1.Gross Morphology: Spike characteristics

Major hexaploid wheat types are categorized into groups with respect to three major gene pairs; viz. Q, C and S1 {1038}.
1. Common wheat Q c S1 v: vulgare group.
2. Club wheat Q C S1 v: compactum group.
3. Shot wheat Q c s1 v: sphaerococcum group.
4. Spelt wheat q c S1 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 c S1 {1049}.
1.1. Squarehead/spelt

Q{881}. [k{1550}]. 5AL{1293}. v: Common wheats. CS. ma: Complete linkage with cDNA clone PtAq22{0127}.

q{881}. [K{1550}]. v: Macha wheats; spelt wheats; vavilovi wheats. s: CS^*8/White Spring Spelt 5A{1048}. ma: Cent - Xrsq805(Empb)-5A - 4.6cM - Q - 4.3cM - Xpsr370-5A{419}; Q was physically mapped in 5AL, fraction length 0.87, bracketed by deletions 5AL-7 and 5AL-23{446}; Q - 9.3cM - Xpsr370-5A{9903}.
The speltoid phenotype of at least some spelts may be caused by genes at other loci {0140}. Fine mapping of the 20cM region possessing Q and delimited by deletions 5AL -7 and -23 is reported in {0324}.
1.2. Club

C{1517}. Although gene C may be present in some forms of group macha {1447}, it is not universally present. Tsunewaki {1500} found that compact spike in one form was controlled by polygenes. [Cd{047}]]. 2D{1192}.2DL{1192,1517}. i: S-615^*11/Elgin{1500}. s: CS^*6/Poso 2D{1304}; CS^*5/Red Egyptian 2D{1304}. v: Club wheats. QTL: Six QTLs for spike compactness were detected in Courtot/Chinese Spring but only 4 on chromosome arms 1AL, 2BS, 2DS and 4AS were consistent for at least two years {0114}.
Although gene C may be present in some forms of group macha {1447}, it is not universally present. Tsunewaki {1500} found that compact spike in one form was controlled by polygenes.
1.3. Sphaerococcum

The naturally-occurring sphaerococcum gene in chromosome 3D and various mutant alleles conferring a similar phenotype form a homoeologous series. The sphaerococcoid alleles are either recessive or incompletely dominant. All three mapped loci are closely linked to the respective centromeres {0030}. The "a" alleles are allocated to Chinese Spring or "normal" wheats.

s2. Partially dominant{1286}. [sp2{1286}]. v: Sphaerococcoid 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.

S-A1{0029}. 3A{0056}. v: CS{0029}.

S-A1a{0029}. v: CS{0029}; common wheats{0029}.

S-A1b{0029}. [S3{0056}]. v: MS 1453{0056}. ma: Xgwm2-3A(S) - 5.1 cM - S-A1 - 6.6 cM - Xgwm720-3A(L){0030}.

S-B1{0029}. 3B{0030}. v: CS{0029}.

S-B1a{0029}. v: CS{0029}; common wheats{0029}.

S-B1b{0029}. [S2{0030}]. v: MSK 2452{0056}; MSK 2454{0056}. ma: Xgwm685-3B(S) - 4.2 cM - S-B1 - 0.5 cM - Xgwm566/Xgwm845/cent{0030}.

S-D1{0029}. 3D{1292,0030}.3DS{1193,1194}.3DL{692}. v: CS{0029}.

S-D1a{0029}. v: CS{0029}; common wheats{0029}.

S-D1b{0029}. [s1, sp1{1286}]. i: S-615^*11/T. sphaerococcum var. rotundatum{1500}. s: CS^*7/T. sphaerococcum rubiginosum 3D{1304}. v: Sphaerococcum wheats{0029}.

S-D1c{0029}. [S1{0056}]. v: MS 3287{0056}. ma: Xgdm72-3D(S) - 8.0 cM - S-D1 - 2.9 cM - Xgwm456-3D/cent{0030}.
1.4. Branched spike

Synonymns: branched head, four-rowed spike, supernumerary spikelet, tetrastichon spikelet.

bh{665}. 2AS{665,9907}. dv: PI 349056{665}.
A chromosome 2B gene of minor effect was identified{9907} and an inhibitor was associated with chromosome 2D{9907}. In a monosomic analysis of the hexaploid line LYB with supernumerary spikelets, Peng et al. {9908} located recessive genes in chromosome 2A and 4A that promote the development of supernumerary spikelets and a gene in chromosome 2D that prevents their expression.
1.5. Elongated glume

Elongated glume is the phenotype associated with the polonicum group of tetraploid wheats. Expression in hexaploid wheat is much reduced compared with tetraploids. Matsumura {911} reported linkage of gene P and a gene for red coleoptiles implicating chromosomes 7A or 7B. A different gene was subsequently located in chromosome 7B {9990}.

P1. [P{911},Eg{922},P-A^pol1{0254},P-A^pet1{0254}]. 7AL{922,1547}.7A or 7B (based on linkage of 0.2 with a gene for red coleoptile){922}. i: Saratovskaya29^*8//Novsibirskaya 67^*2/T. polonicum{922}. itv: P-LD222 = LD222^*11/T. turgidum var polonicum{1546,1547}. tv: T. polonicum{0254}; T. petropavlovskyi{0254}. ma: Xgwm260 - 7A(S) - 2.3cM - P1 - 5.6cM - Xgwm1083-7A(L){0254}; Xgwm890 -7A - 2.1cM - P1{0254}; Xgwm260-7AS - 2.3cM - P1^pol - 5.6cM - Xgwm1083-7AL {0254}; Xgwm890-7AS - 2.1cM - P1^pet {0254}.
Note: The loci determining elongated glumes in T. turanicum and T. durum conv. falcatum are not homoeologous to the P loci in the centromeric region of the group 7 chromosomes{0254}.

P2{9990}. 7BL{9990}. ti: LD222^*7/T. ispahanicum{9990}. tv: T. 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.
1.6. Ear length

QEl.ocs-5A.1{0068}. 5AL{0068}. v: CS(T. spelta 5A)/CS(Cappelle-Desprez 5A) RI mapping population{9903}. ma: Associated with Xbcd9-5A{0068}.
2.Accumulation of Abscisic Acid

A QTL was mapped on 5AL between Xpsr575-5A {proximal} and Xpsr426-5A {distal} {1180}.
3.Alkylresocinols Content in Grain

Ar1{0281}. High alkylresocinols content is dominant {0281}. 5AL{0281}. tv: Langdon{0281}.

ar1{0281}. tv: Ardente{0281}; This cultivar has a low content compared to all tested durum and common wheats{0281}.
4.Aluminium Tolerance

Alt1{234}. v: ET3 = Carazinho/4^*Egret{234}.

alt1{234}. v: ES3 = Carazinho/4^*Egret{234}.

Alt2{848}. [Alt_BH{1213}]. 4DL{848}. su: T. turgidum cv. Langdon 4D(4B){848}. ma: Alt2 was mapped to a 4 cM interval flanked by Xpsr914 and Xpsr1051{848}; on a consensus 4B-4D map of T. aestivum; Alt2 - 1.1 cM - Xbcd1230-4D{1213}. v: BH1146{1213,0115}; IAC-24{0115}; IAC-60{0115}; 13 induced mutants of Anahuac{0115}. ma: Alt2 cosegregated with Xbcd1230-4D and fell within the interval Xgdm125-4D - 4.8cM - Alt2 -1.1cM - Xpsr914-4D{0248}.
5.Anthocyanin Pigmentation
5.1. Purple anthers.

A single, dominant factor was reported {1326}.

Pan1{921}. 7DS{921}. v: Ilyitchevka{921}; Mironovskaya 808{921}; Novosibirskaya 67{921}; Pyrothrix 28{921}; Saratovskaya 210{921}; Strela{921}; Ukrainka{921}. tv: T. polonicum{921}.

Pan2. 7AS{9959}. tv: T. turgidum ssp. dicoccoides acc. MG4343{9959}. ma: Pan2 - 9.2 cM - Rc1 - 12.2 cM - Xutv1267-7A (proximal){9959}.
5.2. Purple/Red auricles. Purple leaf base

For review see {1641}.
Melz and Thiele {983} described a "purple leaf base" phenotype where anthocyanin pigmentation extended to the leaf base as well as auricles. Purple leaf base was expressed only when pigmentation occurred in the coleoptiles.

Ra1. [Ra{1645}]. 1D Gulyeeva{474}.(cited in{983}).2D{1645}. v: Kenya 58{1645}.

Ra2{983}. 4B{983}.

Ra3{983}. 6B{983}.

An5{983}. 5R{983}.
5.3. Red/purple coleoptiles.

There is an orthologous gene series on the short arms of homoeologous group 7. The 'a' alleles confer red coleoptiles.

Rc-A1a{0250}. [Rc1, R{401}]. 7A{769,1293}.7AS{0250}. s: CS^*6/Hope 7A{1293}. v: Hope Rc-B1{1293}. tv: T. turgidum ssp. dicoccoides acc. MG4343{9959}. ma: Pan2 - 9.2cM - Rc1 - 12.2cM - Xutv1267-7A(proximal){9959}; Rc-A1(distal) - 11.9cM - Xgwm913-7A{0250}.

Rc-B1a. [Rc2, R2{401}]. 7B{742}.7BS{401,769,0250}. s: CS^*6/Hope 7B{769}. v: Hope Rc-A1. ma: Xgwm263-7B - 26.1cM - Rc-B1 - 11.0cM - Xgwm1184-7B{0250}.

Rc-D1a{0250}. [Rc3]. 7D{596}.7DS{1241,1444,0250}. v: 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}. ma: Rc-D1 (distal) - 3 cM - Xpsr108-7D{180}; Xgwm44-7D - 6.4cM - Rc-D1 - 13.7cM - Xgwm111-7D{0250}.
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.
5.4. Purple/red culm/straw/stem.

Purple or red colour is dominant.

Pc1{743}. [Pc{743}]. 7B{743}.7BS{768}. s: CS^*6/Hope 7B{743,768}. itv: LD222^*11/CS (Hope 7B}{1546}. ma: Pc (proximal) - 5.7 cM - Xpsr490(Ss1)-7B{110}².

Pc2{921}. 7DS{921}. v: Ilyitchevka{921}; Mironovskaya 808{921}; Novosibirskaya 67{921}; Pyrothrix 28{921}; Saratovskaya 210{921}; Strela{921}; Ukrainka{921}.
5.5. Purple grain/pericarp

Genes for purple pericarp have been transferred from tetraploid wheats to the hexaploid level {112,214,941,1138}. At the hexaploid level duplicate genes {112,941} and complementary genes {112,939,1138,438} were reported. At the tetraploid level, duplicate-gene {941} and single-gene {1327} inheritances were observed. Purple colour is dominant and may be affected by environment and genetic background. Complementary genes were located in chromosomes 3A and 7B {1138}. Possible pleiotropic relationships of genes affecting pigmentation of various tissues have not been studied in detail. Pc2 and Rc-B1a may be the same gene {769}. Also, complementary genes involved in determination of purple pericarp could be related to culm colour {112}.
For review, see {1643}.
Complementary dominant genes.

Pp1{041}. 6A{041}. i: Saratovskaya 29^*8/Purple {Australia} Pp2{040}.

