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-Apol1{0254},P-Apet1{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 - P1pol - 5.6cM - Xgwm1083-7AL {0254}; Xgwm890-7AS - 2.1cM - P1pet {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}. [AltBH{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. 4AmL{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-1Am{0364}. [Eps-Am1]. 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}. [W1I{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}. [ma{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}. [mb{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 12oC - 15oC, but not at 20oC. 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
Ph1I. 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 vrn1B, vrn2B, vrn3B and vrn1M, vrn2M, vrn3M. 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 orthologous to Vrn-H1 {Sh2/Sgh2} of barley {726,419,767} and Vrn-R1{Sp1} of rye {726,1142} based on map locations using common RFLP
Vrn-A1a{1398}. [Vrn1{1172},Sk{002}]. 5AL{775,883}. i: Triple Dirk{1171,1172}. s: Kharkov 22MC*/Rescue 5A{358}; Winalta*8/Rescue 5A{876}; Rescue*/Cadet 5A Vrn-D1a Vrn-B1a{1221}. v: Cadet{1221}; Conley{1171}; Diamant II{885}; Falcon{1172}; Koga II{1611}; Kolben{001,1171,1172}; Konosu 25{460}; Marquis{001}; Reward{1171}; Saitama 27{460}; Saratov 29{883}; Saratovskaya 29{885}; Saratovskaya 210{883}; Shabati Sonora{885}; Thatcher{1171}; WW15{1172}. v2: Shortandinka Vrn-B1a{885}; Takari Vrn-B1a{253}; Hope Vrn-B4a{1424}. ma: Vrn-A1 - 7.5cM - Xwg644-5A{726}; Located mid 5A cosegregating with Xcdo504-5A, Xwg644-5A and Xpsr426-5A{419}; Vrn-A1 - 0.8cM - Xbcd450-5A/Xrz395-5A - 4.2cM - Xpsr426-5A{9903}.
Cultivars possessing Vrn-A1a are insensitive to vernalization. Vrn-A1a is epistatic to other genes. According to {1221}, Vrn-A1a is not always fully dominant and not always epistatic. Kuspira et al. {745} attributed single gene variation in T. monococcum to the Vrn-A1a locus. Multiple recessive alleles were suggested {745}. Vrn-Am1 was mapped on the long arm of chromosome 5Am closely linked to the same RFLP markers as Vrn-A1 {279}. Vrn-Am1 was mapped to the Xcdo504-5A - Xpsr426-5A region {0312}. In the opinion of the curators this location may not be correct
Multiple alleles also were reported in {9930}, and the dominant allele of Novosibirskaya 67 and the weaker dominant allele of Pirotrix 28 were designated Vrn1a and Vrn1b, respectively.
Vrn-B1a{1398}. [Vrn4{1173},Vrn2{1172},Ss{002},Vrn2a = Vrn2{921,920},Vrn2b = Vrn2{921,920}]. 5B{885}.5D{635}.5BL{885}.5B{921,920}.5BL or 7BL{635,0282}. i: Ank-18{921,920}; Triple Dirk B{1172}. s: Diamant 1*8/Mironovskaya 5A{920}; Diamant 1*8/Skorospelka 35 5A{920}; Rescue*/Cadet 5A Vrn-A1 Vrn-D1a{885}; Saratovskaya 29*8/Mironovskaya 808 5A{920}; Saratovskaya 29*8/Odesskaya 51 5A{920}. v: Bersee{557}; Brown Schlanstedt{001,002,1171,1172}; Cadet{1221}; Festiguay{1172}; Magali; Mara{1611}; Milturum 321{885,920}; Milturum 885{885,920}; Noe{002}; Pyrothrix 28{920}; Spica{557}; Ulyanovka 9{920}. v2: Borsum Vrn1-A1a{001}; Dala Vrn1-A1a{001}; Diamant 1 Vrn1{001,920}; Gabo Vrn4{1172}; Halland Vrn-A1a{001}; Harukikari Vrn-A1a{883}; Rubin Vrn-A1a{001}; Saratovskaya 29 Vrn-A1a{920}; Shortandinka Vrn-A1a{1221}; Triple Dirk Vrn-A1a{1173}. ma: A dCAPS marker derived from Xwg644-5B was 1.7cM from Vrn-B1{10006}.
When mapped as a QTL Vrn-B1 showed closest association with Xgwm408-5B {10007}. Two variants of Vrn-B1a were distinguished in {920,921}. Carriers of an earlier designated Vrn2b (characterized by Diamant 1*8/Skorospelka 35 5A) showed accelerated flowering after 15 and 30 days of vernalization, whereas carriers of Vrn-2a, (characterized by Ank-18 and Saratovskaya 29*8/Mironovskaya 808 5A) did not respond to these periods of vernalization. This distinction was not made in the above list.
Vrn-D1a{1398}. [Vrn3{1172}]. 5DL{775,883}. i: Triple Dirk E{1172}. s: Rescue*/Cadet 5A Vrn-A1a{1221}. v: Chinese Spring{1172}; Loro{1172}; Norin 61{460}; Shinchunaga{460}; Shirasagi Komugi{460}; Ushio Komugi{460}. v2: Rescue Vrn-B1a{1221}.

Stock:Genotype:Vernalization Response
Triple Dirk, Kolben: Vrn-A1a Vrn-B1b Vrn-D1b:No
Triple Dirk B, Festiguay:Vrn-A1b Vrn-B1a Vrn-D1b:Yes
Gabo:Vrn-A1b Vrn-B1a Vrn-D1b:Yes
Triple Dirk E, Chinese Spring:Vrn-A1b Vrn-B1b Vrn-D1a:Yes
Vrn-2{1398}. Orthologous series in chromosomes of homoeologous group 4. Vrn-Am2 was located in T. monococcum {279} on chromosome 5Am on the 4Am translocated region. Vrn-Am2 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-Am2 based on comparative maps {279,767}.
Vrn-A2a{279}. Winter habit- dominant in diploid wheat. dv: G1777{279}; G3116{279}.
Vrn-A2b{279}. Spring habit. dv: DV92{279}.
Vrn-3{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}.
Vrn-4{279}. To date, only Vrn-B4 has been detected.
Vrn-B4a{279}. The distal region of 7BS was translocated with a chromosome segment with homoeology to the distal region of 5AL. It is not known if Vrn-4 is located in the region homoeologous to 5L or 7S. [Vrn5{771,769},eHi{771,769}]. 7BS{768,771}. s: CS*/Hope 7B Vrn-D1a{768}. v: Hope Vrn-A1a{1424}.
Vrn4. 5D{10002}.5DL{10004}. i: Triple Dirk F. v2: Gabo Vrn-B1a{1172}; IL47/Vrn-A1a{10005}.
Eight land races with only Vrn4 were detected in {10003}; others combined Vrn4 with other Vrn genes. Stelmakh {1424} doubted the existance of Vrn4. References to additional studies are given in {1424}. QTL: Analysis in Courtot/CS {0132}.


62.Restorers for Cytoplasmic Male Sterility

62.1. Restorers for T. timopheevi cytoplasm
Rf1{823}. 1A{1224,1619,873}.1AS{868}. v: L22{868}; (T. timopheevi/Aegilops squarrosa)// 3*Dirk{1619}. v2: T. timopheevi/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} is unknown.
Rf2{823}. 7D{871}. v: T. timopheevi/3*Marquis Rf1{823}.
Rf3{1453}. 1B{1453}.1BS. v: T. spelta var. duhamelianum{1453}. ma: Xcdo388-1B - 1.2 cM - Xabc156-1B{9934}; RFLP markers Xcdo442-1B and Xbcd249-1B were found to be associated with Rf3 on 1BS{860}.
Rf4{1619}. [Rf2{1619}]. 6B{1619,873}.1BS{868}. v: L3{868}; (T. timopheevi/Aegilops squarrosa) / 3*Canthatch Rf5{1619}; R113 Rf1{873}.
Rf5{1619}. [Rf3{1619}]. 6D{1619}. v: (T. timopheevi/Aegilops squarrosa)/ 3*Canthatch Rf4{1619}.
Rf6{865,859}. 6AS [T6AL.6AS-6U]{865}.6BS [T6BL.6BS-6U]{865}. tr: Line 2114{865}; Lines 040-5; 061-1{865}; 061-4{865}.
Genes Rfc3 in chromosome 6RL and Rfc4 in chromosome 4RL were reported in {225}. Novel Rf genes were identified on 5AL linked to Xcdo786-5A and XksuH1-5A{860}.


62.2. Restorers for T. longissimum cytoplasm
Vi{867}. 1B{870}.1BS{027}. v: T. turgidum{867}.
Probably derived from a cv. Selkirk (T. aestivum) line with Ae. cylindrica cytoplasm {867}.


62.3. Restorers for photoperiod-sensitive Aegilops crassa cytoplasm
Morai & Tsunewaki {1047} described photoperiod sensitive CMS caused by Ae. crassa cytoplasm in wheat cv. Norin 26. Almost complete sterility occurred when plants were grown in photoperiods of 15h or longer.
Rfd1{1047}. 7BL{1047}. v: Chinese 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}.



63.Ribosomal RNA

The 5S-Rrna-1 loci were physically mapped in 1AS, 1BS, and 1DS and the 5S-Rrna-2 loci were physically mapped in 5AS, 5BS and 5DS of Chinese Spring using deletion lines {1043}.
Table 1 in {276} lists the chromosome or chromosome arm locations of rRNA loci in 12 Triticeae species.

63.1. 5S rRNA genes
Within the Triticeae there are basically two sets of 5S rRNA loci. One set, identified by repetitive units 320-468 bp in length, is located on group 1 chromosomes. The other set, identified by repetitive units 469-500 bp in length, is on group 5 chromosomes. Within species the repetitive units at a locus are extremely uniform in size and sequence. They remain stable in foreign genetic backgrounds.
5S-Rrna-A1. [5SDna-A1{295}]. 1AS{295,658}. v: CS{1043}.
5S-Rrna-B1. [5SDna-B1{295}]. 1BS{039,295}. dv: T. monococcum. ma: A PCR marker specific 5S-Rrna-B1 was developed{9974}.
5S-Rrna-D1. [5SDna-D1{295}]. 1D{295,758}.1DS{295}. v: CS{295,758}. dv: T. tauschii{758}. ma: A PCR marker specific for 5S-Rrna-D1 was developed in {9974}.
5S-Rrna-E1. [5SDna-E1{960}]. 1E{1290}. dv: L. elongatum.
5S-Rrna-R1. [5SDna-R1{1206}]. 1RS{039,1206}. al: S. cereale. ma: A PCR marker specific for 5S-Rrna-R1 was developed in {9974}.
5S-Rrna-Sc1. [5SDna-Sc1{960}]. 1Sc{1290}. al: Elymus ciliaris.
5S-Rrna-St1. [5SDna-St1{960}]. 1St{1290}. al: E. trachycaulus.
5S-Rrna-Y1. [5SDna-Y1{960}]. 1Y{1290}. al: E. ciliaris.
5S-Rrna-A2. [5SDna-A2{295}]. 5AS{295,658}. v: CS{295}. dv: T. monococcum{295,658}.
5S-Rrna-B2. [5SDna-B2{295}]. 5BS{295}. v: CS.
5S-Rrna-D2. [5SDna-D2{295}]. 5D{295,758}.5DS{758}. v: CS{295,758}. dv: T. tauschii{758}.
5S-Rrna-R2. [5SDna-R2{1206}]. 5RS{1206}. al: S. cereale.
5S-Rrna-Ht2. [5SDna-Ht2{960}]. 5Ht{1290}. al: E. trachycaulus.
5S-Rrna-U2. [5SDna-U2{295}]. 5U{295}. al: T. umbellulatum.
5S-Rrna-V2. [5SDna-V2{960}]. 5V{1290}. al: D. villosa.
5S-Rrna-H3. [5SDNA-H3{793}]. 2H{710}.2HL{793}. al: Betzes Barley; Sultan barley.
5S-Rrna-H4. [5SDNA-H4{793}]. 3HL{793}. al: Betzes barley; Sultan barley.
5S-Rrna-H5. [5SDNA-H5{793}]. 4HL{793}. al: Betzes barley; Sultan barley.
5S-Rrna-H6. [5SDNA-H6{793}]. 4HS{793}. al: Betzes barley; Sultan barley.



64.Seedling Leaf Chlorosis
sc{149}. 3BS{149}. s: CS*/Hope3B{149}. v: Hartog{149}; Suneca{149}; wheats with Sr2{149}.
Leaf chlorosis is affected by temperature and light and is enhanced by infection with pathogens. sc is completely linked with Pbc (pseudo-black chaff) and Sr2 (reaction to Puccinia graminis).


65.Segregation Distortion
QSd.ksu-1D{9931}. 1DL{9931}. dv: Ae. tauschii var. meyeri acc. TA1691/var. typica acc. TA1704{9925}. ma: Association with Xcmwg706-1D{9931}.
QSd.ksu-3D{9931}. 3DS{9931}. dv: Ae. tauschii var. meyeri acc. TA1691/var. typica acc TA1704{9925}. ma: Association with Xwg177-3D{9931}.
QSd.ksu-4D{9931}. 4DS{9931}. dv: Ae. tauschii var. meyeri acc. TA1691/var. typica acc. TA1704{9925}. ma: Association with XksuF8-4D{9931}.
QSd.ksu-5D.1{9931}. 5D{9931}. dv: Ae. tauschii var. meyeri acc. TA1691/var. typica acc. TA1704{9925}. ma: Association with Xcdo677-5D{9931}.
QSd.ksu-5D.2{9931}. 5DL{9931}. dv: Ae. tauschii var. meyeri acc. TA1691/var. typica acc. TA1704{9925}. ma: Association with Xglk614-5D (synonym 'Xtag614-5D'){9931}.
QSd.ksu-5D.3{9931}. 5DL{9931}. dv: Ae. tauschii var. meyeri acc. TA1691/var. typica acc. TA1704{9925}. ma: Association with Xwg1026-5D{9931}.
QSd.ksu-7D{9931}. 7DS{9931}. dv: Ae. tauschii var. meyeri acc. TA1691/var. typica acc. TA1704{9925}. ma: Association with Xglk439-7D (synonym 'Xtag439-7D'){9931}.


66.Sterol Esterification in Kernels - Synthesis of b-Sitosterol Esters
Two sterol-ester phenotypes, p-L (palmitate + linoleate) and L (linoleate) are inherited as alleles at a single locus.
Pln{428}. [P-L{428}]. 7DS{1476}. v: Aradi{428}; Aragon 03{428}.
pln{428}. [L{428}]. L{428}. v: Mara{428}; Pane 247{428}.


67.Temperature-Sensitive Winter Variegation
This phenotype involves reduced vigour and chlorotic patches on leaves of certain genotypes in Ae. umbellutata cytoplasm when grown at low temperatures {1596}.
Vgw. Variegation is dominant {1596}. [Vg{1021}]. 5BL{1021}. v: Bersee{1596}; Cappelle-Desprez{1596}; Hobbit Sib{1596}; Mara{1596}.
vgw{1021}. v: Besostaya I{1596}; CS{1596}; Poros{1596}; Sava{1596}; T. spelta{1596}.


68.Tenacious Glumes
Tg1{1240}. Derived from T. tauschii. Dominant. [Tg{1240}]. 2DS{1240}. v: Synthetic ABD wheats{652}.
Tg2{0046}. Derived from T. dicoccoides 2BS{0046}. ma: Tg2 is associated with Xrsq805(Embp)-2B and Xpsr899-2B{0046}.


69.Tiller Inhibition
tin1{1212}. Restricted tiller number is recessive{1212} [Tin{1212}]. 1AS{1212}. v: Israel Uniculm 494{1212}.
tin2{1212}. Tiller-reducing affect of this allele was dominant {9909}. [Tin{9909}]. 2A{9909}. v: 88 F2 185{9909}.


70.Uniculm Stunt
Stunting is favoured by a combination of long days and low night temperatures {581}. Caused by duplicate recessive genes, us1 and us2, located in chromosomes 4A and 5B, respectively {200}.
Genotypes: Normal v: Us1 us2: Alfa {581}; Jaral {581}.
Normal v: us1 Us2: Mabruk {581}.
Stunted v: us1 us2: Line 492 {581}.


71.Variegated Red Grain Colour
vg{498}. v: Line 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}.


72.Yield Components

72.1. Grain weight

72.1.1. 50-grain weight
QFgw.ocs-4A.1{0047}. 4A{0047}. v: CS/CS(Kanto107 4A) mapping population{0047}. ma: Associated with Xbcd265-4A and Xbcd1738-4A{0047}.


72.1.2. 1000-grain weight
QTL: Two QTLs for 1,000-kernel weight were assigned to chromosome 3A in RSLs from Cheyenne*7/Wichita 3A {0025}. QTLs for grain size were identified on chromosome arms 1DS, 2DL and 6BL in a RIL population from RS111/CS {0236}. Eight QTLs for 1,000-kernel weight (54 % of the variation) were mapped in Forno/ Oberkulmer spelt {0280}.
QGw1.ccsu-1A{0165}. 1AS{0165}. v: RS111/CS mapping population{0165}. ma: Associated with Xwmc333-1A{0165}.
QTgw.ipk-5A{0255}. 5AL{0255}. v: Opata/W-7984 (ITMI) RI mapping population{0255}; The higher yielding allele was contributed by W-7984{0255}. ma: Associated with Xfba351-5A and Xcdo1312-5A{0255}.



72.2. Grain weight/ear
QGwe.ocs-4A.1{0047}. 4AS{0047}. v: CS/CS(Kanto107 4A) mapping population{0047}. ma: Associated with Xbcd1738-4A{0047}.
QGwe.ipk-2D{0255}. 2DS{0255}. v: Opata/W-7984 (ITMI) RI mapping population{0255}; Higher grain weight was contributed by Opata{0255}. ma: Associated with Xcdo1379-2D and Xbcd1970-2D{0255}.
QGwe.ipk-4A{0255}. 4AL{0255}. v: Opata/W-7984 (ITMI) RI mapping population{0255}; Higher grain weight was contributed by Opata{0255}. ma: Associated with Xmwg549-4A, Xabg390-4A and Xbcd1670-4A{0255}.
QGwe.ipk-4A coincided with QTLs for height (QHt.ipk-4A), spike length (XEl.ipk-4A) and grain number (QGnu.ipk-4A) {0255}.


72.3. Grain number per spike
QGnu.ipk-4A{0255}. 4AL{0255}. v: Opata/W-7984 (ITMI) RI mapping population{0255}; Higher grain number was contributed by Opata{0255}. ma: Associated with Xmwg549-4A, Xabg390-4A and Xbcd1670-4A{0255}.
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}.
QTL: Three QTLs for kernel number per spike were assigned to chromosome 3A in RSLs from Cheyenne*7/Wichita{0025}.


72.4. Plant yield
QYld.ocs-4A.1{0047}. 4AS{0047}. v: CS/CS(Kanto107 4A) mapping population{0047}. ma: Associated with Xbcd1738-4A{0047}.


72.5. Spikelet number/ear
QSpn.ocs-4A.1{0047}. 4AS{0047}. v: CS/CS(Kanto107 4A) mapping population{0047}. ma: Associated with Xbcd1738-4A{0047}.


72.6. Spike number per square metre
QTL: A QTL for spike number per square metre was assigned to chromosome 3A in RSLs from Cheyenne*7/Wichita 3A {0025}.


72.7. Spike length
QEl.ipk-1B{0255}. 1BL{0255}. v: Opata/W-7984 (ITMI) RI mapping population{0255}; Longer ear was contributed by Opata{0255}. ma: Associated with Xbcd388-1B and Xwg605-1B{0255}.
QEl.ipk-4A{0255}. 4AL{0255}. v: Opata/W-7984 (ITMI) RI mapping population{0255}; Longer ear was contributed by Opata{0255}. ma: Associated 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{0255}. 5AL{0255}. v: Opata/W-7984 (ITMI) RI mapping population{0255}; Longer ear was contributed by W-7984{0255}. ma: Associated with Xmwg522-5A{0255}.
QTL: Five QTLs for spike length were detected in Courtot/Chinese Spring{0114} but only one on chromosome arm 5AL was consistent for at least two years.


72.8. Tiller number/plant
QTn.ocs-4A.1{0047}. 4AS{0047}. v: CS/CS(Kanto107 4A) mapping population{0047}. ma: Associated with Xpsr163-4A{0047}.



73.Yellow Berry Tolerance
QTL : A QTL for yellow berry tolerance, contributed by RS111, was associated with Xgwm190-5D and Xgwm174-5D in a RIL population from RS111/CS {0237}. A tolerance QTL contributed by CS, the susceptible parent, was detected on 6B {0237}.


74.Proteins

74.1. Grain protein content
Pro1{777}. 5DL{777}. s: CS*6/Hope 5D{777}.
May be identical to Vrn3.
Pro2{777}. 5DS{777}. s: CS*6/Hope 5D{777}.
QGpc.ccsu-2D{0015}. 2DL{0015}. ma: Association with microsatellite locus Xwmc41-2D accounted for 19% of the variation in grain protein content between PH132 and WL711{0015}.
QGpc.ndsu-6B{623}. 6BS{623}. tv: Langdon{623}.
QGpc.ndsu-6Ba{623}. tv: Langdon{623}.
QGpc.ndsu-6Bb{623,0071}. tv: Langdon-T. dicoccoides 6B{623}. v: Glupro{0179}. ma: QGpc.ndsu-6B was associated (LOD score = 18.9) with the interval Xmwg79-6B - Xabg387-6B. These loci were mapped in 6BS: Xmwg79-6B - 5.9 cM - Xabg387-6B - 9.0 cM - centromere{623}; Located in the 4 cM interval flanked by Xmwg79-6B and Xcdo365-6B{0244}; Flanking microsatellite markers and PCR-specific markers for Glupro are available{0179}.
QPro.mgb-4B. Associated with Gai1 and Xpsr622-4B {110}2.
QPro.mgb-5A. Associated with Xpsr911-5A {110}2 and Xcdo412-5A{0343}*.
QPro.mgb-6A.1. Associated with Xpsr167-6A and XksuG8-6A {110}2.
QPro.mgb-6A.2. Associated with Xmgb56-6A {110}2 and Xpsr627-6A{0343}*.
QPro.mgb-6B. Associated with Gli-B2-6B {110}2 and Nor-2{0343}*. ma: QGpc.ndsu-6B was associated (LOD score =18.9) with the interval Xmwg79-6B - Xabg387-6B. These loci were mapped in 6BS: Xmwg79-6B - 5.9cM - Xabg387-6B - 9.0 cM - centromere{623}.
Qpro.mgb-7A. Associated at P<=0.01 with Pan2{0343}*.
QPro.mgb-7B. Associated with Xpsr490(Ss1)-7B, Pc {110}2 and Xutv913-7B{0343}*.

QTLs for grain protein content were detected on chromosome arms 6AS (associated AFLP marker, XE38M60200) and 1BL (associated RFLP marker, Xcdo1188-1B) in Courtot/Chinese Spring {0141}.
Nine QTLs (51% of the variation) were mapped in cross 'Forno'/ 'Oberkulmer' spelt {0280}.
A QTL for grain and flour protein content, contributed by CS, was associated with XTri-1D/Centromere in a RSL population from the cross Cheyenne (high quality wheat)/CS (low quality wheat) {0251}.

74.2. Enzymes

74.2.1. Acid phosphatase
Acph-A1{504}. [Acph2{516},Acph3{516},Acph-B1{936}]. 4AS{504,516}. v: CS.
Acph-B1{504}. [Acph4{516},Acph8{516},Acph-A1{936}]. 4BL{504,516}. v: CS.
Acph-D1. [Acph5{516},Acph6{516}]. 4DL{504,516}. v: CS.
Acph-H1{1153}. 4H{1153}. ad: CS/Betzes.
Acph-Mv1{237}. [Aph-v{237},Acph-Mv1{985}]. 4Mv{237}. tr: H-93-33{984}.
Acph-R1. 7R{1457}.7RS{506}. ad: CS/Imperial.
Acid phosphatase gene loci were reported for 7RL in S. cereale {1251}, chromosomes L1 (= 7Agi) and L4 (= 4Agi) of Thin. intermedium {361}, and chromosome E of Ae. umbellulata {69}. Two loci on 7R were separated by 25 +or- 5.2cM {1534}.Wehling {1559} identified four acid phosphatase loci in S. cereale, three of which were located in 7R.
Acph-Ss1{1140}. 4Ss{1140}. ad: CS/T. searsii.


74.2.2. Alcohol dehydrogenase (Aliphatic)
Adh-A1{502}. [AdhB{502},Adh-B1{504}]. 4A{502}.4AL{504,516}. v: CS.
Adh-B1{501,502}. [Adh1{501},AdhA{502},Adh-A1{504}]. 4B{502}.4BS{504,516}. v: CS.
Adh-B1a{1442}. [Adh11{501},Adh-A1a{1442}]. v: CS. tv: PI 226951{501}; Malavika{1442}.
Adh-B1b{1442}. [Adh12{501},Adh-A1b{1442}]. v: Rageni derivative{1443}. tv: CI 4013{501}; Bijaga Yellow{1442}.
Adh-B1b was the only variant Adh-1 allele detected in study of a large number of T. aestivum and T. turgidum accessions {503}.
Adh-C1{1278}. [G{1278}]. ad: T. aestivum cv.Alcedo / Ae. caudata line G.
Adh-D1{504}. [AdhD{502}]. 4D{502}.4DS{504,516}. v: CS. ma: Adh-D1 [Adh1, Adh2] was mapped 4cM distal to Xpsr163-4D and closely proximal to Xcsiha114-4D.1 [XcsIHA114-1a'] {757}.
Adh-Agi1{560},{374}. [Adh-X1{361}]. 4Agi{560}. ad: Vilmorin 27/Thin. intermedium; Caribo/Thin. intermedium.
Adh-E1{518}. 4ES{518}. ad: CS/E. elongata.
Adh-H1{520}. 4H{520}. ad: CS/Betzes.
Adh-Mv1{984}. [ADHmu{984},Adh-Mv1{985}]. 4Mv{984}. v: H-93-33.
Adh-R1{1457}. [AdhR2{582}]. 4R{582,1457}.4RS{506}. ad: CS/Imperial{1457,506}; FEC28/Petkus{043}; Holdfast/King II{582}.
Adh-V1{1026,242}. 4V{1026}. ad: CS/D. villosum.

