1s is common in tetraploid wheats 1080.
Ne1w
VarietiesBobin group 532: Kenya Farmer 532; The Bobin selection used in breeding Gabo 532 and its sister selection, Timstein {532, 1556} was in fact Gular. Hence Gular, not Steinwedel, is the presumed source. The Sydney University accession Bobin W39 was the parent of Gabo and Timstein, whereas "true" Bobin carried the accession number W360. The particular accession tested by Hermsen is not clear. According to Metgzer 1000 Steinwedel is a non-carrier; Federation group 532; Cadia 532; Cleveland 971; Minister group 532; Rieti group 532: Mentana 532; Mara 532.
Ne1m
iS-615[*] 11/Prelude 1500.
VarietiesCarpo 532; Eskisehir 220-39 532; Garnet 532; Klein Aniversario 532; Koga 532; Mus XII/80/22 532; Prelude {1491, 532}.
Unknown Ne1 alleles.
Ne2
SynonymF 971, Le2 1491, 550, 155
Chromosome2B
Chromosome2BS
Marker associationsXgwm148-2B – 6.7 cM – Ne2 – 3.2 cM – Xbarc55-2B 10334. Xbarc7-2B – 3 cM – Ne2 – 6 cM – Xwmc344-2B 11343. Xgwm148-2B – 5.2 cM – Xwgrc1713/Xwgrc1736-2B – 1.3 cM – NE2/ 3 markers 11518; Xgwm148-2B – 5.4 cM – Ne2/Xwrgc1774/1775/1739 – 3.0 cM – Xwmc474-2B 11537.
Ne2s
iS-615[*] 11/Kharkov 1500.
VarietiesCrimean group 532: Blackhull 550; Chiefkan 550; Clarkan 550; Kharkov 1491; Liaochun 10 11530; Zhoumai 22 11531; Michigan Amber 532; Minhardi 155; Red Chief 550; Stepnaja 135 1264; Turkey 532.
cEncodes a CCNBS-LRR protein {11531; 11532; 11533}. One of two Ne2m haplotypes is Lr13 11531. GenBank MW756036 11532.
cAdd: N22m is an allele of the YR27/LR13 locus {M22053}.
Ne2m?
VarietiesBarleta group 532: South American wheats, e.g. Klein Titan 532; La Prevision 25 532; Lin Calel 532.
Ne2ms
VarietiesMediterranean group 532: Dawson 550, 155; Fultz 550; Fulcaster 550; Fulhard 550; Honor 550; Jones Fife 1491; Shepherd 550, 971; Trumbull 155; Vermillion 530; Wabash 155. (Although placed in this group on basis of pedigree, the last three stocks, as well as Fultz selection of CI 19293, appear to have the stronger allele of the Crimean group 532; Mironovskaya 10630; Noe group 532: Vilmorin 27 532
Ne2m
VarietiesAlsen 10334; Squarehead group 532: European wheats 532; Fronteira group 532: Sonalika 1475; South American wheats and derivatives, e.g. Atlas 40 532: Wheats possessing Lr13 939, e.g. Manitou 939; HD2329 10985.
Marker associationsXbarc55-2B – 1.1 cM – Xkwh37 – 4.9 cM – Lr13/Ne2 – 5.8 cM; Xgpw1109 – 3.7 cM – Xbarc18-2B 11068.
Ne2w
VarietiesVakka 532; Varma 532.
Unknown Ne2 allele 532;
ne1 ne2
VarietiesChancellor 531; Elgin 1491; Gladden 155; Leap 155; Purkof 155; Red Bobs 1491; Red Egyptian 1491; Steinwedel 1000; S-615 1491; Wichita 531.
Genotype lists in 531, 532, 535, 640, 696, 698, 1093, 1135, 1264, 1381, 1473, 1474, 1475, 1492, 1496, 1497, 1502, 1503, 1512, 1505, 1506, 1507, 1508, 1509, 1510, 1630, 1631, 1632, 1633, 1637, 1638, 1639, 0112, 10985}. The incompletely dominant Els2 mutant found in an EMS-treated M2 population of H261 was likely ne2 11472; A similar situation was described in 11423 where a claimed a putatively novel gene was designated as yglw-1 . Rye line 1R136-2 carries Ner1 1210 that complements wheat gene Ne2 {1289, 1210} and rye gene Ne2 1210 to produce necrosis. Rye lines L155 and L256 carry Ne2 1210 that complements Ne1 {630,} 1210 and Ne1 1210.
Ner1
Chromosome5RL
alS. cereale 1R136-2 1210.
Ner2
Chromosome7RL
alS. cereale L155, L256 1210.

CH1

Ch1
Synonymm[a ] 1245
2A 939, 538.

CH2

Ch2
SynonymNe3 1504, m[b ] 1245
3DL 692, 939. 3D {1504, 1495}.
ch1 ch2
VarietiesAlbit {1509, 1000}; Burt {1509, 1000}; Chancellor 1000; Garra 1549; Kharkof 535; Steinwedel 1549.
suTAP 67 (= Pawnee 3Ag(3D)) 1644.
Lists in 535, 697, 1381, 1473, 1474, 1475, 1496, 1497, 1502, 1503, 1512, 1505, 1506, 1507, 1508, 1509, 1510}. A gene, Chr1 , in rye produces chlorosis symptoms in hybrids with wheats such as C306, HD2939 and NI5439 possessing Ch2 1472. Evidence for multiple alleles of Chr1 was also presented 1472.
Chr1
dvCereal rye lines, EC179188 = WSP527A 1472; EC143825 = WSP506A 1472; EC338685 = Blanco 1472; others 1472.
chr1
dvEC179178 1472; EC179185 = SAR/SWPY5 1472.

CS1

Cs1
SynonymChl[1]
Chromosome5A
VarietiesT. dicoccum cv. Hokudai 1511. Occurs at high frequency in the T. paleocolchicum group of emmers.

CS2

Cs2
SynonymChl[2 ] 1501
Chromosome4G
Type varietiesMany accessions of T. timopheevii and T. araraticum 1511,637. Multiple allelism at the Cs2 locus is discussed in 637.
A gene named NetJingW176 (after Ae. tauschii accession Jing Y176) was located in chromosome 2DS: Xgwm-102- 2D – 4.5 cM – Nec2 – 3.8 cM – Xgwm515-2D 11307.
Apd1
VarietiesWR95 = Kalyansona/Gigas//HD1999/Sonalika/3/ T. carthlicum 10492.
Apd2
VarietiesHD2009 10492; HW2041 10492; Lok-1 10492; others 10492.
Apd1 Apd2
VarietiesAtila 10492; Kalyansona 10492; others 10492.
apd1 apd2
Lethal genotype. Uniculm plants occured as heterozygous segregates among progenies, but homozygous uniculm lines could not be established 10492.

NEC1

Nec1
Chromosome7DS
Varieties( T. durum cv. Langdon x Ae. tauschii KU-2828) amphiploid 11158.
alAe. tauschii KU-2828 11158.
Marker associationsXbarac352-7D – 5.3 cM – Lr34Xgwm295-7D – 4.0 cM – Xbarc154-7D – 1.7 cM – Nec1 – 13.2 cM – Xcfd-7D 11158.
Although this form of hybrid necrosis is caused by complementary genes, mapping of Nec1 was based on a cross of necrotic and non-necrotic Langdon x Ae. tauschii amphiploids. Consequently only Nec1 was mapped 11158.

1.45. Iron deficiency⌂ Home

Fe1
Chromosome7DL
VarietiesSaratovskaya 29 921.
Fe2
Chromosome7BS
VarietiesCS 927.

1.46. Lack of ligules⌂ Home

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. An Imperial rye chromosome 2R addition restored the liguled condition to a liguleless CS derivative 939. An erect leaf mutant involving TaSPL8 (SQUAMOSA promoter-binding protein-like transcription factor), a homolog of LG1 in maize and rice and was located in chromosome 2D. Knockout mutants of TaSPL8 orthologs led to a fully liguleless phenotype. The gene in 2D was identified as TraesCS2D01G502900. TaSPL8 transcript was highly expressed in the laminar joint region and young spike. TaSPL8- 2D transcript was produced at much higher levels than TaAPL-2B whereas TaSPL-2A was produced at a minimal level 11401.

LG1

lg1
Chromosome2B
Chromosome2BS
iANK33 = Novosibirskaya 67*10/K59990 10061.
VarietiesEligulate W1342 lg2 lg3 942, 10133; K31289 10133; K59990 10061; K53660 10133; Liguleless partial backcross derivative of CS 939; Partial backcross derivative of CS 939.
Type varietiesK17769 10133; K17784 10133.
**LG2** TraesCS2D01G502900 .
lg2
Chromosome2D
iANK33 = Novosibirskaya 67*10/K59990.
VarietiesEligulate W1342 lg1 lg3 ; Liguleless partial backcross derivative of CS 939.
A dominant mutant allele for liguleless phenotype is reported in Ae. tauschii – this locus is located in chromosome 2DL but is independent of LG2 .

LG3

lg3
Chromosome2A
iANK33 = Novosibirskaya 67*10/K59990 10061.
VarietiesPresent in all hexaploid cultivars.
LG[t] [11220].
Lg2[t]
dvLiguleless mutants of Ae. tauschii accession KU20-9 11220.
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.

1.47. Leaf characteristics⌂ Home

QLer.ipk-2A
Chromosome2AS
VarietiesOpata/W-7984 (ITMI) RI mapping population 0255; The erect leaf phenotype was contributed by Opata 0255.
Marker associationsAssociated with Xbcd348-2A 0255. Mutants lacking ligules are known to have erect leaves. However, the QTL for leaf erectness reported here is not related to liguleless mutants 0255.

LTN1

Ltn1
SynonymLtn 1361
Chromosome7D
v : Wheats with Lr34/Yr18 301, 1361;

LTN2

Ltn2
Chromosome1B
VarietiesWheats with Yr29/Lr46 10281; See Yr29, Yr46 .
Varieties (alt.)Parula Ltn1 10281.
Marker associationsXwmc44-1B – 1.4 cM – Xbac24prot – 9.5 cM – Ltn2 – 2.9 cM – Xbac17R ........ Xgwm1401B 10281; Xgwm44-1B – 3.6 cM – Ltn2 – 2.1 cM – XtG818/XBac17R.....Xgwm140-1B 10281.
According to Messmer et al. 0031 LTN may be caused by several QTLs and is affected by genetic background and environment.