Pp2{041}. 7A{041}. tv: T. durum Desf. subsp. abyssinicum Vav{040}.
Piech and Evans{1138} located complementary genes on chromosomes 3A and 7B.
6.Awnedness

hd b1 b2. Bearded or fully awned genotype
6.1. Dominant inhibitors
6.1.1. Hooded

Hd{1551}. 4AS{1195,1293}. i: S-615^*11/CS{1500}. v: Chinese Spring B2{1293}. ma: Xcdo1387-4A - 8.2cM - Hd - 7.2cM - Xpsr163 - 4A{0047}.
was mapped as a QTL with a peak on Xfba78-4A in {0309}.

hd. s: CS^*6/Hope 4A; CS^*5/Thatcher 4A; CS^*6/Timstein 4A.
6.1.2. Tipped 1

B1{1551}. 5AL{1293,0242}. i: S-615^*11/Jones Fife{1500}. v: Timstein{741}; Redman{160}.
B1 was mapped as a QTL with a peak on Xwmc182-6B {0309}.

B1a{041}. s: Saratovskaya 29^*8/Festiguay 5A{041}.

B1b{041}. s: Saratovskaya 29^*8/Aurora 5A{041}.

B1c{041}. s: Saratovskaya 29^*8/Mironskaya 808 5A{041}.
In a common genetic background, carriers of B1a have the shortest tip-awned phenotype; carriers of B1b and B1c have awns 2 to 3 times longer depending on environment. In F1 hybrids, differences between the substitution line combinations are significant. The postulation of B1 in both CS and Courtot {0309} based on the phenotype of a CS deletion stock is not supported by genetic observations
6.1.3. Tipped 2

B2{1551}. 6BL{1293,1297}. i: S-615^*11/CS{1500}. v: Chinese Spring Hd{1293}.

b2. s: CS^*6/Hope 6B; CS^*5/Thatcher 6B; CS^*9/Timstein 6B.
6.1.4. Awnless

Genotypes Hd B2 (e.g., Chinese Spring) and B1 B2 (e.g., Federation) are awnless. Presumably Hd B1 is awnless. Watkins & Ellerton {1551} noted the probability of a third allele "b1a" leading to a half-awned condition, and in discussion they consider the possibility of a similar third allele at the B2 locus. In view of more recent cytogenetic analyses, it seems that the half-awned condition could result from epistatic interactions between the alleles B1 and/or B2 and various promotor genes.
Although hooded, half-awned, tip-awned and awnless variants occur among tetraploid wheats, these are relatively infrequent. It has not been established with certainty that the above inhibitors are involved.
The inhibitor alleles have a pleiotropic effect on glume-beak shape {1348}. Acuminate beak is associated with full beardedness and occurs only in b1 b2 types. B2 reduces beak length producing an acute beak shape. B1 reduces beak length producing an obtuse beak shape. In this effect B1 is epistatic to B2.
6.2. Promotors

The effects of (recessive) awn-promoting genes were documented in a number of studies, mainly through monosomic and disomic F1 comparisons, and in tetraploids, whereas Heyne & Livers {549} provided genetic evidence of their effects. A series of "a" genes was documented, but the evidence supporting the existence of at least some of these was not well supported. Hence symbols for this gene series are not recognized.
6.3. Smooth awns

Smooth-awned tetraploid wheats were reported {016,045,690,1259} and genetic analyses {016,045,690} suggested a single recessive factor, with modifiers in most instances, relative to rough awns. The phenotype has not been reported in hexaploid wheats. No gene symbol is applied.
7.Basal Sterility in Speltoids

The presence of gene Q ensures the fertility of the first and subsequent florets in wheat spikelets {378}. In speltoids lacking Q, fertility of the second and subsequent florets is ensured by the dominant allele Bs (designated A in {378}) located on chromosome 5D {377}. In the presence of Bs the fertility of the first floret is under polygenic control.
In bs bs speltoids floret development is under polygenic control, and stocks with varying levels of basal fertility were isolated.
All group vulgare genotypes so far studied carry Bs.
The following stocks were described {378}:

Genotype Approx. sterile-base score

Group vulgare ---- QQ Bs Bs 0.00

Speltoids StFF qq Bs Bs 0.00

StF qq Bs Bs 0.08

St1A qq Bs Bs 0.39

St1 qq Bs Bs 0.96

St2 qq bs bs 1.41
8.Blue Aleurone

The Ba allele in T. monococcum spp. aegilopoides acc. G3116 determines a half-blue seed phenotype and is different from the allele present in Elytrigia pontica that determines a solid blue phenotype {282}. They are treated as different genes.
For review see {1643}.

Ba1{643}. Derived from Elytrigia pontica (2n=70). [Ba{643}]. 4B[4BS-4el₂]{643}. tr: UC66049B{594}.

Ba2. 4A^mL{282}³. dv: G3116{282}. ma: Ba2 cosegregated with Xcdo1387-4A, Xmwg677-4A and Xbcd1092-4A {282}.
9.Brittle Rachis

Br1{9970}. 3DS{9970}. v: T. aestivum var. tibetanum{9970}.

Br2{0130}. 3A{0130}. sutv: LDN(DIC 3A){0130}.

Br3{0130}. 3B{0130}. sutv: LTN(DIC 3B){0130}.
Evidence for a homoeologous series extending to many related species is discussed in {0130}.
10.Boron Tolerance

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

Bo1{1111,1113}. 7B{177}. v2: Halberd Bo2Bo3.

Bo2{1111,1113}. v2: (W1^*MMC)/Warigal Bo3. Halberd Bo1 Bo3.

Bo3{1111,1113}. 4A{0012}. v2: Warigal Bo2. Halberd Bo1 Bo2.
Very sensitive genotype: Kenya Farmer bo1 bo2 bo3.
11.Cadmium Uptake
11.1. Low cadmium uptake

Cdu1{963}. [Cdu1{1128}]. dv: Biodur{1128}; Hercules{1128}; Nile{1128}.

cdu1{963}. [cdu 1{1128}]. dv: Kyle{1128}. ma: Cdu1 - 4.6cM - OPC-20{1128}; Cdu1 - 21.2 cM - UBC-180{1128}.
12.Chlorophyll Abnormalities
12.1. Virescent

V1. 3B{122,1311,1294}.3BS{1423}. v: CS.

v1a. [v{1294}]. i: S-615^*11/Neatby's Virescent{1500}. s: CS^*9/Neatby's Virescent{1304}. v: Neatby's Virescent{1055}.

v1b. i: CS^*/Hermsen's Virescent v2b{1304}. v: Hermsen's Virescent v2b{1311}.

V2. 3A{1311,1545}. v: CS.

v2a. v: Viridis 508{1545}.

v2b.Expressed only when combined with v2b i: CS^*/Hermsen's Virescent v1a{1304}. v: Hermsen'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.
12.2. Chlorina

Cn-A1. 7A{1132}.7AL{1131,1304,1311}. v: CS.

cn-A1a. [cn1a]. i: Chlorina-1{1311}.

cn-A1b. [cn1b]. i: Cornell Wheat Selection 507aB-2B-21/6^*CS{1133}.

cn-A1c. [cn2]. i: Chlorina-448. (CS background){1545}.

cn-A1d{665}. dv: CDd6{665,666}.

Cn-B1. 7BL{1131}. v: Chinese Spring{1131}.

cn-B1a{665}. dv: CDd1{665,666}; CBC-CDd1{665}.

cn-B1b{665}. dv: CDd2{665,666}.

Cn-D1. [Cn3]. 7D{1545}.7DL{1131}. v: Chinese Spring{1131}.

cn-D1a. [cn-D1,cn3]. i: Chlorina-214{1545}. v: CD3{1583}.
12.3. Striato-virescens

A mutant of this type was described {376} but has been lost.
13.Cleistogamous Flowering in Durums

Cleistogamy, a rare flowering habit in durum wheats, is controlled by a single recessive gene relative to chasmogamy {191}.
Cleistogamous genotypes clcl.tv: HI8332 {191}; WH880 {191}.
Chasmogamous genotypes ClCl .tv: IWP5308 {191}; PWB34 {191}; WH872 {191}.
14.Copper Efficiency

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

Ce{1276}. 4BL = T4BL.5RL{1276}. v: Cornell Selection 82a1-2-4-7{462}; Backcross derivatives of Cornell Selection to Oxley, Timgalen, Warigal{464}; Hairy necked Viking{1276}.
5BS = T5BS.5RL.ad:CS+5R{463}. su: CS 5R{5D}{463}. v: Sears' stock HN-2{464}; Backcross derivatives to Warigal and Timgalen{464}.
15.Corroded

co1. [co{1297}]. 6BS{1293}. v: Sears' corroded mutant.

co2. 6D{1570}. v: Kurrachee{1570}.
A gene(s) in chromosome 6A acts as an inhibitor of corroded {1039,1570}.
16.Crossability with Rye and Hordeum and Aegilops spp.
16.1. Common wheat

High crossability of some wheats, particularly those of Chinese origin, viz. Chinese 446 {790}, Chinese Spring {1216}, and TH 3929 {939}, with cereal rye, weed rye (S. segetale L.) {1646}, and other species, e.g., Aegilops squarrosa {691}, Hordeum bulbosum {1387,1397,1469} and H. vulgare {349,693], is determined by additive recessive genes. The kr genes influence crossability with H. vulgare. Allele kr1 is more potent in suppressing crossability than Kr2 which is stronger in effect than Kr3 {1387}. According to Zheng et al. {1649}, the effect of Kr4 falls between Kr1 and Kr2.

Kr1. 5B{1216}.5BL{762}.

kr2. 5A{1216}.5AL{1387}.

kr3. 5D.

kr4. 1A{1649}.

kr1 kr2. v: Chinese 446{790}; Chinese Spring{762,1216,1025}; Martonvarsari 9^*4/CS{1016}.

Kr1 kr2. s: CS^*6/Hope 5B{762,1216}. v: Blausamtiger Kolben{790}.

kr1 Kr2. s: CS^*6/Hope 5A{1216}.

Kr1 Kr2. v: Marquis{790}; Peragis{790}.

kr1 kr2 kr3 kr4. v: J-11{1649}.

Kr1 Kr2/Kr1 kr2. (heterogeneous). v: Martonvarsari 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 also occur {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 appears to coincide with known locations of Kr genes.
16.2. Tetraploid wheat

The Chinese tetraploid, Ailanmai, possesses recessive crossability genes on chromosomes 1A, 6A and 7A with the 6A gene being the least effective {0017}.
17.Dormancy (Seed)

Phs{9960}. Semi-dominant {9960}. 7D{9960}. v: Soleil{9960}. ma: Associated with Xpsr1327-4A{0346}.
QTL: Several QTL for falling number and alpha-amylase activity, two indicators for pre-harvest sprouting resistance, were identified in {0169}. The most significant were associated with Xglk699-2A and Xsfr4(NBS)-2A, Xglk80-3A and Xpsr1054-3A, Xpsr1194-5A and Xpsr918-5A, Xpsr644-5A and Xpsr945-5A, Xpsr8(Cxp3)-6A and Xpsr563-6A, and Xpsr350-7B and Xbzh232(Tha)-7B {0169}.
In cross AC Domain/Haruyutaka, one major QTL in chromosome 4AL and two lesser possibly homomeologous QTLs for dormancy in 4BL and 4DL {0226} were found. Tolerance to preharvest sprouting (PHS) in the cross SPR8198/HD2329 was associated with Xwmc104-6B and Xmst101-7D {0032}. QTL for preharverst sprouting were identified on chromosomes 3A (associated with Xfbb293-3A at P=0.01), 3B (associated with Xgwm403-3B and Xbcd131-3B at P=0.001), 3D (associated with Xgwm3-3D at P=0.001) and 5A (associated with Xbcd1871-5A at P=0.001) in the population Renan/Recital {0347}. The resistant alleles on the group 3 chromosomes and on 5A were contributed by Renan and Recital, respectively. All QTL for preharvest sprouting co-located with QTL for grain colour {0347}
18.Ear Emergence

QEet.ocs-4A.1{0047}. 4AL{0047}. v: CS/CS(Kanto107 4A) mapping population. ma: Associated with Wx-B1{0047}.