Three Adh genes were identified in Hordeum vulgare and H. spontaneum {144,490,493,520}. Two of these were tightly linked at the Adh-H1 locus {144}. The third gene was tentatively located in 5H {490}.
A low-level of aliphatic alcohol dehydrogenase activity is commonly observed on zymograms in the absence of added substrate {513}; this may account for the observation of wheat lactate dehydrogenase that was reported in {1465}.
The gene series formerly designated Adh-2 and Adh-3 appear under 2.20. Aromatic Alcohol Dehydrogenase

74.2.3. Aminopeptidase
Amp-A1{504}. 6AS{504,516}. v: CS.
Amp-A1a. v: CS{1533}.
Amp-A1b. v: Vitka{1533}.
Amp-B1{504}. 6BS{504,516}. v: CS.
Amp-B1a. v: CS{1533}.
Amp-B1b. v: Iskra{1533}.
Amp-B1c{703,1244}.Null allele v: T. spelta IPSR 1220017{703}; Sinvalocho M.A{1244}.
Amp-C1{1278}. 6D{1278}. ad: Alcedo/Ae. caudata line D.
Amp-D1{504}. 6DS{504,516}. v: CS.
Amp-D1a{703}. v: CS.
Amp-D1b{703}. v: Sears' Synthetic IPSR1190903.
Amp-Age1{1575}. 6Age{1575}. ad,su: Rescue/Thin. elongatum.
Amp-Agi1{703}. 6Agi{703}. ad: Vilmorin 27/Thin. intermedium.
Amp-E1{518}. 6E{518}. ad: CS/E. elongata.
Amp-H1{520}. 6H{520}. ad: CS/Betzes.
Amp-R1{1457}. 6R{1457,1280}. ad: CS/Imperial{1457}; Holdfast/King II{1280}.
Amp-A2{703}. 4AL{703}. v: CS.
Amp-A2a{703}. v: CS.
Amp-A2b{703}. v: T. spelta IPSR 1220017.
Amp-B2{703}. 4BS{703}. v: CS.
Amp-B2a{703}. v: CS.
Amp-B2b{703}. v: Timstein.
Amp-B2c{703}. v: Hope.
Amp-D2{703}. 4DS{703}. v: CS.
Amp-D2a{703}. v: CS.
Amp-D2b{703}. v: Sears' Synthetic IPSR 1190903.
Amp-D2c{703}. v: Bersee.
Amp-Agi2{703}. 4Agi{703}. ad: Vilmorin27/Ag. intermedium.
Amp-E2{703}. 4E{703}. ad: CS/E. elongata.
Amp-H2{703}. 4H{703}. ad: CS/Betzes.
Amp-Hch2{703}. 4Hch{703}. ad: CS/H. chilense.
Amp-J2{703}. 4J{703}. ad: CS/Thin. junceum.
Amp-Mv2{235}. 4Mv{235}. su: H-93-33{235}.
Amp-R2{703}. 4R{703}.4RS{702,093}. ad: CS/Imperial.
Amp-Sl2{703}. 4SlL{703}. ad: CS/Ae. sharonensis{180}. tr: 4DS.4DL-4SlL{660}.
Amp-V2{703}. 4V{703}. ad: CS/D. villosum.
Amp-A3{703}. 7AS{703}. v: CS.
Amp-A3a{703}. v: CS.
Amp-A3b{703}. v: Timstein.
Amp-H3{703}. 7H{703}. ad: CS/Betzes.


74.2.4. Alpha-amylase
a-Amy-A1{007}. [Amy6A{1082}]. 6AL{412,1082}. v: CS.
a-Amy-A1a{007}. [Amy 6A1{1084}]. v: CS.
a-Amy-A1b5{007}. v: Bezostaya 1; Kavkaz.
a-Amy-A1c5. [Amy 6A1m{1084}]. v: Aka.
a-Amy-B1{007}. [Amy6B{1082}]. 6BL{412,1082}. v: CS.
a-Amy-B1a{007}. [Amy 4{1084},Amy 6B1{1084},Amy 6B2o{1084}]. v: CS{007}; Rare.
a-Amy-B1b{007}. [Amy 4m{1084},Amy 6B1o{1084},Amy 6B2{1084}]. v: Mara{007}.
a-Amy-B1c{007}. [Amy 4{1084},Amy 6B1{1084},Amy 6B2{1084}]. v: Sava{007}; Rare.
a-Amy-B1d{007}. [Amy 4m{1084},Amy 6B1o{1084},Amy6B2o{1084}]. v: Sicco{007}; Rare.
a-Amy-B1e{007}. [Amy 4m{1084},Amy 6B14'{1084},Amy 6B2o{1084}]. v: Cappelle-Desprez{007}.
a-Amy-B1f {007}. [Amy4m{1084},Amy 6B14{1084},Amy 6B2o{1084}]. v: Sappo{007}.
a-Amy-B1g{007}. [Amy 4{1084},Amy 6B14{1084},Amy 6B2o{1084}]. v: Cheyenne{007}.
a-Amy-B1h{007}. [Amy 4{1084},Amy 6B1o{1084},Amy 6B2o{1084}]. v: T. macha Line 1{007}; Rare.
Two types of nomenclature were assigned to the genes encoding the a-AMY-1 isozymes. In one, allelic states were defined for individual isozymes {1084} whereas in the other, several isozymes were considered to be the products of compound loci {007,412}. This listing shows the 'alleles' described in {1084} which are assumed in {007}
to be synonymous with the a-Amy-B1a through a-Amy-B1h nomenclature. Amy 4 and Amy 41 are unmapped alternatives {1084} which appear to be identical to zymogram bands [bands 9 and 9b {007}] forming part of the a-Amy-B1phenotype. Amy 6B1 [with forms Amy 6B1o, and Amy 6B14', considered to be mutually exclusive {1084}] and Amy 6B2 [with forms Amy 62 and Amy 6B2o {1084}] describe further aspects of a-Amy-B1 {007}. See a-Amy1 below for further consideration of Amy 6B2 {1084}.
a-Amy-D1{007}. [Amy6D{1082}]. 6DL{412,1082}. v: CS.
a-Amy-D1a{007}. [Amy6D1{1084},Amy 6D2{1084}]. v: CS.
a-Amy-D1b{007}. [Amy6D1{1084},Amy 6D2{1084}]. v: Prelude{1082}; Capelle-Desprez{007}.
a-Amy-D1c. [Amy6D1m{1084},Amy 6D2{1084}]. v: T. spelta var. duhamelianum.
a-Amy-Agi1{374}. 6Agi{374}. ad: Vilmorin 27/Thin. intermedium.
a-Amy-E1{013}. 6E{013}. ad: CS/E. elongata.
a-Amy-H1. [a-Amy1{146}]. 6H{146,1051}. ad: CS/Betzes.
a-Amy-R1{013}. 6RL{013}. su,ad: CS/Imperial; CS/King II; Holdfast/King II.
a-Amy-Rm1{013}. 6RmL{013}. ad: CS/S. montanum.
a-Amy-S1{598}. 6SS{598}. v: Wembley derivative 31. al: Ae. speltoides.
It was estimated {902} that there are two a-Amy-1 genes in 6A and five or six in both 6B and 6D, and three or four a-Amy-2 genes at each of the 7A, 7B, and 7D loci.
a-Amy-A2{007}. [Amy7A{1082}]. 7AL{412,1082}. v: CS.
a-Amy-B2{007}. [Amy7B{1082}]. 7BL{412,1082}. v: CS.
a-Amy-B2a{412}. [Amy 7B1{1084},Amy 7B2{1084}]. v: CS.
a-Amy-B2b{412}. [Amy 7B1{1084},Amy 7B2m{1084}]. v: Hope.
The alternative states of Amy 7B2, namely, Amy 7B2 and Amy 7B2m {1084}, are identical to the variation in band 2 {412}. The complete description of the a-Amy-B2 variation also includes variation in band 11 {412}.
a-Amy-D2. [Amy7D{1082}]. 7DL{412,1082}. v: CS.
a-Amy-D2a{412}. [Amy 7D1{1084}]. v: CS.
a-Amy-D2b{417}. [Amy 7D1o{1084}]. v: Largo{007}; Sears' Synthetic{007}; VPM1{417}.
a-Amy-Agi2{374}. 7Agi{374}. ad: Vilmorin 27/Thin. intermedium.
a-Amy-E2{013}. 7EL{013}. ad: CS/E. elongata.
a-Amy-H2. [a-Amy2{146}]. 7HL{146,1051,793}. ad: CS/Betzes.
a-Amy-Hch2{1015}. 7Hchbeta{1015}. su,ad: CS/H. chilense.
a-Amy-R2{013}. 7RL{013}. su,ad: CS/Imperial; CS/King II; Holdfast/King II.
a-Amy-Sb2{013}. 7Sb{013}. ad: Holdfast/Ae. bicornis.
a-Amy-U2{013}. 7U{013}. ad: CS/Ae. umbellulata.
Three other a-Amy loci, namely, Amy 6B2, Amy 6D2, and Amy 7B2, were reported {1084}. No variation was observed for the products of Amy 6D2 and Amy 7B2, although nullisomic analysis located the genes in 6DL and 7B, respectively. In accordance with the Guidelines, these genes are assumed to be part of the a-Amy-D1 and a-Amy-B2 loci, respectively. Amy 6B2 was observed to produce alternative phenotypes {1084}. In a test of the segregation of these phenotypes relative to two alternative products of Amy 6B1, the two loci were found to be linked with a recombination frequency of 20.6% {1084}. However, an attempt to confirm the presence of more than one a-Amy locus in 6BL was unsuccessful {007}.
a-Amy1{1084,1083}. [Amy 6B2{1084},Amy-B2{1083}]. 6BL{1084,1083}. v: CS.
a-Amy1a{1083}. [a-Amy-B1a]. v: CS.
a-Amy1b{1083}. [a-Amy-B1b]. v: CS.
a-Amy1c{1083}. [a-Amy-B4]. tv: T. durum ssp. georgicum.
The presence of a-Amy1 reported in {1084} was confirmed by segregational tests in a CS/Jones Fife population and in a population derived from a tetraploid cross {1083}. The recombinations with a-AmyB1 were 9.3% and 22.3%, respectively.
A further set of a-amylase genes, Xa-Amy-5 [a-Amy3], was identified in 5A, 5B and 5D by cross-hybridization with a-AMY-1 and a-AMY-2 probes {080}. Only one gene copy appears to be present at each locus. In rye, evidence was obtained for three a-Amy-1 genes, two or three a-Amy-2 genes and three a-Amy-3 genes {907}.
Synthesis of a-amylase isozymes controlled by a-Amy-1 genes on chromosomes 6A and 6D is reduced in DT6BS compared to euploid CS. This result suggests the presence of a gene(s) on the long arm of chromosome 6B, which is (are) required for GA-induced alpha-amylase synthesis in the aleurone {0072}.


74.2.5. Beta-amylase
b-Amy-A1{008,227}. [b-Amy-A2{008},b-Amy-B1{1331}]. 5AL{008,227}. v: CS{008}. s: CS/Federation{227}.
b-Amy-A1a{008}. [b-Amy-A2a{008},b-B1a{936}]. v: CS.
b-Amy-A1b{008}. [b-Amy-A2b{008},b-B1b{936}]. v: Koga II..
b-Amy-A1c{008}. [b-Amy-A2c{008},b-B1c{936}]. v: T. macha IPSR 1240005.
b-Amy-A1d{008}. [b-Amy-A2d{008},b-B1d{936}]. v: Holdfast.
b-Amy-A1e{008}. [b-Amy-A2e{008},b-B1e{936}]. v: Bezostaya I.
b-Amy-B1{628}. [b-Amy-A1{008}]. 4BL{008,628}. v: CS.
b-Amy-B1a{1330}. [b-Amy-A1a{008,1330}]. v: CS.
b-Amy-B1b{1330}. [b-Amy-A1b{008,1330}]. v: Sears' Synthetic IPSR 1190903.
b-Amy-B1c{1330}. [b-Amy-A1b{008},b-Amy-A1c{1330}]. v: Ciano 67.
b-Amy-B1d{1330}. [b-Amy-A1c{1330,400}]. v: Manella.
b-Amy-C1{1278}. B{1278}. ad: Aestivum cv. Alcedo /Ae. caudata line B.
b-Amy-D1{008}. 4DL{008,628}. v: CS.
b-Amy-D1a{008}. v: CS.
b-Amy-D1b{008}. v: Bersee.
b-Amy-D1c{008}. v: Sears' Synthetic. Rare.
Previously listed alleles b-Amy-D1d and -D1e were found to be b-Amy-B1 alleles {400}.
Two b-Amy-Dt1 alleles were predominant in 60 accessions of T. tauschii {1578}.
b-Amy-Agi1{168,013}. 4Agi{168}. ad: Vilmorin27/Thin. intermedium.
b-Amy-Eb1{661}. 5EbL{661}. tr: 5AS.5EbL.
b-Amy-H1. 4H{1153}. ad: CS/Betzes.
b-Amy-Hch1{013}. 4Hch{013}. ad: CS/H. chilense.
b-Amy-R1. [b-Amy-R2{013},b-AmyR1{043}]. 5R{103,1280}.5RL{043}. ad: FEC 28/Petkus{043,82}; Holdfast/King II{043,1280}. tr: CS/Imperial 5BL-5RL{043}.
b-Amy-Sl1{013}. 4Sl{013}.D{013}. ad: CS/Ae. sharonensis D{013}. su: CS/Ae. sharonensis. ad: CS/Ae. longissima.
b-Amy-U1{013}. [b-Amy-U2{013}]. 5U{013}. su: CS/Ae. umbellulata.
A second set of loci with homology to b-Amy-1 genes was identified in 2AS, 2BS and 2DS and designated the Xb-Amy-2 [b-Amy-2 {1331}] set. Evidence for these genes derives from cross-hybridization with a b-AMY-H1 cDNA probe {1331}. Further members of the same set were identified in 2H {732}, and 2R and 2U {1331}.
Sixty Ae. tauschii lines revealed two b-Amy-Dt1 alleles {1578}.


74.2.6. Endopeptidase
Ep-A1{516}. 7AL{516}. v: CS.
Ep-A1a{516,708}. v: CS.
An EP isozyme encoded by Ep-A1a of CS is visible on zymograms following starch gel electrophoresis {516}. The product of this allele is not observable, however, on zymograms following isoelectric focusing {708}.
Ep-A1b{708}. v: Cappelle-Desprez{708}; Hobbit{704}; Rendezvous{1603}.
Ep-A1c{708}. v: Sears' Synthetic.
Ep-A1d{894}.Isozyme 6. v: PI 294994{894}.
Ep-B1{516}. [Ep1{516}]. 7BL{516}. v: CS.
Ep-B1a{708}. v: CS.
Ep-B1b{708}. v: Cappelle-Desprez.
Ep-B1c{708}. v: Ciano 67.
Ep-B1d{708}. v: Bersee.
Ep-B1e{708}. v: Sears' Synthetic.
Ep-D1{516}. 7DL{516}. v: CS.
Ep-D1a{708}. v: CS.
Ep-D1b. [EP-V1{973}]. v: 5L 219{1521}; H-93-70{1521}; Hyak{021}; Madsen{020}; Rendezvous{708}; VPM1{973}.
Ep-D1c{708}. v: Sears' Synthetic.
Ep-D1d{1587}. Null allele. v: Wheats with Lr19{1587}.
Ep-D1e{894}.Isozyme 5. v: PI 294994{894}.
Ep-E1{518}. 7EL{518}. al: CS/E. elongata.
Ep-H1{520}. 7HL{520}. al: CS/Betzes.
Ep-Hch1{708}. 7Hch{708}. su: CS/H. chilense.
Ep-Ht1{1037}. 7Htp{1037}. ad: CS/E. trachycaulus.
Ep-Mv1{985}. [Ep-Mv1{985}]. 7MvL. su: 7Mv{7D}.
Ep-R1{092,266,708}. 6RL{092}. ad: CS/Imperial.
Ep-Sb1{708}. 7Sb{708}. su: Holdfast/Ae. bicornis.
Ep-Sl1{517}. 4Sl{517}. ad: CS/Ae. longissima.
Ep-Ss1{1140}. 7Ss{1140}. ad: CS/T. searsii.
Ep-U1{708}. 7U{708}. su: CS/Ae. umbellulata.
Ep-V1{708}. 7V{708}. ad: CS/D. villosum.
Ep-B2{599}. 6BS{599}.
An Ep locus was located in 4RS in King II {1280}, using Holdfast/King II addition lines and in 4R in Imperial {266} using Chinese Spring/Imperial addition lines.


74.2.7. Esterase
Genetic control of esterases [carboxylic ester hydrolases (E.C.3.1.1.1)] was the subject of a comparative study {814}.

74.2.7.1. EST-1
EST-1 is a dimeric enzyme that electrofocuses around pH4.0 and is expressed in all tissues except endosperm {814}.
Est-A1. [EstA{061}]. 3AS{060}. v: CS.
Est-B1. [EstB{061}]. 3B{060}.3BS{100}. v: CS.
Est-D1. [EstD{061}]. 3D{060}.3DS{100}. v: CS.
Est-E1{518}. 3ES{518}. ad: CS/E. elongata.
Est-H1{814}. 3H{814}. ad: CS/Betzes.
Est-R1. [EstR{061}]. 3R{060,1254}. ad: CS/Imperial{060}; Holdfast/King II{100}; Kharkov/Dakold{100}.
Est-S11{814}. 3S1{814}. ad: CS/Ae. longissima.
Each of 208 hexaploid accessions carried the same Est-1 alleles except accessions of T. compactum var. rubriceps, each of which carried an Est-B1 or Est-D1 electrophoretic mobility variant {585}.


74.2.7.2. EST-2
EST-2 is a coleoptile-specific monomeric enzyme that electrofocuses at low pI.
Est-A2. [Est-2A{585}]. 3A{585}. v: CS.
Est-B2. [Est-2B{585}]. 3BL{585}. v: CS.
Est-D2. [Est-2D{585}]. 3DL{585}. v: CS.
Among 208 hexaploid accessions, an apparent Est-B2 null allele occurred frequently in accessions of T. macha and T. sphaerococcum and occasionally in accessions of T. compactum. The allele was not observed in T. aestivum and T. spelta accessions {585}.


74.2.7.3. EST-3
EST-3 is a monomeric enzyme that is expressed in young seedlings (this enzyme was not observed in {814}.
Est-B3. [Est-3B{585}]. 7BS{585}. v: CS.
Est-D3. [Est-3D{585}]. 7DS{585}. v: CS.
Est-H3{520}. 7H{520}. ad: CS/Betzes.
One accession carrying an apparent Est-B3 null allele and one carrying an apparent Est-D3 null allele were found among 208 hexaploid accessions {585}.
A 7AS locus encodes three esterase isozymes in immature grains {009}.


74.2.7.4. EST-4
EST-4 is a monomeric, leaf-specific enzyme that electrofocuses around pH 4.5.
Est-A4. [Est-4A{585}]. 6AL{585,919}. v: CS.
Est-B4. [Est-4B{585}]. 6BL{585,919}. v: CS.
Est-D4. [Est-4D{585}]. 6DL{585,919}. v: CS.
Probable Est-A4 and Est-D4 null alleles were detected in several accessions of T. compactum var. rubriceps {585}; otherwise, no Est-4 variant occurred among 208 hexaploid accessions {585}.
An esterase gene was located in chromosome L7 (= 6Agi) of Thin. intermedium {361}.


74.2.7.5. EST-5
EST-5 consists of 20 or more monomeric, grain-specific isozymes that electrofocus between pH 5.6 and 7.0.
Est-A5{009}. 3AL{009}. v: CS.
Est-A5a{009}. v: CS.
Est-A5b{009}. v: Kalyansona{009}; T. compactum AUS12084{756}.
Est-B5{009}. 3BL{009}. v: CS.
Est-B5a{009}. v: CS.
Est-B5b{009}. v: Big Club.
Est-B5c{009}. v: Timstein.
Est-B5d{009}. v: Sears' Synthetic.
Est-D5{009}. 3DL{009}. v: CS.
Est-D5a{009}. v: CS.
Est-D5b{009}. v: T. macha.
Est-D5c{009}. v: Hobbit 'S'.
Est-D5d{009}. v: T. macha Line 1.
Est-D5e{756}. v: T. macha WJR 38548.
Encoding of the endosperm esterases of hexaploid wheat by 12-15 genes in five compound loci located in 3AL, 3BL, 3DL, 3AS and 3DS was postulated in {1204}.
Three and six alleles at Est-Dt5 (in T. tauschii) were reported in {756} and {1578}, respectively.
In S. cereale, in addition to Est-R1, genes encoding leaf esterases were located in three chromosomes {1561}. These included a gene designated Est8 in 6R in cvs. Imperial and King II, a gene designated Est2 and two genes, designated Est6 and Est7, which are part of a separate compound locus {1560}, in 5RL in Imperial, and a gene designaged Est10 in 4R of King II and 4RL of Imperial. In Hordeum vulgare, genes encoding leaf esterases were located in 3H {1071; see also, 520,580} and 7H {520}.
Est-Agi5{374}. 3Agi{374}. ad: Vilmorin 27/Thin. intermedium.
Est-H5{010}. 3H{010}. ad: CS/Betzes.
Est-Hch5{010}. 3Hch{010}. ad: CS/H. chilense.
Est-R5{010}. [EstA{737}]. 6R{043,1280}. ad: CS/Imperial{010,043}; Kharkov/ Dakold 6RL{010,1280}; CS/King II{010}; Holdfast/King II{043,1280}.
A second S. cereale gene encoding grain esterases, designated EstB, was located in 4RL in King II and Petkus and in 7RS in Imperial {737}.
Est-Rm5{010}. [EstB{737}]. 6Rm{010}.6RmL{737}. ad: CS/S. montanum.
Est-Sb5{010}. 3Sb{010}. su,ad: CS/Ae. bicornis.
Est-Sl5{010}. 3Sl{010}. ad: CS/Ae. longissima.
Sixty Ae. tauschii lines revealed six Est-Dt5 alleles {1578}.


74.2.7.6. EST-6
EST-6 is a dimeric enzyme that electrofocuses around pH 7.6 and is specific to endosperm.
Est-A6{1130}. 2AS{1130}. v: CS.
Est-A6a{1130}. v: CS.
Est-A6b{1130}. v: Ceska Previvka.
Est-B6{1130}. 2BS{1130}. v: CS.
Est-B6a{1130}. v: CS.
Est-B6b{1130}. v: Hope.
Est-D6{1130}. 2DS{1130}. v: CS.
Est-D6a{1130}. v: CS.
Est-D6b{1130}. v: Sears' Synthetic IPSR 1190903.
Est-M6{1130}. 2MS{1130}. su: CS/Ae. comosa.
Est-R6{370}. 2RS{370}. al: DS2 x RxL10 rye popn.
EST-7 is a monomeric enzyme that electrofocuses in the same region as EST-6 but is specific to green tissues.
A group of leaf esterase isozymes controlled by the long arms of the homoeologous group 3 chromosomes were reported {919}. The relationship of these esterases to EST-2 and to the leaf esterases designed EST-6 reported in {629} has not been determined.

A group of leaf esterase isozymes controlled by the long arms of the homoeologous group 3 chromosomes were reported {919}. The relationship of these esterases to EST-2 and to the leaf esterase designed EST-6 reported in {629} has not been determined.

74.2.7.7. EST-7
EST-7 is a monomeric enzyme that electrofocuses in the same region as EST-6 but is specific to green tissues.
Est-A7{812}. 2AL{812}. v: CS.
Est-B7{812}. 2BL{812}. v: CS.
Est-D7{812}. 2DL{812}. v: CS.
Est-D7a{812}. v: CS.
Est-D7b{812}. v: Synthetic {IPSR 1190903}.
Est-E7{812}. 2E{812}. ad: CS/E. elongata.
Est-H7{812}. 2HL{812}. ad: CS/Betzes.
Est-R7{812}. 2RL{812}. ad: CS/Imperial. su: Holdfast/KingII.
Est-Rm7{812}. 2Rmalpha{812}. ad: CS/S. montanum.
Est-U7{812}. 2U{812}. ad: CS/Ae. umbellulata.
Est-V7{812}. 2V{812}. ad: CS/D. villosum.


74.2.7.8. EST-8
EST-8 consists of about 10 isozymes that electrofocus between pH 4.5 and 6.5 and are expressed only in vegetative tissues. EST-8 is likely to be the enzyme previously described in {919} and {629}.
Est-A8{629,814}. [Est-A6{629}]. 3AL{629}. v: CS.
Est-B8{613,814}. [Est-B6{629}]. 3BL{629}. v: CS.
Est-D8{629,814}. [Est-D6{629}]. 3DL{629}. v: CS.
Est-R8{613,814}. 6RL{629}. ad: CS/Imperial, CS/King II.


74.2.7.9. EST-9
EST-9 is a monomeric enzyme that electrofocuses around pH 5.0 and is expressed only in embryos.
Est-A9{814}. 3AS{814}. v: CS.
Est-B9{814}. 3BS{814}. v: CS.
Est-D9{814}. 3DS{814}. v: CS.


EST-2, EST-5 and EST-8 are controlled by genes on 3L and where a recombination test was possible between Est-D5 and Est-D8, no segregation was observed. The different gene symbols were retained because of the different tissue specificities and polymerisation profiles of the enzymes. The same arguments surround the EST-1 and EST-6 genes located in the 3S arms {814}.
The Est-6 gene of rye was mapped {249}. The Est-6 genes of wheat were mapped comparatively in the proximal regions of chromosomes 2S {256}. The Est-2, Est-5 and Est-8 were mapped to the extreme distal regions in the 3L arms {247}.

74.2.8. Glucosephosphate isomerase
Gpi-A1{507}. 1AS{195,507}. v: CS.
Gpi-B1{507}. 1BS{195,507}. v: CS.
Gpi-D1{507}. 1DS{195,507}. v: CS.
Gpi-D1a{195}. v: CS.
Gpi-D1b{195}. v: CS variant and certain CS aneuploids. Rare.
Varietal differences in GPI zymograms were noted in {1127}.
GPI zymogram phenotypes observed in Triticum and Aegilops species are reported in {456,457}.
No allelic variation at Gpi-Dt1 was found in 60 accessions of Ae. tauschii {1578}.
Gpi-Agi1{361},{374}. [Gpi-X1{361}]. 1Agi{361}. ad: Vilmorin 27/Thin. intermedium..
Gpi-E1{518}. 1ES{518}. ad: CS/E. elongata.
Gpi-H1{1153}. 1HS{1153}. ad: CS/Betzes.
Gpi-Hch1{195}. 1Hch{195}. ad: CS/H. chilense.
Gpi-R1{195}. 1R{195}.1RS{779}. ad: CS/King II{195}. al: 2a, 2b, and R14{779}.
Gpi-Rm1{195}. 1R{195}. ad: CS/S. montanum.
Gpi-Sl1{1228}. 1S1{517}.1S1S{1228}. ma: In Ae. longissima 2 x Ae. longissima 10, Gpi-Sl1, two glutenin loci, and three gliadin loci were mapped relative to one another as follows: Glu-Sl1 - 15.9 cM - Gpi-Sl1 - 38 cM - Gli-Sl4 - 7.1 cM - Glu-Sl3 - 0.9 cM - Gli-Sl1 - 5.6 cM - Gli-Sl5{1228}; Glu-Sl1 is located in 1SlL and the other loci are in 1SlS.
Gpi-Ss1{1140}. 1Ss{1140}. ad: CS/Ae. searsii.
Gpi-U1{195}. 1U{195}. ad: CS/Ae. umbellulata.
Gpi-V1{1026}. 1V{1026,241}. ad: CS/D. villosum.