LTN3

Ltn3
iRL6077 11070.
VarietiesChapingo 48 11070.
cThis multiple disease resistance/necrosis locus was identified as a hexose transporter most similar to the STP13 family and containing 12 predicted transmembrane helices 11070.
QLtn.sfr-1B
Chromosome1BS
VarietiesForno/ T. spelta var. Oberkulmer mapping population 0050.
Marker associationsAssociated with Xgwm18-1B and Xglk483-1B 0050.
QLtn.sfr-3A
Chromosome3A
VarietiesForno/ T. spelta var. Oberkulmer mapping population 0050.
Marker associationsAssociated with Xpsr570-3A and Xpsr543-3A 0050.
QLtn.sfr-4B.1
Note4B00 50.
VarietiesForno/ T. spelta var. Oberkulmer mapping population 0050.
Marker associationsAssociated with Xpsr921-4B and Xpsr593-4B 0050.
QLtn.sfr-4B.2
Chromosome4B
VarietiesForno/ T. spelta var. Oberkulmer mapping population 0050.
Marker associationsAssociated with Xpsr593-4B and Xpsr112-4B 0050.
QLtn.sfr-4D
Chromosome4DL
VarietiesForno/ T. spelta var. Oberkulmer mapping population0050.
Marker associationsAssociated with Xpsr302-4D and Xpsr1101-4D 0050.
QLtn.sfr-5A
Chromosome5A
VarietiesForno/ T. spelta var. Oberkulmer mapping population 0050.
Marker associationsAssociated with Xpsr549-5A and Xglk163-5A 0050.
QLtn.sfr-6A
Chromosome6A
VarietiesForno/ T. spelta var. Oberkulmer mapping population 0050.
Marker associationsAssociated with Xpsr563-6A and Xpsr966-6A 0050.
QLtn.sfr-7B.1
Chromosome7B
VarietiesForno/ T. spelta var. Oberkulmer mapping population 0050.
Marker associationsAssociated with Xpsr350 and Xbzh232(Tha)-7B 0050.
QLtn.sfr-7B.2
Chromosome7B
VarietiesForno/ T. spelta var. Oberkulmer mapping population 0050.
Marker associationsAssociated with Xglk750-7B and Xmwg710-7B 0050.
QLtn.sfr-7D
Chromosome7DS
VarietiesForno/ T. spelta var. Oberkulmer mapping population 0050.
Marker associationsAssociated with Xpsr160-7D and Xgwm44-7D 0050.

SC

Sc
Chromosome3BS
Sources / synonymsCS[*] /Hope3B 149.
VarietiesHartog 149; Suneca 149; wheats with Sr2 149.
Leaf chlorosis is affected by temperature and light and is enhanced by infection with pathogens. The sc allele is completely linked with Pbc (pseudo-black chaff) and Sr2 (reaction to Puccinia graminis ).

ELS1

els1
Chromosome2BS
VarietiesZK331 / Xiangmai 99171 // 2*Lumai 30 Line 114 11326.
Marker associationsWGGB305 – 0.3 cM – els1/WGGB302 – 1.2 cM – WGGB303/WGGB304/WGGB306 – 0.6 cM – Xbarc92-2B 11326.
The els1 ‘mutant’ was detected in an F4 population. Since the parents had normal phenotype complementary genes were likely involved. The similar location of ELS1 to the NE1 locus in chromosome 2BS and similar phenotype suggests that this gene may be Ne2 . See 49, Hybrid Weakness; 49.1, Hybrid necrosis

ELS2

Els2
Chromosome2BL
VarietiesLF2099 {11472 }.
Marker associationsXgpw4043-2B – 8.87 cM – Els2 – 22.27 cM Xwmc149-2B 11472.
The incompletely dominant Els2 mutant was found in an EMS-treated M2 population of H261 11472. ELS3 . Candidate gene CS2D02G332700 .
Els3
VarietiesYanzhan 4110 11772.
cThe candidate encodes an LRR-RLK -like gene located on the cell membrane 11772.
els3 . v: els3 mutant 11772. The mutant was identified in an EMS-mutagenized population of Yanzhan 4110 11772.

1.48. Lesion Mimicry⌂ Home

Add introductory sentence: Lesion mimic phenotypes are characterised by spontaneous hypersensitivity not unlike disease resistance. Lesions are often not produced when leaf sectors are protected from light, and disease levels on mutant individuals may be lower than on non-mutant sibs. Lesion mimics that resemble the response of plants to infection by pathogens occur in many species (10743 for examples).

LM

lm
Chromosome1BL
Chromosome binC1BL6-0.32 10743.
VarietiesNing 7840 10743.
Marker associationsProximal to Xgwm264.1-1B 10743.
LM was positively associated with QLr.pser.1BL 10743.

LM1 and LM2

lm1
VarietiesZaosui 30 Lm2 11572.
LM2
Chromosome4BL
Marker associationsXgwm513-4B – 1.5 cM – LM2 – 2.9 cM – Xksum154-4B 11572.
lm2
VarietiesYanzhan 1 Lm1 11572.
LM3
Chromosome3BL
Marker associationsMapped to a 3 cM proximal region of chromosome 3BL 11573.
Lm3
The mutant phenotype appeared in an F1 plant of cross Line 3-1/Jing 411. The plant was then backcrossed 6 times with Line3-1 and the selected line was named Lm3.
LM4
Chromosome2DS
Marker associationsFine mapped within a 8.06 cM interval flanked by Xgwm210 -2D and Xgwm455-2D using specifically developed markers m401cib and lm402cib 11577.
Lm4 . Wild type allele. Recessive.
lm4
NoteThe allele named lm4 and conferring the lesion phenotype was described as dominant 11577. This lesion mutant appeared in a Yanzhan 1/Neixiang 188 RIL population in which the segregation of mutant versus normal phenotype was 1:1. LM5 . Semi-dominant.
Chromosome2AL
Lm5
VarietiesMC21, an EMS-induced mutant in Chuannong 16 11576.
Marker associationsKASP-4211 (630.3 Mbp) – 0.6 cM – Lm5 – 3.7 cM – KASP5353 (703.53 Mbp) 11576.

1.49. Lodging⌂ Home

QLd.crc-3D
Chromosome3D
Marker associationsLinked to Xgwm191-3D (LOD 3.7) in RL4452/AC Domain 10287.
QLd.sfr-1B
Chromosome1BS
VarietiesForno/ T. spelta var. Oberkulmer mapping population 0052.
Marker associationsAssociated with Xpsr949-1B and Xgwm18-1B 0052.
This QTL coincided with QTL for reduced height, increased culm stiffness and broader leaf width 0052.
QLd.sfr-2A
Chromosome2AS
VarietiesForno/ T. spelta var. Oberkulmer mapping population0052.
Marker associationsAssociated with Xpsr958-2A and Xpsr566-2A 0052.
This QTL coincided with QTL for reduced height, increased culm stiffness, broader leaf width, more erect growth habit, later ear emergence and increased culm thickness 0052.
QLd.sfr-2D
Chromosome2D
VarietiesForno/ T. spelta var. Oberkulmer mapping population 0052.
Marker associationsAssociated with Xpsr933-2D and Xglk529-2D 0052.
QLd.sfr-3A
Chromosome3AS
VarietiesForno/ T. spelta var. Oberkulmer mapping population0052.
Marker associationsAssociated with Xpsr598-3A and Xpsr570-3A 0052.
This QTL coincided with QTL for increased culm stiffness and reduced culm thickness 0052.
QLd.sfr-4A
Chromosome4AS
VarietiesForno/ T. spelta var. Oberkulmer mapping population 0052.
Marker associationsAssociated with Xgwm397-4A and Xglk315-4A 0052.
This QTL coincided with QTL for reduced height, increased culm stiffness and more erect growth habit 0052.
QLd.sfr-5A
Chromosome5AL
VarietiesForno/ T. spelta var. Oberkulmer mapping population 0052.
Marker associationsAssociated with Xpsr918-5A and Xpsr1201-5A 0052. This QTL coincided with QTL for reduced height, increased culm stiffness, reduced leaf width, more erect growth habit, later ear emergence and increased culm thickness 0052.
QLd.sfr-5B
Chromosome5BL
VarietiesForno/ T. spelta var. Oberkulmer mapping population 0052.
Marker associationsAssociated with Xpsr370-5B and Xpsr580-5B 0052. This QTL coincided with QTL for increased culm stiffness, broader leaf width and more erect growth habit 0052.
QLd.sfr-6B
Chromosome6BL
VarietiesForno/ T. spelta var. Oberkulmer mapping population 0052.
Marker associationsAssociated with Xpsr964-6B and Xpsr142-6B 0052.
QLd.sfr-7B
Chromosome7BL
VarietiesForno/ T. spelta var. Oberkulmer mapping population 0052.
Marker associationsAssociated with Xpsr927-7B and Xpsr350-7B 0052.
This QTL coincided with QTL for reduced height and later ear emergence 0052.

1.50. Male sterility⌂ Home

Ms1376
NoteSterility is dominant.
VarietiesTR1376A 10814. Male fertile counterpart: TR1376B 10814.
Ms1376 was discovered among progenies of a transgenic family of Xinong 1376 containing leaf senescence-inhibiting gene P SAG12-IPT 10814. MS1 4B 268. 4BS 64. TraesCS4B02G017900 .
ms1
Recessive alleles for sterility
ms1a
VarietiesBriggle's Chancellor Derivative 268; Pugsley's Male Sterile 268.
cTerminal deletion 11422.
ms1b
VarietiesProbus mutant 268. c Interstitial deletion 11269.
ms1c
VarietiesCornerstone 64.
cTerminal deletion 11269.
ms1d
VarietiesMutant FS2 619; Ningchen mutants msd.1 and msd.2 11421.
cG329A in exon 1 {11421, 11269}.
ms1e
VarietiesMutant FS3 619.
cG1431A del1432 11421; C1435T + 16 bp del in exon 2 11269.
ms1f
VarietiesMutant FS24 619.
cG155A in exon 111269.
ms1g
Chromosome4BS
VarietiesLanzhou Mutant 257A {10355, 10354}; Male sterile line 257A10546.
cDeletion of entire sequence 11421.
ms1h
VarietiesNingchun 4 mutant 11421.
cC1762T 11421.
ms1i
VarietiesNingchun 4 mutant 11421.
cG1603A 11421.
ms1j
VarietiesNingchun 4 mutant 11421.
cC1775A 11421.
ms1k
VarietiesNingchun 4 mutant 11421.
cG1397A 11421.
ms1l
VarietiesNingchun 4 mutant 11421.
cC226T 11421.
ms1m
VarietiesNingchun 4 mutant 11421.
cC1472T 11421.
ms1n
VarietiesNingchun 4 mutant 11421.
cT164A 11421.
ms1o
VarietiesNingchun 4 mutant 11421.
cG281A 11421.
ms1p
VarietiesNingchun 4 mutant 11421.
cG155A 11421.
ms1q
VarietiesNingchun 4 mutant 11421.
cC148T 11421. ms1r {11269]. [ Ms1h 11269].
VarietiesTilling mutant in Qual2000 11269.
cG178A 11269.
Ms1 orthologs in the A and D genomes are epigenetically silenced 11421.
Shw
Synonym1HL 331
adAdditions of 1H and 1HL to wheat and certain translocation lines 331.
Marker associationsLocated in a 16.4 cM interval flanked by Xmwg800-1H and Xmwg943-1H . A possible relationship with Ncc genes is discussed 331.
tmsBS20T
Chromosome2BL
VarietiesBS20-T 11157.
Marker associationsXgwm403-2B – 2.2 cM – tmsBS20T – 4.5 cM – Xgwm374-2B 11157.