QEet.ocs-5A.1{0068}. 5AL{0068}. v: CS(T. spelta 5A)/CS(Cappelle-Desprez 5A) RI mapping population{9903}. ma: Associated with Xcdo584-5A and morphological locus Q{0068}.

QEet.ocs-5A.2{0026}. 5AL{0026}. ma: Xcdo 412-5A - Xbcd9-5A region{0026}.

QEet.ipk-2D{0255}. QEet.ipk-2D coincides with a QTL for flowering time, QFlt.ipk-2D. Both QTLs may correspond to Ppd-D1 {0255}. 2DS{0255}. v: Opata/W-7984 (ITMI) RI mapping population{0255}; Lateness was contributed by W-7984{0255}. ma: Associated with Xfba400-2D and Xcdo1379-2D{0255}.

QEet.ipk-5D{0255}. QEet.ipk-5D coincides with a QTL for flowering time, QFlt.ipk-5D. Both QTLs probably correspond to Vrn-D1 {0255}. 5DL{0255}. v: Opata/W-7984 (ITMI) RI mapping population{0255}; Lateness was contributed by W-7984{0255}. ma: Associated with Xbcd450-5D{0255}.
19.Earliness Per Se

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

Eps-1A^m{0364}. [Eps-A^m1]. 1AL{0364}. dv:: T. monococcum. DV92 allele for late flowering, G3116 early flowering.{0364}. ma:: 0.8 cM distal to Xwg241 {0364}.

Eps-A1a{0024}. 3A{0023}.3AL{0024}. v: Chinese Spring{0024}.

Eps-A1b{0024}. v: Timstein{0024}.

epsCnn{0025}. v: Cheyenne{0025}.

EpsWi{0025}. 3A{0025}. su: Cheyenne^*7/Wichita 3A{0025}. ma: Linked to QTLs for plant height, kernel number per spike, and 1,000-kernel weight in RSLs derived from CNN/CNN(WI3A){0025}.
QTL: Analysis in Courtot/CS {0132}.
20.Flowering Time

QFlt.ipk-3A{0255}. 3AL{0255}. v: Opata/W-7984 (ITMI) RI mapping population{0255}; Lateness was contributed by W-7984{0255}. ma: Associated with Xbcd451-3A{0255}.
21.Flour Colour

Loci controlling flour colour were identified and mapped in a recombinant inbred population derived from hexaploid wheat cultivars Schomburgk and Yarralinka {9936}. Regions in 3A and 7A accounted for 13% and 60% of the genetic variation, respectively, and Xbcd828-3A, Xcdo347-7A and Xwg232-7A.1 were significantly associated with flour colour. The association was highly significant in all three replicates only for the 7A QTL. Symbols were not assigned to the flour colour loci.
22.Free-threshing Habit

QFt.mgb-5A{0046}. 5AL{0046}. tv: Messapia/T. dicoccoides MG4343 mapping population{0046}. ma: Associated with XksuG44-5A{0046}.

QFt.mgb-6A{0046}. 6A{0046}. tv: Messapia/T. dicoccoides MG4343 mapping population{0046}. ma: Associated with Xpsr312-6A{0046}.
23.Frost Resistance

Fr1{1446}. 5AL{1446}. v: Hobbit{1446}. ma: Mapped to the mid-region of 5AL, 2.1 cM distal from Xcdo504-5A and Xwg644-5A and proximal to Xpsr426-5A{419}; Mapped 2cM proximal to Xwg644-5A and Vrn-A1{0291}; and flanked by deletion points 0.67 and 0.68{0292}.

Fr2{0291}. 5DL{0291}. s: CS^*7/Cheyenne 5D{0291}. ma: Fr2 mapped 10 cM proximal to Vrn-D1{0291}.

QWin.ipk-6A. 6AS{0255}. v: Opata/W-7984 (ITMI) RI mapping population{0255}; Winter hardiness was contributed by W-7984{0255}. ma: Associated with Xfba85-6A and Xpsr10(Gli-2)-6A{0255}.
Responses to cold exposure and their genetics are reviewed in {0020,0274}.
24.Gametocidal Genes
24.1. Gametocidal activity

Gc1-B1a{1485}. [Gc1a{1490},Gc1{1487}]. 2B{1490}. i: CS^*8/Aegilops speltoides. subsp. aucheri{1487}.

Gc1-B1b{1485}. [Gc1b{1490}]. 2B{1490}. i: S^*8/Ae. Speltoides subsp. ligustica{1490}.

Gc1-C1{0188}. 2CL{0189}. ad: CS/2C{0189}. su: CS2C(2A), CS2C(2B), CS2C(2D){0189}.

Gc1-Sl1{1485}. [Gc-S¹3{1485}]. 2S¹{334}. ad: CS/Ae. sharonensis{334}.

Gc2-Sl1a{1485}. [Gc-S¹1{1485}]. 4S¹{866}. ad: CS/Ae. longissima{866}.

Gc2-Sl1b{1485}. [Gc-S¹2{1485}]. 4S¹{1013}. ad: S/Ae. sharonensis{1013}.

Gc3-C1{1485}. [Gc-C{1485}]. 3C{333}. ad: CS/Ae. triuncialis{338}.
Gc1-B1a, Gc1-B1b and Gc1-S¹, classified in the same functional group, are hypostatic to the genes Gc2-S¹1a and Gc2-S¹1b. 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} and 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.

Sd1{1647}. 7D{1647}. v2: Agatha Sd2{1647,1161}.

Sd2{1161}. 7BL{1163}. v: 88M22-149{1163,1161}.
24.2. Suppression of gametocidal genes

Igc1{1489}. Causes suppression of the 3C chromosome gametocidal gene of Ae. triuncialis. This alien gametocidal factor also promotes chromosome breakage {1486}. 3B{1488}. v: Norin 26{1483,1488}; Nineteen wheats listed in{1483,1488}.

igc1. v: Chinese Spring{1483,1488}; Forty wheats are listed in{1483,1488}.
25.Gibberellic Acid Response (insensitivity)

Gai1. [GAI1{565,1246}]. 4B{406}.4BS{980}. i: See{408}. v: Norin 10 Der.{565; List in{407}. ma: Xpsr622-4B (distal) - 1.9 cM - Gai1 - 8.3 cM - Xbcd110-4B (proximal){9959}. tv: Messapia{9959}.

Gai2. [GAI2{565,1246}]. 4D{411}.4DS{980}. i: See{408}. v: Maris Hobbit{411}; Norin 10 Der.{565}; List in{407}.

Gai3. [GAI3{565,1246}]. 4B{413}.4BS{980}. i: See{408}. v: Minister Dwarf{413}; Selection D6899{359}; Tom Thumb{405}; Tom Thumb Der.{565,567}.
In wheats with Gai3, the aleurone layer fails 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.
26.Glaucousness (Waxiness/Glossiness)

Glaucousness refers to the whitish,wax-like deposits that occur on the stem and leaf-sheath surfaces of many graminaceous species. The expression of glaucousness depends on the arrangement of wax deposits rather than the amount of wax {603}. Non-glaucous variants also occur and genetic studies indicate that non-glaucousness can be either recessive or dominant. Recessive forms of non-glaucousness are apparently mutants of the genes that produce the wax-like deposits. Dominant non-glaucous phenotypes (as assessed visually) appear to be due to mutations that affect the molecular structure, and reflectance, of the wax-like substances {10001}. The genes involved in wax production and the "inhibitors" are duplicated in chromosomes 2B and 2D. There appear to be independant genes for wax production and "inhibitors" {912,1493,10001}. In earlier issues of the gene catalogue the two kinds of genes were treated as multiple alleles {1432}. All forms of wild and cultivated einkorn are non-glaucous {10001}. Orthologous loci occur in barley chromosome 2HS (gs1, gs6, gs8) {467}, rye chromosome 7RL (wa1) {725} and maize (gl2) {211}. A gene for spike glaucousness, Ws, was mapped distally on the short arm of chromosome 1B in the cross T. durum cv. Langdon / T. dicoccoides acc. Hermon H52 {0171}.
26.1. Genes for glaucousness

W1. 2BS{267,1493}. i: Chinese Spring mono-2D/S615//10*wS615{10001}. v: Chinese Spring{1493}. itv: LD222^*11/T. turgidum var. pyramidale recognitum{1546}. v2: S615 W2{10001}.

w1. Recessive allele for reduced glaucousness. 2BS{1432}. v: CS mono-4B mutant{1064}; Mentana{1432}; Salmon{1493}.

W2. i: Chinese Spring mono-2B/S615//11*wS615{10001}. v: T. compactum cv. No 44{10001}. v2: S615 W1{10001}.

W2a. dv: Glaucous forms of Ae. tauschii.

W2b. v: Chinese Spring - weak hypomorph recognized at increased dosage{1432}.
A non-glaucous spike phenotype in line L-592, a 7S(7A) substition line, is described in {0113}.

w1w2{10001}. i: w-S615 = S615*11/Salmon{10001}. v: Salmon{10001}; Mentana{1432}; CS mono-4B mutant{1064}.
26.2. Epistatic inhibitors of glaucousness

Each inhibitor inhibits all genes for glaucousness.

Iw1{10001}. [W1^I{1493},I1-W{1493}]. 2BS{10001}. i: S615/Cornell 5075//10*S615{10001}.

Iw2{10001}. [I2-W{1493}]. 2DS{10001}. i: S615/Golden Ball Synthetic//10-*S615{10001}. v: Golden Ball Synthetic{10001}; Vernal Synthetic{10001}. dv: Non-glaucous forms of Ae. tauschii{1493}.

Iw3{277}. [IW3{277},I3-W{277}]. 1BL{277}. tv: T. turgidum var. dicoccoides{277}.
A non-glaucous spike phenotype in line L-592. A 7S(7A) substitution line, is described in {0113}.
27.Glume Colour and Awn Colour
27.1. Red (brown/bronze) glumes

The majority of studies report a single dominant gene for red glume colour. A few papers report two factors {1009,1477,1520}. Red glume colour in Swedish land cultivars is apparently associated with hairy glumes {1277} suggesting, because Hg is located in chromosome 1A, that a red glume factor different from Rg1 is involved in the Swedish stocks. Nothing was known of the possible association of such a gene with Bg, another glume colour gene on 1A. See {1640} for review. A 1A gene, Rg3, was eventially identified by linkage with Gli-A1 {1405} and shown to cosegregate with Hg {624}.

Rg1. [Rg]. 1B{1517}.1BS{369}. s: CS^*5/Red Egyptian 1B{1304}. v: Diamant I{9906}; Federation 41{1517}; Highbury{1121}; Red Egyptian{1304}; T. petrapavlovskyi{9906}. v2: Milturum 321 Rg3{9906}; Milturum 553 Rg3{9906}; Strela Rg3{9906}. tv: Ward{792}. ma: Xytv1518-1B (distal) - 7.7cM - Rg1 - 0.8cM - G1i-B1 (proximal){9959}.

Rg2. Derived from Aegilops squarrosa. 1DL{769,1241}.1DS. i: Saratovskaya 29*5/T. timopheevii/T. tauschii{9906}. v: (Triticum dicoccoides/Ae. squarrosa){769}; (Tetra Canthatch/Ae. squarrosa var. strangulata RL 5271), RL 5404{1240}; (Tetra Canhatch/Ae. squarrosa var. meyeri RL 5289), RL 5406{648,1240}. dv: Aegilops squarrosa accessions. QTL: QRg.ipk.1D was mapped in the Opata/W-7984 (ITMI) mapping population{0255}; Linkage with Gli-D1 implied Rg2. This QTL coincided with a QTL for awn colour, QRaw.ipk-1D{0255}.