74.2.9. Glutamic oxaloacetic transaminase
Got-A1{505}. 6AS{505}. v: CS.
Got-B1{505}. 6BS{505}. v: CS.
Got-D1{505}. 6DS{505}. v: CS.
Got-A2{505}. 6AL{505}. v: CS.
Got-B2{505}. 6BL{505}. v: CS.
Got-D2{505}. 6DL{505}. v: CS. ma: Cent - Got-D2 - 2cm - Xpsr154-6D{757}.
Got-Age2{1575}. 6Age{1575}. ad,su: Rescue/Thin. elongatum.
Got-E2{518}. 6EBeta{518}. ad: CS/E. elongata.
Got-H2{520}. 6H{520}. ad: CS/Betzes.
Got-R2{1457}. [Got3{1559}]. 6R{1457}.6RL{1280}. ad: CS/Imperial 6R{1457}; Holdfast/King II 6RL{1280}.
Got-V2{1026,242}. 6V{1026}. ad: Creso/D. villosum.
Got-Ht2{1037}. 6Ht{1037}. ad: CS/E. trachycaulus.
Got-A3{505}. 3AL{505}. v: CS.
Got-B3{505}. 3BL{505}. v: CS.
Got-C3{1278}. F{1278}. ad: T. aestivum cv. Alcedo /Ae. caudata line C.
Got-D3{505}. 3DL{505}. v: CS.
Got-Age3{521}. 3AgeL{521}. ad: CS/TAP 67. su: CS/TAP 67. tr: Certain CS 3D/Ag lines.
Got-E3{518}. 3EL{518}. ad: CS/E. elongata.
Got-H3. [Got-b3{090}]. 3H{090}. ad: CS/Betzes.
Got-Hch3{351}. 3Hch{351}. ad: MA/H. chilense.
Got-R3{1457}. [Got3{1559}]. 3R{1457}. ad: CS/Imperial{1457}; Holdfast/ King II{1253}; Kharkov/Dakold{1253}.
Got-Ss3{1140}. 3Ss{1140}. ad: CS/Ae. searsii.
Got-V3{1518,242}. 3VL{1518}. ad: Creso/D. villosum.
Got-R4. [Got1/7R{1203},Got2{1559}]. 7RL{1203}. al: S. cereale.

Wehling {1559} identified a GOT locus designated Got1 in 4RL of S. cereale.

74.2.10. Hexokinase
Hk-B1{006}. 1BS{006}. v: CS.
Hk-D1{006}. 1DS{006}. v: CS.
Hk-A2{810}. 3A{810}. v: CS.
Hk-A2a{810}. v: CS.
Hk-A2b{810}. s: CS*/Sears' Synthetic 3A. v: Sears' Synthetic IPSR 1190903.
Hk-B2{006}. 3BS{006,810}. v: CS.
Hk-D2{810}. 3DS{810}. v: CS.
Hk-D2a{810}. v: CS.
Hk-D2b{810}. v: Sears' Synthetic IPSR 1190903.
Hk-E2{006}. 3ES{006}. ad: CS/E. elongata.

Allelic variation was observed in three of 55 hexaploid accessions {006}.

74.2.11. Lipoxygenase
Lpx-A1{516}. [Lpx-B1{516}]. 4AL{516}. v: CS. ma: Xksu919(Lpx-1)-4A{0091}.
Lpx-B1{516}. [Lpx-A1{516}]. 4BS{516}. v: CS. ma: Xcn110(Lpx-1)-4B{0269,0367}.
Lpx-B1a{1533}. [Lpx-A1a{936}]. v: CS.
Lpx-B1b{1533}. [Lpx-A1b{936}]. v: Bosanka{1533}.
Lpx-D1{516}. 4DS{516}. v: CS.
Lpx-E1{518}. 4ES{518}. ad: CS/E. elongata.
Lpx-H1{716}. 4H{716}. ad: CS/Betzes.
Lpx-A2{516}. 5AL{516}. v: CS. ma: Xksu919(Lpx-2)-5A{0091}.
Lpx-B2{516}. 5BL{516}. v: CS. ma: Xksu919(Lpx-2)-5B{0091}; Xcn111(Lpx-2)-5B{0269}.
Lpx-D2{516}. 5DL{516}. v: CS.
Lpx-E2{518}. 5EL{518}. ad: CS/E. elongata.
Lpx-H2{716}. 5H{716}. ad: CS/Betzes.
Lpx-Ss2{1140}. 5Ss{1140}. ad: CS/Ae. searsii.
Lpx-V2{242}. 5V. ad: CS/D. villosum.


74.2.12. Malate dehydrogenase
Mdh-A1. [Mdh2A{087}]. 1AL{087}. v: CS.
Mdh-B1. [Mdh2B{087}]. 1BL{101,087}. v: CS.
Mdh-D1. [Mdh2D{087}]. 1DL{087}. v: CS.
Mdh-H1{1153}. 1HL{1153}. ad: CS/Betzes.
Mdh-Hch1{352}. 1Hch{352}. ad: MA/H. chilense.
Mdh-R1. [Mdh2-1{1252}]. 1RL{1252}. ad: CS/Imperial 1R; Kharkov/Dakold 1R; Holdfast/King II 1RL.
Mdh-Ss1{1140}. 1Ss{1140}. ad: CS/T. searsii.
Mdh-H2. [Mdh2-b2{090}]. 3H{090}.
Mdh-R2. [Mdh2-2{1252}]. 3R{1252}. ad: CS/Imperial.
A third set of dimeric MDH isozymes identified in mature grain are separable from MDH-1 and MDH-2 by their higher pI's in IEF {811}.
Mdh-A3{811}. 5AS. v: CS.
Mdh-A3a{811}. v: CS.
Mdh-A3b{811}. v: Bersee.
Mdh-B3{811}. 5BS. v: CS.
Mdh-B3a{811}. v: CS.
Mdh-B3b{811}. v: Hope.
Mdh-D3{811}. 5DS. v: CS.
Mdh-D3a{811}. v: CS.
Mdh-D3b{811}. v: Sears' Synthetic.
Mdh-E3{811}. 5ES. ad: CS/E. elongata.
Mdh-H3{811}. 5H. ad: CS/Betzes.
Mdh-U3{811}. 5U. ad: CS/Ae. umbellulata.
Mdh-R4{360}. 1RL{360}. v: Various crosses.


74.2.13. Peroxidase
Peroxidase (EC1.11.1.7) isozymes have high tissue specificity. Staining and electrophoretic systems are reviewed in {118}. PER-1, -2, -3, -4 and -5 are all reported in {816}.

74.2.13.1. PER-1
PER-1 is expressed in leaf {012} and coleoptile {816} tissues.
Per-B1{012}. 1BS{012,919}. v: CS.
Per-D1{012}. 1DS{012,919}. v: CS.
Per-D1a{012}. v: CS.
Per-D1b{012}. v: Sears' Synthetic.
Per-Hch1{012}. 1Hch{012}. ad: CS/H. chilense.
Per-R1{012}. [Prx{1561}]. 1RS{012,1561}. ad: CS/King II{012}; Holdfast/King II{1561}. tr: Veery 'S'{012}.
Per-V1{241}. 1V{241}. ad: Creso/D. villosum.


74.2.13.2. PER-2
PER-2 is expressed in young leaf {118}, coleoptile and root {816} tissues.
Per-A2. 2AS. v: CS.
Per-A2a{816}. v: CS.
Per-A2b{816}. v: Timstein.
Per-B2{118}. 2BS{118}. v: CS.
Per-B2a{816}. v: CS.
Per-B2b{816}. v: Sears' Synthetic IPSR1190903.
Per-D2{118}. 2DS{118}. v: CS.
Per-H2{118}. [Per-5{095}]. 2H{118}. ad: CS/Betzes.
Per-R2{118}. 2RS{118}. ad: CS/Imperial; Kharkov/Dakold.


74.2.13.3. PER-3
PER-3 is expressed in embryo {119,816} and scuteller {119} tissues.
Per-A3{119}. 3AL{119}. v: CS.
Per-A3a{816}. v: CS.
Per-A3b{816}. v: Timstein.
Per-A3c{816}. v: Hobbit 'S'.
Per-B3{086},{119}. [Per4{961}]. 3BL{086,119}. v: CS.
Per-B3a{816}. v: CS.
Per-B3b{816}. v: Hope.
Per-B3c{816}. v: T. macha IPSR1240005.
Per-B3d{816}. v: Timstein.
Per-B3e{816}. v: Sears' Synthetic IPSR1190903.
Per-D3{086},{119}. [Per5{961}]. 3DL{086,119}. v: CS.
Per-D3a{816}. v: CS.
Per-D3b{816}. v: Hope.
Per-D3c{816}. v: Timstein.
Per-D3d{816}. v: T. macha IPSR 142005.
Per-D3e{816}. v: Sava.
Per-D3f{816}. v: Cheyenne.
Per-D3g{816}. v: Sears' Synthetic IPSR 1190903.
Varietal variation for Per-3 was reported in {094}.


74.2.13.4. PER-4
PER-4 is expressed in endosperm tissue {086,119}.
Per-A4{695},{119}. [Per3{961}]. 7A{695}.7AS{694,086,119}. v: CS.
Per-A4a{816}. v: CS.
Per-A4b{816}. v: Hope.
Per-A4c{816}. v: Sicco.
Per-B4{695},{119}. [Per2{961}]. 4A{695}.4AL{086,119,694}. v: CS.
Per-B4a{816}. v: CS.
Per-B4b{816}. v: Hope.
Per-B4c{816}. v: Thatcher.
Per-D4{695},{119}. [Per1{961}]. 7D{695}.7DS{694,086,119}. v: CS.
Per-D4a{816}. v: CS.
Per-D4b{816}. v: Thatcher.
Per-Age4. 7AgeS{694}. tr: Certain CS 7D/Age lines.
Per-Agi4. [Per-Agi3{374}]. 7Agi{168}. ad: Vilmorin 27/Thin. intermedium.
Cultivar variation for Per-4 was reported in {094}.


74.2.13.5. PER-5
PER-5 is expressed in roots {816}.
Per-D5{816}. 2DS{816}. v: CS.
Per-Sl5{816}. 2Sl{816}. ad: CS/Ae. longissima.



74.2.14. Phosphodiesterase
Pde-A1. [Pde-A3{1590}]. 3A{1590}.3AS{1589}. v: CS.
Pde-B1. [Pde-B3{1590}]. 3B{1590}.3BS{1589}. v: CS.
Pde-D1. [Pde-D3{1590}]. 3DS{1590}. v: CS.
Pde-Sl1. 3SlS{172}. ad: CS/Ae. longissima.
Pde-V1{1518}. 3VS{1518}. ad: CS/D. villosum.



74.2.15. Phosphogluconate dehydrogenase
Pgd1{282}. [Pgd3{282},Pgd-A3{963}]. 7AmS{282}. v: T. monococcum{664}.
PgdR1. 4RL{1191}. ad: CS/Imperial; Holdfast/King II.
PgdR2. 6RL{1191}. ad: CS/Imperial; Holdfast/King II.

Loci were also identified in 6B {1435}, 1EL {1435}, 1HL {147,1072}, 1Hch {352} and 1RL {779}.

74.2.16. Phosphoglucomutase
Pgm-A1{088}. [Pgm-B1{088}]. 4AL{088}. v: CS.
Pgm-D1{088}. 4DS{088}. v: CS.
Pgm-Agi1{361},{374}. [Pgm-X1{361}]. 4Agi{361}. ad: Vilmorin 27/Thin. intermedium.
Pgm-H1. [Pgm-b1{090}]. 4H{090}. ad: CS/ Betzes.
Pgm-Hch1{351}. 4Hch{351}. ad: MA/H. chilense.
Pgm-R1. 4RS{1561,1253}. ad: CS/Imperial 4RS{1253,1561}; Kharkov/Dakold 4R{1253}; Holdfast/King II 4RS{1253,1561}.


74.2.17. Shikimate dehydrogenase
Skdh-A1{706,1065}. 5AS{706,1065}. v: CS.
Skdh-B1{706,1065}. 5BS{706,1065}. v: CS.
Skdh-D1{706,1065}. 5DS{706,1065}. v: CS.
Skdh-H1{085}. 5H{085}. ad: CS/Betzes.
Skdh-Ht1{1037}. 5Ht{1037}. ad: CS/E. trachycaulus.
Skdh-Mv1{985}. [Skdh-Mv1{985}]. 5Mv. su: 5Mv(5A), 5Mv(5D).
Skdh-R1{706}. 5RS{706}.5R{085}. ad: CS/King II{085}; CS/Imperial{706}; Kharkov/Dakold{085}. tr: CS 4AS-5RL; CS 5BL-5RL.
Skdh-Sl1{085}. 5SlS{085}. ad: CS/ Ae. longissima.
Skdh-Ss1{1140}. 5Ss{1140}. ad: CS/Ae. searsii.
Skdh-V1{085}. 5V{085}. ad: CS/D. villosum.
Skdh-U1. 5U{706}. ad,su: CS/Ae. umbellulata.


74.2.18. Superoxide dismutase
Sod-A1{1066}. 2AL{1066}. v: CS.
Sod-B1{1066}. 2BL{1066}. v: CS.
Sod-D1{1066}. 2DL{1066}. v: CS.
Sod-E1{808}. VI E{808}. ad: CS/E. elongata.
Sod-H1{716}. 2H{716}. ad: CS/Betzes.
Sod-R1{1066}. [Sod-3{586}]. 2R{1066}. ad: CS/Imperial.
Sod-Ss1{1140}. 2Ss{1140}. ad: CS/Ae. searsii.
Sod-VI{1026}. 7V{1026}. ad: CS/D. villosum.


74.2.19. Triosephosphate isomerase
Tpi-A1{1139}. 3AS{1139}. v: CS.
Tpi-B1{1139}. 3BS{1139}. v: CS.
Tpi-D1{1139}. 3DS{1139}. v: CS.
Tpi-E1{1139}. 3E{1139}. ad: CS/E. elongata.
Tpi-H1{1139}. 3H{1139}. ad: CS/Betzes.
Tpi-R1{1139}. 3R{1139}. ad: CS/Imperial; Kharkov/Dakold.
Tpi-Sl1{1139}. 3Sl{1139}. ad: CS/ Ae. longissima.
Tpi-A2{1139}. 5AL{1139}. v: CS.
Tpi-B2{1139}. 5BL{1139}. v: CS.
Tpi-D2{1139}. 5DL{1139}. v: CS.
Tpi-H2{1139}. 5H{1139}. ad: CS/Betzes.
Tpi-R2{1139}. 5R{1139}. ad: CS/Imperial; Kharkov/Dakold.
Tpi-Sl2{1139}. 5Sl{1139}. ad: CS/Ae. longissima.
Tpi-U2{1139}. 5U{1139}. ad: CS/Ae. umbellulata.
Tpi-Agi2{374}. 5Agi{374}. ad: Vilmorin 27/Ag. intermedium.


74.2.20. Aromatic alcohol dehydrogenase
Aadh-A1. [Adh-A2{584}]. 5AL{584}. v: CS. ma: XksuG44-5A(proximal) - 6.9cM - Aadh-A1 - 24.7cM - Xcdo412-5A(distal){9959}.
Aadh-A1a. v: CS; 133 other accessions{584}.
Aadh-A1b. v: T. spelta; K-24696; other accessions{584}.
Aadh-B1. [Adh-B2{584}]. 5BL{584}. v: CS.
Aadh-B1a. v: CS{1533}.
Aadh-B1b. v: Drina {1533}.
Aadh-C1{1278}. C{1278}. ad: Alcedo/Ae. caudata line C.
Aadh-D1. [Adh-D2{584}]. 5DL{584}. v: CS.
Aadh-E1. [Adh-E2{518}]. 5EL{518}. ad: CS/E. elongata.
Aadh-R1. 5RL{1280}. ad: Holdfast/King II.
Aadh-A2. [Adh-A3{508}]. 6A{1279}.6AL{513,587}. v: CS{513}; Carola{1279}.
Aadh-B2. [Adh-B3{508}]. 6B{1279}.6BL{513}. v: CS{513}; Carola{1279}.
Aadh-D2. [Adh-D3{508}]. 6D{1279}.6DL{513}. v: CS{513}; Carola{1279}.
Aadh-Age2{1575}. 6Age{1575}. ad,su: Rescue/Thin. elongatum.
Aadh-E2. [Adh-E3{518}]. 6EBeta{518}. ad: CS/E. elongata.
Aadh-R2. 6RL{1280}. ad: Holdfast/King II.
Aadh-V2{241}. 6V{241}. ad: CS/D. villosum.

The Aadh-1 and Aadh-2 loci were designated with the synonyms Adh-2 and Adh-3, respectively, in a number of publications in addition to {508,518,584}. These include: {510,509,511,519,517,587,1066,1139}.

74.2.21. Aconitase
Aco-A1{189}. 6AL{189}. v: CS.
Aco-A1a. v: CS{1533}.
Aco-A1b. v: Dubravka{1533}.
Aco-B1{189}. 6BL{189}. v: CS.
Aco-B1a. v: CS{1533}.
Aco-B1b. v: Dubravka{1533}.
Aco-B1c. v: Slavonka{1533}.
Aco-D1{189}. 6DL{189}. v: CS.
Further alleles at Aco-A1 and Aco-B1 are listed in {1127}; these have not been tested against those found in {1533}.
Aco-Age1{1575}. 6Age{1575}. ad,su: Rescue/Thin. elongatum.
Aco-E1{189}. 6Ebeta{189}. ad: CS/E. elongata.
Aco-H1{147},{189}. [Aco-1{147}]. 6H{147}.6HL{189}. ad: CS/Betzes.
Aco-R1{189}. 6RL{189}. ad: Sturdy/PI 252003.
Aco-Sl1{189}. 6Sl{189}. ad: CS/Ae. longissima.
Aco-Ss1{1140}. 6Ss{1140}. ad: CS/Ae. searsii.
Aco-U1{189}. CSU-31{189}. ad: CS/Ae. umbellulata.
Aco-A2{189}. 5AL{189}. v: CS.
Aco-B2{189}. 4BL{1513}. v: CS.
Aco-B2a{1513}. v: CS.
Aco-B2b{1513}. v: PI 278437.
Aco-B2c{1513}. v: PI 182575.
Aco-B2d{1513}. v: PI 157589.
Aco-D2{189}. 4DL{1513}. v: CS.
Aco-E2{189}. 4EL{189}. ad: CS/E. elongata.
Aco-Mv2{1341}. [Aco-Mv2{985}]. 5Mv. ad: 5Mv(5A),5Mv(5D).
Aco-R2{189}. 5RL{189}. ad: CS/King II 5R; Holdfast/ King II 5RL.
Aco-Ss2{1140}. 4Ss{1140}. ad: CS/Ae. searsii.


74.2.22. NADH dehydrogenase

74.2.22.1. Ndh-1
Ndh-A1{513},{1037}. [Ndh-B1{513}]. 4AL{513}. v: CS.
Ndh-A1a{1533}. [Ndh-B1a{936}]. v: CS.
Ndh-A1b{1533}. [Ndh-B1b{936}]. v: Sutjeska.
Ndh-A1c{1533}. [Ndh-B1c{936}]. v: Fruskogorka.
Ndh-A1d{1037}. [Ndh-A1b{1037}]. v: Hope, Timgalen.
Ndh-B1{513}. [Ndh-A1{513}]. 4BS{513}. v: CS.
Ndh-D1{513}. 4DS{513}. v: CS.
Ndh-E1{362}. 4E{362}. ad: CS/Ag. elongatum.
Ndh-H1{147},{513}. [Nadhd-1{147}]. 4H{147}.4HS{813}. ad: CS/Betzes.
Ndh-Hch1{813}. 4Hch{813}. ad: CS/H. chilense.
Ndh-V1{241}. 4V{241}. ad: CS/D. villosum.
Ndh-R1{813}. 4R{362}.4RS{813}. ad: CS/Imperial, CS/King II{813,362}; CS/Dakold{362}.
Ndh-S11{813}. 4Sl{813}. ad: CS/Ae. longissima.
Ndh-U1{362}. A{362}. ad: CS/Ae. umbellulata.

Based on the correspondence of the electrophoretic patterns, isoelectric points (pIs) and chromosomal location, it was proposed that the Ndh1 (NADH dehydrogenase) and Dia3 (diaphorase) represent the same locus {0356}.

74.2.22.2. Ndh-2
Ndh-A2{813}. 7A{813}. v: Hope.
Ndh-D2{813}. 7DS{813}. v: CS.
Ndh-R2{813}. 7RS{813}. ad: CS/Imperial, CS/King II, Holdfast/King II (7R).

Based on the correspondence of the electrophoretic patterns, isoelectric points (pIs) and chromosomal location, it was proposed that the Ndh-2 (NADH dehydrogenase) and Dia2 (diaphorase) represent the same locus {0356}.

74.2.22.3. Ndh-3
Ndh-A3{813}. 3AL{813}. v: CS.
Ndh-B3{813}. 3BL{813}. v: CS.
Ndh-B3a{813}. v: CS.
Ndh-B3b{813}. v: Carmen.
Ndh-D3{813}. 3DL{813}. v: CS.
Ndh-H3{813}. 3HL{813}. ad: CS/Betzes.
Ndh-R3{813}. 6RL{813}. ad: Holdfast/King II, CS/Imperial (6R), CS/King II (6R).
Ndh-Sl3{813}. 3SlL{813}. ad: CS/Ae. longissima; CS/Ae. sharonesis (3Sl).

Based on the correspondenc of the electrophoretic patterns, isoelectric points (pIs) and chromosomal location, it was proposed that the Ndh-3 (NADH dehydrogenase), Dia1 (diaphorase) and Mnr1 (menadione reductase) represent the same locus {0356}.

74.2.22.4. Ndh-4
Ndh-A4{813}. 3AS{813}. v: CS.
Ndh-B4{813}. 3BS{813}. v: CS.
Ndh-E4{813}. 3ES{813}. ad: CS/E. elongata.
Ndh-H4{813}. 3HS{813}. ad: CS/Betzes, CS/King II.
Ndh-R4{813}. 3RS{813}. ad: CS/King II, CS/Imperial (3R).



74.2.23. Dipeptidase
Dip-A1{454},{700}. [Pept-A1{454}]. 6AL{454,700}. v: CS.
Dip-A1a{700}. v: CS.
Dip-A1b{700}. v: Cheyenne.
Dip-B1{454},{700}. [Pept-B1{1533}]. 6BL{454,700}. v: CS.
Dip-B1a{700}. v: CS.
Dip-B1b{700}. v: Cappelle-Desprez.
Dip-D1{700}. 6DL{700}. v: CS.
Dip-H1{147},{700}. [Pept-1{147},Dip 1{145}]. 6H{147,145,700}. ad: CS/Betzes.
Dip-J1{700}. 6J{700}. ad: CS/Thin. junceum.
Dip-V1{700}. 6V{700}. ad: CS/D. villosum.


74.2.24. Malic enzyme
A dimeric enzyme extractable from mature grains.
Mal-A1{809}. 3AL. v: CS.
Mal-B1{809}. 3BL. v: CS.
Mal-B1a{809}. v: CS.
Mal-B1b{809}. v: T. spelta IPSR line 1.
Mal-B1c{809}. v: Sears' Synthetic.
Mal-D1{809}. 3DL. v: CS.
Mal-E1{809}. 3E. ad: CS/E. elongata.
Mal-H1{809}. 3H. ad: CS/Betzes.
Mal-R1{809}. 3R. ad: CS/Imperial.


74.2.25. Adenylate kinase
Adk-A1{091}. [Adk-a{091}]. 7AL{091}. v: CS.
Adk-B1{091}. [Adk-b{091}]. 7BL{091}. v: CS.
Adk-D1{091}. [Adk-d{091}]. 7DL{091}. v: CS.
Adk-E1{091}. 7E{091}.7E{1435}. ad: CS/E. elongata.
Adk-H1{091}. 7H{091}.7HS{1435}. ad: CS/Betzes.
Adk-Mv1{985}. [Adk-Mv1{985}]. 7MvL. su: 7Mv(7D).
Adk-R1{091}. 7RL{091}. ad: CS/Imperial; Holdfast/King II.
Adk-U1{091}. E{091}. ad: CS/Ae. umbellulata.
Adk-AgiI{091}. 7Agi{091}. ad: CS/Thin. intermedium.
Adk-H2. 6HL{1435}. ad: CS/Betzes.


74.2.26. Glutamate-pyruvate transaminase
Gpt-A1{1435}. 1AS{1435}. v: CS.
Gpt-B1{1435}. 1BS{1435}. v: CS.
Gpt-D1{1435}. 1DS{1435}. v: CS.
Gpt-E1{1435}. 1ES{1435}. ad: CS/E. elongata 1E.
Gpt-H1{1435}. 1H{1435}. dv: H. vulgare cv. Betzes.


74.2.27. Catalase
Cat-B1{1466}. [Cat-A1{1466}]. 4BL{1466}. v: CS.


74.2.28. Beta-glucosidase
b-Gls{282}. 2AmL{282}. dv: DV92.
b-Glsa{282}. dv: DV92.
b-Glsb{282}.Null allele dv: G3116.


74.2.29. Starch branching enzyme
SbeI1{9937}. 1DL{9937}. v: CS{9937}.
SbeI2{9937}. 7BL{9937}. v: CS{9937}.



74.3. Endosperm storage proteins

74.3.1. Glutenins
These are heterogeneous mixtures of proteins comprising subunits linked by disulfide bonds. 'A' are high-molecular-weight (HMW) and 'B', 'C' and 'D' are low-molecular-weight (LMW) subunits.

Using proteomic analysis of 2D gels of seed storage proteins in 39 ditelocentric lines of cv. CS, 105 protein spots were resolved {03129}. Locations of structural genes controlling 26 spots were identified in 10 chromosomal arms (4 on 1BL, 5 on 1BS, 4 on 1DL, 4 on 1DS, 2 on 6AS, 3 on 6BS, 1 on 6DL, 1 on 6DS, 1 on 3BS and 1 on 3BL). Multiple regulators of the same protein located on various chromosome arms were observed. Two novel subunits, named 1Bz and 1BDz, were found to have very similar structures to HMW glutenin subunit 12 (encoded by Glu-D1-2a - see the relevant list below) and were located to chromosome arms 1BL and 1DL, respectively.