MS2

ms2
NoteDominant allele for sterility.
SynonymTa1 240
Chromosome4DS
VarietiesTaigu = Line 223 240, 806, 807; ms2 confers sterility when present in octaploid triticale 597.
Marker associationsMapped to a 0.05 cM region
flanked by Xsauw27-4D and Xsdauw29-4D 11388.

MS3

ms3
NoteDominant allele for sterility.
Chromosome5AS
iChris derivative 872; KS87UP9 219.
Marker associationsXwg341-5A – 0.8 cM – ms3 .......cent 0289; Xcdo-677-5A and Xbcd1130-5A also cosegregated with Xwg341-5A but were located in a different region in the physical map 0289.

MS4

ms4
NoteDominant allele for sterility, distinguished from ms2 on the basis of different degrees of recombination with the 4D centromere.
Chromosome4DS
VarietiesKonzak's male sterile.
MS5 . TraesCS3A02G217000 . 3A 619, 3AL 11427. Ms5 : Encodes a glycosylphosphatidylinositol-anchored lipid transfer protein that is required for pollen exine development 11427. TraesCS3A02G217000 11427. GenBank MK577897.
ms5
VarietiesMutant FS20 619; H45 11427. Complete sterility conferred by ms5 is dependent on the homoeologous ms-D genotype – Ecalibur, Gladius and RAC 875 have an ms-D allele that restores fertility to ms5 genotypes 11427.
Two non-functional ms-B alleles (Chinese Spring and Synthetic W7984 types) were identified 11427.

WTMS1

wtms1
Chromosome2B
VarietiesBNY-S 10332.
Marker associationsE: AAG/M:CTA163 – 6.9 cM – wtms1 – 4.8 cM – Xgwm374-2B 10332. Described as a thermo-sensitive gene (TGMS), giving complete sterility at less than 10C, but fertile at higher temperatures 10332.

WPTMS1

wptms1
Chromosome5B
VarietiesLine 337S wptms2 10333.
Marker associationsXgwm335-5B – 4.2 cM – wptms1 – 24.4 cM – Xgwm371-5B 10333.
wptms1 produces sterility only in the presence of wptms2 . Chromosome 5B was also implicated in spontaneous mutant line Xinong 291S: a second gene was not located 11143.

WPTMS2

wptms2
Chromosome2B
VarietiesLine 337S wptms1 10333.
Marker associationsXgwm374-2B – 6.9 cM – wptms2 – 20.9 cM – Xgwm120-2B 10333.
wptms2 produces sterility only in the presence of wptms1 . wptms1 and wptms2 were analysed and mapped under long photoperiod/high temperatures, but an earlier study indicated a single gene for male sterility under short photoperiod/low temperatures. Although mapping data are different a possible relationship between wtms1 and wptms2 needs to be resolved.

1.52. Maturity time⌂ Home

RL4452 / AC Domain:

QMat.crc-3B
Chromosome3B
Marker associationsLinked to Xwmc231-3B (LOD 3.0) 10287.
QMat.crc-4A
Chromosome4A
Marker associationsLinked to Wx-B1 (LOD 6.1) 10287.
QMat.crc-7D
Chromosome7D
Marker associationsLinked to Xgwm130-7D (LOD 17.5) 10287.

1.53. Megasporogenesis⌂ Home

Msg
Chromosome7AS
Type varietiesLangdon 625.
TaASY1
Encodes an axial-element-associated protein that is essential for synapsis and crossover formation in Arabidopsis and rice 11766. Mutants affect synapsis and distribution of chiasmata along the chromosome 11767.
TaASY-A1
TaASY-B1
TaASY-D1
QTug.sau-3B
Chromosome bin5BS5-0.07-0.03.
Marker associationsXgwm285-3B – 1.0 cM – Xcfp11012-3B 11471. Identified in T. turgidum / Ae. tauschii hybrids involving Langdon durum (high unreduced gamete formation) crossed with AS313 and AS2225 (low unreduced gamete formation) topcrossed with Ae. tauschii AS60. The QTL was located near Ttam , a homologue of the TAM (tardy asynchronous meiosis/CYCA1;2) cyclin gene from Arabidopsis 11471.

LTP

Ltp
VarietiesChinese Spring 527. Expressed in the absence of chromosomes 5D at 12[o] C - 15[o] C, but not at 20[o] C. A contrasting allele, Ltp , for normal pairing at the lower temperature range was demonstrated in T. dicoccum .

PH1. Traes………

Ph1
Ta-Zip4-B2
Chromosome5BL
VarietiesHexaploid wheat.
Type varietiesTetraploid wheat.
Marker associationsPCR-based assays for presence and absence of Ph1 were described 359, 0217, 9965, 0214; The Ph1 factor(s) was restricted to a region flanked by Xrgc846-5B and Xpsr150-5B 0219; Ph1 was physically mapped in 5BL to fraction length 0.55, bracketed by deletions 5BL-1 and ph1b 446. A complex Ph1 candidate structure comprising at least one 5B-specific member of the cdc2 complex multigenic cluster (involved in chromosome condensation), a unique repeat structure with similarities to repeats on chromosome 3B, and a heterochromatic sub-telomeric insertion from chromosome 3AL was identified 10240. Ta-Zip4-B2 is embedded within this complex 11765.
A set of homoeologous set of Ta-Zip-1 genes ( Ta-Zip-A1 , TaZip-B1 and TaZip-D1 genes is present in Group 3 chromosomes but their expression levels are very low compared to Ta-Zip-B2 {11765).
ph1a
Not applicable - see ph2b 1303.
ph1b
VarietiesSears' high pairing mutant 1301.
Marker associationsA PCR-based detection system for ph1b ph1b individuals is described in 9965. Dualplex marker Xwgc2111 + Xwgc2049 behaves like a co-dominant marker 11359. The Ph1b deletion involves a region of at least 60,014,523 bp 11359.
ph1c
Type varietiesCappelli ph1 mutant 593,449; This mutant is deficient for a terminal portion of chromosome 5BL449.
Marker associationsMutant lines with ph1b and ph1c carry deletions of the chromosome segment possessing Ph1 in the respective parent lines {593, 447}. Several ph1 mutants are described in 0219.
**PH2** TraesCS3D02G119400 .
Ph2
Chromosome3DS
TaMSH703D 11527.
ph2a
VarietiesSears' intermediate pairing mutant {1302, 1301}. ma : ph2a is a 120-125 Mb deletion 11526.
ph2b
Synonymph1a 1537
VarietiesChinese Spring mutant 10/13 1537.
cContains a G to A transition at position 74,359.312 in the TaMSH7-3D gene 11527. TaMSH7 is a plant-specific member of the DNA mismatch repair (MMR) famil y 11527.
Wide cross hybrids involving ph2 mutants have a 5.5-fold increase in homoeologous pairing 11516.

1.55. Nitrate reductase activity⌂ Home

NRA

Nra
VarietiesUC44-111 424.
nra
VarietiesAnza 424.

1.56. Nuclear-cytoplasmic compatability enhancers⌂ Home

SCS

Scs
NoteDerived from T. timopheevii 869.
Synonymscs[ti ] 10878
1AL 870, 27.

1.57. Nucleolus organizer regions⌂ Home

Nor1a and Nor2a
VarietiesMaris Huntsman 1399.
Nor1b and Nor2b
VarietiesBezostaya 1 1399.
Nor1c and Nor2c
VarietiesCappelle-Desprez, Maris Ranger 1399.

NOR-A1

Nor-A1
Chromosome1AS
VarietiesT. spelta {1012, 221, 367, 835}.
dvT. monococcum 658.

NOR-B1

Nor-B1
SynonymNor1 1120
Chromosome1BS
Chromosome1B
VarietiesCS 288. Deletion mapping divided the Nor-B1 in a proximal subregion Nor-B1p (short repeat) and a distal subregion Nor-B1d (long repeat) 0275
Nor-B1a
VarietiesCheyenne, Chinese Spring, Hope, Kite, Oxley, Teal, Timstein288, 37; Vasco, 8 others 288.
Nor-B1a-
VarietiesA derivative allele of Nor-B1a with a significantly reduced amount of spacer. Condor 64-1918; Sonora 64-1918.
Nor-B1b
VarietiesOlympic, Robin, Shortim 917.
Nor-B1c
VarietiesBanks 917; Corella 917; Warigal 917; 5 others 917.
Nor-B1c-
VarietiesRosella 918.
Nor-B1d
VarietiesMaris Huntsman 918.
Nor-Ag[i] 1 374. 1Ag[i ] 374.
Nor-H1
SynonymNor-I1 794
Chromosome1HS
dvSultan barley 794.
Nor-R1
Chromosome1RS
adCS/Imperial 39.
Nor-S1
Chromosome1SS
alAe. speltoides 294.
Nor-U1
Chromosome1U
suCS/ Ae. Umbellulata 906.
Nor-V1
Chromosome1V
adCS/ D. Villosum 241.
Nor-B2
SynonymNor2 1120
6BS 1041, 221, 366, 835.
Nor-B2a
Chromosome6B
VarietiesCS 37, 917.
Nor-B2a-
VarietiesBlueboy 918; Sonora 64-1 918.
Nor-B2b
NoteT
Chromosome6B
VarietiesBanks, Oxley, Shortim, Timstein 37; 12 others 917.
Nor-B2c
VarietiesCorella, Robin, Teal, 1 other 917.
Nor-B2d
NoteH6B 288.
VarietiesHope 37; Olympic 917; Warigal 917.
Nor-B2d-
VarietiesHarrier 918; Kite {918, 917}.
Nor-B2e
VarietiesVasco 917.
Nor-B2f
NoteCh6B 288.
VarietiesCheyenne {37, 917}.
Nor-B2g
VarietiesFalcon; Gluclub; La Prevision 918.
Nor-B2h
VarietiesYaktana 918.
Nor-B2i
VarietiesMaris Huntsman; Thatcher 918.
More detailed listings for allelic variation at Nor-B1 and Nor-B2 are given in 917, 918.
Nor-E2
Chromosome6ES
adCS/ E. elongate 294.
Nor-G2
Chromosome6G
Type varietiesT. timopheevii IPSR (PBI) No. 1 294.
Nor-H2
SynonymRnr1 1248
6H 1070, 39, 1248. 6HS 794.
Nor-S2
Chromosome6SS
alAe. speltoides 294.
Nor-A3
Note5AS 658, 1014.
dvT. monococcum, T. urartu IPSR (PBI) Acc. A.

NOR-D3

Nor-D3
Note5DS 221, 835.
VarietiesCS; most wheats 288, 37, 917.
Nor-Ag[i] 3. 5Ag[i ] 374.
Nor-E3
Chromosome5ES
adCS/ E. elongate 294.
Nor-H3
SynonymRnr2 1248
Chromosome5HS
5H 1070, 39, 1248.
Nor-U3
Chromosome5U
ad,su: CS/ Ae umbellulata 906.