Rg3{924,562}. 1AS{924,562,9906}. i: Saratovskaya 29^*3 //F2 CS mono 1A/Strela{924}. v: CS/Strela Seln{9906}; Iskra{9906}; L'goskaya-47{1405}; L'govskaya-47{1405}; Zhnitstra{9906}. v2: Milturum 553 Rg1{9906}; Milturum 321 Rg1{9906}; Strela Rg1{9906,924}; Sobko & Sozinov reported a further group of 30 international wheats which, by inference from their Gli-A1 alleles, probably carry Rg3. A linkage order of Glu-A1 - cent - Hg - Rg3 was reported Strela Rg1 {9906,924}{1405}.
Kovel {729} described a brown or smokey-grey glume phenotype in T. aestivum var caesium K-28535. This phenotype was also present in accession K-40579 and botanical varieties cinereum, columbina and albiglaucum. Close linkage to Gli-D1 was shown and a gene designated Brg was assumed to be an allele of Rg2 present in Ae. tauschii and synthetic hexaploid wheats. v: K-28535 {729}. i: ANK-23 = Novosibirskaya 67*10/ K-28535 {729}.
A 1B gene controlling red glume colour was mapped in a cross between durum wheat cv. Messapia and T. turgidum ssp. dicoccoides acc. MG4343. ma: Xutv1518-1B (distal) - 7.7 cM - Rg1 - 0.8 cM - Gli-B1 (proximal) {9959}.

QRg.ipk-1D{0255}. This QTL coincides with a QTL for awn colour, QRaw.ipk-1D {0255}. 1DS{0255}. v: Opata/W-7984 (ITMI) RI mapping population{0255}; The glume colour was contributed by W-7984{0255}. ma: Associated with Gli-D1{0255}.
27.2. Black glumes

Bg{282,1304}. 1A{282,1304}. s: CS^*7/Indian 1A{1304}. dv: G1777{282}; G3116{282}. ma: Bg and Nor9 co-segregated in T. monococcum{282}³; Xutv1391-1A (distal) - 3cM - Bg - 1.6cM - Hg - 2.4cM - Gli-A1 (proximal){9959}.

Bga{282}. [Bg(a){282}]. dv: G1777.

Bgb{282}. [Bg(b){282}]. dv: G3116.

bg{282}. dv: DV92, G2528.

Bga and Bgb are dominant and cause a solid black glume and a black line at the margins of the glume, respectively. bg is recessive and confers non-black glumes.
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.
27.3. Pseudo-black chaff

This is a blackening condition transferred from Yaroslav emmer to Hope wheat by McFadden at the same time as stem-rust resistance was transferred. The association of this condition with mature-plant stem-rust reaction (Sr2) has been noted in a number of papers. According to {742}, the condition is recessive. Pan {1102} considered linkage with stem-rust reaction could be broken, but this seems unlikely.

Pbc. 3B{742}.3BS. s: CS^*6/Hope 3B{742}; CS^*6/Ciano 5B{939}.
27.4. Black-striped glumes

This phenotype was reported in group dicoccon. v: E4225 {1417}.
27.5. Inhibitor of glume pigment

An inhibitor of glume pigment was reported on chromosome 3A {106}.
27.6. Chocolate chaff

cc{719}. 7B{719}.7BS{665}. tv: Langdon mutant{719}; PI 349056{665}. dv: CBC-CDd1{665}.
The chocolate chaff phenotype was suppressed by a gene(s) in chromosome 7D {719}.
27.7. Awn colour

The literature on awn colour is not clear. In general, awn colour is associated with glume colour {045}. Occasionally, however, awn colour and glume colour may be different. According to Panin & Netsvetaev {1103}, black awns were determined by three complementary genes designated Bla1, Bla2, Bla3. Bla1 was located in chromosome 1A and linked with Gld 1A (= Gli-A1) and Hg.

QRaw.ipk-1A{0255}. 1AS{0255}. v: Opata/W-7984 (ITMI) RI mapping population{0255}; The awn colour was contributed by W-7984{0255}. ma: Associated with Gli-A1{0255}.

QRaw.ipk-1D{0255}. 1DS{0255}. v: Opata/W-7984 (ITMI) RI mapping population{0255}; The awn colour was contributed by W-7984{0255}. ma: Associated with Gli-D1{0255}.
28.Grain Hardness/Endosperm Texture

Grain hardness or endosperm texture significantly influences flour milling, flour properties and end-use. The difference in particle size index between a hard wheat (Falcon) and a soft wheat (Heron) was reported by Symes {1452} to be due to a single major gene. Symes {1452} also found evidence for "different major genes or alleles" which explained differences amongst the hard wheats Falcon, Gabo and Spica. Using Cheyenne (CNN) substitution lines in CS and a Brabender laboratory mill, Mattern et al. {915} showed that the hard wheat milling and flour properties of Cheyenne were associated with 5D. Using Hope 5D substitution line in CS [CS(Hope 5D)] crossed to CS, and CS(Hope 5D) crossed to CS ditelosomic 5DL, Law et al. {777} showed that grain hardness was controlled by alleles at a single locus on 5DS. The dominant allele, Ha, controlling softness was present in Chinese Spring and the allele for hardness, ha, was present in the other varieties mentioned. A similar study using CS (CNN5D) x 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) {0380}, 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 section 5.8.

Ha{777}. Soft phenotype. 5DS{777}. i:: Falcon/7*Heron, Heron/7*Falcon{03109}; Paha*2//Early Blackhull/5*Paha{0203,0298}; Early Blackhull Derivative/5*Nugaines{0203,0298}. v:: Chinese Spring {777,03106}; Cappelle Desprez {470}; Heron {1452,470}; Paha, Nugaines {0203,0298}; NY6432-18 {0241}.

ha{777}. Hard phenotype i: Falcon/7*Heron, Heron/7*Falcon {03109}; Paha*2//Early Blackhull/5*Paha {0203,0298}; Early Blackhull Derivative/5*Nugaines {0203,0298}. s: CS*6/Cheyenne 5D {915}; CS*6/Hope 5D {777}; Cappelle Desprez/Besostaya 5D {470}. v: Falcon {1452,470}; Holdfast {470}; Early Blackhull, Early Blackhull Derivative{0203,0298}; Cheyenne {03106}; Clark's Cream {0241}. ma: Ha was closely linked to Xmta9(Puil) - 5D {1414}.
Single factor effects on hardness were found for chromosome 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 are thought to have soft texture.
Two genes for grain hardness were reported in {055}.
Hard and soft NILs are listed in {0298}.
QTL: In a DH population of Courtot/CS 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}.
Ten QTLs for kernel hardness (54 % of the variation) were mapped in a cross 'Forno'/ 'Oberkulmer' spelt {0280}.
29.Grain Quality Parameters
29.1. Sedimentation value

Qsev.mgb-6A{9920}. 6AL{9920}. tv: Messapia/T. dicoccoides MG4343 mapping population{9920}. ma: Associated with Xrsq805-6A{9920}.

Qsev.mgb-7A{9920}. 7BS{9920}. tv: Messapia/T. dicoccoides MG4343 mapping population{9920}. ma: Associated with Xpsr103-7A{9920}.
QTL: Associated with Glu-1 on chromosome arms 1AL and 1DL and Gli-1/Glu-3 on 1BS in RSLs from the cross Cheyenne (high quality)/CS (low quality) {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}.
29.2. Flour, semolina and pasta colour

QTL: A QTL was detected on chromosome 7A {9936}. Cultivar Schomburgk contributed the yellow colour allele in a cross Schomburgk/Yarralinka {9936}. Markers Xcdo347-7A and Xwg232-7A accounted for 60% of the genetic variation {9936}. A Sequence Tagged Site PCR marker is available {0180}.
QTL: A major QTL was detected in the distal region of chromosome 7BL in the cross Omrabi 5/ T. dicoccoides 600545. The QTL explained 53% of the variation and was completely linked to microsatellite marker Xgwm344-7B. Omrabi 5 contributed the allele for high level of yellow pigment. Two additional small QTLs were detected on 7AL {0365}. Other references to flour colour are given under Lr19 and Sr25.
29.3. Amylose content

Amylose content has a significant effect on industrial quality; for example, reduced amylose wheats perform better in some types of noodles. The waxy protein genes have an important influence, but other genes are also involved.

QAmc.ocs-4A.1{0047}. 4AS{0047}. v: CS/CS(Kanto107 4A) mapping population{0047}. ma: Associated with Xbcd1738-4A and Xcdo1387-4A{0047}.
29.4. Milling yield

QTL: A QTL was detected on chromosome 3A {0181}. Cultivar Schomburgk contributed an allele for the higher milling yield in cross Schomburgk/Yarralinka {0181}. RFLP markers Xbcd115-3A and Xpsr754-3A were associated with this QTL at LOD>3 {0181}.
A QTL associated with Pinb on chromosome arm 5DS was detected in RILs from the cross NY6432-18/Clark's Cream {0241}. Cultivar Clark's Cream contributed the higher flour yield allele {0241}. This QTL coincided with QTL for hardness, hydration traits (dough water absorption, damaged starch and alkaline water retention capacity (AWRC)), and baked product traits (cookie diameter and cookie top grain) {0241}.
29.5. Alveograph dough strength W

QTL: QTLs for W were detected on chromosome arms 5DS (associated with Xmta10-5D), 1AS (associated with Xfba92-1A), and 3B (associated with XksuE3-3B) in cross Courtot/Chinese Spring {0141}. The first two QTLs coincided with those for hardness.
Ten QTL for W (39% of the variation), nine QTL for P (48% of the variation) and seven QTL for P:L (38% of the variation) were mapped in Forno/Oberkulmer spelt {0280}.
29.6. Mixograph peak time

QTL : A QTL associated with Glu-Dy1 on chromosome arm 1DL was detected in RILs from the cross NY6432-18/Clark's Cream {0241}. Clark's Cream contributed the higher mixograph peak time allele {0241}. This QTL coincided with a QTL for bread mixing time {0241}.
29.7. Starch characteristics

QTL: QTLs for starch viscosity and swelling were associated with the Wx-B1 locus in the cross Cranbrook (Wx-B1a) x Halberd (null Wx-B1b). An additional QTL for starch viscosity was found on 7BL between markers Xgwm344-7B and Xwg420-7B in the first cross. This QTL disappeared when amylase activity was inhibited indicating that it was determined by the late maturing a-amylase activity contributed by Cranbrook. A QTL for starch viscosity was associated with the Wx-A1 locus in the cross CD87/Katepwa {0362}.
30.Grass-Clump Dwarfness/Grass Dwarfness

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

D1{534}. [G{972}]. 2D{534,939,1595}.2DS{942}. s: CS^*7/Kenya Farmer 2D{1000}; CS^*6/Timstein 2D{534}. v: Big Club{534}; Burt{1000}; Federation{942}; Mus{534}; Ramona 50{358}; Selection 1403{1000}. v2: Hermsen's pure-breeding dwarf D2{1000}; Falcon D3{1172}; Gabo D3{944}; Timstein D3{534}; Metzger's pure-breeding dwarf D2 D3{1000}.

D2{534}. [Bi{972}]. 2B{536,574}.2BL{944}. s: CS^*7/Cheyenne 2B{1000}; CS^*4/Red Egyptian 2B{1000}. v: Bezostaya 1{1595}; Crete-367{1029}; Desprez 80{1595}; Florence{1000}; Kenya W744{944}; Loro{1172}; Mara{1595}; Marquis{1000}; Poros{1595}; Redman{534,574,1001}; Riebesel{534}; Tobari 66{358}. v2: Hermsen's pure-breeding dwarf D1{534,1000}; Amby D3{358}; Cedar D3{1000}; Mendel D3{534}; Plantahof D3{534}; Spica D3{944}; Cappelle-Desprez D4{1595}; Brevor D4{1000}; Cheyenne D4{1000}; Metzger's pure-breeding dwarf D1D3{1000}.