74.3.1.1. Glu-1
The Glu-1 loci, all of which are compound, encode HMW glutenin subunits.
Each Glu-1 locus in hexaploid wheat contains two genes, the products of which were described as 'x-type' and 'y-type' based on differences in molecular weight and isoelectric point {1118}.
Other evidence has shown that these gene products differ in electrophoretic fingerprint pattern {1124} and cysteine content {1028}, and the genes themselves differ in nucleotide sequence {1470,1433,373}.
Although early evidence suggested up to 6 genes in total at each locus {1471,373], it appears likely that only a single copy of each gene is present at the 1AL, 1BL, and 1DL loci {495}.
No 'y-type' protein from the Glu-A1 locus has been demonstrated in hexaploid wheat {1118}, although they are found in diploid wheats {1535,798}, and sequencing experiments have shown the presence of a terminating sequence inside the transcribed portion of the gene {373}. Definitive evidence that subunit 21* {602}, which has a mobility close to that of subunit 21, is a 'x-type' protein rather than a 'y-type' protein has not been obtained. The gene coding for 'x-type' proteins within Glu-A1 is also often silent {1118,420}.
The symbols for the genes within the Glu-1 loci coding for 'x-type' and 'y-type' proteins will be Glu-1-1 and Glu-1-2, respectively, rather than Glu-1x and Glu-1y {1470}. The genes are closely linked but recombination has been observed between Glu-B1-1 and Glu-B1-2 with a frequency of 3 in 3,450 {1117}. The gene order, relative to the centromere, has not been ascertained.
The subunit nomenclature used is that devised in {1116}; however, an alternative system based upon molecular weight was proposed in {1068}. A system of naming the Glu-A1-1, Glu-A1-2, Glu-B1-1 and Glu-B1-2 alleles in T. turgidum var. dicoccoides is given in {796}.
In {00116}, a comparison between spelt wheats (T. spelta) and bread wheat was carried out for the glutenins using a nomenclature system described in {00117}.
The Glu-1 loci may be recognised by the DNA probe pTag1290 {1471} and probe pwhe1(Dy10) {{030}}. Individual Glu-1-1 loci on 1A, 1B and 1D and the Glu-1-2 loci may be recognised by specific primers {263}.
In {00105}, the evolution of the high molecular weight glutenin loci of the A, B, D and G genomes of wheat was explored; 30 partial allele sequences were compared, designated by Greek letters (alpha, beta, gammma, etc.) (5 of which were cited as Schlumbaum, pers. comm.; the remaining 25 were deposited as GenBank, accession nos. X98583-X98592, X98711-X98715 and Y12401-Y12410). These partial alleles derive from all six Glu-1-1 and Glu-1-2 loci in current-day samples taken from seven species of wheat, as well as from DNA extracted from charred grain of two samples from archaeological excavations, dated 3000 and 5000 years old, respectively.
Following the first listing which considers the Glu-1 set for hexaploid wheat as a single locus, there is a provisional listing based on x- and y- type glutenins. These are not referenced.
The importance of the HMW glutenin subunits for bread-making quality was first noted from observations in wheat cultivars of related pedigree on the effects of the presence of subunit 1 encoded by Glu-A1a {0197}, effects that have repeatedly been confirmed since (for example {0198,0199,01100}).

A nomenclature system for prolamin banding patterns of triticale was proposed in {03139}. Extensive allelic variation in triticale at Glu-A1, Glu-B1, Glu-R1 and Gli-R2 loci was reported in {03121}.
Glu-A1{780,1125}. [Glt-A1{420},Glt-A2{420},Glu 1A{1415}]. 1A{780}.1AL{781,1125}. s: CS*/Hope 1A{1125}. v: CS{780,781}; various{420}.
Glu-A1a{1116}. 1{1116}. v: Hope.
Glu-A1b{1116}. 2*{1116}. v: Bezostaya 1.
Glu-A1c{1116}. Null allele{1116}. v: CS.
Glu-A1d{1535}. v: V74, Spain{1115}.
Glu-A1e{1535}. v: 132c, Poland{1115}.
Glu-A1f{1535}. v: 112-29, Sudan{1115}.
Glu-A1g{1535}. v: Landrace 1600.
Glu-A1h{1527}. [GLU-A1-I{1527}]. tv: PI 94683, USSR, T. dicoccum.
Glu-A1i{1527}. [GLU-A1-II{1527}]. tv: CI 12213, India, T. dicoccum; Lambro{1523}.
Glu-A1j{1527}. [GLU-A1-III{1527}]. 1'{125}. tv: PI 352359, Germany, T. dicoccum.
Glu-A1k{478}. 26{478}. v: BT-2288{478}.
Glu-A1l{847}. tv: Chinook, Canada.
Glu-A1m{1069}. tv: Nugget Biotype 1, Canada, T. durum.
Glu-A1n{1526}. [Glu A1-IV{1526},Glu-A1m{959}]. 1'{125}. tv: Corado, Portugal{1526}.
Glu-A1o{1526,125}. [Glu A1-V{1526,125},Glu-A1n{959}]. 2**{125}. tv: Aric 581/1{125}; PI 61189{1525}; USSR.
Glu-A1p{1146}. 3*{1146}. v: David 1.
Glu-A1q{125}. [Glu A1VI{125}]. 2***{125}. tv: Melianopus 1528.
Glu-A1r{1232}. 39+40{1232}. i: T. thaoudar IPSR 1020006/6*Sicco.
Glu-A1s{1231}. 41+42{1231}. i: T. thaoudar G3152/6*Sicco.
Glu-A1t{602}. 21*{602}. v: W29323, W3879, W31169.
Glu-A1u{02106}. 2*B{02106}. v: Bankuti 1201.
The allele designated Glu-A1u and Glu-A1-1u in the appropriate list below encodes a high molecular weight glutenin subunit (denominated 2*B) that is identical to subunit 2* apart from one amino acid difference involving the exchange of serine for cysteine (which itself is due to a C to G point mutation at the 1181 bp point of the coding region of 2*). The authors of {02106} suggest that the additional cysteine residue facilitates the formation of further disulphide bonds (cf. the 1Dx5 subunit) which might lead to an improvement in gluten quality characters.
Glu-A1v{03137}. [Glu-A1-VII{03137}]. VII{03137}. v: PI-308879 emmer {03137}.
There is a possibility that Glu-A1 alleles i, j {1527} and k {478} correspond to alleles d, e, f or g {1535} that were published shortly earlier. Glu-A1m [{1526}] was changed to n, because the m allele in {1069} has precedence. Allele n [{1526}] was changed to o. An earlier reference to an allele designated Glu-A1d {1411} was withdrawn {1114}. There appears to be a minor band associated with subunit 2* encoded by Glu-A1b {1516}; this may be the same as a band named A5 in {420}.
Six combinations involving 5 HMW subunits [1A (u-z)] are listed in {420}, from a study of 109 genotypes including representatives of botanical varieties. A number of alleles in T. turgidum var. dicoccoides populations, 12 at Glu-A1-1 and 3 at Glu-A1-2, were described in {798}. In a further study using different germplasm of this species {205}, 14 alleles at Glu-A1 were observed, including 12 not previously found; the 15 alleles included up to 15 alleles at Glu-A1-1 (with up to 10 not previously observed), and 5 alleles at Glu-A1-2 (with 4 not previously observed) (numbers take the null allele into account). The uncertainty in numbers is due to the very similar electrophoretic mobilities of some of the subunits compared with others observed either in this study or previously.
In a study including emmers (T. dicoccon) {00115}, new subunits named 1+ and 2- were found in accessions MG4378/1 and MG5380/1, respectively, and provisionally assigned to Glu-A1. Until confirmed, they are not included in the Glu-A1 list.
Glu-B1{107},{1125}. [Glt-B1{420},Glt-B2{420},Glt-B3{420},Glu 1B{1415}]. 1BL{107,780,1125}. v: CS.
Glu-B1a{1116}. 7{1116}. v: Flinor.
Glu-B1b{1116}. 7+8{1116}. v: CS.
Glu-B1c{1116}. 7+9{1116}. v: Bezostaya 1.
Glu-B1d{1116}. 6+8{1116}. v: Hope.
Glu-B1e{1116}. 20{1116}.20x+20y{03133}. v: Federation.
Glu-B1f{1116}. 13+16{1116}. v: Lancota (rare).
Glu-B1g{1116}. 13+19{1116}. v: NS 335 (rare).
Glu-B1h{1116}. 14+15{1116}. v: Sappo (rare).
Glu-B1i{1116}. 17+18{1116}. v: Gabo.
Glu-B1j{1116}. 21{1116}.21x+21y{03116}. v: Dunav (rare); Foison{03116}.
Glu-B1k{1116}. 22{1116}. v: Serbian (rare).
Glu-B1l{778}. 23+24{778}. v: Spica D.
Glu-B1m{1527}. [GLU-B1-I{1527}]. tv: PI 94640, Iran, T. dicoccum.
Glu-B1n{1527}. [GLU-B1-II{1527}]. tv: PI 355505, Germany, T. dicoccum.
Glu-B1o{1527}. [GLU-B1-III{1527}]. tv: PI 352354, Ethiopia, T. dicoccum.
Glu-B1p{1527}. [GLU-B1-IV{1527}]. 23+18{125}. tv: Dritto{1523}; Ethiopia, PI 94655, T. dicoccum{1527}.
Glu-B1q{1527}. [GLU-B1-V{1527}]. tv: PI 94633, Morocco, T. dicoccum.
Glu-B1r{1527}. [GLU-B1-VI{1527}]. 19{125}. tv: PI 946669, Bulgaria, T. dicoccum{1527}; Lambro{1523}.
Glu-B1s{478}. 7+11{478}. v: BT-2288.
Glu-B1t{847}. v: Supreza, Canada.
Glu-B1u{1069}. 7*+8{1146}. v: Owens{1069}; Fiorello{1146}.
Glu-B1v{1069}. v: Mondor.
Glu-B1w{1069}. 6*+8*{1146}. v: Dawbull{1069}; Sieve{1146}.
Glu-B1x{1526}. [Glu-B1-VII{1526},Glu-B1t{959}]. tv: Canoco de Grao Escuro, Portugal, T. turgidum.
Glu-B1y{1526}. [Glu-B1-VIII{1526},Glu-B1u{959}]. tv: Tremez Mollez, Portugal, T. durum.
Glu-B1z{1524}. [Glu-B1-IX{1524},Glu-B1v{959}]. 7+15{125}. tv: Roccia, Italy, T. durum {1523,125}.
Glu-B1aa{1524}. [Glu-B1-X{1524},Glu-B1w{959}]. tv: Quaduro, Italy, T. durum.
Glu-B1ab{1523}. [Glu-B1-XI{1523},Glu-B1x{959}]. tv: Athena, Italy, T. durum.
Glu-B1ac{125}. [Glu B1XIII{125}]. 6+16{125}. tv: Espa l8914, T. durum.
Glu-B1ad{125}. [Glu B1XIV{125}]. 23+22{125}. tv: Greece 20, T. durum.
Glu-B1ae{1146}. 18*{1146}. v: David.
Glu-B1af{1146}. 26+27{1146}. v: Cologna 1.
Glu-B1ag{1146}. 28+29{1146}. v: Forlani.
Glu-B1ah{782}. null allele{782}. v: Olympic mutant.
Glu-B1ai{714}. 7'{714}. v: Adonis.
Glu-B1aj{759}. 8{759}. v: AUS 14444, Afghanistan.
Glu-B1ak{899}. 7*+8*{899}. v: Norstar.
Glu-B1al{899}. 7+8*{899}. v: Glenlea.
Glu-B1am{1229}. 18{1229}. v: Royo.
Glu-B1an{1229}. 6{1229}. v: BG-2013.
Glu-B1ao{1229}. 7+16{1229}. v: BG-3545.
Glu-B1ap{1229}. 30+31{1229}. v: Marinar.
Glu-B1aq{1229}. 32+33{1229}. v: BG-1943.
Glu-B1ar{1229}. 34+35{1229}. v: Jeja Almendros.
Glu-B1as{1229}. 13{1229}. v: PI 348435.
Glu-B1at{1229}. 13+18{1229}. v: PI 348449.
Glu-B1au{1032}. 37{1032}. v: Shedraya Polesja.
Glu-B1av{03116}. [Glu-B1r{03116}]. 7-18{03116}. v: Triticor hexaploid triticale{03116}.
Glu-B1aw{03116}. [Glu-B1s{03116}]. 6.8-20y{03116}. v: Carnac hexaploid triticale{03116}.
Glu-B1ax{03137}. [Glu-B1-XV{03137}]. XV{03137}. tv: PI-190922, BG-012302 emmers{03137}.
Glu-B1ay{03137}. [Glu-B1-XVI{03137}]. XVI{03137}. tv: PI-277681 emmers{03137}.
Glu-B1az{03137}. [Glu-B1-XVII{03137}]. XVII{03137}. tv: PI-348620 emmers{03137}.
Glu-B1ba{03122}. [Glu-B1-XVIII{03122}]. 13*+16{03122}. v: PI-348767 spelt {03122}.
Glu-B1bb{03122}. [Glu-B1-XLX{03122}]. 6+18'{03122}. v: PI-348631 spelt wheat accession{03122}.
Glu-B1bc{03138}. 6+17{03138}. v: ICDW 20975{03138}.
Glu-B1bd{03140}. 20+8{03140}. v: Abadja{03140}.
The alleles formerly designated t to x in {959} were renamed x to ab because allele t in {847} and alleles u, v and w in {1069} had precedence. Subunit 8 of Glu-B1b (7+8) is more acidic in isoelectric focusing than subunit 8 of Glu-B1d (6+8) {555}. Variation in the mobility of subunits designated 7 has been observed {1118}, which accords with more recent observations {714,1069}. The subunit encoded by Glu-B1v {1069} has the same mobility as subunit 7 of Glu-B1c (7+9); it could be the same subunit as 7' encoded by Glu-B1ai [{714}].
Variation in the staining intensity of subunit 7 in different lines was observed {1069}; a duplication of the gene encoding subunit 7 probably occurred in cultivar 'Red River 68', as evidenced by increased intensity of the subunit in SDS-PAGE and by approximately doubled intensity of restriction fragments carrying the gene in Southern blotting {9989}. Possible low gene expression at Glu-B1 was noted for Glu-B1w, where subunits 6*+8* stain very faintly {1146}.
One of the Glu-B1af subunits was numbered 26 in {1146}; 26 was previously used to number the subunit encoded by Glu-A1k {478}. Subunit 28 of Glu-B1ag (28+29) {1146} is referred to as subunit 19* in {1068}. Subunit 23 of Glu-B1p {23+18} and Glu-B1ad (23+22) {125} may not be the same subunit as that numbered 23 of Glu-B1l {23+24} {778}. Glu-Blz carried by Roccia was numbered (7+15) and named Glu-B1-XII in {125}; however, the earlier name, Glu-B1-IX {1523}, has precedence; also, {1523} states that the Glu-B1-IX subunit of faster mobility is slightly slower than subunit 15. Subunit 11 of Glu-B1s {7+11} was so numbered in {478} because its mobility is the same as one of the subunits encoded by a Glu-D1 allele (2+11) described in {755}.
Eight alleles at Glu-B1-1 and 10 alleles at Glu-B1-2 in T. turgidum var. dicoccoides populations were described in {798}. In a further study using different germplasm of this species {205}, 19 alleles at Glu-B1 were observed, including 15 not previously observed; the 19 alleles included 11 alleles at Glu-B1-1 and 14 alleles (including the null allele) at Glu-B1-2, although, as the authors pointed out, it was not conclusively clear how many of these alleles were distinct from each other, or from others previously observed.
In a study including emmer wheats (T. dicoccon) {00115}, new subunits named 7+ (in accessions MG5400/5 and MG30835/1), 8- (in accessions MG5400/5, MG30835/1, MG5333/1 and MG5507) and 13- (in accession MG5282/2) were found and provisionally assigned to Glu-B1. Until confirmed, they are not included in the Glu-B1 list.
Glu-D1{1100,1125}. [Glt-D1{420},Glt-D2{420},Glu 1D{1415}]. 1DL{107,150,780,1100,1125}. v: CS.
Glu-D1a{1116}. 2+12{1116}. v: CS.
Glu-D1b{1116}. 3+12{1116}. v: Hobbit.
Glu-D1c{1116}. 4+12{1116}. v: Champlein.
Glu-D1d{1116}. 5+10{1116}. v: Hope.
Glu-D1e{1116}. 2+10{1116}. v: Flinor (rare).
Glu-D1f{1116}. 2.2+12{1116}. v: Danchi (rare).
Glu-D1g{478}. 5+9{478}. v: BT-2288.
Glu-D1h{1145}. 5+12{1145}. v: Fiorello, Italy.
Glu-D1i{107}. null{107}. v: Nap Hal, Nepal.
Glu-D1j{1146}. 2+12*{1146}. v: Tudest.
Glu-D1k{421}. 2{421}. s: CS/Timstein 1D.
Glu-D1l{759}. 12{759}. v: AUS 10037, Afghanistan.
Glu-D1m{759}. 10{759}. v: AUS 13673, Afghanistan.
Glu-D1n{759}. 2.1+10{759}. v: AUS 14653, Afghanistan.
Glu-D1o{755}. 2.1+13{755}. v: AUS 14519, T. macha.
Glu-D1p{1233}. 36{1233}. i: Iranian landrace accession 3048/5* Sicco.
Glu-D1q{124}. 2+11{124}. v: Flinor.
Glu-D1r{1229}. 2.3+12{1229}. v: PI 348465.
Glu-D1s{1032}. 38{1032}. v: Leningradka.
Glu-D1t{668}. 43+44{668}. i: Ae. tauschii accession TA2450/2*.
Glu-D1u{836}. 2+10'{836}. v: Coker 68-15.
Glu-D1v{755}. 2.1+10.1{755}. dv: Ae. tauschii.
Glu-D1w{03124}. 5*+10{03124}. v: Fiorello{03124}.
Glu-D1x{755}. 2+T2{755}.2t+12.2t{03124}. dv: Ae. tauschii.
Glu-D1y{755}. 3+T2{755}.3t+12.2t{03124}. dv: Ae. tauschii.
Glu-D1z{755}. 3+10{755}. dv: Ae. tauschii.
Glu-D1aa{755}. 3+10.3{755}. dv: Ae. tauschii.
Glu-D1ab{755}. 4.1+10{755}. dv: Ae. tauschii.
Glu-D1ac{755}. 4+10{755}. dv: Ae. tauschii.
Glu-D1ad{755}. 5.1+10.2{755}. dv: Ae. tauschii.
Glu-D1ae{1578}. 2.1+T2{1578}.2.1t+12.2t{03124}. dv: Ae. tauschii.
Glu-D1ag{1578}. 1.5+T2{1578}.1.5t+12.2t{03124}. dv: Ae. tauschii.
Glu-D1ah{1578}. 1.5+10{1578}. dv: Ae. tauschii.
Glu-D1ai{1578}. 2.1+10.5{1578}. dv: Ae. tauschii.
Glu-D1aj{1578}. 1.5+12{1578}. dv: Ae. tauschii.
Glu-D1ak{1578}. 3+10.5{1578}. dv: Ae. tauschii.
Glu-D1al{02107}. 2.2*{02107}. v: MG315.
Glu-D1am{03122}. [Glu-D1-I{03122}]. 2+12'{03122}. v: PI-348495 spelt {03122}.
Glu-D1an{03122}. [Glu-D1-II{03122}]. 2+12*{03122}. v: PI-348672 spelt {03122}.
Glu-D1ao{03122}. [Glu-D1-III{03122}]. 2.4+12{03122}. v: PI-348473 spelt {03122}.
Glu-D1ap{03122}. [Glu-D1-IV{03122}]. 2.5+12{03122}. v: PI-348572 spelt {03122}.
Glu-D1aq{03124}. 1.5t+10.1t{03124}. dv: Ae. tauschii.
Glu-D1ar{03124}. 2t+10.1t{03124}. dv: Ae. tauschii.
Glu-D1as{03124}. 1.5t+10.2t{03124}. dv: Ae. tauschii.
Glu-D1at{03124}. 3t+10.1t{03124}. dv: Ae. tauschii.
Glu-D1au{03124}. 2.1t+10.2t{03124}. dv: Ae. tauschii.
Glu-D1av{03124}. 2t+12.3t{03124}. dv: Ae. tauschii.
Glu-D1aw{03124}. 1t+10t{03124}. dv: Ae. tauschii.
Glu-D1ax{03124}. 1t+12t{03124}. dv: Ae. tauschii.
Glu-D1ay{03124}. 1t+10.1t{03124}. dv: Ae. tauschii.
Glu-D1az{03124}. 4t+12.2t{03124}. dv: Ae. tauschii.
Glu-D1ba{03124}. 1t+12.3t{03124}. dv: Ae. tauschii.
Glu-D1bb{03124}. 1.5t+11t{03124}. dv: Ae. tauschii.
Glu-D1bc{03124}. 1.5t+10.3t{03124}. dv: Ae. tauschii.
Glu-D1bd{03124}. 1t+11t{03124}. dv: Ae. tauschii.
Glu-D1be{03124}. 2.1t+12.4t{03124}. dv: Ae. tauschii.
Glu-D1bf{03124}. 2t+12.1t{03124}. dv: Ae. tauschii{03124}.
Glu-D1bg{03124}. 3t+10.2t{03124}. dv: Ae. tauschii.
Glu-D1bh{03124}. 4t+10.1t{03124}. dv: Ae. tauschii.
Glu-D1bi{03124}. 4t+10.2t{03124}. dv: Ae. tauschii.
Glu-D1bj{03124}. 5t+11t{03124}. dv: Ae. tauschii.
Glu-D1bk{03124}. 5t+10.1t{03124}. dv: Ae. Tauschii.
Glu-D1bl{03124}. 5t+12.2t{03124}. dv: Ae. tauschii.
Glu-D1bm{03124}. 5*t+null{03124}. dv: Ae. tauschii.
Glu-D1bn{03124}. 5*t+12{03124}. dv: Ae. tauschii.
Glu-D1 {421} appears to have arison as the result of a deficiency of subunit 12 from Glu-D1a (2+12); subunits 2 and 12 are referred to as D1 and D5 in {421}. One of the Glu-D1o subunits has been numbered 13 in {755}; 13 was previously used to number a subunit encoded by Glu-B1f {13+16} and Glu-B1g {13+19} {1116}. Subunit 9 of Glu-D1g (5+9) was so numbered in {478} because its mobility is the same as one of the subunits encoded by Glu-B1c (7+9). Five combinations involving 6 HMW subunits [1D {p-t}] are listed in {420}. Eleven additional Glu-D1 alleles in T. tauschii were described {755}. Seven transfers of Glu-D1a and 10 of Glu-D1d (5+10) from chromosome 1D to chromosome 1A in triticale were described {846}. The subunit 2.2* encoded by Glu-D1al above and Glu-D1-1m in the appropriate list below has an unusually high Mr comparison of its N-terminal sequence and amino acid composition with those of subunit 2 (encoded by Glu-D1-1a indicated that its greater Mr, could be due to the presence of a larger central repetitive domain, although further evidence suggested that this does not affect the conformational properties of the subunit {02107}. The alleles originally designated Glu-D1w (encoding 'subunits' 2 (or 2t denoting its origin in the Ae. tauschii genome) +T1+T2), Glu-D1af (encoding 3 9or 3t)+T1+T2) and Glu-D1ag (encoding 1.5 (or1.5t)+T1+T2) share the component T1 that was originally classified as a HMW glutenin. However, it has since been shown {02108} that this protein is soluble in aqueous ethanol, casting doubt upon this classification. More recently, it has been shown {02109}, from one and two dimensional gel electrophoresis based upon SDS-PAGE and A-PAGE, and from N-terminal sequencing, that this protein is an omega-gliadin of unusually low electrophoretic mobility in SDS-PAGE, encoded by a locus located on the short arm of chromosome 1D, though distant 913.18cM) from the principle gliadin-encoding locus on 1D, Gli-D1, and 40.20cM from the high molecular weight encoding locus, Glu-D1. The authors named the locus Gli-DT1 (see below, section Gliadins). Reference to T1 was consequently removed from the Glu-D1 list. As a consequence of this finding, allele Glu-D1w was reused for a distinct allele, and Glu-D1af was omitted and will be reused for a future allele, since the combinations of subunits that these alleles originally represented are no longer unique. In {03124}, null alleles were observed for both Glu-D1-1 and Glu-D1-2,which, naturally, are not necessarily the same as those previously reported for this locus, meaning that composite alleles involving them in this study and corresponding to combinations apparently already listed in the Catalogue, may, in fact, represent novel alleles. It is also found that certain subunits of apparently identical relative mobility in SDS-PAGE showed different surface hydrophobocities in RP-HPLC; and the reverse situation was also observed (the same hydrophobicity, but different electrophoretic mobilities). It was shown {03126} that the relatively small size of a y-type HMW glutenin subunit, named 12.4t (encoded by Glu-D1-1t {03124} - see the relevant list below) and carried by accession CPI110750 of Ae. tauschii, is due to the deletion of blocks of repetitive motifs, amounting to approximately 200 amino acids, probably caused by unequal crossing-over. Note that the cultivar Fiorello is given as a standard for Glu-D1h encoding subunits 5+12 and for Glu-D1w encoding subunits 5*+10. An attempt to resolve this apparent conflict will be made in a future update. Alleles and subunits at Glu-A1-1 and Glu-A1-2: Glu-A1-1 encodes X-type glutenins and Glu-A1-2 encodes y-type glutenins.
Glu-A1-1.
Glu-A1-1a. Null. v: CS.
Glu-A1-1b. 1. v: Hope.
Glu-A1-1c. 2*. v: Bezostaya 1.
A PCR marker specific for the Glu-A1-1c (Ax2*) allele was developed in {0147}.
Glu-A1-1d. v: V74, Spain.
Glu-A1-1e. v: 132c, Poland.
Glu-A1-1f. v: 112-29, Sudan.
Glu-A1-1g. v: Landrace 1600.
Glu-A1-1h. tv: PI 94683, USSR, T. dicoccum.
Glu-A1-1i. tv: CI 12213, India, T. dicoccum.
Glu-A1-1j. 1'. tv: PI 352359, Germany, T. dicoccum; Lambro.
Glu-A1-1k. 26. v: BT-2288.
Glu-A1-1l. tv: Chinook, Canada.
Glu-A1-1m. tv: Nugget Biotype 1, Canada.
Glu-A1-1n. 1". tv: Corado, Portugal.
Glu-A1-1o. 2**. tv: PI 61189, USSR, Aric 581/1.
Glu-A1-1p. 3*. v: David 1.
Glu-A1-1q. 2***. tv: Melianopus 1528.
Glu-A1-1r. 39. i: T. thaoudar IPSR 1020006/6* Sicco.
Glu-A1-1s. 41. i: T. thaoudar G3152/6*Sicco.
Glu-A1-1t{602}. 21*{602}. v: W29323, W 3879, W 31169.
Glu-A1-1t is a provisional designation; definitive evidence that subunit 21*, which has a mobility similar to that of subunit 21, is a 'x-type' and not a 'y-type' protein has not been obtained.
Glu-A1-1u{02106}. 2*B{02106}. v: Bankuti 1201.
Glu-A1-2.
Glu-A1-2a. Null. v: CS.
Glu-A1-2b. 40. i: T. thaoudar IPSR1020006/6* Sicco.
Glu-A1-2c. 42. i: T. thaoudar G3152/6*Sicco.
Glu-B1-1.
Glu-B1-1a. 7. v: CS.
A PCR marker (2373 bp) for the Glu-B1-1a (Bx7) allele was developed in {0145}.
Glu-B1-1b. 7,7*. v: Flinor, Bezostaya 1, Owens, Norstar.
Glu-B1-1c. 7'. v: Adonis.
Glu-B1-1d. 6. v: Hope.
Glu-B1-1e. 20. v: Federation.
Glu-B1-1f. 13. v: Lancota.
Glu-B1-1g. 14. v: Sappo.
Glu-B1-1h. 17. v: Gabo.
Glu-B1-1i. 21.21x{03116}. v: Dunav; Foison{03116}.
Glu-B1-1j. 23. v: Spica D.
Glu-B1-1k. tv: PI 94640, Iran, T. dicoccum.
Glu-B1-1l. tv: PI 355505, Germany, T. diccocum.
Glu-B1-1m. tv: PI 352354, Ethiopia, T. dicoccum.
Glu-B1-1n. tv: PI 94633, Morocco, T. dicoccum.
Glu-B1-1o. v: Supreza, Canada.
Glu-B1-1p. v: Mondor.
Glu-B1-1q. tv: Canoco de Grao Escuro, Portugal.
Glu-B1-1r. tv: Tremez Mollez, Portugal.
Glu-B1-1s. tv: Quaduro, Italy.
Glu-B1-1t. tv: Athena, Italy.
Glu-B1-1u. 26. v: Cologna 1.
Glu-B1-1v. 28. v: Forlani.
Glu-B1-1w. Null. v: Olympic mutant.
Glu-B1-1x. 30. v: Marinar.
Glu-B1-1y. 32. v: BG-1943.
Glu-B1-1z. 34. v: Jeja Almendros.
Glu-B1-1aa. 37. v: Shedraya Polesja.
Glu-B1-1ab. 6*. v: Dawbill.
Glu-B1-1ac{03116}. 6.8{03116}. v: Carnac hexaploid triticale{03116}.
Glu-B1-1ad{03122}. 13*{03122}. v: PI-348767 spelt {03122}.
Glu-B1-2.
Glu-B1-2a. 8. v: CS.
Glu-B1-2b. 9. v: Bezostaya 1.
Glu-B1-2c. 16. v: Lancota.
Glu-B1-2d. 19. v: NS 335.
Glu-B1-2e. 15. v: Sappo.
Glu-B1-2f. 18. v: Gabo.
Glu-B1-2g. 22. v: Serbian.
Glu-B1-2h. 24. v: Spica D.
Glu-B1-2i. tv: PI 355505, Germany, T. dicoccum.
Glu-B1-2j. tv: PI 352354, Ethiopia, T. dicoccum.
Glu-B1-2k. tv: PI 94633, Morocco, T. dicoccum.
Glu-B1-2l. 11. v: BT-2288.
Glu-B1-2m. v: Supreza, Canada.
Glu-B1-2n. v: Mondor.
Glu-B1-2o. 8*. v: Dawbull.
Glu-B1-2p. tv: Canoco de Grao Escuro, Portugal.
Glu-B1-2q. tv: Tremez Mollez, Portugal, T. durum.
Glu-B1-2r. tv: Quaduro, Italy, T. durum.
Glu-B1-2s. 18*. v: David.
Glu-B1-2t. 27. v: Cologna 1.
Glu-B1-2u. 29. v: Forlani.
Glu-B1-2v. Null. v: Olympic mutant.
Glu-B1-2w. 31. v: Marinar.
Glu-B1-2x. 33. v: BG-1943.
Glu-B1-2y. 35. v: Jeja Almendros.
Glu-B1-2z{03116}. 20y{03116}. v: Carnac hexaploid triticale{03116}.
Glu-B1-2aa{03122}. 18'{03122}. v: PI-348631 spelt {03122}.
Glu-B1-2ab{03116}. 21y{03116}. v: Foison{03116}.
Glu-D1-1.
Glu-D1-1a. 2. v: CS.
Glu-D1-1b. 3. v: Hobbit.
Glu-D1-1c. 4. v: Champlein.
Glu-D1-1d. 5. v: Hope.
PCR markers specific for the Glu-D1-1d (Dx5) allele were developed in {0145} and {0147}.
Glu-D1-1e. 2.2. v: Danchi.
Glu-D1-1f. Null. v: Nap Hal, Nepal.
Glu-D1-1g. 2.1. v: AUS 14653, Afghanistan.
Glu-D1-1h. 2.3. v: PI 348465.
Glu-D1-1i. 38. v: Leningradka.
Glu-D1-1j{668}. 43{668}. i: Ae. tauschii accession TA2450/2*.
Glu-D1-1k{755}. 4.1{755}. dv: Ae. tauschii.
Glu-D1-1r{755}. 5.1{755}. dv: Ae. tauschii.
This allele was previously designated Glu-D1-1j in the 1998 Catalogue.
Glu-D1-1l{1578}. 1.5{1578}. dv: Ae. tauschii.
Glu-D1-1m{02107}. 2.2*{02107}. v: MG315.
Glu-D1-1n{03122}. 2.4{03122}. v: PI-348473 spelt wheat accession{03122}.
Glu-D1-1o{03122}. 2.5{03122}. v: PI-3484572 spelt wheat accesion{03122}.
Glu-D1-1p{03124}. 1t{03124}. dv: Ae. tauschii{03124}.
Glu-D1-1q{03124}. 5*t{03124}. dv: Ae. tauschii{03124}.
Glu-D1-2.
Glu-D1-2a. 12. v: CS.
A PCR marker (612 bp) for the Glu-D1-2a (Dy12) allele was developed in {0145}.
Glu-D1-2b. 10. v: Hope.
PCR markers (576 bp and 2176bp) for the Glu-D1-2b (Dy10) allele were developed in {0145} and {0147}, respectively.
Glu-D1-2c. 9. v: BT-2288.
Glu-D1-2d. Null. v: Nap Hal, Nepal.
Glu-D1-2e. 12*. v: Tudest.
Glu-D1-2f. 13. v: AUS 14519, T. macha.
Glu-D1-2g. 36. i: Iranian landrace 3048/5* Sicco.
Glu-D1-2h. 11. v: Flinor.
Glu-D1-2i{668}. 44{668}. i: Ae. tauschii TA2450/2*.
Glu-D1-2j{836}. 10'{836}. v: Coker 68-15.
Glu-D1-2k{755}. T1{755}. dv: Ae. tauschii.
Glu-D1-2l{755}. T2{755}. dv: Ae. tauschii.
Glu-D1-2m{755}. 10.1{755}. dv: Ae. tauschii.
Glu-D1-2n{755}. 10.2{755}. dv: Ae. tauschii.
Glu-D1-2o{755}. 10.3{755}. dv: Ae. tauschii.
Glu-D1-2p{1578}. 10.5{1578}. dv: Ae. tauschii.
Glu-D1-2q{03122}. 12'{03122}. v: PI-348495 spelt wheat accession{03122}.
Glu-D1-2r{03124}. 12.1t{03124}. dv: Ae. tauschii.
Glu-D1-2s{03124}. 12.3t{03124}. dv: Ae. tauschii.
Glu-D1-2t{03124}. 12.4t{03124}. dv: Ae. tauschii.
Glu-Agi1{374}. 1Agi{374}. ad: Vilmorin 27/Thin. intermedium.
Glu-E1{781}. 1ES{781}. ad: CS/E. elongata.
Glu-H1{781}. [Hor 3{1337}]. 1H{781}.1HL{1337}. ad: CS/Betzes{781}. al: Various barley cultivars{1337}.
Glu-Hch1. 1Hch{1123}. ad: CS/H. chilense.
Glu-Hch1a{03114}. 1Hch{03114}. al: H. chilense Accession H1{03114}.
Glu-Hch1b{03114}. 2Hch{03114}. al: H. chilense Accession H11{03114}.
Glu-Hch1c{03114}. 3Hch{03114}. al: H. chilense Accession H7{03114}.
Glu-Hch1d{03114}. 4Hch{03114}. al: H. chilense Accesion H16{03114}.
Glu-Hch1e{03114}. 5Hch{03114}. al: H. chilense Accession H47{03114}.
Glu-Hch1f{03114}. 6Hch{03114}. al: H. chilense Accession H220{03114}.
Glu-Hch1g{03114}. 7Hch{03114}. al: H. chilense Accession H293{03114}.
Glu-Hch1h{03114}. 8Hch{03114}. al: H. chilense Accession H297{03114}.
Glu-Hch1i{03114}. 9Hch{03114}. al: H. chilense Accession H252{03114}.
Glu-Hch1j{03114}. 10Hch{03114}. al: H. chilense Accession H210{03114}.
38 accessions (natural populations) of Hordeum chilense carrying the following 10 subunits were used as the maternal parents of 121 lines of primary tritordeum, and evaluations for associations with bread-making quality initiated {03114}. Subunits 1Hch, 2Hch and 3Hch were previously referred to as Hcha, Hchb and Hchc {03112}.
Glu-Ht1{1037}. 1HtL{1037}. ad: CS/E. trachycaulum.
Glu-R1{781,1356}. [Sec 3{1336}]. 1R{781,1336}.1RL{1356,1340}. ad: CS/Imperial; Holdfast/ King II{1340}. tr: CS Imperial 1DS.1RL{1356}.
Glu-R1a{03116}. 1r-4r{03116}. v: Indiana hexaploid triticale{03116}.
Glu-R1b{03116}. 2r-6.5r{03116}. v: Graal hexaploid triticale{03116}.
Glu-R1c{03116}. 6r-13r{03116}. v: Almao hexaploid triticale{03116}.
Glu-R1d{03116}. 2r-9r{03116}. v: Olympus hexaploid triticale{03116}.
Glu-R1e{03116}. 6.5r{03116}. v: Clercal hexaploid triticale{03116}.
Glu-R1f{03115}. 0.8r-6r{03115}. v: Carmara hexaploid triticale{03115}.
Glu-R1g{03115}. 5.8r{03115}. v: Arrayan hexaploid triticale{03115}.
There is a difficulty in the assignment of subunit 6r in the Glu-R1-1 and Glu-R1-2 lists, since it appears as an x-type subunit in allele Glu-R1c and as a y-type subunit in allele Glu-R1f. It is currently provisionally assigned to the Glu-R1-1 list since, based upon its relative electrophoretic mobility, it is considered more likely to be an x-type subunit. Some of the remaining designations should also be considered as provisional since they too are not free of ambiguity.