NOR-D4

Nor-D4
Chromosome7DL
VarietiesCS 1042.
dvAe squarrosa 1042.
Nor-H4
SynonymNor-I4 794
Chromosome7HS
alSultan barley 794.
Nor-H5
SynonymNor-I5 794
Chromosome2HS
alSultan barley 794.

NOR-B6

Nor-B6
Chromosome1BL
VarietiesCS; Cheyenne, Wichita 601.
Type varietiesLangdon 601.

NOR-A7

Nor-A7
Chromosome5AL
VarietiesCS; Cheyenne, Wichita 601.
Type varietiesLangdon 601.

NOR-D8

Nor-D8
Chromosome3DS
VarietiesWitchita 601.

NOR-A9

Nor-A9
SynonymNor-A1 1012, 221, 367, 835
1AS 276, 282.

NOR-A10

Nor-A10
SynonymNor-A3 658, 1014
5AS 276, 282.

1.58. Osmoregulation⌂ Home

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.

OR

Or
NoteLow osmoregulation.
Sources / synonymsCS (Red Egyptian 7A).
VarietiesCappelle Desprez; Condor[*] 4/3Ag14 1030; Red Egyptian.
Marker associationsOr (proximal in 7AS) – 13 cM – Xpsr119-7A 1031.
Or
NoteHigh osmoregulation.
Chromosome7AS
Chromosome7A
VarietiesCS, Condor, Songlen, Takari 1030.

1.59. Phenol colour reaction of kernels⌂ Home

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).

TC

Tc1
Chromosome2AL
suVarious substitutions of chromosomes 2A into CS 10131.
sutvLangdon*/dicoccoides 2A 10130.
Type varietiesGolden Ball 10130.
Tc2
Chromosome2BL
sutvLangdon*/Golden Ball 2B 10130.
Type varietiesGolden Ball 10130.
Tc3
SynonymTc 10131
Chromosome2DL
VarietiesChinese Spring (intermediate response) 10130.
Varieties (alt.)Timstein Tc1 10131.
suCS/*Timstein 2D 10131.
Type varietiesCocorit 71 10130; Langdon 10130.
sutvLangdon*/CS 2D(2A); Langdon*/CS 2B(2D) 10130; T. dicoccoides Israel A 10130. Lines with a negative phenol colour reaction.

1.60. Pollen killer⌂ Home

KI

Ki
NoteKilling allele is dominant.
Chromosome6BL
VarietiesChinese Spring1306; Mentana 929.
ki
VarietiesProbably the majority of wheats including Timstein, Gabo and Yalta 1306. Modifiers also appear to be involved as Luig {840, and unpublished} found variation among kiki parents. Some F2 and F3 Sr11sr11 plants from Yalta/Chinese Spring crosses segregated with less than 50% Sr11 - phenotypes among the progeny indicating that killing extended to eggs as well as pollen. See also, Gametocidal Activity.
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 :

1.61. Polyphenol oxidase (PPO) activity⌂ Home

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

QTL

Chara (mod high) / WW2449 (low): DH population: PPO activity Associated with Xgwm294b-2A (R[2] =0.82), Xwmc170-2A , Xgwm312-2A and Xwmc178-2A (R[2] >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 . – 0.6 cM – XPPOLDOPA/XPPO18/BQ161439 10484.

M6 (high activity)/ Opata 85 (low activity): A QTL on 2D, associated with Xfba314-2D was identified using the L-DOPA assay 0344.

NY18 / Clark's Cream: Markers significantly associated with PPO activity were also detected on chromosomes 2A, 2B, 3B, 3D and 6B in the population 0344. A multiplex of markers PPO33 and PPO16 was reliable for selecting genotypes with low PPO activity 10418.

Zhongyou 9507 / CA9632: PPO18 explained 28-43% of the variation in PPO activity 10290.

Tetraploid wheat

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 Xgwm312-2AL and Xgwm294-2AL .

Messopia / T. dicoccoides : RIL population: Associated with RFLP Xutv1427-2A 10411.

R-1

R-A1
SynonymR2 548
3AL 1003, 957.
R-A1a
Marker associationsBased on Tamyb10-A1 sequences this allele in CS lacks the ability to bind DNA due to deletion of the first half of the R2 repeat of the MYB domain 10838; the R-A1a allele in Norin 17 has a 2.2 bp insertion in the second intron that appears to prevent transcription 10838.
R-A1b
SynonymR2
Tamyb10-A1 10107.
R-B1
SynonymR3 548
3BL 1003, 370.
R-B1a
Marker associationsBased on the Tamyb10-B1 sequence this allele in CS has a 19 bp deletion of the CCG repeat region causing a frameshift mutation 10838.
R-B1b
Note[ R3 ], Tamyb-10-B1 10107.
iNovosibirskaya 67[*] 9/k-28536 730.
VarietiesCanon 370; Dollar 370; Grana 370; Supreme 370.
cGenBank AB191459.
R-D1
SynonymR1 548
Chromosome3DL
VarietiesCS.
Marker associationsXbcd131-3D/R-D1 – 15 cM – Xabc1743D 410; Xgwm2-3D –.4 cM – R-D1 – 3.2 cM – Xgwm4306-3D 10839.
R-D1b
Note[ R1 ], Tamyb10-D1 10107.
iNovosibirskaya 67[*] 9/CS 730.
VarietiesAlexandria 370; Apollo 370; Axona 370; CS 1293; Dwarf A 370; Fortress 370; Jerico 370; Longbow 370; Luna 370; Mardler 370; Maris Huntsman 370; Minaret 370; NFC 75/93/27A; Rapier 370; Pawnee 549; Voyage 370; Vuka 370.
cGenBank AB19160.
R-N1
Chromosome3N
suCS/ Ae. Uniaristata 1018.
R-R1
Chromosome6RL
adHoldfast/King II 1011.
R-V1
Chromosome3VL
adCreso/ D. villosum 1518. A 3Ag chromosome from decaploid Ag. elongatum carries an allele for red grain colour which was transferred to Agent and the majority of Sears' 3D-3Ae#1 translocations 939. Other studies have identified wheats carrying either one or two, unidentified R-1 alleles: {056, 437, 549, 631, 654, 659, 1078, 1148, 1333, 1349, 1454, 370}. See also Variegated Red Grain Colour.
R-A1b R-B1b R-D1a
SynonymR3, R2
VarietiesRed Chief 548; Avalon 370; Bersee; Cappelle Desprez; Feuvert; Mission; Parade; Rendezvous; Yuri 370.
R-A1b R-B1a R-D1b
SynonymR1, R2
VarietiesBezostaya 1 370; Brigand 370; Broom 370; Brock 370; Kronjuwel 370.
R-A1a R-B1b R-D1b
SynonymR3, R1
VarietiesFenman 370; Kharkov 1003; Norman 370; Pastiche 370; Riband 370; Sperber 370; Squadron 370; Urban 370.
R-A1b R-B1b R-D1b
SynonymR2, R1, R3
VarietiesBowie; Frondoso 1148; Frontiera 437; Hope {206, 204}; Japanese Bearded 1548; Kanred {1078, 1426}; Lin Calel 1078.
Functional markers based on Tamyb10 sequences are given in 10838. A fourth QTL for red seed colour in hexaploid wheat was detected on chromosome 1B in a GWAS of U.S winter genotypes 11409.
R-N1
Chromosome3N
suCS/ Ae. Uniaristata 1018.
R-R1
Chromosome6RL
adHoldfast/King II 1011.
R-V1
Chromosome3VL
adCreso/ D. villosum 1518.
A 3Ag chromosome from decaploid Ag. elongatum carries an allele for red grain colour which was transferred to Agent and the majority of Sears' 3D-3Ae#1 translocations 939. Other studies identified wheats carrying either one or two, unidentified R-1 alleles: 056, 437, 549, 631, 654, 659, 1078, 1148, 1333, 1349, 1454, 370. See also Variegated red grain colour.
Vg
VarietiesLine 10859 498.
vgvg genotypes in Line 10859 are variegated. The Vg/vg locus was independent of the single red gene locus in Line 10859. In a cross to Selkirk ( R-A1b, R-B1b, R-D1b) vgvg was expressed only in plants with one R gene 498. Variegated red pericarp was also studied in crosses of cv. Supreme. In this case, two red colour genes were present 0136.

1.63. Response to photoperiod⌂ Home

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".

PPD-A1

Ppd-A1a
Type varietiesGS100 10612; GS105 10612. Ppd-A1a was present in 39% of Chinese landraces and 97% of improved cultivars 10622. GS100 and GS105 had different deletions relative to GS101 and GS104, respectively and both were consistently a few days earlier flowering than their near-isogenic counterparts with Ppd-A1b 10612
Ppd-A1b
Type varietiesGS101 10612; GS104 10612.

PPD-B1

Ppd-B1
Chromosome2BS
Marker associationsXwhs2002-2B/Xgwm257-2BPpdB1Xgwm148-2B . Actual linkage value varied between crosses 10129; Xpsr666-2B – 1.2 cM – Xpsr109-2B – 4.4 cM – Ppd-B1 – 4.8 cM – Xpsr8042B ...Cent 0062.
According to 10611 the Ppd-B1 allele from Japanese wheats has a stronger effect than the allele from CS.
Ppd-B1a
SynonymPpd2 1566
iH(C) = Haruhikari*5 / Fukuwasekomugi 10611; H(D) = Haruhikari*/5/ Fukuwasekomugi Ppd-D1a 10611.
Sources / synonymsCappelle-Desprez*/CS 2B 0058.
VarietiesCS 1268; Spica 557; Timstein 1269.
Varieties (alt.)Fukuwasekomugi Ppd-D1a 10611; Sharbati Sonora Ppd-A1a 887.
cVarieties with the photoperiod insensitive allele have more than one Ppd-B1 copy per chromosome 2B: two copies in Recital, three copies in Sonora 64, Timstein and C591, and 4 copies in Chinese Spring 10881.
Ppd-B1b
VarietiesBeaver 10881; Cappelle-Desprez 10881; Cheyenne 10881; Norstar 10881; Paragon 10881; Renan 10881.
Varieties (alt.)Haruhikari Ppd-D1b 10611.
cVarieties with the photoperiod sensitive allele have a single Ppd-B1 copy 10881.