D3{534}. [A{972}]. 4A{534,1595}.4AL{939}. s: CS^*6/Timstein 4A{534,1000}; CS^*7/Kenya Farmer 4A{534,1000}. v2: Amby D1{358}; Falcon D1{1172}; Gabo D1{944}; Kenya Farmer D1{1000}; Timstein D1{534}; Metzger's pure-breeding dwarf D1 D2{1000}.

D4{1000}. 2D{1000,1595}.2DL{1598}. s: CS^*7/Cheyenne 2D{1000}. v2: Cappelle-Desprez D2{1595}; Cheyenne D2{1000}; Brevor D2{1000}.

d1d2d3d4. v: Chinese Spring{534,1000}.
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.
31.Grain Weight

QTL : Variation at locus QGw1.ccsu-1A, associated with Xwmc333-1A, accounted for 15% of the variation in a RIL population from RS111/CS {0143}.
32.Hairy/Pubescent Auricles

Pa{886,042}. 4BS{886,042}. s: Saratovskaya 29^*9/Yanetzkis Probat 4B{886}; Saratovskaya 29^*5/Shabati Sonora 4B{886}; Saratovskaya 29^*4/Siete Cerros 4B{886}. v: Diamant 1{886}; Magali{886}; Pirotrix 28{886}; Shabati Sonora{886}; Siete Cerros{886}; Ulyanovka 9 {886}.

pa. v: Gabo{886}; Saratovskaya 29{886}; This phenotype is expressed in Diamant ditelo 4BL{886}.
33.Hairy Glume

Hg{1494}. 1A{1293}.1AS{947}. i: S-615^*11/Jones Fife{1500}. s: CS^*7/Indian 1A{1293}. v: A well-known, widespread and easily identified dominant marker - few examples will be listed. Indian{1293}; Jones Fife{1494}; Prelude{1494}. itv: LD222^*11/T. Turgidum var. durum melanops{1546}. tv: Golden Ball{1342,1494}. dv: T. monococcum lines{1494}. ma: Xutv1391-1A (distal) - 3cM - Bg - 1.6cM - Hg - 2.4cM - Gli-A1 (proximal){9959}.
A 1A gene controlling hairy glumes was mapped in a cross between durum cv. Messapia and T. turgidum ssp. dicoccoides acc. MG4343.

Hg1{1405}. v: Ulyanovkn{1405}; Pionerskaya{715,1405}.
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}.
34.Hairy Leaf

Hl1{0316}. Weakly hairy. [Hl{884}]. 4B{884}.4BL{760}. v: Artemovka{925}; Caesium 111{925}; Lutescens 53/12{925}; Lutescens 62{925}; Milturum 321{884}; Poltavka{925}; Pyrothrix 28{925}; Saratov 321{884}; Saratovskaya 29{884,760}; Sarrubra{925}.

hl1 hl2. v: Chinese Spring{884}.
Kuspira et al. {744} provided evidence for at least three alleles at an Hl locus in T. monococcum.

Hl2{0316}. 7BS{0316}. v: Hong-mang-mai {0316}.
35.Hairy Leaf Sheath

Levy & Feldman {795} concluded that complementary genes determined hairy leaf sheath in T. dicoccoides.

Hs{795}. [Hls{761}]. v: Certain hexaploid derivates of G25 produced in Israel{939}. tv: T. dicoccoides G25{761}.

hs. v: Most hexaploid wheats{939}. tv: T. dicoccoides G7{761}.
36.Hairy Neck/Pubescent Peduncle

Hp{275}. Derived from Secale cereale
4BL{T4B.5R}{274,275}.i:S-615^*11/CS Derivative{1500}.
5BS{T5B-5R}{1298}.v:HN-2 (CS type){1298}.
6D{T6D-5R}{1298}.v:HN-1(CS type){1298}.
4BL{T4B.5R}{274,275}.v:CS Derivative{1304}.
37.Hairy Node/Pubescent Node

Inheritance of hairy (glabrous) node versus non-hairy node was attributed to a single, dominant gene difference {396,837,910,914} and the Hn/hn locus was shown to be linked with B1 (awn inhibitor). Observations on 5A trisomics and telosomics of Chinese Spring confirmed this location. Love & Craig {837} studied a cross involving Velvet Node CI 5877, and Gaines & Carstens {396} studied an offtype single plant designated Velvet Node Wash. No. 1981.

Hn. 5AL. v: Aurore{722}; Fylgia{722}; Extra-Kolben II{722}; Marquis{910}; Tammi{765}; T. vulgare erythrospermum{910}. tv: T. polonicum vestitum{910}.

hn. v: Garnet{722}; Kimno{722}; Pika{722}; Timantii{722}.
Levy & Feldman {795} concluded that complementary genes determined hairy leaf sheath in T. dicoccoideds.
Multiple alleles were reported in T. monococcum{744}.
38.Heat tolerance

QTL: QTLs contributing to grain-filling duration (GFD) under high temperatures were associated with Xgwm11-1BS (11% of variability) and Xgwm293-5AS (23% of variability) in Ventnor (tolerant) // Karl 92 (Non-tolerant) {0327}.
39.Height

Ht is the general symbol.
39.1. Reduced Height : GA-insensitive

Rht-1{371,0019}.
The Rht-1 homoeoloci are orthologous with the D8 locus in maize and the GAI locus in Arabidopsis. They encode proteins resembling nuclear transcription factors and are involved in sensing gibberellin levels {0019}. Common wheat and durum NIL pairs are listed in {02102}.

Rht-A1a{0019}. v: Chinese Spring{0019}; All wheats are assumed to be monomorphic.

Rht-B1{116}. 4B{109,406,1040}.4BS{089,116}. ma,tv: Gai1/Rht-B1b - 1.8cM - Xpsr622-4B{110}.

Rht-B1a{116}. v: Tall wheats{116}; e.g. Chinese Spring{0019}.

Rht-B1b{116}.Partially recessive {024}, recessive {357}, semi-dominant {408}. [Rht1{015},Sd1{015}]. i: See{408,414,02102}. v: Frontier{1597}; Guardian{1597}; Selection 14-53/Burt, 5{015}; Siete Cerros{407}; Wren{1174}; WW15{407}. v2: Norin 10-Brevor, 14 Rht-D1b{015}; Oleson Rht-D1b{357}; Selection D6301 Rht-D1b{357}; Shortim Rht-D1b{243}; See{407,415,1062,1386}. tv: Cocorit 71{109,416}; Creso{109,416,451}; Malavika{1442}; Mida{450}; Sansone{109}; Valgerado{109,416}; Valnova{450}; Valselva{450}.
The development of allele-specific primers for Rht-B1b was reported in {0378}.
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 (dwarf) / Halberd (tall). The dwarfing effect underlying the QTL is caused by the Rht-B1b allele {0379}.

Rht-B1c{116}.Semi-dominant {1040}. [Rht3{565},Sd3{565}]. i: Tom Thumb/7^* Kharkov// Lancer{1040}; See{408}. v: Minister Dwarf{404}; Selection D6899 (Tom Thumb-Sonora 64/Tacuari){357}; Tom Thumb{405}; Tom Pouce Blanc{407,1634}; Tom Pouce Barba Rouge{407,1634}; Topo; Tordo. ma: Xmwg634-4B (distal) - 30.6 cM - Rht-B1c - 11.9 cM - Xpsr144-4B (proximal){117}.

Rht-B1d{116}.Semi-dominant {1599,116}. [Rht1S{1599}]. v: Saitama 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.

Rht-B1e{116}. [RhtKrasnodari1{452},Rht1(B-dw){1600}]. v: Krasnodari 1 (a spontaneous GA-insensitive offtype of Bezostaya 1){1600}.

Rht-B1f{116}.Semi-dominant {116}. [RhtT. aethiopicum{116}]. tv: T. aethiopicum accessions W6824D{116}; W6807C{116}.

Rht-B1g{0019}. v: Highbury 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}.

Rht-D1{116}. 4D{411,583,1544}.4DS{980,1266,116}. i: Common wheat and durum NIL pairs are listed in {02102}. ma: Xpsr1871(Pki)-4D - 4cM - Rht-D1 - 6 cM - Xubc821(PhyA)-4D{410}; Rht-D1 - 2.8cM - Xglk578-4D{9966}; Xpsr1871 - 1cM - Rht-D1b - 4cM - Xpsr821(PhyA){0019}.

Rht-D1a{116}. v: Tall wheats{116}; e.g. Chinese Spring.

Rht-D1b{116}.Partially recessive {024}, recessive {357}, semi-dominant {408}. [Rht2{015},Sd2{015}]. 4D{411}.4DS{980}. i: See{408,414,02102}. v: Combe{567}; Era{407}; Gaines Sib 2{015}; Jaral{407}; Kite{1174}; Maris Hobbit {411}; Pitic 62 {567}; Songlen{243}; Oleson Rht-B1b{357}; Norin 10-Brevor 14 Rht-B1b{015}; Selection D6301 Rht-B1b{357}; List in{1386}.
The development of allele-specific primers for Rht-D1b was reported in {0378}.

Rht-D1c{116}.Dominant {114}. [Rht10{1266}]. v: Ai-bian{1544,1266}. ma: Xpsr921-4D (4DS) - 0.8 cM - Rht-D1c - 28 cM - Xgwm165-4D (4DL){117}.

Rht-D1d{116}.Semi-dominant {116}. [RhtAi-bian 1a{115}]. v: Ai-bian 1a (spontaneous mutant of Ai-bian 1){115}.
The 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}.
39.2. Reduced Height : GA-sensitive

Borner et al. {116} found no evidence of orthologous GA-sensitive genes in rye, but reviewed evidence for orthologous GA-insensitive gene. The close linkage of Rht8 and Xgwm261-2D permitted the use of the microsatellite as a marker for the detection of allelic variants at the Rht8 locus{9962}.

Rht4{568}. Recessive. v: Burt ert 937, CI 15076{566,717}.

Rht5{717}. v: Marfed ert 1, M1, CI 13988{717,718,1593}.

Rht6{718}. Recessive. v: Brevor{569}; Burt{569,718}. v2: Norin 10-Brevor, 14 Rht-B1b Rht-D1b{569}.

Rht7{1602}. 2A{1602}. v: Bersee Mutant A{1602}; Bersee Mutant C{1602}.

Rht8. 2D{772,1601,1598}.2DL. s: Cappelle-Desprez^*/ Mara 2D{1601}. v: Novasadska Rana 1{1604}; Sava{1601,414}. v2: Akakomugi Rht9{1191}; Mara Rht9{1191}. ma: Xgwm484-2D (proximal) - 19.9 cM - Rht8 - 0.6 cM - Xgwm261-2D (distal){727}.
The close linkage of Rht8 and Xgwm261-2D permitted the use of the microsatellite as a marker for the detection of allelic variants at the Rht8 locus{9962}.

Rht8a.Associated with a 165-bp fragment of WMS 261 {9962}. v: Autonomia{9962}; Bobwhite{9962}; Brevor{9962}; Chaimite{9962}; Ciano 67{9962}; Chris{9962}; Dugoklasa{9964}; Federation{9962}; Frontana{9962}; Glennson 81{9962}; Jupateco 73{9962}; Kenya{9962}; Klein 32{9962}; Lerma Rojo{9962}; Lusitano{9962}; Maringa{9962}; Mentana{9962}; 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}; Hope{0243}; Marquis{0243}; Michigan Amber{0243}.