From study of chromosome substitutions in bread wheat {03117}, it was found that a chromosome 1R carrying HMW secalin subunit 6.5r (Glu-R1e), originally derived from the 'Petkus' rye population, was associated with bread making quality (i) intermediate between chromosome 1A carrying the null allele Glu-A1c and chromosome 1A carrying HMW glutenin subunit 2* encoded by Glu-A1b; (ii) equivalent to a chromosome carrying HMW glutenin subunit 7 encoded by Glu-B1a; and (iii) inferior to chromosomes 1D with distinct alleles.
Glu-R1-1.
Glu-R1-1a{03116}. 1r{03116}. v: Indiana hexaploid triticale{03116}.
Glu-R1-1b{03116}. 2r{03116}. v: Graal hexaploid triticale{03116}.
Glu-R1-1c{03116}. 6r{03116}. v: Alamo hexaploid triticale{03116}.
Glu-R1-1d{03115}. 0.8r{03115}. v: Carmara hexaploid triticale{03115}.
Glu-R1-1e{03115}. 5.8r{03115}. v: Arrayan hexaploid triticale{03115}.
Glu-R1-2. 1R, 1RL.
Glu-R1-2a{03116}. 4r{03116}. v: Indiana hexaploid triticale{03116}.
Glu-R1-2b{03116}. 6.5r{03116}. v: Graal hexaploid triticale{03116}.
Glu-R1-2c{03116}. 13r{03116}. v: Alamo hexaploid triticale{03116}.
Glu-R1-2d{03116}. 9r{03116}. v: Olympus hexaploid triticale{03116}.
Glu-Rm1{1339}. 1RmL{1339,1340}. ad: CS/S. montanum{1339,1340}.
Glu-Sl1{1228}. 1SlL{1228}. ma: In Ae. longissima 2 /Ae. longissima 10,Glu-Sl1, Glu-Sl3, one glucose phosphate isomerase locus, and three gliadin loci were mapped relative to one and other {1228} as follows: Glu-Sl1 - 15.9 cM - Gpi-Sl1 - 38 cM - Gli-Sl4 - 7.1 cM - Glu-Sl3 - 0.9 cM - Gli-Sl1 - 5.6 cM - Gli-Sl5. Glu-Sl1 is located in 1SlL and the other loci are in 1SlS.
Glu-U1{150}. 1U{150,781}. ad: CS/Ae. umbellulata {150,781}.
Glu-V1{111,242,1026}. 1V{1026,111}. ad: CS/D. villosum; Creso/D. villosum.
Glu-V1a{1651}. 71{1651}. al: D. villosum.
Glu-V1b{1651}. 72{1651}. al: D. villosum.
Glu-V1c{1651}. 73{1651}. al: D. villosum.
Glu-V1d{1651}. 74{1651}. al: D. villosum.
Glu-V1e{1651}. 75{1651}. al: D. villosum.
Glu-V1f{1651}. 76{1651}. al: D. villosum.
Glu-V1g{1651}. 77{1651}. al: D. villosum.
Glu-V1h{1651}. 78{1651}. al: D. villosum.
Glu-V1i{1651}. 79{1651}. al: D. villosum.
Glu-V1j{1651}. 80{1651}. al: D. villosum.
Glu-V1k{1651}. null{1651}. al: D. villosum.
Glu-V1l{1651}. 81+82{1651}. al: D. villosum.
Glu-V1m{1651}. 83+84{1651}. al: D. villosum.
Glu-V1n{1651}. 85+86{1651}. al: D. villosum.
Alleles and subunits at Glu-V1 and GLU-V1-2 : The following is analogous to the Glu-1-1 and Glu-1-2 lists given earlier to identify x-type and y-type subunits in wheat. It was assumed that where an allele at Glu-V1 produces only a single subunit, it is an x-type subunit and so encoded by Glu-V1-1 rather than by Glu-V1-2; the electrophoretic mobilities of the subunits are all greater, though some only marginally so, than subunit 7 encoded by Glu-B1-1a (an x-type subunit), and extend beyond the mobility of subunit 12 encoded by Glu-D1-2a (a y-type subunit) {1651}; therefore, it is quite possible that any one of the subunits designated as encoded by Glu-V1-1 is, in fact, encoded by Glu-V1-2. The designation given here is intended to be the most practically useful until the identities of the genes encoding the alleles are directly established.
Glu-V1-1.
Glu-V1-1a{1651}. 71{1651}. al: D. villosum.
Glu-V1-1b{1651}. 72{1651}. al: D. villosum.
Glu-V1-1c{1651}. 73{1651}. al: D. villosum.
Glu-V1-1d{1651}. 74{1651}. al: D. villosum.
Glu-V1-1e{1651}. 75{1651}. al: D. villosum.
Glu-V1-1f{1651}. 76{1651}. al: D. villosum.
Glu-V1-1g{1651}. 77{1651}. al: D. villosum.
Glu-V1-1h{1651}. 78{1651}. al: D. villosum.
Glu-V1-1i{1651}. 79{1651}. al: D. villosum.
Glu-V1-1j{1651}. 80{1651}. al: D. villosum.
Glu-V1-1k{1651}. null{1651}. al: D. villosum.
Glu-V1-1l{1651}. 81{1651}. al: D. villosum.
Glu-V1-1m{1651}. 83{1651}. al: D. villosum.
Glu-V1-1n{1651}. 85{1651}. al: D. villosum.
Glu-V1-2.
Glu-V1-2a{1651}. null{1651}. al: D. villosum.
Glu-V1-2b{1651}. 82{1651}. al: D. villosum.
Glu-V1-2c{1651}. 84{1651}. al: D. villosum.
Glu-V1-2d{1651}. 86{1651}. al: D. villosum.
A Chinese cultivar of T. aestivum, Xiaoyanmai 7, carries a subunit with electrophoretic mobility in 10% SDS-PAGE well beyond that of subunits so far observed in T. aestivum. It may derive from Agropyron elongatum, which was used in the breeding programme that led to the variety {1538}. It has not been given a subunit number or allelic designation, because its genetic control has not been elucidated.


74.3.1.2. Glu-2
Glu-B2{819,277}. [XGlu-B2{277}]. 1BS. s: CS*/Cheyenne 1B{277}. stv: Langdon*/T. turgidum var. dicoccoides 1B{277}.
Glu-B2a{00114}. 12{00114}. tv: Mexicali.
Glu-B2b{00114}. Null{00114}. tv: Langdon.
Gli-B3 was designated Glu-B2 {589} until the name of the locus was changed in {1119}.


74.3.1.3. Glu-3

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

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

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

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

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

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

A naming system in which roman numerals are assigned to whole banding patterns for the LMW glutenin subunit is given in {03131} as an alternative to the LMW-1/-2 system described in {03136}. A further system naming whole banding patterns from LMW-1 to LMW-23 in emmer wheat is described in {03137}. In {00111}, in a study of common and durum wheats from Portugal, the authors used the nomenclature system described in {00113} for the LMW subunits in common wheat, and that described in {00114} for the LMW subunits in durum wheat. The latter system was updated according to {02110}, but has been changed herein to new alleles with the earlier durum designation {00114} given as synonyms.In {03116}, it was suggested that Glu-B3d (common wheat standard genetic stock) is equivalent to Glu-B3r (durum wheat standard genetic stock), and that (referring to article {03127}) LMWsubunits observed in some Portugese triticales could be of the durum type.
Glu-A3{1358}. 1AS{1358}. v: CS.
Glu-A3a{481}. v: CS.
Glu-A3b{481}. v: Gabo.
Glu-A3c{481}. v: Cheyenne.
Glu-A3d{481}. v: Cappelle Desprez, Orca.
Glu-A3e{481}. v: Hope, Insignia.
Glu-A3f{481}. v: Rescue.
Glu-A3g{00113,00114}. 6+10+20{00114}. tv: Claro de Balazote.
Glu-A3h{00114,03116}. [Glu-A3d'{03116}]. null{00114}. v: Magistral hexaploid triticale{03116}.
Glu-A3i{02110}. 8*+11{02110}. tv: Mourisco Fino.
In 112 common wheat cultivars from Argentina, 11 microsatellite alleles plus a null allele were found at the Glu-A3 locus {03123}.
Glu-A3j{00114}. [Glu-A3a{00114}]. 6{00114}. tv: Mexicali.
Glu-A3k{00114}. [Glu-A3b(00114)]. 5{00114}. tv: Langdon.
Glu-A3l{00114}. [Glu-A3c{00114}]. 6+10{00114}. tv: Cocorit.
Glu-A3m{00114}. [Glu-A3d{00114}]. 6+11{00114}. tv: Alaga.
Glu-A3n{00114}. [Glu-A3e{00114}]. 11{00114}. tv: Blatfort.
Glu-A3o{00114}. [Glu-A3f{00114}]. 6+11+20{00114}. tv: Clarofino.
Glu-A3p{00114}. [Glu-A3h{00114}]. Null{00114}. tv: Jiloca.
Glu-B3{1358}. 1BS{1358}. v: CS.
Glu-B3a{481,}. v: CS.
Glu-B3b{481}. v: Gabo, Timstein, Hope.
Glu-B3c{481}. v: Insignia, Halberd.
Glu-B3d{481}. v: Orca.
Glu-B3e{481}. v: Cheyenne.
Glu-B3f{481}. v: Radja.
Glu-B3g{481}. v: Kharkov, Bungulla.
Glu-B3h{481}. v: Thatcher, Rescue.
Glu-B3i{481}. v: Norin-61.
Glu-B3j{476,02110}. 4+6*+15+19{02110}. tv: Duramba-B, Duramba-D, Langdon; Mourisco Fino.
Glu-B3k{476,02110}. 8+9+13+16+19{02110}. tv: ALP-153, Dural, Durati, Edmore; Faisca.
Glu-B3l{476}. tv: Gionp-1954.
Glu-B3m{03120}. [Glu-B3b'{03120}]. v: Soissons{03120}.
Glu-B3n{03120}. [Glu-B3c'{03120}]. v: Courtot{03120}.
Glu-B3o{03116}. [Glu-B3i'{03116}]. v: Olympus hexaploid triticale{03116}.
Glu-B3p{03116}. [Glu-B3k{03116}]. v: Alamo hexaploid triticale{03116}.
Glu-B3q{03115}. [Glu-B3h'{03115}]. v: Torote hexaploid triticale{03115}.
Glu-B3r{00114}. [Glu-B3a{00114}]. 2+4+15+19{00114}. tv: Mexicali.
Glu-B3s{00114}. [Glu-B3b{00114}]. 8+9+13+16{00114}. tv: Langdon.
Glu-B3t{00114}. [Glu-B3c{00114}]. 2+4+14+15+19{00114}. tv: Jiloca.
Glu-B3u{00114}. [Glu-B3d{00114}]. 2+4+15+17+19{00114}. tv: Mundial.
Glu-B3w{00114}. [Glu-B3f{00114}]. 2+4+15+17{00114}. tv: Ardente.
Glu-B3v {00114}. [Glu-B3e{00114}]. 2+4+15+16+18{00114}. tv: Granja Badajoz.
Glu-B3x{00114}. [Glu-B3g{00114}]. 2+4+15+16{00114}. tv: Claro de Balazote.
Glu-B3y{00114}. [Glu-B3h{00114}]. 1+3+14+18{00114}. tv: Alaga.
Glu-D3{1358,707}. 1DS{707,1358}. v: CS.
Glu-D3a{481}. v: CS.
Glu-D3b{481}. v: Gabo.
Glu-D3c{481}. v: Insignia, Cappelle Desprez.
Glu-D3d{481}. v: Norin-61A.
Glu-D3e{481}. v: Orca, Thatcher.
Glu-E3{480}. 1ES{480}. su: CS/E. elongata.
Glu-Sl3{480,1228}. 1Sl{480}.1S 1S{1228}. su: CS/Ae. longissima{480,1228}. ma: In Ae. longissima 2 /Ae.longissima 10 glucose phosphate isomerase locus, and three gliadin loci were mapped relative to one another in {1228} as follows: Glu-Sl1 - 15.9 cM - Gpi-Sl1 - 38 cM- Gli-Sl4 - 7.1 cM - Glu-Sl3 - 0.9 cM - Gli-Sl1 - 5.6 cM - Gli-Sl5. Glu-Sl1 is located in 1SlL and othe other loci are in 1S1S.
Glu-U3{480}. 1U{480}. su: CS/Ae. umbellulata.
A series of papers {00106, 00107, 00108 and 00109} describe considerable variation in primitive wheats not present in bread wheat (A genome species T. boeoticum, T. urartu, T. thaoudar, T. aegilopoides, T. monococcum, and D-genome species T. tauschii) for the low molecular weight subunits, sufficient to use them as a source for potentially changing flour properties in bread wheat.
In {00110}, variants for LMW glutenin subunits have been reported from study of twenty-four accessions of einkorn wheat (T. monococcum ssp. monococcum). Nine of these showed two electrophoretic bands for LMW subunits, arbitrarily designated 'a' and 'b', that appeared to be associated with good bread-making quality.The isolation of a new low-molecular-weight glutenin subunit gene, located on chromosome 1D, was reported in {0350}.



74.3.1.4. Glu-4
The following loci, Glu-D4 and Glu-D5, encoding low molecular weight subunits of glutenin (30-32 kDa) were described in {02111}; the proteins encoded by them were first observed earlier {02114, 02115}, and the former was later tentatively assigned the symbol Glu-4 {02116}, before its chromosomal location was established and the locus definitively named as Glu-D4 in {02111}. While this locus is located on chromosome 1D (in accordance with the position on the group 1 chromosomes of the remaining glutenin encoding loci found to date), the locus Glu-D5 is located on chromosome 7D. In SDS-PAGE, the proteins from both loci are detected only in the presence of 4-vinylpyridine added to the sample extract. Their amino acid compositions do not match those of the major prolamin groups; nonetheless, they classify as glutenins based upon solubility, immunological behaviour and N-terminal amino acid sequence (the latter suggesting an evolutionary link with the major (B and C) low molecular weight glutenin subunits).
Glu-D4{02111}. 1D{02111}. su: CS/Langdon 1D(1A); CS/Langdon 1D(1B){02111}.
Glu-D4a{02111}. v: J 24.
Glu-D4b{02111}. v: PBW 154.
Glu-D4c{02111}. Null allele. v: NI 4.


74.3.1.5. Glu-5
Glu-D5{02111}. 7D{02111}. su: CS/Langdon 7D(7A); CS/Langdon 7D(7B){02111}.
Glu-D5a{02111}. v: PBW 154.
Glu-D5b{02111}. Null allele. v: K 68.

A collection of 173 Ae. tauschii accessions were analysed for low molecular weight glutenin subunits by SDS-PAGE {02112}. Thirty three different patterns for B-subunits and 43 for C-subunits were identified, some of which were of identical electrophoretic mobility to those observed in common wheat. Also observed were subunits with the same mobilities as the D-subunits and as the subunits encoded by the Glu-D4 and Glu-D5 loci. This variation represents a source of novel germplasm of potential value for breeding programmes aimed at improving the D-genome of common wheat in the context of bread-making quality.

PCR amplification of genomic DNA has been used to isolate three LMW glutenin genes in cultivar Chinese Spring, named LMWG-MB1, LMWG-MB2 and LMWG-MB3 {01101}. The deduced amino-acid sequences showed a high similarity between these ORFs and with those of other LMW glutenin genes. The authors state that the study provided direct evidence that insertions and/or deletions provide a mechanistic explanation for the allelic variation, and hence the resultant evolution, of prolamin genes, and comment on relationships with gamma-secalins and beta-hordein families. Single-base substitutions at identical sites generate premature stop codons in both LMWG-MB2 and LMWG-MB3, indicating that these clones are pseudogenes.

74.3.2. Gliadins
These are heterogeneous mixtures of alcohol-soluble polypeptides without quaternary structure. The Gli-1 loci are compound and are now considered to comprise the omega-gliadin and gamma gliadin {982,1415} multigene families {494}, which in some circumstances may be divided into Gli-1-1 and Gli-1-2, respectively. The LMW glutenin multigene families, which are closely linked to the Gli-1 loci {588}, are listed separately as the Glu-3 set {1358}; information on map distance and gene order in relation to Glu-3 and the centromere is given in the preamble for the Glu-3 loci. There is evidence that a few of the omega-gliadin genes are separated from the main omega-gliadin gene cluster {993}.
Variation at the Gli-1 loci was described earlier {634,996,1126} and applied in mapping experiments {1243,1125,196,422,1120}. A rational system of naming the alleles was produced by Dr. E.V. Metakovsky{988}. This nomenclature is reproduced below. A considerable number of alleles were added to the original list given in {988}, and referenced here accordingly. A few alleles have been deleted, because, following much detailed comparison, there is now doubt that they can be reliably distinguished from existing alleles {9981}. The allelic letter in these cases has not been reused. To facilitate practical use of the list, the aim was to give at least three standard cultivars from a range of countries for each allele {9981}. This was achieved for the vast majority of entries and is a change from the original list compiled from {988}, where up to two standards were given. While the three or more standards described almost always include the original standards, some have been replaced for various reasons, such as international awareness of the cultivar, availability of seed, or the ease with which an allele can be identified in a particular genetic background {9981}. In the original list, where two cultivars were given as prototypes for an allele, the first named was from the USSR and the second from elsewhere; this is no longer the case, although care was taken to include a Russian cultivar where possible, to maintain a wide base of germplasm in which the alleles are available, as well as to acknowledge the research groups in the country where much of the pioneering work was carried out.
For discussion of null alleles at the Gli-1 and Gli-2 loci, see {9984}.
Recombination was observed within the gliadin multigene family at XGli-A1 {277}. These closely linked genes may correspond to Gli-A1 and Gli-A5, but they were temporarily designated XGli-A1.1 and XGli-A1.2 until orthology with Gli-A1 and/or Gli-A5 is established.