PPD-D1

Ppd-D1
Chromosome2DS
Comparative mapping showed that Ppd-D1 was co-linear with barley Ppd-H1 - a member of the pseudo-response regulator (PRR) gene family 10466. Jagger amplified the 414 bp band 10466 associated with daylength sensitivity whereas 2174 amplified the 288 bp band associated with insensitivity 10722.
Ppd-D1a
SynonymPpd1 1566
iH(C) = Haruhikari*5/Fukuwasekomugi 10611; Haruhikari*5/Saitama 27 10611; H(D) Haruhikari*5/Fukuwasekomugi Ppd-B1a 10611.
Sources / synonymsCapelle Desprez[*] /Ciano 2D 1598; Capelle-Desprez[*] /Mara 2D 1598; CS[*] /Ciano 2D Ppd-B1a 1268.
VarietiesAkakomugi 1604; Bezostaya 1 1604; Festival 10466; Kavkaz; Mara 1604; Orqual 10466; Recital 10466; Saitama 27 10466; Sava 1604; Sideral 10466; Soissons 10466; Sonora 64 1566; Talent 10466; Texel 10466.
Varieties (alt.)Sharbati Sonora Ppd-D1a 887; Fukuwasekomugi PpdD1a 10611.
Marker associationsStocks with Ppd-D1a had a 2,089-bp deletion upstream of the coding region leading to mis-expression of the 2D PRR gene 10466.
Ppd-A1b Ppd-B1b Ppd-D1b
VarietiesCheyenne 1141; Diamont 1 887; Lancer 638; Saratovskaya 29 887; Warrier 638.
Two genes controlled photoperiod response in T. turgidum 788. Gene Ppd-H2 on barley chromosome 2HS may be a member of the Ppd-1 orthologous series 766.

PPD-B2

Ppd-B2
Chromosome7BS
suFavorit (F26-70 7B) 10628.
VarietiesF26-70 0093.
Marker associationsXgwm255-7B – 20.7 cM – Ppd-B2 – 4.4 cM – Xgwm537-7B 10628. This gene confers earlier flowering under long photoperiod conditions 10628.
QPpd.zafu-4AL
Chromosome4AL
Sources / synonymsCASL 4AL 11443.
Type varietiesT. dicoccoides TTD140 11443.
Marker associationsFlanked by M576 and Xwmc468-4AL in a 1.2 cM region 11443. Other publications reporting Ppd genes/QTL in the same region are reviewed in 11443.
Trident (early)/Molineux (late): In addition to an effect associated with chromosome 2B, three QTLs were designated as: QPpd.agt-1AL ( Xwmc304Xgwm497 region), QPpd.agt-7AS ( Xbarc154Xbarc108 ) and QPpd.agt-7BS ( Xgwm46Xgwm333 ) 10382. The QTL in chromosome 1A was possibly orthologous to Ppd-H2 in barley. QTL A QTL was detected in chromosome 4BS in Courtot/CS 0132.

CONSTANS 1

CO-A1
Chromosome7A
VarietiesChinese Spring.
cTraesCS7A02G211300 11295.
Type varietiesKronos 11495. c GenBank MT043302 11495.
CO-B1
Chromosome7B
VarietiesChinese Spring. c TraesCS7B02G118300 11495.
Type varietiesKronos 11495.
cGenBank MT043303 11495
CO-D1
Chromosome7D
VarietiesChinese Spring.
cTraesCS7D02G212900 11495.

CONSTANS 2

CO-A2
Chromosome6A
VarietiesChinese Spring.
cTraesCS6A02G289400 11495.
Type varietiesKronos.
cGenBank MT04330411495.
CO-B2
Chromosome6B
VarietiesChinese Spring.
cTraesCS6B02G319500 11495.
Type varietiesKronos.
cGenBank MT043305 11495.
CO-D2
Chromosome6D
VarietiesChinese Spring.
cTraesCS6D02G269500 11495.
Triticum turgidum ssp. durum cv. Kronos: accession numbers MT043302 ( CO-A2 ) and MT043305 ( COB2 ) (11495}.

1.65. Response to salinity⌂ Home

KNA1

Kna1
Chromosome4DL
Chromosome4BS
4BL-4DL 283. 4BS.4BL-4DL-4BL 849.

NAX1

Nax1
Chromosome2AL
itvTamaroi*6/Line 149 = P06306 10453.
Type varietiesLine 149 Nax2 = 126775b 10452.
dvAUS 90382 Nax2 = C68.101 10455 = JIC T. aegilopoides no. 3.
Marker associationsNax1 was mapped as a QTL in the region Xpsr102-2A – 5.4 cM – Xwmc170-2A – 0.9 cM – Xksud222A/Xksu16-2A – 0.8 cM – Xgwm312-2A with R[2] = 0.38 in Tamaroi/Line 149 10452; TmHKT7-A2 was identified as a putative candidate Na[+] transporter 10454.
Nax1 promotes withdrawal of Na[+] from xylem in leaf bases and roots 10453.

NAX2

Nax2
Chromosome5AL
itvTamaroi*6/Line 149 = P05603 10453.
Type varietiesLine 149 Nax1 = 126775b {10452, 10453}.
dvAUS 90382 Nax1 = C68.101 10455 = JIC T. aegilopoides no. 3.
Marker associationsCo-segregation with Xgwm291-5A/Xgwm140-5A/Xgpw2181-5A 10455; TmHKT1;5-A was identified as a candidate for Nax2 10455.
Nax2 is a likely orthologue of Kna1 10455. QTL Berkut / Krichauff: QTL for Na[+] exclusion and seedling biomass under salt stress were detected in chromosomes 2A ( Nax1 region) and 6A ( Xcfd080-Xbarc171-6A ) 10917.

1.66. Response to tissue culture⌂ Home

Qtcr.ipk-2B.1
SynonymTcr-B1 84
Marker associationsWeakly associated with Xpsr102-2B 84.
Qtcr.ipk-2B.2
SynonymTcr-B2 84
Marker associationsClosely linked and distal to Ppd-B1 84.
Qtcr:ipk-2B.3
SynonymTcr-B3 84
Marker associationsLinked with Yr7/Sr9g 84.
QGpp.kvl-2A
Chromosome2AL
v: Ciano / Walter DH mapping population. The green plant percentage was contributed by Ciano 0253.
QGpp.kvl-2B.1
Chromosome2BL
v: Ciano / Walter DH mapping population. The green plant percentage was contributed by Ciano 0253.
QGpp.kvl-2B.2
Chromosome2BL
v: Ciano / Walter DH mapping population. The green plant percentage was contributed by Ciano 0253.

1.67. Response to vernalization⌂ Home

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 a 290 kb 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.

VRN-1
Synonymous with TaVRT-1 10019 Orthologous series in long arms of chromosomes of homoeologous group 5. VRN-1 is a MADS-box gene similar to Arabidopsis APETALA1 10014. Spring types are associated with mutations in the promoter or the first intron {10014, 10198, 10202, 10288}. Reduction of VRN-1 transcripts in transgenic hexaploid spring wheat delays flowering 10300.

VRN-A1

Vrn-A1a
SynonymVrn1 1172, Sk 2
5AL 775, 883.
Vrn-A1b
VarietiesMarquis PI94548 (GenBank AY616461) 10198.
Type varietiesT. turgidum var. durum ST36 10198.
Vrn-A1c
NoteThis allele has a promoter similar to recessive vrn-A1a from Triple Dirk C 10198 and a large deletion in intron 1 10202.
VarietiesIL162 10198; IL369 10198 has a 5.5 kb deletion in Vrn-A1 intron 1 10202.
Type varietiesAldura PI 486150 10202; Leeds CI 13796 10202; Mexicali 75 PI 433760 10202; Minos CI 15161 10202. Most durum genotypes have a 7.2 kb deletion in intron 1 10202.
Vrn-A1d
Type varietiesT. turgidum var. dicoccoides Amrim 34 10198; FA15 (GenBank AY616462) 10198; Iraq 8736 10198; Tabigha 15 10198.
Vrn-A1e
Type varietiesT. turgidum var. dicoccum ST27 = Vernal (GenBank AY616463) 10198.
vrn-A1
Copy number variation for vrn-A1 was detected in IL369 (2 copies) 10202, Malacca (2 copies) and Hereward (3 copies). Higher copy number was associated with later flowering or with increasing requirement for vernalization (i.e. longer exposure to cold is needed to achieve full vernalization) 10881.
vrn-A1a
VarietiesClaire 10880; Triple Dirk C 10880.
Varieties (alt.)Chinese Spring Vrn-D1a 10880.
cGenBank AY616455 10198.
vrn-A1b
VarietiesIL369 10202; Malacca 10881.
cGenBank JF965396 10881. This allele has two copies of the gene, possibly arranged in tandem although the physical structure is unknown. Both copies are distinguished from Chinese Spring vrn-A1a by a SNP in exon 7 (T in Malacca, C in Chinese Spring). One copy also has a SNP in exon 4 (T in Malacca, C in Chinese Spring). Sequenced cDNAs from Malacca show that both copies are expressed 10881.
vrn-A1c
VarietiesHereward 10881.
cGenBank JF965397 10881. A comparison of Claire ( vrn-A1a ), Malacca ( vrn-A1b) and Hereward ( vrn-A1c ) indicated that increasing gene copy number is associated with lateness 10881. Two winter alleles were identified based on an SNP in exon 4 10656.
vrn-A1v
VarietiesDon Ernesto INTA 10916; Jagger 10916; Norin 61 10916; Opal 10916.
vrn-A1w
VarietiesBezostaya 10916; Bavicora M 92 10916; Kavkaz 10916; Gennson 81 10916; Seri M 82 10916; Wichita 10916.
A polymorphism between Jagger and 2174 was associated with vrn-A1a . A point mutation was present in exon 4 10695; 17 of 19 genotypes surveyed, including Jagalene, carried the 2174 mutation and only Jagger and Overley carried the Jagger allele 10695. All accessions of T. aestivum ssp. petropavlovsky shared the Vrn-A1a sequence ( Vrn-A1L ) of Langdon (7,222 bp deletion 391-7,612 bp) and some other tetraploid wheat accessions 11668.

VRN-B1

Vrn-B1
Marker associationsTsn1 – 14.8 cM – Vrn-B1 – 0.7 cM – Xwmc75-5B 10880.
vrn-B1
cGenBank AY747604.110695.
Vrn-B1a
SynonymVrn2 a = Vrn2 920, 921, Vrn2 1172, Vrn4 1173, Vrn2b = Vrn2 920 921, Ss 2
Chromosome5BL
5B 885, 920, 921; 5BL or 7BL 635, 282. 5D 635.
Vrn-B1b
VarietiesAlpowa 10695; Ciano 67 10991; Polo 10991; Yaktana 54 10991.
cGenBank FJ766015. Relative to Vrn-B1a (Triple Dirk B), Vrn-b1b has a G-C SNP at position 1656 and a 36 bp deletion at 1661-1696 10695.
Vrn-B1c
Type varietiesT. turgidum ssp. carthlicum PI 94749 10880. Much more common in durum cultivars from Russia and Ukraine than from Europe 11771.
cGenBank JN817430 contains a 5,463 retrotransposon insertion in the 5' UTR region 10880.
Vrn-B1d [11520]. Vrn-B1c 11520.
Vrn-B1f
VarietiesBarta 11523.
cHas a partially duplicated 837 bp sequence in intron 1 11523.