Rht8b.Associated with a 174-bp fragment of WMS 261 {9962}. s: Cappelle Desprez^*/Mara 2D{1601}. v: Arthur{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/Brevior 14{9962}; Oasis{0243}; 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.Associated with a 192 bp fragment of WMS 261 {9962}. v: Akakomugi{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}; 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}; Kaloyan{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{119,9962}; 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{0243}; Odesskaya 51{0243}; Odesskaya 117{0243}; Odesskaya 132{0243}; Odesskaya Krasnokolosaya{0243}; Odesskaya Polukarlikovaya{0243}; Orlandi{9962}; 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{1601,414,9962}; Siete Cerros{9962}; Sinvalocho{9962}; Simvol Odesskii{0243}; Sivka{0243}; Strumok{0243}; Skopjanka{9962}; Skorospelka 3B{9962}; Slavonija{9964}; Somorka{9964}; 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}.

Rht8d.Associated with a 201-bp fragment of WMS 261 {9962}. v: Pliska{9962}; Courtot{9962}.

Rht8e.Associated with a 201-bp fragment of WMS 261 {9962}. v: Chino{9962}; Klein Esterello{9962}; Klein 157{9962}.

Rht8f.Associated with a 215-bp fragment of WMS 261 {9962}. v: Klein 49{9962}.

Rht8g.Associated with a 196-bp fragment of WMS 261 [{0243}]. v: Mirleben{0243}.

Rht8h.Associated with a 206-bp fragment of WMS 261 [{0243}]. v: Weihenstephan M1{0243}.

Rht9. 7BS{772,1601}. s: Cappelle-Desprez^*/Mara 5BS-7BS{1601}. v: Acciao{718}; Forlani{718}; Akakomugi Rht8{1601}; Mara Rht8{1601}.

Rht11{718}. v: Karlik 1{718}.

Rht12{718}. Dominant. 5A{1445,1606}. v: Karcagi 522M7K{721}. ma: Rht12 is located distally on 5AL cosegregating with B1 and closely linked to b-Amy-A1{1606}; Xgwm291-5A - 5.4cM - Rht12{726}.
Rht12 delayed ear emergence by 6 days{1606}. Rht12 delayed ear emergence by 6 days {1606}.

Rht13{718}. v: Magnif 41M1 CI 17689{718}.

Rht14{718}. v: Cp B 132 {123} = Castelporziano PI 347331{718}.

Rht15{718}. tv: Durox{718}.

Rht16{718}. v: Edmore M1{718}.

Rht17{718}. v: Chris Mutant CI 17241{1129}.

Rht18{718}. tv: Icaro{718}.

Rht19{718}. tv: Vic M1{718}.

Rht20{718}. v: Burt M860{718}.

Rht21{0230}. The existence of this gene was not confirmed {0231}.
39.3. Reduced Height : QTL

QHt.fra-1A{9957}. ma: Linkage with Xfba393-1A.

QHt.fra-1B{9957}. ma: Linkage with Xcdo1188-1B.2.

QHt.fra-4B{9957}. ma: Linkage with Xglk556-4B.

QHt.fra-7A{9957}. ma: Linkage with Xglk478-7A.

QHt.fra-7B{9957}. ma: Linkage with XksuD2-7B.

QHt.ocs-4A.1{0047}. 4AL{0047}. v: CS/CS(Kanto107 4A) mapping population{0047}. ma: Associated with Xpsr119-4A and Wx-B1{0047}.

QHt.ocs-4A.2{0047}. 4AS{0047}. v: CS/CS(Kanto107 4A) mapping population{0047}. ma: Associated with Xbcd1738-4A and Hd{0047}.

QHt.ocs-5A.1{0068}. [Qt.ocs-5A.1{0068}]. 5AL{0068}. v: CS(T. spelta 5A)/CS(Cappelle-Desprez 5A) RI mapping population{9903}. ma: Associated with the intervalXcdo1088 - 5A - Xbcd9 - 5A{0068}.
This QTL coincided with a QTL for culm length, QCl.ocs-5A.1 {0068}.

QHt.ipk-4A{0255}. 4AL{0255}. v: Opata/W-7984 (ITMI) RI mapping population{0255}; the height is contributed by Opata{0255}. ma: Associated with Xmwg549 - 4A, Xabg390 - 4A and Xbcd1670 - 4A{0255}.
QHt.ipk-4A coincided with QTLs for ear length (QEl.ipk-4A), grain number (QGnu.ipk-4A) and grain weight per ear (QGwe.ipk-4A) {0255}.

QHt.ipk-6A{0255}. 6A{0255}. v: Opata/W-7984 (ITMI) RI mapping population{0255}; The height is contributed by W-7984{0255}. ma: Associated with Xcdo29 - 6A and Xfba234 - 6A{0255}.
QHt.ipk-6A coincided with QTLs for peduncle length (QPdl.ipk-6A) and ear length (QEl.ipk-6A) {0255}.
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}.
40.Herbicide Response
40.1. Difenzoquat insensitivity

Dfq1{1396}. Insensitive. 2B{1396}.2BL{789}. v: CS{1396}.

dfq1. Sensitive. s: CS^*6/Ciano 67 2B{1396}; CS^*7/Marquis 2B{789}; CS^*/Sicco 2B{1396}. v: Ciano 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.
40.2. 2,4-D tolerance

Randhawa et al. {1190} reported a single dominant gene in each of WL711, CPAN1874 and CPAN1922 controlling tolerance. HD2009 and PBW94 were described as susceptible.
40.3. Chlortoluron Insensitivity

Su1{1402}. Insensitive. 6B{1402}.6BS{799}. v: Cappelle-Desprez{1402}. tv: B-35{735}.

su1. Sensitive. v: Chinese Spring{1402}; Poros{1402}. tv: B-7{735}. ma: Xpsr312-6B - 5.3cM - Su1 - 6.8cM - Xpsr477(Pgk2)-6B{736}. ma,tv: Nor2 (6BS) - 2.7cM - Su1{1401}; Su1 - 5.2cM - 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.
41.Hybrid Weakness
41.1. Hybrid necrosis

[Progressive lethal necrosis {155}; Firing {971}].
Complementary dominant genes. Descriptive alleles w (weak), m (medium) and s (strong) were allocated by Hermsen {532}. Phenotype is affected by modifying genes (and/or genetic background) and environment {566}. According to Dhaliwal et al. {257} progressive necrosis is suppressed at 28C.

Ne1{530}. [Le{155,550},F{971},Le1{1491}]. 5B{1491}.5BL{1636}.

Ne1m{530}. i: S-615^*11/Prelude{1500}. v: Carpo{532}; Eskisehir 220-39{532}; Garnet{532}; Klein Aniversario{532}; Koga{532}; Mus XII/80/22{532}; Prelude{532,1491}.

Ne1s{530}. v: Big Club{155,532,550}; C306{1475}; Felix{531}; Gaza 141 PI 220429 {532}; Luteseens 1163{1264}; Marquillo{115,532,550}; Ponca{532}; Spica{939}. tv: Gaza 1E PI 133460; Gaza PI 189262{532}; Iumillo{532}; Kubanka{532}; PI 94587{155,532}; Quanah{532}.
Ne1s is common in tetraploid wheats {1080}.
Unknown Ne1 allele. tv: HW75 {697}; HW178 {697}. Chinese Spring carries the weakest allele {532} and its effect can be observed in CS*7/Atlas 66 2B {939} relative to CS.

Ne1w{530}. v: Bobin 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 acession 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}.

Ne2. [Le2{155,550,1491},F{971}]. 2B{1491}.2BS{1085}.

Ne2m{530}. v: 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}.

Ne2m?{530}. v: Barleta group{532}: South American wheats, e.g. Klein Titan{532}; La Prevision 25{532}; Lin Calel{532}.

Ne2ms{530}. v: Mediterranean group{532}: Dawson{155,550}; 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}; Noe group{532}: Vilmorin 27{532}; Unknown Ne2 allele{532}; Harvest Queen{532}. tv: Acme{532}; Arnautka{532}; Carleton{532}; Langdon{1498}; Mindum{532}; Stewart{532}.
However, Ne2 was stated to be absent or rare in tetraploid wheats {1080}.
The Chinese Spring 2BS telosome carries an Ne2 allele that is not present in Chinese Spring {1085}.

Ne2s{530}. i: S-615^*11/Kharkov{1500}. v: Crimean group{532}: Blackhull{550}; Chiefkan{550}; Clarkan{550}; Kharkov{1491}; Michigan Amber{532}; Minhardi{155}; Red Chief{550}; Stepnaja 135{1264}; Turkey{532}.

Ne2w{530}. v: Vakka{532}; Varma{532}.

ne1 ne2. v: Chancellor{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}.

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{1210}. 5RL{1211}. al: S. cereale 1R136-2{1210}.

Ner2{1210}. 7RL{1211}. al: S. cereale L155, L256{1210}.
41.2. Hybrid chlorosis type 1

Ch1{535}. [m^a{1245}]. 2A{538,939}. i: Steinwedel^*2/Khapli{939}; T. macha var. colchicum{535}. v: T. macha var. subletschumicum{1245,1493}. tv: Khapli{1080,1549}; T. dicoccoides var. kotschyanum{535}; T. dicoccoides var. straussianum{535}.
36 group dicoccon wheats are listed in {697}.

Ch2{535}. [m^b{1245},Ne3{1504}]. 3D{1495,1504}.3DL{692,939}. v: Chinese Spring{535,1504}; T. vavilovi.
Extremely widespread, very few wheats lack this gene.
Allelic variation at the Ch2 locus was suggested {537,1000}. Prelude, Reward and Red Bobs were exceptional in producing severe symptoms and death at an early stage. Konosu 25 may carry a weak allele {1000}. Different alleles in C306 (strong) and Sonalika (medium) were suggested in {697}.

ch1 ch2. v: Albit{1000,1509}; Burt{1000,1509}; Chancellor{1000}; Garra{1549}; Kharkof{535}; Steinwedel{1549}. su: TAP 67 (= Pawnee 3Ag(3D)){1644}.

Lists appear 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{1472}. dv: Cereal rye lines, EC179188 = WSP527A{1472}; EC143825 = WSP506A{1472}; EC338685 = Blanco{1472}; others{1472}.

chr1{1472}. dv: EC179178{1472}; EC179185 SAR/SWPY5{1472}.
41.3. Hybrid chlorosis (type 2) {1511}.

Cs1{1511}. [Chl¹]. 5A{1498}. v: T. dicoccum cv. Hokudai{1511}.
Occurs at high frequency in the T. paleocolchicum group of emmers.

Cs2{1511}. [Chl²{1501}]. 4G{1498}. tv: Many accessions of T. timopheevi and T. araraticum{637,1511}.
Multiple allelism at the Cs2 locus is discussed in {637}.
42.Iron Deficiency

Fe1{926}. 7DL{927}. v: Saratovskaya 29{926}.

Fe2{926}. 7BS{927}. v: CS{927}.
43.Lack of Ligules

The liguleless character is controlled by complementary recessive genes in hexaploid wheat {077,738,942} and by a single recessive in tetraploid wheat {047,050,939}. One gene at the tetraploid level is allelic with one of those in the hexaploid {939}. Evidence for orthology of lg1 and lg2 with lg of rice {170}, lg1 of maize {004}, li of barley {1155} and al of rye was presented in {725}. al: Imperial rye chromosome 2R restored the liguled condition to a liguleless CS derivative {939}.

lg1{047}. 2B{942}. v: Partial backcross derivative of CS{939}.

lg2. 2D{942}. v: Eligulate W1342 lg1{942}; Partial backcross derivative of CS{939}.
Because diploid wheats are liguled, polyploid wheats presumably carry a third recessive factor in chromosome 2A.
44.Leaf Erectness

QLer.ipk-2A{0255}. 2AS{0255}. v: Opata/W-7984 (ITMI) RI mapping population{0255}; The erect leaf phenotype was contributed by Opata{0255}. ma: Associated 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}.
45.Leaf Tip Necrosis

Ltn{1361}. 7D{1361}. v: Wheats with Lr34/Yr18{301,1361}; See Lr34, Yr18.