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

1B and 1D sulphur-poor omega-gliadins in cultivar Butte 86 were characterised by RP-HPLC, SDS-PAGE, two-dimensional PAGE, amino acid composition determination and sequencing, matrix assisted laser desorption ionisation-time of flight mass spectrometry and circular dichroism spectroscopy to reveal the detailed nature of the peptides belonging to the two groups, and showing that the complexity of mixtures of the peptides of the 1B group was greater than that of the 1D group {01105}. Although circular dichroism spectra were similar for the two groups of peptides, and suggested a mainly flexible random structure, there was evidence for a significant amount of left-handed polyproline II helical conformation in the case of the 1D components. The authors placed some of the results in the context of the possible ancestor of the B-genome and relationships with the barley C-hordeins and rye omega-secalins.
Eleven new gliadin alleles were found in a collection of 52 Spanish landraces of common wheat {03141}. These will be added to the Gli-1 and Gli-2 allelic lists in the next Supplement.

74.3.2.1. Gli-1
Gli-A1{1334,1125}. [Gld 1A{1415}]. 1AS{150,634,1334,1607}. s: CS*/Cheyenne{634}. v: CS{150,1334,1607}.
Gli-A1a{988}. v: Castan{991}; CS{988}; Mara{9986}; Mentana{9986}; Millewa{00119}.
Gli-A1b{988}. v: Bezostaya 1, Mercia{988}; Tracy{991}.
Gli-A1c{988}. v: Ukrainka{998}; Gazul{9985}; Sava{994}; Hopps{00119}.
Gli-A1d{988}. v: Dankowska{988}; Cabezorro{9985}.
Gli-A1e{988}. v: Falchetto{988}; Open{991}; Touzelle{991}.
Gli-A1f{988}. v: Mironovskaya 808, Maris Freeman{988}; Arminda{991}.
Note: An allele Gli-A1f* is mentioned in {03130}.
Gli-A1g{988}. v: Gabo{988}; Adalid{9985}.
Gli-A1h{988}. v: Sadovo I{988}; Predela{9981}; Krajinka{9981}.
Gli-A1i{988}. v: Saratovskaya 36{988}.
Gli-A1j{988}. v: Lutescens 62{988}.
Gli-A1k{988}. v: Courtot{991}; Skala (heterogeneous){988}; Soissons{991}; Spada{9986}.
Gli-A1l{988}. v: Lesostepka 75{988}; David{9986}; Salmone{9986}; Mura{9981}.
Gli-A1m{988}. v: Marquis{988}; Dneprovskaya 521{988}; Carat{991}; Liocorno{9986}.
Gli-A1n{988}. v: Intensivnaya{988}.
Gli-A1o{988}. v: Odesskaya 16 (heterogeneous){988}; Oderzo{9986}; Cappelle-Desprez{991}; Capitole{991}.
Gli-A1p{988}. v: Pyrotrix 28{988}; Zagore{9981}.
Gli-A1q{988}. v: Akmolinka 1{988}.
Gli-A1r{988}. v: Ranniaya 73{988}; Barbilla{9985}.
Gli-A1s.
Although reported {9986}, this gene is omitted because it requires further confirmation {9981}.
Gli-A1t{9985}. v: Jeja del Pais{9985}; Milturum 553{9981}; Strela{9981}.
Gli-A1u{9985}. v: Candeal Alcala{9985}.
Gli-A1v{9981}. v: Japhet{9981}; Rouge de Bordeaux{9981}.
Gli-A1null{9984,9987}. v: Saratovskaya 29 (mutant){9987}; E. Mottin{9981}.
Gli-B1{1607,1125}. [Gld 1B{1243,1415},Gld-B1{420},Gld-B2{420},Gld-B3{420},Gld-B4{420},Gld-B5{420},Gld-B6{420}]. 1B{1607}.1BS{150,634}. s: CS*/Cheyenne{634}. v: CS{1607,150}.
Gli-B1a{988}. v: CS{988}.
Gli-B1b{988}. v: Bezostaya 1{988}; Carat{991}; Marquis{988}; Liocorno{9986}; Soissons{991}.
Gli-B1c{988}. v: Siete Cerros 66{988}; Prinqual{991}; Loreto{9986}.
Gli-B1d{988}. v: Dneprovskaya 521{988}; Chopin{991}; Petrel{991}; Tiberio{9986}; Yecora{9985}; Neepawa{995}; Suneca{00119}.
Gli-B1e{988}. v: Apexal{991}; Fournil{991}; Lutescens 62{988}; Oderzo{9986}.
Gli-B1f{988}. v: Capitole{991}; Cappelle-Desprez{991}; Dankowska{988}; Maris Freeman{988}; Mercia{998}.
Gli-B1g{988}. v: Champtal{991}; Galahad{988}; Mara{9986}; Sadovo 1{988}; Tracy{991}.
Gli-B1h{988}. v: Cabezorro{9985}; Krasnodonka{988}; Pepital{991}; Rudi{991}Tincurrin{00119}.
Gli-B1i{988}. v: Ghurka{988}; Insignia{988}.
Gli-B1j{988}. v: Cluj 650{988}.
Gli-B1k{988}. v: Crverkapa{994}; De Carolis{9986}; Kremena{988}; Mentana{9986}.
Gli-B1l{988}. v: Avrova{9981}; Clement{991}; Damier{991}; Fiocco{9986}; Kavkaz{9981}.
Gli-B1l encodes secalins ssociated with the 1BL.1RS translocation.
Gli-B1m{988}. v: Costantino{9986}; Et.d'Choisy{991}; Pyrotrix 28{988}.
Gli-B1n{988}. v: Intensivnaya{988}.
Gli-B1o{988}. v: Aragon 03{9985}; Levent{988}; Pippo{9986}; San Rafael{9985}.
Gli-B1p{988}. v: Inia 66{9985}; New Pusa 834{988}.
Gli-B1q{9986}. v: Gallo{9986}; Goelent{991}; Goya{991}.
Gli-B1r{995}. v: Chinook{995}; Gazul{9985}; Sevillano{9985}.
Gli-B1s{9986}. v: Salmone{9986}; Resistente{9986}; E.Mottin{9981}.
Gli-B1t{9985}. v: Jeja del Pais{9985}.
Gli-B1u{9985}. v: Negrillo{9985}.
Gli-B1v{9985}. v: Montjuich{9985}.
Gli-B1w{9981}. v: Ardica{9981}; Barbilla (MCB-1017){9981}.
Gli-B1null{9984,9987,991}. v: Touzelle{991}; Florence Aurora{9985}.
In 112 bread wheat cultivars from Argentina, 12 microsatellite alleles plus a null allele were found at the Gli-B1 locus tightly linked to Glu-B3 {03123}.
Gli-D1{121,1125}. [Gld 1D{1415},Gld-D1{420},Gld-D2{420},Gld-D3{420}]. 1DS{121,150,634,1334,1607}. s: CS*/Cheyenne{634}. v: CS{121,150,1334,1607}.
Gli-D1a{988}. v: CS{988}; Marquis{988}; Mentana{9986}; Prinqual{991}; Saratovskaya 36{988}.
Gli-D1b{988}. v: Bezostaya 1{988}; Cappelle-Desprez{991}; Et.d'Choisy{991}; Galahad{988}.
Gli-D1c{988}. v: Skorospelka Uluchshennaya (biotype){988,9982}.
Gli-D1d{988}. v: De Carolis{9986}; Solo{988}.
Gli-D1e{988}. v: Gerek 79{988}.
Gli-D1f{988}. v: Carlos{991}; Gabo{988}; Maris Freeman{988}; Orso{9986}.
Gli-D1g{988}. v: Fournil{991}; Ghurka{988}; Mironovskaya 808{988}; Open{991}.
Gli-D1h{988}. v: Sadovo I{988}; Zlatostrui{9981}.
Gli-D1i{988}. v: Insignia{988}; Napayo (biotype){995}; San Rafael{9985}; Tselinogradka{988}.
Gli-D1j{988}. v: Aubain{991]; Chinook{995}; Inia 66{9985}; Petrel{991}; Promin{988}.
Gli-D1k{988}. v: Cargimarec{991}; Kremena{988}; Mara{9986}; Pippo{9986}.
Gli-D1l{988}. v: Artaban{991}; Corin{991}; Longbow{988}.
Gli-D1m{991}. v: Heurtebise{991}.
Gli-D1n{981}. v: Blanquillo de Toledo (MCB-0950){9981}.
Gli-D1null{9984,9987,991}. v: Darius{991}; Touzelle{991}; Saratovskaya 29 (mutant){9987}.
Gli-Agi1. 1Agi{168}. ad: Vilmorin 27/Thin. intermedium.
Gli-DT1{02109}. 1DS{02109}. dv: AUS18913 Ae. tauschii. v: L/18913 (synthetic).
A locus designated Gli-DT1 controlling an omega-gliadin of T. tauschii was mapped on the short arm of chromosome 1D between loci Gli-D1 (strictly Gli-Dt1) and Glu-D1 (strictly Glu-Dt1), 13.18 cM proximal to the former and 40.20 cM from the latter {02109}. The only omega-gliadin to date identified as being encoded by this locus, namely T1, is of unusually low electrophoretic mobility in SDS-PAGE gels and was formally thought to be a high molecular weight glutenin encoded by the Glu-Dt1 locus of T. tauschii (see note following the Glu-D1 list in section '74.3.1 Glutenins'). The authors speculate that, due to their similar relative map positions, the loci Gli-A4, Gli-D4, Gli-R3, Gli-Sl4 and this locus, Gli-DT1, form a series of 'Gli-4' orthologous loci. However, this should be interpreted in the light of the above discussion on Gli-A3 and Gli-A4.
Gli-DT1a{02109}. T1. dv: AUS18913 Ae. tauschii. v: L/18913 (synthetic).
Gli-E1{781}. 1ES{781}. ad: CS/E. elongata.
Gli-Ht1{1037}. 1Htp{1037}. ad: CS/E. trachycaulum.
In barley, the B and C hordeins are controlled by the Hor2 and Hor1 loci, respectively, which are linked {1341} on chromosome 1HS {1063,1153}. The map distances and homology of the proteins indicate that Hor1, the locus closest to the centromere, is equivalent to the omega-gliadins (Gli-1-1) in Gli-1 {1338}.
Gli-R1{1334}. [SecR1{1356},Sec1{1336}]. 1RS{781,1334,1336,1340}. ad: CS/Imperial{781,1334,1336,1340}; Holdfast/King II{1334,1340}. tr: CS 1DS. Imperial 1RL{1356}.
Sec-12 and Sec13 are given as allelic alternatives in 1BL.1RS translocation lines by {03132}.
Gli-Rm1{1340}. 1RmS{1340}. ad: CS/S. montanum.
Gli-Sl1{573}. 1Sl{573}. ad: CS/Ae. longissima.
Gli-U1{1335}. 1U{1335,150}. ad: CS/Ae. umbellulata.
Gli-V1{1026,111}. 1V{1026,111}. ad: CS/D. villosum{1026}; Creso/D. villosum{111}.

Three alleles at each of the Gli-1-1 (omega gliadin) loci were noted {1358}. The complexity of the Gli-1 compound loci is further emphasized by a report of individual genes being separable by recombination, where G1d-1A (a block of gamma and omega genes) is separable by 0.3% from Gld4-1A (omega gliadins) which is in turn, separable by 1.5% from Gld3-1A (omega gliadins) {1103}.
Elsewhere, variation was described {634,996,1126} and applied in mapping experiments {107,196,422,1120,1125,1243}. Sixteen combinations of Gli-B1 and 4 combinations of Gli-D1 subunits are listed in {420}. Multiple alleles described in {996}, number 15 at Gli-A1, 18 at Gli-B1, and 8 at Gli-D1.
The Gli-1 alleles present in 57 Yugoslav wheat varieties were reported in {994}.

74.3.2.2. Gli-2
Gli-A2{1334,1125}. [Gld 6A{1415}]. 6A{1334}.6AS{1122}. v: CS.
Gli-A2a{988}. v: Cabezorro{9985}; CS{988}; Insignia{988}; Rieti DIV{9986}.
Gli-A2b{988}. v: Aradi{9985}; Bezostaya 1{988}; Rivoli{991}; Tiberio{9986}.
Gli-A2c{988}. v: Eagle{00119}; Escualo{9985}; Loreto{9986}; Prinqual{991}; Siete Cerros 66{988}.
Gli-A2d{988}. v: Dneprovskaya 521{988}; Kenyon (biobype){995}; Mocho Sobarriba{9985}.
Gli-A2e{988}. v: Cobra{991}; Mentana{9986}; Resistente{9986}; Sadovo 1{988}; Sevillano{9985}.
Gli-A2f{988}. v: Adalid{9985}; Gala{991}; Maris Freeman{988}; Sistar{9986}.
Gli-A2g{988}. v: Cappelle-Desprez{991}; Ducat{988}; Mahissa 1{9985}; Mara{9986}.
Gli-A2h{988}. v: Apollo{991}; Basalt{9981}; Hereward{988}; Montjuich{9985}; N. Strampelli{9986}.
Gli-A2i{988}. v: Krasnodonka{988}; Lesostepka 75{988}.
Gli-A2j{988}. v: Avalon{9981}; Camp Remy{991}; E. Mottin{9981}; Recital{991}.
Gli-A2k{988}. v: Akmolinka 1{988}; Estica{991}; Pyrotrix 28{988}; Renan{991}; Zena{9986}.
Gli-A2l{988}. v: Chamorro{9985}; Champlein{991}; Longbow{988}.
Gli-A2m{988}. v: Marquis{988}; Rex{991}; Suneca{00119}.
Gli-A2n{988}. v: Mironovskaya 808{988}.
Gli-A2o{988}. v: Calatrava{9985}; Castan{991}; Glenwari{9981}; Lontra{9986}; Touzelle{991}.
Gli-A2p{988}. v: Cajeme 71{9985}; Capitole{991}; Clement{991}; Pliska{988}; S. Lorenzo{9986}; Yecora{9985}.
Gli-A2q{988}. v: Candeal Alcala{9985}; Montcada{9985}; Saratovskaya 39{988}.
Gli-A2r{988}. v: Genial{991}; Open{991}; Riband{988}.
Gli-A2s{988}. v: Saratovskaya 36{998}.
Gli-A2t{988}. v: Courtot{991}; Prostor{9981}; Rinconada{9985}; Soissons{991}.
Gli-A2u{988}. v: Aragon 03{9985}; Kirgizskaya Yubileinaya{988}; Saunders{995}; Titien{991}.
Gli-A2v{988}. v: Kzyl-Bas{988}.
Gli-A2w{988}. v: Bezenchukskaya 98 (biotype){988}.
Gli-A2x{988}. v: Solo{988}.
Gli-A2y{9981}. v: Gentil Rosso 202{9981}; PI 191245{9981}.
Gli-A2z{9986}. v: Gallo{9986}; Giuliana{9986}.
Gli-A2aa{9985}. v: Navarro 122{9985}.
Gli-A2ab{9985}. v: Navarro 150{9985}.
Gli-A2ac{9981}. v: Blanquillo de Barcarrota (MCB-0893){9981}.
Gli-A2ad{9981}. v: Hembrilla Soria (MCB-1298){9981}.
Gli-A2ae{9981}. v: Candeal de S.Lorenzo Parrilla (MCB-0932){9981}.
Gli-A2af{9981}. v: Barbilla de Leon (MCB-1292){9981}.
Gli-A2ag{9981}. v: Gluclub{9981}; Tincurrin{9981}.
Gli-A2ah{9981}. v: Candeal de Nava del Rey (MCB-0892){9981}.
Gli-A2ai{9981}. v: Blanquillo (MCB-0908){9981}.
Gli-A2null{9984,9987}. v: Saratovskaya 29 (mutant){9987}.
Gli-B2{1607,1125}. [Gld 6B{1415}]. 6B{1607}.6BS{1122}. v: CS.
Gli-B2a{988}. v: CS{988}.
Gli-B2b{988}. v: Bezostaya 1{988}; Cobra{991}; Gladio{9986}; Sideral{991}.
Gli-B2c{988}. v: Courtot{991}; Escuala{9985}; Gabo{988}; Loreto{9986}; Manital{9986}; Prinqual{991}; Siete Cerros 66{988}; Sinton{995}; Yecora{9985}.
Gli-B2d{988}. v: Akmolinka 1{988}; Cesar{9981}; Friedland{991}; Tselinnaya 20{988}.
Gli-B2e{988}. v: Arsenal{991}; Veronese{9986}; Zlatna Dolina{994}.
Gli-B2f{988}. v: Basalt{9981}; Maris Freeman{988}; Master{991}.
Gli-B2g{988}. v: Capitole{991}; Cappelle-Desprez{991}; Galahad{988}; Forlani{9986}.
Gli-B2h{988}. v: Castan{991}; Mentana{9986}; Pane 247{9985}; Partizanka{994}; Sadovo 1{988}; Sistar{9986}.
Gli-B2i{988}. v: Insignia{988}; Robin{9981}.
Gli-B2j{988}. v: Farnese{9986}; Funo R250{9986}; Novosadska Rana 1{994}.
Gli-B2k{988}. v: Skala{988}.
Gli-B2l{988}. v: Clement{991}; Longbow{988}; Tracy{991}.
Gli-B2m{988}. v: Mironovskaya 808{988}; Open{991}; Renan{991}.
Gli-B2n{988}. v: Japhet{9981}; Rouge de Bordeau{9981}; Solo{988}.
Gli-B2o{988}. v: Hardi{9981}; Mara{9986}; Odesskaya 16{988}; Pippo{9986}; Rivoli{991}; Slavjanka{9981}.
Gli-B2p{988}. v: Pliska{983}; Champtal{991}; Oderzo{9986}; Recital{991}; Gazul{9985}.
Gli-B2q{988}. v: Saratovskaya 39{988}.
Gli-B2r{991}. v: Arminda{991}; Estica{991}; Genial{991}.
Gli-B2s{988}. v: Aquila{9981}; Saratovskaya 36{988}.
Gli-B2t{988}. v: Tselinogradka{988}.
Gli-B2u{988}. v: Kirgizskaya Yubileinaya{988}.
Gli-B2v{988}. v: Declic{991}; Garant{991}; Libellula{9986}; Mahissa 1{9985}; Poljarka{988}.
Gli-B2w{995,9986}. v: Palata{9986}; Pembina{995}; Rieti DIV{9986}.
Gli-B2x{994}. v: Super Zlatna (biotype){994}; Prostor{9981}; 251/83{9981}.
Gli-B2y{9986}. v: Centauro{9986}; E. Morandi{9986}.
Gli-B2z{9985}. v: Maestro{9985}.
Gli-B2aa{9986}. v: Salmone{9986}; E. Mottin{9981}.
Gli-B2ab{991}. v: Bordier{9981}; Orepi{991}.
Gli-B2ac{991}. v: Scipion{991}; Artaban{991}; Riol{991}; Lontra{9981}.
Gli-B2ad{991}. v: Champion{991}; Chopin{991}.
Gli-B2ae{991}. v: Priam{991}; Et.d'Choisy{991}; Campeador{9985}; Krajinka (biotype){994}.
Gli-B2af{9985}. v: Montjuich{9985}; Mocho Sobarriba{9985}.
Gli-B2ag{9981}. v: Jeja del Pais{9985}; Barbilla de Leon (MCB-1292){9981}.
Gli-B2ah{9981}. v: Rojo de Humanes (MCB-1262){9981}; Grano de Miracolo{9981}.
Gli-B2ai{9981}. v: Blanquillo (MCB-0908){9981}.
Gli-B2aj{9981}. v: Negrete de Malaga (MCB-1754){9981}.
Gli-B2ak{9981}. v: HY320{9981}; Leader{9981}.
Gli-B2al{9981}. v: Dankowska{991}.
Gli-B2am{9981}. v: TM-275{9981}; Uralochka{9981}.
Gli-B2an{9981}. v: Eagle{9981}; Glenwari{9981}.
Gli-B2ao{9981}. v: Olympic{9981}; Mokoan{9981}.
Gli-B2ap{9981}. v: Veda{9981}; Magnif 27{9981}.
Gli-B2aq{9981}. v: Winglen{9981}; Isis{9981}.
Gli-B2ar{9981}. v: Arbon{9981}; Roazon{9981}.
Gli-B2as{9981}. v: Strela{9981}; Sredneuralskaya{9981}.
Gli-B2at{9981}. v: Ranee{9981}; Javelin 48{9981}.
Gli-B2null{9984,9987}. v: Saratovskaya 29{9987}.
Gli-D2{1334,1125}. [Gld 6D{1415}]. 6D{1334}.6DS{1122}. v: CS.
Gli-D2a{988}. v: CS{988}; Maris Freeman{988}; Sistar{9986}; Tracy{991}.
Gli-D2b{988}. v: Bezostaya 1{988}; Cobra{991}; Farnese{9986}; Partizanka{994}.
Gli-D2c{988}. v: Escualo{9985}; Eridano{9986}; Rieti DIV{9986}; Siete Cerros 66{988}.
Gli-D2d{988}. v: Dneprovskaya 521{988}.
Gli-D2e{988}. v: Dollar{9985}; Lada{9981}; Mironovskaya 808{988}; Open{991}.
Gli-D2f{988}. v: Creneau{991}; Kirgizskaya Yubileinaya{988}; Rempart{991}.
Gli-D2g{988}. v: Capprelle-Desprez{991}; Futur{991}; Galahad{988}; Ghurka{988}; Mec{9986}.
Gli-D2h{988}. v: Capitole{991}; Chinook{995}; Eagle{00119}; Garant{991}; Sadovo 1{988}; Thatcher{995}.
Gli-D2i{988}. v: Insignia 49{00119}; Lario{9986}.
Gli-D2j{988}. v: Arcane{991}; Gallo{9986}; Gazul{9985}; Inia 66{9985}; Mentana{9986}.
Gli-D2k{988}. v: Crvencapa{944}; Kzyl-Bas{988}; Skala{988}.
Gli-D2l.
Omitted. No reliable differences compared to existing alleles {9981}.
Gli-D2m{988}. v: Marquis{988}; Rex{991}; Rinconada{9985}; Suneca{00119}; Veronese{9986}; Yecora{9985}.
Gli-D2n{988}. v: Castan{991}; Champlein{991}; Mahissa 1{9985}; Mercia{988}; Pippo{9986}.
Gli-D2o{988}. v: Omskaya 12{988}.
Note: cultivars Salmone and Resistente, which carry Gli-D2aa {9981}, were erroneously given as standards for allele Gli-D2o in {9986}.

Gli-D2p{988}. v: New Pusa {988}.
Gli-D2q{988}. v: Cook{9981}; E. Mottin{9981}; Fournil{991}; Volshebnitsa (biotype){988}; Winglen{9981}; Soissons{991}.
Gli-D2r{988}. v: Kremena{988}; Mara{9986}; Montcada{9985}.
Gli-D2s{988}. v: Akmolinka 1{988}; Bezenchukskaya 98{988}; Selkirk (biotype){995}.
Gli-D2t{9986}. v: Golia{9986}; Gabo{9981}; Manital{9986}; Bokal{9981}.
Gli-D2u{9986}. v: Loreto{9986}; Martial{991}; Cibalka{9981}.
Gli-D2v{991}. v: Epiroux{991}; Arbon{991}.
Gli-D2w{9985}. v: Navarro 150{9985}; Javelin{9981}; Hopps{9981}; Canaleja{9985}.
Gli-D2x{9985}. v: Montjuich{9985}; Blanquillo{9985}.
Gli-D2y{9985}. v: Candeal Alcala{9985}.
Gli-D2z{9985}. v: Aragon 03{9985}.
Gli-D2aa{9981}. v: Salmone{9981}; Resistente{9981}.
Gli-D2ab{9981}. v: Rojo de Boadilla de Campos (MCB-1031){9981}.
Gli-D2ac{9981}. v: Albatros{9981}.
Gli-D2ad{9981}. v: Hembrilla Soria (MCB-1298){9981}.
Gli-D2null{9984,9987}. v: Saratovskaya 29 (mutant){9987}.
Gli-Agi2{374}. 6Agi{374}. ad: Vilmorin 27/ Thin. intermedium.
Gli-R2{781}. [Sec 2{1336}]. 2R{781,1336}.2RS{1340}. ad: CS/Imperial{781,1336,1340}; Holdfast/King II{1340}.
Gli-R2a{03116}. d1{03116}. v: Carnac hexaploid triticale{03116}.
Gli-R2b{03116}. d2{03116}. v: Mostral hexaploid triticale{03116}.
Gli-R2c{03116}. t1{03116}. v: Alamo hexaploid triticale{03116}.
Gli-R2d{03116}. null{03116}. v: Triticor hexaploid triticale{03116}.
Gli-R2e{03115}. t2{03115}. v: Tornado hexaploid triticale{03115}.
Gli-Rm2{1339}. 6Rm{1339,1340}. ad: CS/S. montanum.
The location of Gli-R2 in S. cereale is thought to have evolved from S. montanum {1339} via a translocation between 2R and 6R {1530}.
Gli-Sl2{573}. 6Sl{573}. ad,su: CS/Ae. longissima.
Gli-U2{1335}. 6U{1335}. ad: CS/Ae. umbellulata.
Gli-V2{111}. 6VS{111}. ad: Creso/D. villosum.

Prior to the publication of {988}, allelic variation was demonstrated at all of the wheat Gli-2 loci, including 13 alleles at Gli-A2, 11 at Gli-B2, and 10 at Gli-D2, in a study of 39 cultivars {996}.
The Gli-2 alleles present in 57 Yugoslav wheat varieties were determined {994}.

74.3.2.3. Gli-3
A Gli-3 set of loci coding for omega-type gliadins are located 22 to 31cM proximal to Gli-1 on the short arms of group 1 chromosomes {422,1403,589}.
Gli-A3{1403,1119}. [Gld-2-1A{1416}]. 1AS{1403}. v: Bezostaya 1.
Each of the following Gli-A3 alleles, apart from Gli-A3d, which is a null, controls one minor omega-gliadin with molecular mass about 41k that occurs in the middle of the omega-region of APAGE fractionation. Gliadins controlled by these alleles differ in their electrophoretic mobility in APAGE in that the fastest of three known Gli-A3-gliadins is controlled by Gli-A3a and the slowest by Gli-A3c {9983}.
Gli-A3a{9983}. v: CS, Prinqual, Courtot, Tselinogradka, Bezenchukskaya 98.
Gli-A3b{9983}. v: Bezostaya 1.
Gli-A3c{9983}. v: Anda.
Gli-A3d{9983}. Null{9983}. v: Saratovskaya 210, Kharkovskaya 6, Richelle.
Gli-B3{422,1119}. [Gld-B6{422},Glu-B2{589}]. 1BS{422,589}. s: CS*/Thatcher{422}. v: Sicco{589}.
Gli-B3a{422,589,1119}. v: CS.
Gli-B3b{589}. v: Sicco.
Gli-B3c{422,1119}. s: CS*/Thatcher.
Gli-R3{164}. 1RS{164}. al: Four inbred lines (R2, J14, 8t, E2666).
Gli-Sl3{1228}. 1S1S{1228}. ad,su: CS/Ae. longissima. ma: In Ae. longissima 2/Ae. longissima 10, three gliadin loci, one glucose phosphate isomerase, and two glutenin loci were mapped relative to one another {1228} as follows: Glu-Sl1 15.9 cM - Gpi-Sl1 - 38 cM - Gli-Sl4 - 7.1 cM - Glu-Sl3 - 0.9 cM - Gli-Sl1 - 5.6 cM - Gli-Sl5. Glu-Sl1 is located in 1SlL and the other loci are in 1SlS.
Gli-V3{111}. 4VL{111}. ad: Creso/D. villosum.