VRN-D1

Vrn-D1
NoteVrn3 1172
Chromosome5DL
Vrn-D1a, Vrn-d1b and Vrn-D1 were present in 27.3, 20.6 and 52.1% of 689 Chinese wheat accessions 11072.
Vrn-D1a
iTriple Dirk E 1172.
Sources / synonymsRescue* / Cadet 5A Vrn-A1a 1221.
VarietiesChinese Spring 1172; Loro 1172; Norin 61 460; Shinchunaga 460; Shirasagi Komugi 460; Ushio Komugi 460.
Varieties (alt.)Rescue Vrn-B1a 1221.
All the common wheat genotypes carrying Vrn-D1a studied so far have a 6.8 kb deletion in intron 1 (Triple Dirk E, Chinese Spring, Norin 61, Shinchunaga, Shirasagi Komugi, Ushio Komugi) 10202.
Vrn-D1b
VarietiesAdditional Chinese germplasm11072.
A Vrn-D1 variant named Vrn-D1s contained a 844 bp TE insertion in the first intron of the wild type vrnD1 allele 11628. GenBank KF800714. Identified in three T. spelta and one T. compactum accessions.
Vrn1
NoteSpring type. d
VarietiesG2528 10014.
vrn1
NoteWinter type. d
VarietiesDV92 10014; G1777 10014; G3116 10014.
cVrn1 was completely linked to MADS-box genes AP1 and AGLG1 . AP1 was considered a better candidate than AGLG1 and differences between winter and spring genotypes appeared to be related to differences in the promoter region of AP1 10014; The involvement of AP1 in vernalization response conditioned by Vrn1 was also reported in 10019.
VRN-1 genotypes in Pacific Northwest USA wheats are listed in 10695.
VRN-2
Orthologous series in chromosomes of homoeologous group 4. Vrn-A[m] 2 was located in T. monococcum 279 on chromosome 5A[m] on the 4A[m] translocated region. Vrn-A[m] 2 was mapped to the distally located Xwg114-5A - Xwec87-5A region 0312. Vrn-H2 ( sh / sgh1 ) occurs in barley chromosome 4H 1455 and is probably orthologous to Vrn-A[m] 2 based on comparative maps {279, 767}. Vrn-2 is a zinc-finger/CCT domain transcription factor ( ZCCT1 ) 10299, and repressor of flowering down-regulated by vernalization and short days 10301. Reduction of Vrn-2 transcripts in transgenic hexaploid winter wheat accelerates flowering 10299. A triple Vrn2 mutant (PI 676269), synthetic vrn2-null ) is available in hexaploid wheat combining the non-functional vrn-A2 allele present in most polyploid wheats with a Vrn-B2 deletion from tetraploid wheat, and a non-functional vrn-D2 allele from Ae. tauschii 11124.

VRN-A2

Vrn-A2a
NoteWinter habit, dominant in diploid wheat 279.
dvG1777 279; G3116 279.
Vrn-A2b
NoteSpring habit, recessive in diploid wheat.
dvDV92 279; PI 355517 10299; PI 345242 10299; PI 352475 10299; PI 277137 10299. Contains a non-functional mutation in the CCT domain 10299.
Vrn-A2c
NoteSpring habit, recessive in diploid wheat.
dvPI 352484 10299; PI 323437 10299; PI 286068 10299; PI 591871 10299; PI 542474 10299; PI 428175 10299; PI 237659 10299; PI 221329 10299; PI 225164 10299; PI 377662 10299; PI 377648 10299;
PI 362610 10299. Complete deletion of the ZCCT1 gene 10299.

VRN-B2

Vrn-B2
Chromosome4BL
A study of winter wheats 2174 and Jagger showed that 2174 has a tandem repeat of Vrn-B2 whereas Jagger has a deletion of this gene 11163. Identical apparently functional sequences of Vrn-B2 were found in contig sequences of Chinese Spring obtained from chromosomes 4BS, 2BS and 5DL 11163.
Vrn-D2
Chromosome4DL
VRN3 [ Vrn3 1398]. Orthologous series in chromosomes of homoeologous group 1 predicted from orthology with Vrn-H3 ( Sh3 ) in barley chromosome 1H 1455,1316. Aneuploid and whole chromosome substitution experiments showed that all group 1 chromosomes of wheat carry genes affecting response to vernalization 773. In both wheat and barley VRN-3 is completely linked with a flowering promoter gene homologous to Arabidopsis FLOWERING LOCUS (FT) 10421. Polymorphisms in the A and D genome copies of TaFT are associated with variation of earliness components in hexaploid wheat 10533.
Vrn-A3
Chromosome7AS
An earlier variant of T. dicoccum line TN28 was caused by a novel allele. Line TN26 lacked a 7 bp insertion, including a cis-element GATA box, in the Vrn-A3 promoter region 11370.

VRN-B3

Vrn-B3
SynonymVrn-4B 279, Vrn5, eHi 769, 779, 771
7BS 769, 768, 771.
vrn-B3
VarietiesChinese Spring Vrn-D1 (GenBank DQ890162) 10421. TaFT are associated with variation of earliness components in hexaploid wheat 10533.
Vrn-H3
SynonymSh3
Marker associationsCompletely linked to HvFT and 1 cM distal to Xabc158 on 7HS. Originally mapped incorrectly on 1H based on loose linkage {1455, 1316}.

VRN4

Vrn4
SynonymVrn5 769, 771, Vrn-D5 10004
Chromosome5DL
Chromosome5D
iTriple Dirk F 10711. v: v2: CS (Hope 7B) VrnD1a 768; Gabo Vrn-B1a 1172; IL47/ Vrn-A1a 10005; Common in T. sphaerococcum accessions 11123.
Chromosome binCentromeric region.
Marker associationsXgdm3-5D – 11.5 & 4.5 cM – Vrn4 10004; Located in a 1.8 cM interval flanked by markers Xcfd78-5D and Xbarc205-5D 10711.
cVrn4 is located in a translocated segment from chromosome 5A containing a modified copy of Vrn-A1 11123.
Eight land races with only Vrn4 were detected in 10003; others combined Vrn4 with other Vrn genes. Stelmakh 1424 doubted the existence of Vrn4 . Goncharov 10108 confirmed the existence of Vrn4 but failed to confirm its location on chromosome 5D. References to additional studies are given in 1424. Vrn4 was mapped on the centromeric region of 5D. Incorrect TDF seed stocks generated confusion about Vrn-D4 existence 10711. Eight land races with only Vrn4 were detected in 10003; others combined Vrn4 with other Vrn genes. Stelmakh 1424 doubted the existence of Vrn4 . Goncharov 10108 confirmed the existence of Vrn4 but failed to confirm its location on chromosome 5D.

VRN-D5

Vrn-D5a
SynonymVrn4 1172, Vrn-D5 10004
Chromosome5D
Chromosome5DL
iTriple Dirk F 1172.
Varieties (alt.)Gabo Vrn-B1a 1172; IL47 Vrn-A1a 10005.
Marker associationsXgdm3-5D – 11.5 & 4.5 cM – VrnD5a 10004.
Eight landraces with only Vrn-D5a were detected in 10003; others combined Vrn-D5a with other Vrn genes. Stelmakh 1424 doubted the existence of Vrn-D5a . Goncharov 10108 confirmed the existence of Vrn-D5a but failed to confirm its location on chromosome 5D. References to additional studies are given in 1424. Three genes up-regulated by vernalization were cloned from T. monococcum 10531. These were VIN3like genes similar to Arabidopsis VIN3.
Vil-1
NoteGenBank DQ886919 10531.
Marker associationsT. monococcum chromosome 5A[m ] 10531.
Vil-2
NoteGenBank DQ886917 10531.
Marker associationsT. monococcum chromosome 6A[m ] 10531.
Vil-3
NoteGenBank DQ886918 10531.
Marker associationsT. monococcum chromosome 1A[m ] 10531.
A QTL analysis of Courtot / CS is reported in 0132. A QTL on chromosome 5BL was linked to Xgwm604-5B (this QTL explained 11% of the variance in flowering time) 10075 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.ocs-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 R[2] 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.

1.68. Restorers for cytoplasmic male sterility⌂ Home

RF1

Rf1
Note1A 1619, 873, 1224}.
Chromosome1AS
VarietiesL22 868; ( T. timopheevii / Aegilops squarrosa) // 3[*] Dirk 1619.
Varieties (alt.)T. timopheevii / 3[*] Marquis Rf2 823; R113 Rf4 873. The second gene referred to as Rf4 1619 in the last stock was located in chromosome 7D, but its relationship to Rf2 in 823 was unknown.

RF2

Rf2
Chromosome7D
VarietiesT. timopheevii / 3[*] Marquis Rf1 823.

RF3

Rf3
Chromosome1BS
Chromosome1B
VarietiesR18 10222; R9034 10222; T. spelta var. duhamelianum 1453.
Varieties (alt.)Primepi Rf7 11476.
Marker associationsXcdo388-1B – 1.2 cM – Xabc156-1B 9934; RFLP markers Xcdo4421B and Xbcd249-1B were found to be associated with Rf3 on 1BS860; Mapped as a QTL in the region Xbarc207-1BSXgwm131-1BLXbarc61-1BL in crosses R18/ND36 and R9034/ND36 10222; Xbarc128-1Bs – 5.3 cM – IWB14060 – 2.0 cM – Rf3/IWB72107/IWB73447 – 14.5 cM – Xwmc406-1B 11476.

RF4

Rf4
SynonymRf2 1619
Chromosome1BS
6B 1619, 873.

RH5

Rf5
SynonymRf3 1619
Chromosome6D
Varieties( T. timopheevii / Aegilops squarrosa ) / 3[*] Canthatch Rf4 1619.

RF7

Rf7
SynonymRf6 11473
Chromosome5DS
Varieties (alt.)Primepi Rf3 11474.

RF9

Rf9
Chromosome6AS
VarietiesGerek 79 11475.
Marker associationsIWB72413-6A – 4.3 cM – Rf9 – 4.7 cM – IWB1550- 6A 11475.
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.
Vi
Chromosome1B
Chromosome1BS
VarietiesT. turgidum 867. Probably derived from a cv. Selkirk ( T. aestivum ) line with Ae. cylindrica cytoplasm 867.
Rfd1
Chromosome7BL
VarietiesChinese Spring 1047.
A different system of restoration occurs in cv. Norin 61 where at least four chromosomes, 4A, 1D, 3D and 5D, appear to be involved 1046. Several Japanese wheats carry a similar or equally effective gene combination 0335.
Rf[multi]
Chromosome1BS
VarietiesAll common wheat genotypes except T. spelta var. duhamelianum 11477.
Marker associationsAll common wheat genotypes except T. spelta var. duhamelianum 11477.
rf[multi]
VarietiesT. spelta var. duhamelianum 11477.
Sterility in the same plasmons is also found in lines with the 1BL.1RS translocation. Hohn & Lukaszewski 11478 produced a chromosome 1B1:6 translocation with a short rye insert replacing the Rf[multi] region ( rf[multi] equivalent) causing partial sterility that can be restored by most common wheat genotypes.

1.69. Ribosomal RNA⌂ Home

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.