QLtn.sfr-1B{0050}. 1BS{0050}. v: Forno/T. spelta var. Oberkulmer mapping population{0050}. ma: Associated with Xgwm18-1B and Xglk483-1B{0050}.

QLtn.sfr-3A{0050}. 3A{0050}. v: Forno/T. spelta var. Oberkulmer mapping population{0050}. ma: Associated with Xpsr570-3A and Xpsr543-3A{0050}.

QLtn.sfr-4B.1{0050}. 4B{0050}. v: Forno/T. spelta var. Oberkulmer mapping population{0050}. ma: Associated with Xpsr921-4B and Xpsr593-4B{0050}.

QLtn.sfr-4B.2{0050}. 4B{0050}. v: Forno/T. spelta var. Oberkulmer mapping population{0050}. ma: Associated with Xpsr593-4B and Xpsr112-4B{0050}.

QLtn.sfr-4D{0050}. 4DL{0050}. v: Forno/T. spelta var. Oberkulmer mapping population{0050}. ma: Associated with Xpsr302-4D and Xpsr1101-4D{0050}.

QLtn.sfr-5A{0050}. 5A{0050}. v: Forno/T. spelta var. Oberkulmer mapping population{0050}. ma: Associated with Xpsr549-5A and Xglk163-5A{0050}.

QLtn.sfr-6A{0050}. 6A{0050}. v: Forno/T. spelta var. Oberkulmer mapping population{0050}. ma: Associated with Xpsr563-6A and Xpsr966-6A{0050}.

QLtn.sfr-7B.1{0050}. 7B {0050}. v: Forno/T. spelta var. Oberkulmer mapping population{0050}. ma: Associated with Xpsr350 and Xbzh232(Tha)-7B{0050}.

QLtn.sfr-7B.2{0050}. 7B{0050}. v: Forno/T. spelta var. Oberkulmer mapping population{0050}. ma: Associated with Xglk750-7B and Xmwg710-7B{0050}.

QLtn.sfr-7D{0050}. 7DS{0050}. v: Forno/T. spelta var. Oberkulmer mapping population{0050}. ma: Associated with Xpsr160-7D and Xgwm44-7D{0050}.
46.Lodging

QLd.sfr-1B{0052}. 1BS{0052}. v: Forno/T. spelta var. Oberkulmer mapping population{0052}. ma: Associated 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{0052}. 2AS{0052}. v: Forno/T. spelta var. Oberkulmer mapping population{0052}. ma: Associated 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{0052}. 2D{0052}. v: Forno/T. spelta var. Oberkulmer mapping population{0052}. ma: Associated with Xpsr933-2D and Xglk529-2D{0052}.

QLd.sfr-3A{0052}. 3AS{0052}. v: Forno/T. spelta var. Oberkulmer mapping population{0052}. ma: Associated with Xpsr598-3A and Xpsr570-3A{0052}.
This QTL coincided with QTL for increased culm stiffness and reduced culm thickness {0052}.

QLd.sfr-4A{0052}. 4AS{0052}. v: Forno/T. spelta var. Oberkulmer mapping population{0052}. ma: Associated 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{0052}. 5AL{0052}. v: Forno/T. spelta var. Oberkulmer mapping population{0052}. ma: Associated 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{0052}. 5BL{0052}. v: Forno/T. spelta var. Oberkulmer mapping population{0052}. ma: Associated 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{0052}. 6BL{0052}. v: Forno/T. spelta var. Oberkulmer mapping population{0052}. ma: Associated with Xpsr964-6B and Xpsr142-6B{0052}.

QLd.sfr-7B{0052}. 7BL{0052}. v: Forno/T. spelta var. Oberkulmer mapping population{0052}. ma: Associated with Xpsr927-7B and Xpsr350-7B{0052}.
This QTL coincided with QTL for reduced height and later ear emergence {0052}.
47.Male Sterility
47.1. Chromosomal

ms1. Recessive alleles for sterility 4B{268}.4BS{064}.

ms1a{268}. v: Briggle's Chancellor Derivative{268}; Pugsley's Male Sterile{268}.

ms1b{268}. v: Probus mutant{268}.

ms1c{064}. v: Cornerstone{064}.

ms1d{0290}. v: Mutant FS2{0290}.

ms1e{0290}. v: Mutant FS3{0290}.

ms1f{0290}. v: Mutant FS24{0290}.

ms2{806}. Dominant allele for sterility. [Ta1{240}]. 4DS{806}. v: Taigu = Line 223{240,807,806}; ms2 confers sterility when present in octaploid triticale{597}.

ms3{872}. Dominant allele for sterility. 5AS{872}. i: Chris derivative{872}; KS87UP9{219}. ma: Xwg341-5A - 0.8cM - 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{0293}. Dominant allele for sterility, distinguished from ms2 on the basis of different degrees of recombination with the 4D centromere. 4DS{0293}. v: Konzak's male sterile.

ms5{0290}. 3A{0290}. v: Mutant FS20{0290}.
47.2. Sterility in hybrids with wheat

Shw{0331}. [1HL{0331}]. ad: Additions of 1H and 1HL to wheat and certain translocation lines {0331}. ma: Located in a 16.4 cM interval flanked by Xmwg800-1H and Xmwg943-1H. A possible relationship with Ncc genes is discussed {0331}.
48.Manganese Efficiency

QTL: Variation associated with Xcdo583-4B explained 42% of the variation for Mn efficiency in the durum cross Stojocri 2 (Mn efficient)/Hazar (MN inefficient) {0320}.
49.Megasporogenesis
49.1. Control of megasporogenesis

Msg{625}. 7AS{625}. tv: Langdon{625}.
50.Meiotic Characters
50.1. Low-temperature pairing

ltp{527}. v: Chinese Spring{527}.
Expressed in the absence of chromosomes 5D at 12^oC - 15^oC, but not at 20^oC. A contrasting allele, Ltp, for normal pairing at the lower temperature range was demonstrated in T. dicoccum.
50.2. Pairing homoeologous

Ph1{1537}. 5BL{1301}. ma: PCR-based assays for presence and absence of Ph1 have been described{0214,0217,9965,0359}; 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}.

ph1a.- Not applicable - see ph2b {1303}.

ph1b{1301}. v: Sears' high pairing mutant{1301}. ma: A PCR-based detection system for ph1b ph1b individuals is described in {9965}.

ph1c{593}. tv: Cappelli ph1 mutant{449,593}; This mutant is deficient for a terminal portion of chromosome 5BL{449}. ma: Mutant 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{1302}. 3DS{1302}.

ph2a{1302}. v: Sears' intermediate pairing mutant{1301,1302}.

ph2b{1304,1303}. [ph1a{1537}]. v: Chinese Spring mutant 10/13{1537}.
50.3. Inhibitor of pairing homoeologous

Ph1^I. al: Aegilops speltoides{1218,439}.
51.Nitrate Reductase Activity

Nra{424}. v: UC44-111{424}.

nra{424}. v: Anza{424}.
52.Nuclear-Cytoplasmic Compatability Enhancers

scs{869}. Derived from T. timopheevii {869}. 1AL{870,027}. v: T. timopheevii{869}. ma: A number of completely linked RAPD makers were identified{044}.
Asakura et al. {044} used the symbol Ncc as a synonymn for scs pointing out that the effects of the gene are not limited to a single species.
53.Nucleolus Organizer Regions
53.1. 18S - 5.8S - 26S rRNA genes

NORs have been observed as secondary constrictions associated with nucleoli on satellited chromosomes {e.g., 221}, and by in situ hybridization to chromosome spreads {039,294,1014} of 18S-5.8S-26S ribosomal-DNA probes {038,433}. Allelic variation in gene number has been demonstrated at all wheat Nor sites and at Nor-R1 by filter {367} and in situ hybridization {1012}. Allelic variants of the Nor loci are detected by hybridization of rDNA probes to restriction endonuclease-treated DNA on Southern blots {037,288,917,1399}. Alleles Nor-B2a to Nor-B2f were identified using Taq1 digests of genomic DNAs hybridized to derivatives of the plasmid pTa250 {433} containing spacer-DNA fragments pTa250.4 {367,917} and pTa250.15 {288}.
Other variants may have been isolated {1399} using BamH1/EcoR1 double digests and pTa71 {433}. The variants may or may not be equivalent to those described below.

Nor1a and Nor2a. v: Maris Huntsman{1399}.

Nor1b and Nor2b. v: Bezostaya 1{1399}.

Nor1c and Nor2c. v: Cappelle-Desprez, Maris Ranger{1399}.

Nor-A1. 1AS{221,367,835,1012}. v: T. spelta{221,367,835,1012}. dv: T. monococcum{658}.

Nor-B1. [Nor1{1120}]. 1B{037,288}.1BS{221,367,835,1041}. v: CS{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{918}. v: Cheyenne, Chinese Spring, Hope, Kite, Oxley, Teal, Timstein{037,288}; Vasco, 8 others{288}.

Nor-B1a-{918}. v: A derivative allele of Nor-B1a with a significantly reduced amount of spacer. Condor 64-1{918}; Sonora 64-1{918}.

Nor-B1b. v: Olympic, Robin, Shortim{917}.

Nor-B1c{918}. v: Banks{917}; Corella{917}; Warigal{917}; 5 others{917}.

Nor-B1c-{918}. v: Rosella{918}.

Nor-B1d{918}. v: Maris Huntsman{918}.

Nor-Agⁱ1{374}. 1Agⁱ{374}. ad: Vilmorin27/Ag. intermedium{374}.

Nor-H1. [Nor-I1{794}]. 1HS{794}. dv: Sultan barley{794}.

Nor-R1. 1RS{039}. ad: CS/Imperial{039}.

Nor-S1. 1SS{294}. al: Ae. speltoides{294}.

Nor-U1. 1U{906}. su: CS/Ae. umbellulata{906}.

Nor-V1{241}. 1V{241}. ad: CS/D. villosum{241}.

Nor-B2. [Nor2{1120}]. 6BS{1041,221,366,835}. v: CS.

Nor-B2a{918}. 6B{288}. v: CS{037,917}.

Nor-B2a-{918}. v: Blueboy{918}; Sonora 64-1{918}.

Nor-B2b. T6B{288}. v: Banks, Oxley, Shortim, Timstein{037}; 12 others{917}.

Nor-B2c. v: Corella, Robin, Teal, 1 other{917}.

Nor-B2d{918}. H6B{288}. v: Hope{037}; Olympic{917}; Warigal{917}.

Nor-B2d-{918}. v: Harrier{918}; Kite{917,918}.

Nor-B2e. v: Vasco{917}.

Nor-B2f. Ch6B{288}. v: Cheyenne{037,917}.

Nor-B2g{918}. v: Falcon; Gluclub; La Prevision{918}.

Nor-B2h{918}. v: Yaktana{918}.

Nor-B2i{918}. v: Maris Huntsman; Thatcher{918}.

Nor-E2. 6ES{294}. ad: CS/E. elongata{294}.

Nor-G2. 6G{578}. tv: T. timopheevii IPSR (PBI) No. 1{294}.

Nor-H2. [Rnr1{1248}]. 6H{1070,039,1248}.6HS{794}. al: Clipper barley{039}; Sultan barley{794}.