74.3.2.4. Gli-4
It is not clear how Gli-Sl4 and Gli-Sl5 relate to the Gli-4 and Gli-5 sets described below.
Gli-A4{1205}. 1AS{1205}. v: Perzivan biotype 2.
A locus designated Gli-A4 controlling omega-gliadins was mapped at 10 cM proximal to Gli-A1 on the short arm of chromosome 1A {1205}.
However, Metakovsky et al. {9983} have since shown that this locus and Gli-A3 are, in fact, the same locus. Furthermore, Dubcovsky et al. {277} did not find evidence for the simultaneous presence of both Gli-A3 and Gli-A4 in five 1A or 1Am mapping populations and concluded that Gli-A4 should be considered to be Gli-A3 until conclusive evidence for the former is obtained. For these reasons, the locus Gli-A4 is deleted from the catalogue.


74.3.2.5. Gli-5
A locus designated Gli-5 controlling omega-gliadins was mapped to the short arms of chromosomes 1A and 1B, distal to Gli-1 {1147}. The map distance between Gli-B5 and Gli-B1 was estimated as 1.4 cM (recombination value of 1.4 +/- 0.4%), although there was significant variation in recombination values over crosses, ranging from 0 % to 5.9 % over the six crosses analysed. This variation was attributed to genotypic influence on the frequency of recombination.
Gli-A5{1147}. 1AS{1147}. v: Salmone.
Gli-A5a{9983}. Null{9983}. v: CS.
Gli-A5b{9983}. v: Marquis.
Allele Gli-A5b controls two slow-moving, easily-recognizable omega-gliadins. It is present in all common wheat cultivars having alleles Gli-A1m and Gli-A1r (and, probably, in those having Gli-A1e, Gli-A1l and Gli-A1q), because earlier (for example, in {988}) two minor omega-gliadins encoded by Gli-A5b were considered to be controlled by these Gli-A1 alleles {9983}.
Gli-B5{1147}. 1BS{1147}. v: Salmone.
Gli-B5a{1147}. v: CS.
Gli-B5b{1147}. v: Salmone.
In {988}, omega-gliadins controlled by Gli-B5 (allele Gli-B5b) were attributed to alleles at the Gli-B1 locus (alleles Gli-B1c, i, k, m, n and o).


74.3.2.6. Gli-6
Gli-A6{9983,993}. 1AS{9983}.
Gli-A6 was first explicitly described in {9983}, but it is now known that it was first observed without designation in {993}. There is strong evidence that it is distinct from Gli-A3 and Gli-A5, mapping distally to Gli-A1, with which it recombines at a frequency of 2-5%. Currently three alleles are known, of which Gli-A6c is particularly well-described in {9983}: the molecular mass of the omega-gliadin controlled by this allele is slightly lower than those of the omega-gliadins controlled by Gli-A3 alleles. In {988}, the omega-gliadin controlled by Gli-A6c was attributed to Gli-A1f. Gli-A6c is rather frequent in common wheat and may relate to dough quality (preliminary data {9983}). Gli-A6a is null {9983}.

Gli-A6a{9983}. Null{9983}. v: CS, Bezostaya 1.
Gli-A6b{9983}. v: Bezenchukskaya 98.
Gli-A6c{9983}. v: Courtot, Anda, Mironovskaya 808.


Four new classes of low molecular weight proteins related to gliadins, though not sufficiently similar to be classified as such, have been reported in {02113}. One of the classes has no close association to previously described wheat endosperm proteins.

74.3.3. Other endosperm storage proteins
Triticin proteins {1360} [Triplet proteins {1357}].
Storage globulins with homology to pea legumins and related proteins in oats, rice and several dicotyledonous species {1360}.
Tri-A1{1357},{1358}. 1AS{1357}. v: CS.
Tri-A1a. [cs{1358}]. v: CS.
Tri-A1b. [h{1358}]. v: Hope.
Tri-D1{1357},{707,1358}. 1DS{1357}. v: CS.
Tri-D1a. [cs{1358}]. v: CS.
Tri-D1b. [i{1358}]. v: India 115.



74.4. Enzyme Inhibitors

74.4.1. Trypsin inhibition
Ti-H1. [Itc 1{528}]. 3H{528}. ad: CS/Betzes.
Ti-R1. 3R{529}. ad: CS/Imperial.
Ti-A2{699}. 5AL{699}. v: CS.
Ti-B2{699}. 5BL{699}. v: CS.
Ti-D2{699}. 5DL{699}. v: CS.
Ti-D2a{699}. v: CS.
Ti-D2b{699}. v: Champlein.
Ti-D2c{699}. v: Synthetic.
Ti-Agi2{699}. 5Agi{699}. ad: Vilmorin 27/ Thin. intermedium.
Ti-Mt2{699}. 5Mt{699}. ad: CS/Ae. mutica.
Ti-R2{699}. 5RL{699}. ad: CS/Imperial. su: CS/King II.
Ti-Sl2{699}. 5SlL{699}. ad: CS/Ae. sharonensis.
Ti-U2{699}. 1U{699}. ad: CS/Ae. umbellulata.


74.4.2. Subtilisin inhibition
Si-R1{529}. 2R{529}.2RS{701}. ad: CS/Imperial, Holdfast/King II.
Si-H1{528}. [Isa 1{528}]. 2H{528}. ad: CS/Betzes.
Si-B2{701}. 1BS{701}. su: Bersee {Koga II}.
Si-D2{701}. 1DS{701}. v: Koga II.
Si-H2{528},{701}. [Ica 1{528},Ica 2{528}]. 1H{528}. ad: CS/Betzes.
Si-R2{529},{701}. 1R{529}.1RS{701}. ad: CS/Imperial{529}. tr: Gabo 1BL.1RS{701}.
Si-Sl2{701}. 1Sl{701}. ad: CS/Ae. longissima.
Si-U2{701}. 1U{701}. ad: CS/Ae. umbellulata.

Considerable genetic variation for Si-2 was noted in {701}. A chromosome location for Si-H2 on 1HL was inferred in {528} but questioned in {701}.
Three subunits of the wheat tetrameric inhibitor of insect a-amylase, CM1, CM3 and CM16, with homology to the dimeric and monomeric a-amylase inhibitors and the trypsin inhibitors, were located by Southern analysis of cDNAs pCT1, pCT2, and pCT3 to 4A, 4B, 4D; 7A, 7B, 7D; and 4A, 4B, 4D, respectively {427}.
Genes encoding proteins which inhibit the action of mammalian and insect, but not cereal, a-amylases, were located in chromosomes 3BS, 3DS and 6DS of Chinese Spring {1260}. Also, genes encoding inhibitors of insect a-amylases were located in H. chilense chromosomes 4Hch and 7Hch {1262}.

74.4.3. Inhibitors of alpha-amylase and subtilisin
Isa-A1{908}. 2AL{908}. v: CS.
Isa-A1a{908}. v: CS.
Isa-A1b{908}. Null allele. v: Cajeme 71.
Isa-B1{908}. 2BL{908}. v: CS.
Isa-B1a{908}. v: CS.
Isa-B1b{908}. v: Bihar.
Isa-D1{908}. 2DL{908}. v: CS.

Orthologous genes were identified in Ae. speltoides and T. timopheevii {908}. All durum wheats investigated had the genotype Isa-A1b, Isa-B1b.

74.4.4. Inhibitors (dimeric) of heterologous alpha-amylases
Chromosome 3BS has duplicated loci controlling two dimeric inhibitors of exogenous a-amylases, one known as 0.53 or Inh I {1260}, and the other as WDA I-3 {1260}. Chromosome 3DS has a homoeologous locus controlling a dimeric inhibitor of exogenous a-amylases, known as 0.19 or Inh III {1260,0124}, that is closely related to 0.53/Inh I. Intervarietal polymorphism for the WDA-3 protein was identified by isoelectric focussing of water-soluble endosperm proteins {0124}. This was interchromosomely mapped on 3BS using both a DH population of Cranbrook/Halberd, and a set of RILs of Opata 85/W-7984 (ITMI population) {0125}.
Iha-B1.1{1260}. 3BS{1260}. v: CS{1260}.
Iha-B1.2{0124}. 3BS{0124}. v: CS{0124}.
Iha-B1.2a{0124}. v: CS{0124,0125}.
Iha-B1.2b{0125}. Null allele. v: Cadoux{0125}; Cranbrook{0125}; Tasman{0125}.
Iha-D1{1260}. 3DS{1260}. v: CS{1260}.



74.5. Other proteins

74.5.1. Lipopurothionins
Pur-A1{351}. 1AL{351}. v: CS{351}.
A PCR marker specific for Pur-A1 was developed in {9976}.
Pur-B1{351}. 1BL{351}. v: CS{351}.
A PCR marker specific for Pur-B1 was developed in {9976}.
Pur-D1{351}. 1DL{351}. v: CS{351}.
A locus in chromosome 5DS affects the level of lipopurothionin {351}.
PCR marker specific for Pur-D1 was developed in {9976}.
Pur-R1. 1RL{1261}.=IRS,1BL. ad: CS/Imperial. su: Several 1R(1B) lines. tr: Aurora, Kavkaz.
A PCR marker specific for Pur-R1 was developed in {9976}.


74.5.2. Lectins
Lec-A1. 1AL{1427}. v: CS.
Lec-B1. 1B{1427}. s: CS*/Hope 1B.
Lec-D1. 1DL{1427}. v: CS.
Lec-U1. 1U{1427}. ad: CS/Ae. umbellulata.


74.5.3. Iodine binding factor
A monomeric water soluble protein from mature grain which preferentially binds iodine {818}.
Ibf-A1{818}. 5AL{818}. v: CS.
Ibf-A1a{818}. v: CS.
Ibf-A1b{818}. v: Cappelle-Desprez.
Ibf-A1c{818}. v: Hope.
Ibf-A1d{818}. v: Chris.
Ibf-A1e{818}. v: Sears' Synthetic.
Ibf-B1{818}. 5BL{818}. v: CS.
Ibf-B1a{818}. v: CS.
Ibf-B1b{818}. v: Cappelle-Desprez.
Ibf-B1c{818}. v: Ciano 67.
Ibf-B1d{818}. v: Sears' Synthetic.
Ibf-D1{818}. 5DL{818}. v: CS.
Ibf-D1a{818}. v: CS.
Ibf-D1b{818}. v: Cappelle-Desprez.
Ibf-D1c{818}. v: Purple Pericarp.
Ibf-D1d{818}. v: Sears' Synthetic.
Ibf-Agi1{818}. 5Agi{818}. ad: Vilmorin/Thin. intermedium.
Ibf-E1{818}. 5EL{818}. ad: CS/E. elongata.
Ibf-H1{818}. 4H{818}. ad: CS/Betzes.
Ibf-R1{818}. 5RL{818}. ad: CS/Imperial, CS/KingII.
Ibf-Sl1{818}. 5Sl{818}. ad: CS/Ae. sharonensis.
Ibf-U1{818}. 5U{818}. ad: CS/Ae. umbellulata.


74.5.4. Water soluble proteins
WSP-1 are monomeric grain endosperm proteins identified by their high pI's {817}.
Wsp-A1{817}. 7AL{817}. v: CS.
Wsp-A1a{817}. v: CS.
Wsp-A1b{817}. v: Huntsman.
Wsp-A1c{817}. v: Hope.
Wsp-A1d{817}. v: Sicco.
Wsp-A1e{817}. v: Condor.
Wsp-B1{817}. 7BL{817}. v: CS.
Wsp-B1a{817}. v: CS.
Wsp-B1b{817}. v: Hope.
Wsp-B1c{817}. v: Condor.
Wsp-D1{817}. 7DL{817}. v: CS.
Wsp-D1a{817}. v: CS.
Wsp-D1b{817}. v: Sears' Synthetic IPSR 1190903.
Wsp-D1c{893}. v: T4 = Agatha{893,890}; Indis{890,892}.
Wsp-E1{817}. 7E{817}. ad: CS/ E. elongata.
Wsp-H1{817}. 7H{817}. ad: CS/Betzes.
Wsp-Hch1{817}. 7Hch{817}. ad: CS/H. chilense.
Wsp-Sl1{817}. 7Sl{817}. ad: CS/Ae. sharonensis.
Wsp-V1{817}. 7V{817}. ad: CS/D. villosum.


74.5.5. Salt soluble globulins
GLO-1 are endosperm proteins (23-26 kDa) soluble in salt but not in water {455}.
Glo-A1{455}. 1AS{455}. v: CS. ma: Distally located: Glo-A1(distal) - 5.2cM - Gli-A1{1077}.
Glo-B1{455}. 1BS{455}. v: CS.
Glo-D1{455}. 1DS{455}. v: CS. ma: Distally located: Glo-D1(distal) - 2.9cM - Gli-D1{1077}.
Glo-E1{455}. 1ES{455}. ad: CS/E. elongata.
Glo-R1{455}. 1RS{455}. ad: CS/Imperial. su: 1B/(1R), eg., Salzmunde 14/44.


74.5.6. Waxy proteins
Waxy protein (granule-bound starch synthase = ADP glucose starch glycosyl transferase, EC 2.4 1.21 = GBSSI) is tightly bound within endosperm starch granules and is involved in the synthesis of amylose {1616}. Waxy variants, characterised by starch granules containing increased amylopectin and reduced amylose, are preferred for Japaness white salted or "udon" noodles {1650}. Similar waxy phenotypes are controlled by orthologous genes in barley, maize and rice but are not known to occur in rye {725}. All combinations of the null alleles were produced in Chinese Spring {0018}. Partial genomic clones of various diploid, tetraploid, and hexaploid wheats were sequenced {0278,0279}.
Wx-A1{180,1053}. [Xwx-7A{179,180},Wx-B1{1053,1054}]. 7AS{180,1053}. v: CS. ma: Variation in the microsatellite gene Xsun1-7A provides a co-dominant marker for this locus{0116}.
Wx-A1a{1054}. [Wx-B1a{1054}]. v: CS; Hoshuu.
Wx-A1b{1054}. [Wx-B1b{1054}]. Null allele. v: Kanto 79; Kanto 107 Shirodaruma{1617}; Sturdy{1617}. tv: Asrodur{0111}; MG826{03101}; A variant allele was present in one Iranian and one Italian accession{03101}.
Wx-A1c{1617}. v: QT105{1617}; WB6{1617}.
Wx-A1d{1616}. tv: T. dicoccoides KU 8937B{1616}.
Wx-A1e{1616}. tv: T. durum KU 3655 and KU 3659{1616}.
The complete genomic sequences for the Wx-A1a allele from CS {0073} and the cDNA sequence for the Wx-A1b allele from Kanto 107 {0075} were determined.
Wx-B1{180,1053}. [XWx-4B{179,180},XWx-4A{961},Wx-A1{1053,1054}]. 4AL{180,1054}. v: CS. tv: A variant allele was present in three accessions{03101}.
Wx-B1a{1054}. [Wx-A1a{1054}]. v: CS; Joshuu.
The complete genomic sequence for Wx-B1a from CS was determined {0073}.
Wx-B1b{1054}. [Wx-A1b{1054}]. Null allele. v: Kanto 79; Kanto 82; Kanto 107; Norin 98; Gabo{1617}; Satanta{1617}; For list of Australian wheats, see{1650}. tv: Blaquetta (BG-13701){0111}.
Wx-B1c{1617}. v: Cikataba{1617}; Junbuk 12{1617}.
Wx-B1d{1616}. tv: T. durum KU 4213D{1616}; KU 4224C{1616}.
Wx-B1e{0027}. v: Blue Boy II{0027}; Canthatch{0027}; Eureka{0027}; Gotz{0027}; Norin 44{0027}; Turkey Red{0027}.
Wx-B1f{0111}. tv: BG-12413{0111}; BG-12415{0111}.
Wx-D1{180,1053}. [XWx-7D{179,180}]. 7DS{180,1053}. v: CS.
Wx-D1a{1054}. v: CS.
Wx-D1b{1617}. Null allele. v: Bai Huo (Baihuomai){1617}. ma: DHWx12 {0117}; STS marker Xsun1-7D produces a distinct band of about 260bp (compared with the standard 840bp) , indicative of a smaller PCR product, but the gene is non-functional{0116,0117}; Xsun4(Wx)-7D is a perfect marker{0118}.
Wx-D1c{1617}. v: Scoutland{1617}.
Wx-D1d{0118}. v: K107wx1{0118}; EMS mutants{0118}; One Iranian and one Italian accession{03101}.
Wx-D1e{0117}. Null allele{0117}. v: NP150{0117}.
STS marker Xsun1-7D fails to produce a PCR product {0117}
Wx-D1f. [Wx-d1e{0234}]. v: Tanikei A6599-4{0234}; Relative to Kanto 107, Tanikei A6599-4 carries an alanine to threonine substitution at position 258 in the mature protein{0234}.
Various hard and soft wheats with the alleles Wx-A1b, Wx-B1b and Wx-D1b are listed in {0304}.

Lists of cutivars, lines and landraces of tetraploid and hexaploid wheats with different, mostly null, alleles at the Wx loci are given in {9910,9911,9912,1053,1054,9913,9915, 9916,1650,9917}.
The complete genomic sequence for Wx-D1a from CS {0073} and the cDNA sequence for the Wx-D1b allele from Bai Huo {0075} were determined.
Isolation of a wheat cDNA encoding Wx-A1 and Wx-D1 was reported in {0123} and {0167}, respectively. Isolation of genomic sequences for the genes encoding granule-bound starch synthase (GBSSI or Wx) in T. monococcum, Ae. speltoides and T. tauschii was reported in {0168}. Cloning of a second set of GBSSI or waxy genes, GBSSII, which were shown to be located on chromosomes 2AL, 2B and 2D, was reported in {0167}.
Various hard and soft wheats with the alleles Wx-A1b, Wx-B1b and Wx-D1b are listed in {0304}.

74.5.7. Starch granule proteins
The proteins, designated SGP-1, are starch synthases, encoded by SsII-A1, SsII-B1 and SsII-D1 {0042}.
Sgp-A1{1615}. 7AS{1615}. v: CS.
Sgp-A1a{1615}. v: CS.
Sgp-A1b{1615}. Null allele. v: Chosen 30, Chosen 57.
Sgp-A1c{1615}. v: Hua Non 9.
Sgp-B1{1615}. 7BS{1615}. v: CS.
Sgp-B1a{1615}. v: CS.
Sgp-B1b{1615}. Null allele. v: K79.
Sgp-B1c{1615}. v: Gnatruche.
Sgp-B1d{1615}. v: Waratah.
Sgp-D1{1615}. 7DS{1615}. v: CS.
Sgp-D1a{1615}. v: CS.
Sgp-D1b{1615}. Null allele. v: T116.
Sgp-A2{1615}. v: CS.
Sgp-B2{1615}. v: CS.
Sgp-D2{1615}. v: CS.
Sgp-A3{1615}. 7AS{1615}. v: CS.
Sgp-A3a{1615}. v: CS.
Sgp-A3b{1615}. Null allele. v: Norin 61.
Sgp-B3{1615}. 7BS{1615}. v: CS.
Sgp-B3a{1615}. v: CS.
Sgp-B3b{1615}. Null allele. v: Crest.
Sgp-B3c{1615}. v: Spica.
Sgp-D3{1615}. 7DS{1615}. v: CS.
The proteins, designated SGP-3, are identical to wheat starch synthase I, encoded by SsI-A1, SsI-A2 and SsI-D1 {0041}.

A triple null stock (SGP-1 null wheat) is reported in {0137}. Deletion mapping indicated that the gene order on the 7S arms is; centromere - Sgp-1 - Sgp-3 - Wx{1615}.