5S rRNA genes

5S-Rrna-A1
Synonym5SDna-A1 295
Chromosome1AS
VarietiesCS 1043.
5S-Rrna-B1
Synonym5SDna-B1 295
1BS 39, 295.
5S-Rrna-D1
Synonym5SDna-D1 295
Chromosome1DS
1D 295, 758.
5S-Rrna-E1
Synonym5SDna-E1 960
Chromosome1E
dvL. elongatum .
5S-Rrna-R1
Synonym5SDna-R1 1206
1RS 1206, 39.
5S-Rrna-Sc1
Synonym5SDna-Sc1 960
Chromosome1S
alElymus ciliaris .
5S-Rrna-S[t] 1. [ 5SDna-S[t] 1 960]. 1S[t] 1290.
5S-Rrna-Y1
Synonym5SDna-Y1 960
1[Y] 1290.
5S-Rrna-A2
Synonym5SDna-A2 295
5AS 658,{ 295}.
5S-Rrna-B2
Synonym5SDna-B2 295
Chromosome5BS
VarietiesCS.
5S-Rrna-D2
Synonym5SDna-D2 295
Chromosome5DS
5D 295, 758.
5S-Rrna-R2
Synonym5SDna-R2 1206
Chromosome5RS
alS. cereale .
5S-Rrna-H[t] 2. [ 5SDna-H[t] 2 960]. 5H[t ] 1290.
5S-Rrna-U2
Synonym5SDna-U2 295
Chromosome5U
alAe. umbellulata .
5S-Rrna-V2
Synonym5SDna-V2 960
Chromosome5V
alD. villosa .
5S-Rrna-H3
Synonym5SDNA-H3 793
Chromosome2H
Chromosome2HL
alBetzes Barley; Sultan barley.
5S-Rrna-H4
Synonym5SDNA-H4 793
Chromosome3HL
alBetzes barley; Sultan barley.
5S-Rrna-H5
Synonym5SDNA-H5 793
Chromosome4HL
alBetzes barley; Sultan barley.
5S-Rrna-H6
Synonym5SDNA-H6 793
Chromosome4HS
alBetzes barley; Sultan barley.

1.70. Segregation distortion⌂ Home

See also, Gametocidal Genes.

QSd.ksu-1D
Chromosome1DL
dvAe. tauschii var. meyeri acc. TA1691; Ae. tauschii var. typica acc. TA1704 9925.
Marker associationsAssociation with Xcmwg706-1D 9931.
QSd.ksu-3D
Chromosome3DS
dvAe. tauschii var. meyeri acc. TA1691; Ae. tauschii var. typica acc TA1704 9925.
Marker associationsAssociation with Xwg177-3D 9931.
QSd.ksu-4D
Chromosome4DS
dvAe. tauschii var. meyeri acc. TA1691/var. typica acc. TA1704 9925.
Marker associationsAssociation with XksuF8-4D 9931.
QSd.ksu-5D.1
Chromosome5D
dvAe. tauschii var. meyeri acc. TA1691; Ae. tauschii var. typica acc. TA1704 9925.
Marker associationsAssociation with Xcdo677-5D 9931.
QSd.ksu-5D.2
Chromosome5DL
dvAe. tauschii var. meyeri acc. TA1691; Ae. tauschii var. typica acc. TA1704 9925.
Marker associationsAssociation with Xglk614-5D (synonym ' Xtag614-5D ') 9931.
QSd.ksu-5D.3
Chromosome5DL
dvAe. tauschii var. meyeri acc. TA1691; Ae. tauschii var. typica acc. TA1704 9925.
Marker associationsAssociation with Xwg1026-5D 9931.
QSd.ksu-7D
Chromosome7DS
dvAe. tauschii var. meyeri acc. TA1691; Ae. tauschii var. typica acc. TA1704 9925.
Marker associationsAssociation with Xglk439-7D (synonym ' Xtag439-7D ') 9931.
Vsr1
Chromosome5DL
Marker associationsXwmc765-5D – 7.7 cM – Vsr1 – 1.1 cM – Xbarc144-5D 11014; Xwmc765-5D – 1.9 cM – XWL938 – 3.3 cM – XWL2506 – 3.3 cM – Vsr1 – 0.5 cM – XWL954 – 0.5 cM – Xbarc144-5D 11014.
Vsr1a
VarietiesChinese Spring 11014.
Vsr1b
VarietiesTA4152-71 11014.

1.71. Soft glumes⌂ Home

SOG was considered to be an homologue of TG1 and TG2 . See Tenaceous glumes. The Soft Glume locus was not considered to be an orthologue of Tenacious Glumes 10769.

SOG

Sog
dvT. monococcum ssp. boeoticum ID49 10555; T. monococcum ssp. aegilopoides TA4342-96 10769.
sog
Chromosome2AS
Chromosome binC-2AS5-0.78.
dvT. monococcum ssp. monococcum var. sinskajae (syn. aegilopoides ) UD69 10555; Tm-9, a mutant of TA4342-9610769.
Marker associationsCo-segregation with AFLP loci Xe4232041 and Xe373311 10555; Xgwm71-2A – 3.3 cM – sog – 3.5 cM – Xbcd120-2A 10769.

1.72. Sterol esterification in kernels - Synthesis of b-sitosterol esters⌂ Home

Two sterol-ester phenotypes, p-L (palmitate + linoleate) and L (linoleate) are inherited as alleles at a single locus.

Pln
SynonymP-L 428
Chromosome7DS
VarietiesAradi 428; Aragon 03 428.
pln
SynonymL 428
VarietiesMara 428; Pane 247 428.

1.73. Stem solidness⌂ Home

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.

Qsst.msub-3BL
Chromosome3BL
Chromosome bin3BL11-0.81-1.00.
VarietiesFortuna 11230; Genou 11230; Judee 11230; Rampart PI 59388 10206; Rescue 11230; S-615 11230; Vida 11230.
Marker associationsLinked to microsatellite markers Xgwm247-3B, Xgwm340-3B , and Xgwm547-3B . These markers explained 76% of the total variation for stem solidness in Rampart/Jerry 10206.
Haplotype analyses in a range of hexaploid and tetraploid accessions suggested the possibility of multiple alleles or loci in the QTL region {11230, 11239}. Conan with an intermediate level of stem solidness represents a different haplotype from other North American cultivars 11230. Stem solidness in chromosome 3B of Golden Ball was verified in Langdon-Golden Ball disomic substitution lines 10730.
Qsst.msub-3DL
SynonymQss.msub-3DL 10395
Chromosome3DL
Associated with Xgwm645-3DL (R[2] =0.31), Xwmc656-3DL (R[2] =0.1), and Xcfd9-3DL (R[2] =0.13) 10395. This gene acted as an enhancer of Qsst.msub-3BL 10395.
Qsf.spa-3B
Type varietiesKyle*2 / Biodur (solid stem) // Kofa (hollow) DH population: Qsf.spa-3BL was located to a 21.3cM interval flanked by Xgwm247-3B and Xgwm114-3B 10351. Mapped as a single gene, Xgwm247-3B – 6.9cM – Qsf.spa-3B –.4cM – Xgwm114-3B 10351. This location was confirmed in two other crosses involving G9580B-FE1C and Golden Ball as the solid stem parents 10351.

1.74. Temperature-sensitive winter variegation⌂ Home

This phenotype involves reduced vigour and chlorotic patches on leaves of certain genotypes in Ae. umbellulata cytoplasm when grown at low temperatures 1596.

Vgw
NoteVariegation is dominant 1596.
SynonymVg 1021
Chromosome5BL
VarietiesBersee 1596; CappelleDesprez 1596; Hobbit Sib 1596; Mara 1596.
vgw
VarietiesBesostaya I 1596; CS 1596; Poros 1596; Sava 1596; T. spelta 1596.

1.75. Tenacious glumes⌂ Home

TG1

Tg1
NoteDerived from Ae. tauschii . Dominant.
SynonymTg 1240
Chromosome2DS
Chromosome bin2BS-3 1.00-0.84.
VarietiesSynthetic ABD wheats 652.
Marker associationsPlaced in a 12 cM interval between Xwmc112-2D and Xbarc1682D 10497.

TG2

Tg2
NoteDerived from T. dicoccoides
Chromosome2BS
VarietiesTA 3419 = Tetra Canthatch / Ae. tauschii ssp. meyeri TA159910769.
Marker associationsTg2 is associated with Xrsq805(Embp)-2B and Xpsr899-2B 0046; Xgwm261/Xwmc503-2D – 2.3 cM – Tg2 – 5.9 cM – Xfba88/Xfbc400-2D 10769.
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.

1.76. Tiller inhibition / Tiller number⌂ Home

TIN1

tin1
NoteRestricted tiller number is recessive1212
VarietiesIsrael Uniculm 494 1212; Banks + tin 10193; Oligoculm 390 10193; Uniculm 492 10193.
Marker associationsXpsp2999(Glu3)-1A – 3.9 cM – tin1/Xgwm136-1A – 2.4 cM – Xwhs179-1A 10193; the 350 bp allele of Xgwm136-1A was diagnostic of tin1 10193.
cA candidate gene identified by map-based cloning was predicted to encode a cellulose synthase-like (Csl) protein with homology to members of the CslA clade; allelic variation was attributed to dinucleotide repeat-length polymorphism in the 5 ́UTR region of the Csl gene 11657.
A near recessive gene, ftin , was located proximal to the TIN1 locus in putative Agropyron cristatum derivative Pubing355811462: Xgwm136-1A – 4.3 cM – Xpsp2999-1A – 0.7 cM – Xcfa2153-1A – 1.0 cM – FTIN 11462. TIN2. 2A 9909. Tin2 Tiller-reducing affect of this allele was dominant 9909.
tin2
SynonymTin 9909
Varieties88 F2 185 9909.
TIN3. A[m] L 10329.
tin3
dvT. monococcum TA4443 = TA4342-96 mutant 10329.
Marker associationsXbcd131/Xbcd14313A – 9.6 cM – tin3/Xpsr1205-3A – 4.7 cM – Xcfa2076-3A 10329.
TIN4 . 2DL {11574, 11575}.
Tin4
NoteQLtn.sicau-2D 11574. Low tillering.
iH461/Chuannong 16 NIL7A 11575.
VarietiesH461 {11574, 11575}.
tin4 . High tillering. QTL Fukuho-Komugi / Oligoculm: DH population: A QTL of large effect on spike number per plant in a DH population of mapping to the HgXpsp2999(Glu3)-1A region 10218 probably corresponds to Tin1 . Three QTL were located on chromosomes 2DL ( Qltn.siau-2D ), 2BL and 5AL in a RIL population from H461 (low tillering) / CN16 (high tillering) , but only the 2DL QTL was confirmed in H461 / CM107 and H461 / MM37 RIL populations 11465.