Nor-S2. 6SS{294}. al: Ae. speltoides{294}.

Nor-A3. 5AS{1014,658}. dv: T. monococcum, T. urartu IPSR (PBI) Acc. A.

Nor-D3. 5DS{221,835}. v: CS; most wheats{037,288,917}.

Nor-Agⁱ3. 5Agⁱ{374}. ad: CS/Ag. intermedium{374}.

Nor-E3. 5ES{294}. ad: CS/E. elongata{294}.

Nor-H3. [Rnr2{1248}]. 5H{1070,039,1248}.5HS{794}. al: Clipper barley{039}; Sultan barley{794}.

Nor-U3. 5U{906}. ad,su: CS/Ae umbellulata{906}.

Nor-D4{1042}. 7DL{1042}. v: CS{1042}. dv: Ae squarrosa{1042}.

Nor-H4. [Nor-I4{794}]. 7HS{794,793}. al: Sultan barley{794}.

Nor-H5. [Nor-I5{794}]. 2HS{794,793}. al: Sultan barley{794}.

Nor-B6{601}. 1BL{601}. v: CS; Cheyenne, Wichita{601}. tv: Langdon{601}.

Nor-A7{601}. 5AL{601}. v: CS; Cheyenne, Wichita{601}. tv: Langdon{601}.

Nor-D8{601}. 3DS{601}. v: Witchita{601}.

Nor-A9{00120}. [Nor-A1{221,367,835,1012}]. 1AS{282,276}. v: T. spelta{221,367,835,1012}.

Nor-A10{00120}. [Nor-A3{1014,658}]. 5AS{282,276}. dv: T. monococcum{282,276}; T. urartu IPSR (PBI) Acc. A.

More detailed listings for allelic variation at Nor-B1 and Nor-B2 are given in {917,918}.
Two sites designated temporarily as Nor-Ax and Nor-Ay were identified in T. monococcum ssp. boeoticum, but were absent in ssp. urartu.
54.Osmoregulation

Osmoregulation is a specific form of solute accumulation regulating turgor pressure and hydration during periods of stress with positive effects on growth. Wheat lines selected for higher osmoregulation in the greenhouse have greater growth and seed yields under water limited conditions in the field.

Or{1030}. Low osmoregulation. s: CS {Red Egyptian 7A}. v: Cappelle Desprez; Condor^*4/3Ag14{1030}; Red Egyptian. ma: Or (proximal in 7AS) - 13 cM - Xpsr119-7A{1031}.

or{1030}. High osmoregulation. 7A{1030}.7AS{1031}. v: CS, Condor, Songlen, Takari{1030}.
55.Pollen Killer

Ki{1306}. Killing allele is dominant. 6BL{1306}. v: Chinese Spring{1306}; Mentana{929}.

ki. v: Probably 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.
56.Polyphenol Oxidase (PPO) Activity

3,4 dihydroxyphenylalanine (L-DOPA) was used as a substrate in a non-destructive test of polyphenol oxidase activity in seeds. Chromosome 2D was shown to carry PPO gene(s) based on Langdon/Chinese Spring (2D) substitution lines and nullisomic-tetrasomic analysis {0342}.
QTL: A QTL on 2D, associated with Xfba314-2D was identified in an M6 / Opata 85 population using the L-DOPA assay. The high PPO activity was contributed by M6 {0344}. Markers significantly associated with PPO activity were also detected on chromosomes 2A, 2B, 3B, 3D and 6B in the population NY18 / Clark's Cream {0344}.
57.Red Grain Colour

Red colour is dominant to white. At each locus, the white allele is assigned a and the red allele, b. White-grained T. aestivum and amber-grained T. durum wheats carry recessive a alleles at each locus. White-grained CS*7/Kenya Farmer and CS*6/Timstein are considered near-isogenic to CS with R-D1b.

R-A1{548}. [R2]. 3AL{957,1003}. i: Novosibirskaya 67^*9/Solo{730}. ma: (Proximal) Xpsr483(Cxp1)-3A - 28cM - R-A1 - Xpsr904-3A{370} (distal).

R-A1b. [R2]. i: Novosibirskaya 67^*9/Solo{730}. v: Baron{370}; Diamant 2{014}; Hustler{370}; Norin 10- Brevor, 14{017}; Maris Widgeon{370}; Mercia;{370}; Motto{370}; Red Bobs{1003}; Sapphire{370}; Slejpner{370}; Talent{370}; Wembley{370}.

R-B1{548}. [R3]. 3BL{1003,370}. i: Novosibirskaya 67^*9/k-28536{730}. ma: Xbcd131-3B - 5cM - R-B1 - 5cM - Xabc174-3B{410}.

R-B1b. [R3]. i: Novosibirskaya 67^*9/k-28536{730}. v: Canon{370}; Dollar{370}; Grana{370}; Supreme{370}.

R-D1{549}. [R1]. 3DL{1291,1293}. i: Novosibirskaya 67^*9/CS{730}. ma: Xbcd131-3D cosegregation with R-D1 - 15cM - Xabc174-3D{410}. v: CS.

R-D1b. [R1]. i: Novosibirskaya 67^*9/CS{730}. v: Alexandria{370}; Apollo{370}; Axona{370}; CS; Pawnee{549}; Dwarf A{370}; Fortress{370}; Jerico{370}; Longbow{370}; Luna{370}; Mardler{370}; Maris Huntsman{370}; Minaret{370}; NFC 75/93/27A; Rapier{370}; Voyage{370}; Vuka{370}.

R-N1{1018}. 3N{1018}. su: CS/Ae. uniaristata{1018}.

R-R1{1011}. 6RL{1011}. ad: Holdfast/King II{1011}.

R-V1{1518}. 3VL{1518}. ad: Creso/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. [R2,R3]. v: Red Chief{548}; Avalon{370}; Bersee; Cappelle Desprez; Feuvert; Mission; Parade; Rendezvous; Yuri{370}.

R-A1b R-B1a R-D1b . [R2,R1]. v: Broom{370}; Bezostaya 1{370}; Brigand{370}; Brock{370}; Kronjuwel{370}.

R-A1a R-B1b R-D1b. [R3,R1]. v: Kharkov{1003}; Fenman{370}; Norman{370}; Pastiche{370}; Riband{370}; Sperber{370}; Squadron{370}; Urban{370}.

R-A1b R-B1b R-D1b. [R1,R2,R3]. v: Bowie; Frondoso{1148}; Frontiera{437}; Hope{204,206}; Japanese Bearded{1548}; Kanred{1078,1426}; Lin Calel{1078}.
58.Response to Photoperiod

One-gene {1169} and two-gene {638,1137,1170} differences were reported in inheritance studies. In Chinese Spring/Hope substitution lines for chromosomes 1A, 4B and 6B greater sensitivity to short photoperiod was found, whereas substitutions of 3B and 7D were less sensitive {487}.
'a' alleles are dominant.
There is an orthologous gene series on the short arms of homoeologous group 2. The "a" alleles confer the insensitive response {0063}, the contrasting allele may be referred to as "b".

Ppd-A1a{0063}. [Ppd3{1141}]. 2AL{1268}. v: C591{0057}.

Ppd-B1a{0063}. [Ppd2{1566}]. 2BS{1566,1268,1269}. s: Cappelle-Desprez^*/CS 2B{0058}. v: Chinese Spring{1268}; Spica{557}; Timstein{1269}. v2: Sharbati Sonora Ppd-A1a{887}. ma: Xpsr666-2B - 1.2cM - Xpsr109-2B - 4.4cM - Ppd-B1 - 4.8cM - Xpsr804-2B Cent{0062}.

Ppd-D1a{0063}. [Ppd1{1566}]. 2DS{1268}. s: Cappelle Desprez^*/Ciano 2D{1598}; Cappelle-Desprez^*/Mara 2D{1598}; CS^*/Ciano 2D Ppd-B1a{1268}. v: Akakomugi{1604}; Bezostaya 1{1604}; Mara{1604}; Sava{1604}; Sonora 64{1566}. v2: Sharbati Sonora Ppd-D1a{887}.

Ppd-A1b Ppd-B1b Ppd-D1b. v: Cheyenne{1141}; Diamont 1{887}; Lancer{638}; Saratovskaya 29{887}; Warrier{638}.
Two genes control photoperiod response in T. turgidum {788}.
Gene Ppd-H2 on barley chromosome 2HS may be a member of the Ppd-1 orthologous series {766}.
QTL : A QTL was detected in chromosome 4BS in Courtot/CS {0132}.
59.Response to Salinity
59.1. K+/Na+ discrimination

Variation in K+/Na+ discrimination ratios correlate with salt tolerance, high ratios being indicative of higher tolerance.

Kna1{290}. 4DL{290}.4BS.4BL-4DL{283}.4BS.4BL-4DL-4BL{849}. v: Hexaploid wheats{290}. tv,su: Langdon 4D(4B){283}. tv,tr: Various lines{290}; Selection 3^*5-4{849}. ma: Kna1 was completely linked with Xabc305-4B, Xabc305-4D, Xbcd402-4B, Xbcd402-4D, Xpsr375-4D, Xpsr567- 4B, Xpsr567-4D, Xwg199-4B and Xwg199-4D in recombined T. turgidum 4B and T. aestivum 4D chromosomes{283,849}.
Lophopyrum elongatum chromosome arms 1ES, 7ES, and 7EL enhance K⁺/Na⁺ selectivity in wheat under salt stress {0065}.
60.Response to Tissue Culture

Qtcr.ipk-2B.1{084}. [Tcr-B1{084}]. ma: Weakly associated with Xpsr102-2B{084}.

Qtcr.ipk-2B.2{084}. [Tcr-B2{084}]. ma: Closely linked and distal to Ppd-B1{084}.

Qtcr:ipk-2B.3{084}. [Tcr-B3{084}]. ma: Linked withYr7/Sr9g{084}.

QGpp.kvl-2A{0253}. 2AL{0253}. v: Ciano/Walter DH mapping population. The green plant percentage was contributed by Ciano{0253}. ma: Associated with Xpsp3045-2A{0253}.

QGpp.kvl-2B.1{0253}. 2BL{0253}. v: Ciano/Walter DH mapping population. The green plant percentage was contributed by Ciano{0253}. ma: Associated with Xgwm388-2B{0253}.

QGpp.kvl-2B.2{0253}. 2BL{0253}. v: Ciano/Walter DH mapping population. The green plant percentage was contributed by Ciano{0253}. ma: Associated with AFLP markers{0253}.
61.Response to Vernalization

Winter cultivars carry recessive alleles at all Vrn loci. Differences among winter wheats with respect to vernalization requirements seem to be due to multiple recessive alleles {1173,0202}. Two genes may determine differences between winter wheats requiring 20 days and 60-65 days of vernalization {461,1173,9902}.
New combinations of vrn alleles from Mironovskaya 808 with a high vernalization requirement and Bezostaya 1 with a lower requirement gave progenies with higher and lower vernalization requirements than the respective parents {9902}. The allelic variants were designated with subscripted letters vrn1^B, vrn2^B, vrn3^B and vrn1^M, vrn2^M, vrn3^M. Spring and intermediate genotypes carry dominant alleles leading to no or reduced vernalization response.

Vrn-1{1398}. Orthologous series in long arms of chromosomes of homoeologous group 5.
'a' alleles are the dominant alleles for insensitivity or low vernalization response. Vrn-1 should be ortholo

	Genotype		Approx. sterile-base score
Group vulgare	----	QQ Bs Bs	0.00
Speltoids	StFF	qq Bs Bs	0.00
	StF	qq Bs Bs	0.08
	St1A	qq Bs Bs	0.39
	St1	qq Bs Bs	0.96
	St2	qq bs bs	1.41