1.78. Yield and yield components⌂ Home

QGnu.ipk-4A
Note4AL 0255 v: Opata / W-7984 (ITMI) RI mapping population 0255; Higher grain number was contributed by Opata 0255.
Marker associationsAssociated with Xmwg549-4A, Xabg390-4A and Xbcd1670-4A 255.
QGnu.ipk-4A coincides with QTL for height ( QHt.ipk-4A ), spike length ( XEl.ipk-4A ) and grain weight per ear ( QGwe.ipk-4A ) 0255.
QKps.unl-3A.1
Chromosome3AS
v: Cheyenne / Cheyenne(Wichita 3A) RI mapping population10044; a higher kernel number of 0.3 kernels was contributed by Wichita and the QTL explained 15.5% of the phenotypic variation 10044; The QTL coincided with QTLs for grain yield, kernel number per square metre and 1000-kernel weight 10044.
Qkps.unl-3A.2
Notev: Cheyenne / Cheyenne(Wichita 3A) RI mapping population 10044; a higher kernel number of 0.3 kernels was contributed by Cheyenne and the QTL explained 9.5% of the phenotypic variation10044.
Marker associationsAssociated with Xbcd141-3A 10044.
Three QTL for kernel number per spike were assigned to chromosome 3A in RSLs from Cheyene[*] 7/Wichita 0025.
QGvwt.unl-3A.1
Chromosome3A
v: Cheyenne / Cheyenne(Wichita 3A) RI mapping population 10044; higher grain volume weight (+23 kg/hL) was contributed by Wichita and the QTL explained 43.1% of the phenotypic variataion 10044; the QTL coincided with a QTL for spikes per square metre 10044.
QGwe.ocs-4A.1
Chromosome4AS
Marker associationsAssociated with Xbcd1738-4A 0047.
In Opata / W-7984 (ITMI): RI mapping population:
QGwe.ipk-2D
Chromosome2DS
Higher grain weight was contributed by Opata 0255.
QGwe.ipk-4A
Chromosome4AL
Higher grain weight was contributed by Opata 0255.
QYld.ocs-4A.1
Chromosome4AS
Marker associationsAssociated with Xbcd1738-4A 0047.
Cheyenne / Cheyenne(Wichita 3A): RI mapping population:
QGyld.unl-3A.1
Chromosome3AS
Higher grain yield of 32 kg/ha was contributed by Wichita and the QTL explained 6.6% of the phenotypic variation 10044; The QTL coincided with QTLs for kernel number per square metre, 1000-kernel weight and kernels per spike 10044;
QGyld.unl-3A.2
Chromosome3A
Higher grain yield of 82 kg/ha was contributed by Wichita and the QTL explained 28.1% of the phenotypic variation 10044; The QTL coincided with a QTL for kernel number per square metre 10044.
QYld.inra-7D
Marker associationsXcdf69-7D (R[2] =3.7-15.7%) 10071.
RL4452 / AC Domain:
QYld.crc-2A
Marker associationsLinked to Xgwm339-2A (LOD 3.0) 10287.
QYld.crc-2B
Marker associationsLinked to Xgwm257-2B (LOD 9.4) 10287.
QYld.crc-4A
Marker associationsLinked to Xgwm130-4A (LOD 4.4) 10287.
Shamrock / Shango: DH population: Non-glaucous (virescent) lines had higher yields than glaucous sibs 10543; see Glaucousness, subsection Epistatic inhibitors of glaucousness. ## Tetrapoloid wheat Langdon(DIC5B) / Langdon:
QYld.ndsu-5B
SynonymQGy.ndsu-5B 10161
Higher yield contributed by contributed by Langdon 10161.
QKpsm.unl-3A.1
Chromosome3AS
Higher kernel number (170 kernels) was contributed by Wichita and the QTL explained 14.6% of the phenotypic variation 10044; The QTL coincided with a QTL for grain yield10044.
QKpsm.unl-3A.2
Chromosome3A
Marker associationsAssociated with Xbarc673A 10044.
Ful2
NoteLoss of function mutation in gene FUL-A2 (Kronos mutant T4-837) and FUL-B2 (Kronos mutant T4-2911) resulted in significant increases in spikelet number 11384. Flowering Locus T2 FT2 Loss-of-function mutations in FT2 increased spikelet number per spike but reduced but reduced fertility 11604. FT-A2 . TraesCS3A02G143100 .
Chromosome3AL
A natural mutation causing an aspartic acid (
WAPO-A1
NoteTraesCS7A02G481600 11383.
Chromosome7AL
WAPO-A1 is the causal gene for QSNS.ucw-7A 11383 that also affects grain number per spike and spike yield 11603.
WAPO-A1a
NoteAssociated with low SNS; has a 115-bp promoter deletion and a D384N amino acid change 11383.
VarietiesRAC875, Clark, Lancer, CDC Lanmark, Julius, Arina, Jagger, Cadenza, Robigus, and SY_Mattis 11383.
Type varietiesKronos, Ben 11383. Most frequent allele in durum 11383.
WAPO-A1b
NoteAssociated with high SNS; has a C47F amino acid change and no promoter deletion 11383.
VarietiesBerkut, Ning7840, PI 41025, MPV57, Platte 11383. Most frequent allele in T. aestivum .
Type varietiesRare in durum 11383.
WAPO-A1c
NoteAssociated with low SNS; has the ancestral C47 and D384 amino acids and no promoter deletion 11383.
VarietiesT. dicoccoides PI 471033 and PI 355455; T. dicoccum CItr 14135, PI 94638, and PI 298586; T. durum PI 286539 11383.
WAPO-A1d 11383 . Associated with low SNS; has the ancestral C47 and D384 amino acids and no promoter deletion but differs from WAPO-A1c by a C667 and G764A DNA changes 11383. tv : T. durum Rusty and Lang; T. dicoccum CItr14919, PI 193877, PI 193882, PI 217640, PI 221400, PI 225332, PI 273980, and PI 94657 11383.
QEl.ipk-1B
Chromosome1BL
Longer ear was contributed by Opata 0255.
QEl.ipk-4A
Note4AL 0255 Longer ear was contributed by Opata0255.
Marker associationsAssociated with Xmwg549-4A, Xabg390-4A and Xbcd1670-4A 0255. This QTL is likely to be a pleiotropic effect of the gene underlying the height QTL, QHt.ipk-4A 0255;
QEl.ipk-5A
Chromosome5AL
Longer ear was contributed by W-7984 0255.
QTn.ocs-4A.1
Chromosome4AS
Marker associationsAssociated with Xpsr163-4A 0047.

50-grain weight

QFgw.ocs-4A.1
Chromosome4A
v: CS / CS(Kanto107 4A) mapping population0047.

1000-grain weight

TaCKX6-D1
Chromosome3D
Encodes a cytokinin oxidase/dehydrogenase.
TaCwi-A1
Marker associationsXbarc15-2A – 10.9 cM – TaCwi-A1 – STS markers Cwi21 and Cwi22 – 17.6 cM – Xgwm71-2AL 10812.
TaCwi-A1 was cloned based on the rice GIF1 gene encoding a cell wall invertase (GenBank accession EU095553), and STS markers Cwi21 and Cwi22 were developed from the polymorphisms between two allelic variants. On average, TaCwi-A1a had 1,000 grain weights 2.4 g higher than TaCwi-A1b 10812.
TaGW2-6A
Marker associationsXcfd80-6AS.2 – 0.6 cM – TaGW2-CAPS – 0.5 cM – Xbarc1466A.1/Xwms132.4-6A 10781.
Based on its OsGW2 orthologue in rice this gene was characterized and mapped as a CAPS marker in wheat 10781. SNPs in the promoter region allowed distinction of two haplotypes. Hap-6A-A was mainly present in southern Chinese wheats; Hap6A-G was present in varieties from central and eastern Europe. On average Hap-6A-A had 1,000 grain weights more than 3g higher than Hap-6A-G 10781. Encodes an E3 RING ligase 11122. QTL Cheyenne / Cheyenne(Wichita 3A): RI mapping population:
QTkwt.unl-3A.1
Chromosome3AS
Higher kernel weight of 0.27% was contributed by Cheyenne and the QTL explained 12.7% of the phenotypic variation 10044; The QTL coincided with QTLs for grain yield, kernel number per square metre and kernels per spike 10044.
QTgw.ipk-5A
Chromosome5AL
Varieties}; The higher yielding allele was contributed by W-7984 255.
Marker associationsAssociated with Xfba351-5A and Xcdo1312-5A 255.
RS111 / CS: mapping population:
QGw1.ccsu-1A
Chromosome1AS
Marker associationsAssociated with Xwmc333-1A 0165.
RS111/CS: RIL population 0236: QTLs for grain size were identified on chromosome arms 1DS, 2DL and 6BL. RL4452 / AC Domain:
QGwt.crc-3D
Chromosome3D
Marker associationsFlanked by Xgwm341-3DXwmc552-3D (LOD 4.3) 10287;
QGwt.crc-4A
Chromosome4A
Marker associationsFlanked by Xgwm494-4AXgwm162-4A (LOD 6.7) 10287;
QGwt.crc-6D
Chromosome6D
Marker associationsFlanked by Xgwm3256DXgwm55-6D (LOD 3.9) 10287.
QTwt.crc-1B
Chromosome1B
Marker associationsLinked to Xgwm374.1-1B (LOD 3.9) 10287.
QTwt.crc-1D
Chromosome1D
Marker associationsLinked to Xgdm126-1D (LOD 5.8) 10287.
QTwt.crc-2D
Chromosome2D
Marker associationsLinked to Xgwm349-2DXbarc59-2D (LOD 5.2) 10287.
QTwt.crc-3B
Chromosome3B
Marker associationsLinked to Xwmc625-3BXbarc164-3B (LOD 15.4) 10287.
QTwt.crc-3D
Chromosome3D
Marker associationsLinked to Xbarc71-3D (LOD 5.2) 10287.
QTwt.crc-5D
Chromosome5D
Marker associationsLinked to Xgdm63-5DXwmc765-5D (LOD 5.3) 10287.

SVP1 and VRT2

QSns.sau-2DS
Chromosome2DS
VarietiesRIL populations from Line 20828 / Chuanong 16 , Line 20828 / Shumai 51 and Line 20828 / Sy95-71 ; LOD score 3.47 – 38.24, PV 10 – 46% in 8 environments. Located in a 2 cM interval flanked by Ax-109836946 (32.8 Mb) and AX-111956072 (34.43 Mb) 11424.
QSns.ucw-7AL
Chromosome7AL
VarietiesUnderlying gene: WAPO1 .
CS / CS(Kanto107 4A): mapping population:
QSpn.ocs-4A.1
Chromosome4AS
Varieties0047.
Marker associationsAssociated with Xbcd1738-4A 0047.
QSn.sdau-4BL
Marker associationsXwmc657-4B – 4.6 cM – QSn.sdau-4B – 1.6 cM – Xgwm495-4B 10784.
QSn.sdau-4BL was resolved as a single gene in 10784. It was associated with decreased spike length and grain number per spike.

GrainGenes is a product of the Agricultural Research Service of the US Department of Agriculture.