74.5.8. Puroindolines and grain softness protein
Puroindolines a and b are the major components of friabilin, a protein complex that is associated with grain texture (see 'Grain Hardness'). The name 'puroindoline' and the complete amino acid sequence of puroindoline a were given in {0382} from cv Camp Remy. Hard grain texture in hexaploid wheat results from unique changes in the puroindoline amino acid sequence or, currently, four null forms {0295} of the completely linked genes (max. map distance 4.3 cM) {452}. Tetraploid (AABB, AAGG) wheats lack puroindolines and are consequently very hard {03103}. A searchable database of wheat varieties and their puroindoline genotype is available at http://www.wsu.edu/~wwql/php/puroindoline.php. Grain softness protein-1 is a closely related gene which is closely located to the puroindoline genes {03111,1185}. 'GenBank' and 'dbEST' refer to sequence databases available at NCBI (also available throught EMBL and DDB).
Pina-A1{03103,03108,03104}. dv: T. urartu unspecified accession {03103}; TA763(GenBank AJ302094){03108,03104}; TA808(GenBank AJ302095){03108,03104}.
Pina-D1{452}. 5DS{452}. v: CS.
Pina-D1a{452}. v: Bellevue{0249}; Courtot{0249}; Fortuna{0249}; Galaxie{0249}; Heron{1035}; Soissons{0249}. v2: Aurelio Pinb-D1a{0249}; Bezostaja Pinb-D1b{0249}; Bilancia Pinb-D1a{0249}; Bolero Pinb-D1a{0249}; Brasilia Pinb-D1b{0249}; Centauro Pinb-D1a{0249}; Cerere Pinb-D1b{0249}; CS Pinb-D1a{452,0249}; Colfiorito Pinb-D1b{0249}; Cologna 21 Pinb-D1b{0249}; David Pinb-D1b{0249}; Democrat Pinb-D1b{0249}; Etruria Pinb-D1b{0249}; Francia Pinb-D1b{0249}; Gemini Pinb-D1b{0249}; Genio Pinb-D1b{0249}; Gladio Pinb-D1b{0249}; Lampo Pinb-D1a{0249}; Leone Pinb-D1a{0249}; Leopardo Pinb-D1a{0249}; Libero Pinb-D1a{0249}; Livio Pinb-D1a{0249}; Marberg Pinb-D1b{0249}; Mentana Pinb-D1a{0249}; Mieti Pinb-D1b{0249}; Mose Pinb-D1a{0249}; Neviana Pinb-D1a{0249}; Newana Pinb-D1b{0249}; Oscar Pinb-D1a{0249}; Pandas Pinb-D1b{0249}; Pascal Pinb-D1b{0249}; Penawawa Pinb-D1a{03104}; Sagittario Pinb-D1b{0249}; Salgemma Pinb-D1b{0249}; Saliente Pinb-D1b{0249}; Salmone Pinb-D1b{0249}; Serena Pinb-D1a{0249}; Serio Pinb-D1b{0249}; Veda Pinb-D1b{0249}; Zena Pinb-D1b{0249}. dv: Ae. tauschii upspecified accession (GenBank AJ249935){03103}; TA1583 (GenBank AY252029) Pinb-D1a, Gsp-D1b{03105}; TA2475 (GenBank AY252037) Pinb-D1i, Gsp-D1b{03105}; TA1599 (GenBank AY252011) Pinb-D1j, Gsp-D1g{03105}; TA1691 (GanBank AY252013) Pinb-D1j, Gsp-D1h{03105}.
Pina-D1a is present in all soft hexaploid wheats and possibly all hard hexaploid wheats that carry a hardness mutation in puroindoline b {452,1035,0082,0204,0295}.
Pina-D1b{1035}. Null allele. i: Falcon/7*Heron, Heron/7*Falcon{03109}; Gamenya sel.{0298,0203}; Heron/7*Falcon sel.{0298,0203}. v: Butte 86{1035}; Eridano{0249}; Falcon{1035}; Kalyansona{0249}; Super X{0249}; Yecora Rojo{0204}. v2: Amidon Pinb-D1a{0249}; Ciano Pinb-D1a{0249}; Dorico Pinb-D1a{0249}; Fortuna (USA) Pinb-D1a{0249}; Glenman Pinb-D1a{0249}; Golia Pinb-D1a{0249}; Guadalupe Pinb-D1a{0249}; Barra Pinb-D1a{0249}; Inia 66 Pinb-D1a{0249}; Indice Pinb-D1a{0249}; Jecora Pinb-D1a{0249}; Manital Pinb-D1a{0249}; Mendos Pinb-D1a{0249}; Padus Pinb-D1a{0249}; Prinqual Pinb-D1a{0249}; Sibilia Pinb-D1a{0249}.
Present only in some hard hexaploid wheats. Pina-D1b is associated with harder texture than Pinb-D1b {0177,0206}.
Pina-D1c{03105}. dv: Ae. tauschii TA2369 (GenBank AY252031) Pinb-D1h, Gsp-D1c; TA2527 (GenBank AY252015) Pinb-D1h, Gsp-D1e; TA2536 (GenBank AY251998) Pinb-D1i, Gsp-D1d{03105}.
Pina-D1d{03105}. dv: Ae. tauschii PI452131 (GenBank AJ302098) Pinb-D1i{03104}; PI554318 (GenBank AJ302099) Pinb-D1k{03104}; TA1649 (GenBank AY251963) Pinb-D1h, Gsp-D1f{03105}; TA2374 (GenBank AY251948) Pinb-D1i, Gsp-D1d{03105}; TA2521 (GenBank AY252042) Pinb-D1i, Gsp-D1e{03105}; TA2455 (GenBank AY252022) Pinb-D1i, Gsp-D1f{03105}.
Pina-D1e{03105}. dv: Ae. tauschii TA2458 (GenBank AY252034) Pinb-D1i, Gsp-D1d{03105}; TA2495 (GenBank AY252041) Pinb-D1i, Gsp-D1e{03105}.
Pina-D1f{03105}. dv: Ae. tauschii TA2436 (GenBank AY251998) Pinb-D1i, Gsp-D1d{03105}.
Pina-Am1{0083}. 5AmS{0083}. dv: T. monococcum DV92(cultivated), G3116 (spp. aegilopoides) (GenBank AJ242715){0083}; unspecified acession (GenBank AJ249933){03103}; PI277138 (GenBank AJ302093){03104}; PI418582 (GenBank AJ302092){03104}; T. monococcum spp. monococcum TA2025, TA2026, TA2037{03108}; T. monococcum spp. aegilopoides TA183, TA291, TA546, TA581{03108}.
In T. monococcum Pina-Am1 is completely linked to Gsp-Am1{0083}.
Pina-S1{03108}. dv: Ae. speltoides PI393494 (GenBank AJ302096){03104}; PI369616 (GenBank AJ302097){03104}; Ae. speltoides spp. speltoides TA2368, TA1789{03108}; Ae. speltoides spp. ligustica TA1777{03108}.
Pina-Sb1{03108}. dv: Ae. bicornis spp. typica TA1954, TA1942{03108}.
Pina-Sl1{03108}. dv: Ae. longissima spp. longissima TA1912{03108}; Ae. longissima spp. nova TA1921{03108}.
Pina-Ss1{03108}. dv: Ae. searsii TA1837, TA1355{03108}.
Pina-Ssh1{03108}. dv: Ae. sharonensis TA1999{03108}.
Pinb-A1{03108,03104}. dv: T. urartu TA763 (GenBank AJ302103){03104}; TA808 (GenBank AJ302104){03108,03104}.
Pinb-D1. 5DS{452}. v: CS{452}; Capitole (GenBank X69914){03110}.
Pinb-D1a{452}. v: Hill 81{452}. v2: Adder Pina-D1a{0317}; Amidon Pina-D1b{0249}; Aurelio Pina-D1a{0249}; Barra Pina-D1b{0249}; Bilancia Pina-D1a{0249}; Bolero Pina-D1a{0249}; Centauro Pina-D1a{0249}; CS Pina-D1a{452,0249}; Ciano Pina-D1b{0249}; Dorico Pina-D1b{0249}; Fortuna (USA) Pina-D1b{0249}; Glenman Pina-D1b{0249}; Golia Pina-D1b{0249}; Guadalupe Pina-D1b{0249}; Inia 66 Pina-D1b{0249}; Jecora Pina-D1b{0249}; Idice Pina-D1b{0249}; Karl Pina-D1a{0317}; Lampo Pina-D1a{0249}; Leone Pina-D1a{0249}; Leopardo Pina-D1a{0249}; Libero Pina-D1a{0249}; Livio Pina-D1a{0249}; Manital Pina-D1b{0249}; Mendos Pina-D1b{0249}; Mentana Pina-D1a{0249}; Mose Pina-D1a{0249}; Neviano Pina-D1a{0249}; Oscar Pina-D1a{0249}; Padus Pina-D1b{0249}; Penawawa Pina-D1a{03104}; Prinqual Pina-D1b{0249}; Serena Pina-D1a{0249}; Sibilia Pina-D1b{0249}; Sigyn II Pina-D1a{0317}. dv: Ae. tauschii unspecified accession (GenBank AJ249936){03103}; TA1583 (GenBank AY251981) Pina-D1a, Gsp-D1b{03105}.
Pinb-D1a is present in all soft hexaploid wheats and possibly all hard hexaploid wheats carrying the Pinb-D1b, -D1c, -D1d, -D1e, or -D1f mutations {452,1035,0082,0204,0295}.
Pinb-D1b{452}. 5DS{452}. v: Thatcher{0204}; Wanser{452}; hard component of Turkey{0204}. i: Paha*2/Early Blackhull/5*Paha{0298,0203}; Early Blackhull der./5*Nugaines sel.{0298,0203}; hard sib sel. from Weston{03107}. s: CS*7/Cheyenne 5D{452}. v2: Bastion Pina-D1a{0317}; Bezostaya Pina-D1a{0249}; Brasilia Pina-D1a{0249}; Cerere Pina-D1a{0249}; Colfiorito Pina-D1a{0249}; Cologna 21 Pina-D1a{0249}; David Pina-D1a{0249}; Democrat Pina-D1a{0249}; Etruria Pina-D1a{0249}; Francia Pina-D1a{0249}; Gemini Pina-D1a{0249}; Genio Pina-D1a{0249}; Gladio Pina-D1a{0249}; Marberg Pina-D1a{0249}; Mieti Pina-D1a{0249}; Newana Pina-D1a{0249}; Pandas Pina-D1a{0249}; Pascal Pina-D1a{0249}; Sagittario Pina-D1a{0249}; Salgemma Pina-D1a{0249}; Saliente Pina-D1a{0249}; Salmone Pina-D1a{0249}; Serio Pina-D1a{0249}; Veda Pina-D1a{0249}; Zena Pina-D1a{0249}.
Pinb-D1b is a "loss-of-function" mutation resulting from the replacement of a glycine by a serine at position 46 {452}.
Pinb-D1c{0082}. v: Avle{0082}; Bjorke{0082}; Portal{0082}; Reno{0082}; Tjalve{0082}.
Pinb-D1c is a "loss-of-function" mutation resulting from the replacement of a leucine by a proline at position 60 {0082}.
Pinb-D1d{0082}. v: Bercy{0082}; Mjolner{0082}.
Pinb-D1d is a "loss-of-functgion" mutation resulting from the replacement of a tryptophan by a arginine at position 44 {0082}.
Pinb-D1e{0204}. v: Gehun{0204}; Canadian Red{0204}; Chiefkan{0204}.
Pinb-D1e is a "loss-of-function" mutation resulting from the replacement of a tryptophan by a stop codon at position 39 {0204}.
Pinb-D1f{0204}. v: the hard component of Utac{0204}.
Pinb-D1f is a "loss-of-function" mutation resulting from the replacement of a tryptophan by a stop codon at position 44 {0204}.
Pinb-D1g{0204}. v: Andrews{0204}.
Pinb-D1g is a "loss-of-function" mutation resulting from the replacement of a cysteine by a stop codon at position 56 {0204}.
Pinb-D1h{03105}. dv: Ae. tauschii TA2369 Pina-D1c, Gsp-D1c{03105}; TA2527 Pina-D1c, Gsp-D1e{03105}; TA1649 Pina-D1d, Gsp-D1f{03105}.
Pinb-D1i{03105}. dv: Ae. tauschii TA2475 (GenBank AY251989) Pina-D1a, Gsp-D1b{03105}; TA2536 (GenBank AY251993) Pina-D1c, Gsp-D1d{03105}; TA2374 (GenBank AY251948) Pina-D1d, Gsp-D1d{03105}; TA2512 (GenBank AY251992) Pina-D1d, Gsp-D1e{03105}; TA2455 (GenBank AY251972) Pina-D1d, Gsp-D1f{03105}; TA2458 (GenBank AY251986) Pina-D1e, Gsp-D1d{03105}; TA2495 (GenBank AY251991) Pina-D1e, Gsp-D1e; TA2436 Pina-D1f, Gsp-D1d{03105}.
Pinb-D1j{03105}. dv: Ae. tauschii TA1599 Pina-D1a, Gsp-D1g{03105}; TA1691 Pina-D1a, Gsp-D1h{03105}.
Pinb-D1k. dv: Ae. tauschii PI554318 (GenBank AJ302108) Pina-D1d{03104}.
Pinb-Am1{0083}. 5AmS{0083}. dv: T. monococcum DV92 (cultivated), G3116 (spp. aegilopoides (GenBank AJ242716){0083}; unspecified accession (GenBank AJ249934){03103}; is identical to allele Pina-D1h{03105}; PI277138 (GenBank AJ302102){03104}; PI418582 (GenBank AJ302101){03104}.
In T. monococcum Pinb-Am1 is 0.1 cM proximal to Pina-Am1 and both loci are less than 36 kb apart {0083}.
Pinb-S1{03108}. dv: Ae. speltoides PI393494 (GenBank AJ302105){03104}; PI369616 (GenBank AJ302106){03104}; Ae. speltoides spp. speltoides TA2368, TA1789{03108}; Ae. speltoides spp. ligustica TA1777{03108}.
Pinb-Sb1{03108}. dv: Ae. bicornis spp. typica TA1954, TA1942{03105}.
Pinb-Sl1{03108}. dv: Ae. longissima spp. longissima TA1912{03108}; Ae. longissima spp. nova TA1921{03108}.
Pinb-Ss1{03108}. dv: Ae. searsii TA1837, TA1355{03105}.
Pinb-Ssh1{03108}. dv: Ae. sharonensis TA1999{03105}.

Pinb-D1b, Pinb-D1c, Pinb-D1d, Pinb-D1e, Pinb-D1f,
or Pinb-D1g are present in hard hexaploid wheats not carrying the Pina-D1b (null) mutation {452,1035,0082,0204}.

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

Genotypes for a selection of North American wheats are given in {0204}.

74.5.9. Grain softness protein
Gsp-1{1185}.
Gsp-A1{614}. [GSP{614}]. 5A{614,0383}. v: CS{614,0383}; Rosella (GenBank AF177218){0383}.
In {1185} partial-sequence clone TSF61 from cv Soft Falcon (GenBank X80380) is identical to this allele.
Gsp-B1{614}. [GSP{614}]. 5B{614}. v: CS{614}; Glenlea{0385}.
In {1185} sequence of clone TSF33 from cv Soft Falcon (GenBank X80379) is identical to this allele, as are ESTs for cv CS (dbEST BJ235798) and cv CNN (dbEST BE423845).
Gsp-D1{614}. [GSP{614}]. 5DS{614}. v: CS{614}; Glenlea{0385}. dv: Ae. tauschii CPI1110799 (GenBank AF177219){0383}. ma: Co-segregation of Gsp-D1 and Ha{614}.
In {1185} sequence of clone TSF69 from cv Soft Falcon (GenBank S72696) is identical, as are ESTs for cv CS (dbEST BJ237450) and cv CNN (dbEST BE422565).
Gsp-D1b{03105}. dv: Ae. tauschii TA1583 (GenBank AY252079) Pina-D1a, Pinb-D1a{03105}; TA2475 (GenBank AY252087) Pina-D1a, Pina-D1i{03105}.
Gsp-D1c{03105}. dv: Ae. tauschii TA2369 (GenBank AY252081) Pina-D1c, Pinb-D1h{03105}; CPI110799 (GenBank AF177219){0383}.
Gsp-D1d. dv: Ae. tauschii TA2536 (GenBank 252093) Pina-D1c, Pinb-D2i{03105}; TA2374 (GenBank AY252046) Pina-D1d, Pinb-D1i{03105}; TA2458 (GenBank AY252084) Pina-D1e, Pinb-D1i{03105}; TA2436 (GenBank AY252048) Pina-D1f, Pinb-D1i{03105}.
Gsp-D1e. dv: Ae. tauschii TA2527 (GenBank AY252066) Pina-D1c, Pinb-D1h{03105}; TA2512 (GenBank AY252092) Pina-D1d, Pinb-D1i{03105}; TA2495 (GenBank AY252091) Pina-D1e, Pinb-D1i{03105}.
Gsp-D1f. dv: Ae. tauschii TA1649 (GenBank AY252063) Pina-D1d, Pinb-D1h{03105}; TA2455 (GenBank AY252073) Pina-D1d, Pinb-D1i{03105}.
Gsp-D1g. dv: Ae. tauschii TA1599 (GenBank AY252062) Pina-D1a, Pinb-D1j{03105}.
Gsp-D1h. dv: Ae. tauschii TA1691 (GenBank AY252064) Pina-D1a, Pinb-D1j{03105}.


74.5.10. Starch synthase
SsI-A1{0041}. 7A{0041}.
SsI-B1{0041}. 7B{0041}.
SsI-D1{0041}. 7D{0041}.
Starch synthase I proteins are identical to the starch granule proteins SGP-3 {0041}.
SsII-A1{0042}. 7A{0042}.
SsII-B1{0042}. 7B{0042}.
SsII-D1{0042}. 7D{0042}.
Starch synthase II proteins are identical to the starch granule proteins SGP-1 {0042}.


74.5.11. Histone H1 Proteins
HstH1-A1{0215}. 5AL{0215}. v: CS{0215}.
HstH1-B1{0215}. 5BL{0215}. v: CS{0215}.
HstH1-D1{0215}. 5DL{0215}. v: CS{0215}.
HstH1-D1a{0215}. v: CS{0215}; 18 others{0215}.
HstH1-D1b{0215}. v: Grekum 114{0215}; Kirgizsky Karlik{0215}.
HstH1-A2{0215}. 5AL{0215}. v: CS{0215}.
HstH1-A2a{0215}. v: CS{0215}.
HstH1-A2b{0215}. Null allele{0215}. v: Mara{0215}; 10 others{0215}.
HstH1-B2{0215}. 5BL{0215}. v: CS{0215}.
HstH1-B2a{0215}. v: CS{0215}; 19 others{0215}.
HstH1-B2b{0215}. v: Excelsior{0215}.
HstH1-D2{0215}. 5DL{0215}. v: CS{0215}.

The relationship of this gene series with a Hst-A1, Hst-B1, Hst-D1 series in group 5 chromosomes {0216} based on DNA hybridization studies was not established.



75.Reaction to Barley Yellow Dwarf Virus
Bdv1{1363,1379}. 7D{1379}.7DS{1363}. i: Jupeteco 73R (compared to Jupeteco 73S){1363}. v: Anza{1379}; Condor BW3991{1379}; Tyrant BW3872{1379}; Hahn BW4097{1379}; Parrot BW10817{1379}; Siren BW18643{1379}; Many CIMMYT genotypes. Bdv1 is completely linked with Ltn, Lr34 and Yr18. See Ltn, Lr34, Yr18.
Note: BW = CIMMYT wheat accession number.
Bdv2{058}. v: TC14*2/Hartog{0225}; TC14*2/Spear{0201}; TC14*2/Tatiara{0225}.
7Ai#1S{552}.su:TAF2{059}; Lines 5395 & 5395-243AA{552}.
7DL= T7DS.7DL-7Ai#1L{552,0182}.tr:TC14{059,0201}; H960642{0182}.
7D = T7DS-7Ai#1S.7Ai#1L{552}.tr:TC5, TC6, TC8, TC9, TC10{059}.
1B = T1BS-7A#1S.7Ai#1L{552}.tr:TC7{447}. ma: Distal 10% of 7DL, translocation point between RFLP markers Xpsr680 and Xpsr965{0182}; Complete association with Xpsr129-7D, Xpsr548-7D, XksuD2-7D, XcslH81-7D, and Xgwm37-7D selected as a diagnostic marker{0225}.


76.Reaction to Blumeria graminis DC.
Disease: Powdery Mildew.

76.1. Designated genes for resistance
Note: Chancellor, used as a susceptible genetic background, for some near-isogenic lines probably carries Pm10 and Pm15 {1479}.
Pm1.
Pm1a{562}. [Pm1{130},Mlt{1175},Mla{348}]. 7A{1293}.7AL{1305}. i: Axminster/8*Chancellor{132}; CI 14114 = As II/8*Chancellor{132}; CI 13836/8*Chancellor{132}; Kenya C6041/5*Federation{1168}; Norka/8*Chancellor{132}. s: CS*5/Axminster 7A{1293}. v: Anfield{098}; As II{130}; Axminster{130,1175}; Birdproof{165}; Bonus{1554}; CI 13836{130}; Converse{1175}; Fedka{939}; Festival{1554}; Fram I{130}; Huron CI 3315{1175,1554}; Kenora{1554}; Kenya W744 = C6041{130,1175}; Norka{130,1175}; Pika{130}; Sweden W1230{1554}; Thew{1175}; TU 4{130}; Zhengzhou 871124{570}. v2: Anfield Pm9{1287}; BGRC 44514 Pm3a{1628}; Mephisto Pm2 Pm9{540}; Normandie Pm2 Pm9{165}; Pompe Pm9{1287}; Ring Pm9{1287}; Sappo Pm2 Pm4b (Carries Lr20){310}; Solo Pm2 Pm4b{052}. ma: Co-seg. with Xcdo347-7A using NILs{864}; Co-segregation or close linkage with three RAPDs; one RAPD converted to a STS{570}; Note: In Solo, Pm1 is translocated to chromosome 7D{052}; Complete cosegregation of several markers including Xcdo347-7A, Xpsr121-7A, Xpsr680-7A, Xpsr687-7A, Xbzh232(Tha)-7A, Xrgc607-7A and Xsts638-7A with Pm1 and Lr20 was reported in {0323}.
Pm1b{562}. v: MocZlatka{562}.
Pm1c{562}. [Pm18{853,562}]. v: Blaukorn{0011}; M1N{1628,562}; In{540}; M1N was described as an undesignated subline of Weihenstephan M1. ma: AFLP marker 18M2 was diagnostic for Pm1c{0011}.
Pm1d{562}. v: T. spelta var duhamelianum TRI2258{562}. ma: AFLP marker 18M1 - various Pm1 alleles 0.9 cM{0011}.
Pm1e{0322}. [Pm22{1134}]. v: Elia{1134}; Est Mottin{1134}; Ovest{1134}; Tudest{1134}; Virest{1134}.
Pm2{130}. [Mlu{1175},Mlx{1088}]. 5D{1007}.5DS{945}. i: CI 14118 = Ulka/8*Chancellor{132}; CI 14119 = CI 12632/8*Chancellor{132}; Federation*4 /Ulka{1168}. v: Avalon{096}; Bounty{096}; Fenman{096}; Galahad{1531}; H8810/47{130}; Longbow{1531}; Maris Beacon{1592}; Maris Nimrod{1592}; Maris Sportsman{096}; Maris Templar{1592}; Norman{096}; Orestis{1079}; PI 92378{1168}; PI 181374{1168}; Sea Island{130}; Sentry{096}; S2303{945}; Synthetic(Iumillo/T. tauschii){1168}; TP 114/2*Starke deriv{626}; Ulka{130,1175}; XX186 = T. durum Santa Maria/Aeg. squarrosa BGRC 1458 Pm19{853}. v2: Apollo Pm4b Pm8{541}; Brigand Pm6{096}; Brimstone Pm6{1531}; CI 12632 Pm6{130}; CI 12633 Pm6{133}; Compal Pm4b{854}; Crossbow Pm5 Pm6{098}; Gawain Pm6 {1531}; Halle Stamm 13471 Mld{097}; Heiduck Pm6{541}; Hustler Pm6 {096}; Hornet Pm8{1531}; Kinsman Pm6{096}; Mardler Pm6{096}; Maris Dove Mld{1592}; Maris Fundin Pm6{096}; Maris Huntsman Pm6{1592}; Mephisto Pm1 Pm9{540}; Normandie Pm1 Pm9{165}; Parade Pm5 Pm6 {1531}; Rendezvous Pm4b Pm6 {1531}; Solo Pm1 Pm4b {052}; Timmo Pm4b{096}; TP 114 Pm6{626}; Virtue Pm6{096}; Walter Pm4b Pm6 {1428}. dv: Aeg. squarrosa BGRC 1458{853}; Forty accessions of T. tauschii{852}. ma: Pm2 - 3.5cM - Xbcd1871-5D using F2s{864}.
Pm3. 1A. ma: Xgdm33-1A - 2.3 cM - Pm3/Xpsp2999-1A{0313}.
Genotype list: {0313}
Pm3a{130,132}. [Mla{1168}]. 1A{1007}.1AS{943,947}. i: Asosan/8*Chancellor{132}; = CI 14120; Asosan/3*Federation{1168}. v: Asosan{130,1168}; BGRC 44514 Pm1a{1628}; Coker 797{786}; Florida 301{786}; Florida 302{786}; Hadden{097}; Halle Stamm{097}; Norin 3{1134}; Norin 29{1134}; PI 46890{1439}; Saluda{786}; Tyler{1419}. ma: Xbcd1434-1A - 1.3 cM - Pm3 using NILs{864}; Xwhs179-1A - 3.3 cM - Pm3{522}.
Pm3b{130,132}. [Mlc{165}]. 1A{1007}. i: Chul*8/Chancellor{132}; T. sphaerococcum*8/Chancellor = CI 15887{539}. v: Chul{165}. ma: Xbcd1434-1A - 1.3 cM - Pm3b using NILs{864}.
Pm3c{130,132}. [Mls{1175}]. 1A{134,1007}. i: Sonora/8*Chancellor{132}; = CI 14122; Sonora/4* Federation{1168}; Triticale/8*Chancellor{539}. s: CS*7/Indian 1A{134}. v: Borenos{854}; Cawnpore{1628}; CI 3008{130}; CI 4546{130}; Hindukusch{1628}; Indian{1175}; Sonora{130,1168}; Sturgeon{1175}.
Pm3d{1628}. [Ml-k{540},Mlk{434}]. 1A{1628}. v: Axona{0313}; Cornette{0313}; Herold{540}; Indian 4{0313}; Kadett{0313}; Kanzler{0011}; Kleiber{0313}; Kolibri{540,542,1628}; Ralle{540}; Socrates {heterogeneous}{540}; Star {heterogeneous}{540}; Syros{540}. v2: Kadett Pm4b{540}; Turbo Pm4b{540}.
Pm3e{1628}. v: Sydney University Accession W150 =AUS 6449{939,1628}.
Pm3f{1628}. i: Michigan Amber/8*Chancellor{1628}; This allele was distinguished from Pm3c with only one of 13 pathogen cultures.
Pm3g{0070}. [Mlar{854}]. 1A{0070}.1AS{0363}. v: Avo{1629}; Aristide{1629}; Champetre{0313}; Courtot{1629}; Lutin{0313}; Oradian{0313}; Rubens{0313}; Soissons{0313}; Valois{0313}. ma: Pm3g - 5.2 cM - Gli-A5 - 1.9 cM - Gli-A1{0070}; Pm3g was completely linked to microsatellite Xpsp2999{0363}.
Pm4{131}.
Pm4a{1464}. [Pm4{131}]. 2AL{1464}. i: CI 14123 = Khapli/8*Chancellor{131}; CI 14124 = Yuma/8*Chancellor{131}. v: Steinwedel*2/Khapli{939}. tv: Khapli{131}; Valgerado{097}; Yuma{131}. ma: Co-seg with Xbcd1231-2A.2 & Xcdo678-2A using F2s{864}; Xbcd1231-2A.1 - 1.5cM - Pm4 - 1.56cM - Xbcd292-2A{864}; Pm4a - 3.5cM - AFLP markers 4aM1 and 4aM2{0011}; Xbcd1231 was converted to a STS marker{0069}.
Pm4b{1464}. [Mle{1591}]. 2A{052}.2AL{1464}. i: Federation*7/T. carthlicum W804{1464}. v: Achill{540}; Ajax{540}; Arkas{540}; Armada{096}; Atlantis{0011}; Boheme{0011}; Botri (heterogeneous){854}; ELS{1591}; Facta{854}; Factor (heterogeneous){854}; Fakon{854}; Fazit{854}; Hermes{540}; Horizont{540}; Maris Halberd; Max{540}; Olymp{540}; Orbis{540}; RE714{1220}; Renan{0016}; Ronos{1079}; S-25{052}; S-28{052}; TP 229{626,1591}; Weihenstephan M1{1591}; VPM1{097}. v2: Apollo Pm2 Pm8{541,802}; Boxer Pm5{541}; Compal Pm2{854}; Kadett Pm3d{540}; Kronjuwel Pm8{541}; Mission Pm5 {78,541,1531}; Rang Pm1{052}; Rendezvous Pm2 Pm6{1531}; Solo Pm1 Pm2{052,540}; Sorbas Pm6{541}; Timmo Pm2 Pm6{096}; Turbo Pm3d{540}; Walter Pm2 Pm6{1428}. ma: Pm4b - 4.8cM - Xgbx3119b{0272}; Xgwm382-2A - +/- 10 cM - Pm4b - +/- 2cM - XgbxG303{0354}.
Pm5.
Pm5a{0257}.Recessive. [Pm5{787},mlH{771}]. 7B{964}.7BL{771}. i: Hope/8* Chancellor = CI 14125{570}. s: CS*6/Hope 7B{771,964}. v: Alidos{854}; Aotea{964}; Caldwell{786}; Ga 1123{786}; Galaxie{0257}; Glenwari{964}; Hardired{786}; Hope{964}; H-44{964}; Kontrast{854}; Kormoran{1079}; Kutulukskaya{0257}; Lambros{0257}; Lawrence{964}; Navid{0257}; Pagode{0257}; Redcoat{097}; Redman{964}; Regina{0257}; Renown{964}; Selpek{540}; Sicco{096,0257}; Spica{964}; Tarasque{0257}; Warigo{964}; Zolotistaya{0257}. v2: Arthur Pm6{786}; Coker 983 Pm6{786}; Double Crop Pm6{786}; Granada Pm8{541}; Sensor Pm8{541}.
Pm5a was transferred to hexaploid wheat from T. dicoccum via Hope and H-44.
Pm5b{0257}. [Mli{540,558}]. Mli{540,558}. v: Aquila{096,541}; Carimulti{541}; Cariplus{541}; Cucurova{0257}; Dolomit{541}; Falke{541}; Flanders{096}; Fruhprobst{0257}; Ilona{0257}; Ibis{096}; Kirkpinar-79{0257}; Kontrast{0257}; Kormoran{541}; Krata{541}; Markant{541}; Mercia{1531}; Milan{541}; Nadadores{0257}; Reiher{541}; Rektor{541}; Rothwell Perdix{096}; Siete Cerros{0257}; Severin{541}; Sicco{096}; Sperber{541}; Tukan{541}; Una{0257}; Urban{541}; Wattines{541}; Wettiness{0257}. v2: Bert Pm6{541}; Boxer Pm4b{541}; Crossbow Pm2 Pm6{098}; Kristall Pm8{541}; Mission Pm4b{541,1531}; Parade Pm2 Pm6{1531}.
Pm5c{0257}. 7B{0257}. v: T. sphaerococcum cv. Kolandi{0257}.
Pm5d{0257}. 7B{0257}. i: IGV 1-455 = CI 10904/7*Prins{0257}; CI 10904/7*Starke{0257}.
Pm5e{0258}.Recessive and hemizygous effective {0258}. [mlfz{0259}]. v: Fuzhuang 30{0258}. ma: Xgwm1267-7B - 6.6cM - Pm5e - 12.6cM - Xubc405628{0258}.
Pm6{627}. [Mlf{626}]. 2B{1088}. i: CI 13250/7*Prins{0069}; CI 12559/8*Prins{0069}; PI 170914/7*Prins 6 NILs based on Prins {0139}.{0069}. v: 1969 IVGS Line C{626}; Abe{097,1256}; Coker747{786,1079}; Mengavi{097}; Oasis{786}; Timgalen{098}; TP 114/2*Starke deriv. B{626}. v2: Arthur Pm5a{097,786}; Brigand Pm2{096}; Brimstone Pm2{1531}; CI 12632 Pm2{626,1088}; CI 12633 Pm2{626,1088}; Crossbow Pm2 Pm5{098}; Double Crop Pm5a{786}; Garwain Pm2{1531}; Greif Pm5a{0011}; Heiduck Pm2{541}; Hustler Pm2{096}; Kinsman Pm2{096}; Mardler Pm2{096}; Maris Fundin Pm2{096}; Maris Huntsman Pm2{1592}; Parade Pm2 Pm5{1531}; Rendezvous Pm2 Pm4b{1531}; Sorbas Pm4b{541}; Timmo Pm2 Pm4b{096}; TP 114 Pm2{626}; Virtue Pm2{096}; Walter Pm2 Pm4b{1428}. ma: Close linkage with Xbcd135(1.5+-1.4cM), Xbcd307(4.7+-2.5cM) and Xbcd266(4.5+-2.4cM){0069}; Mapped to the interval Xbcd35-2B-Xpsr934-2B{0139}; However, the fact that Timgalen and a 'CI12632/Cc' line lacked the critical T. timopheevii markers {0139} is cause for concern.
Pm7. Derived from S. cereale cv. Rosen. 4BL{270,271,389}=T4BS.4BL-5RL{543}, but more recently revised to T4BS.4BL-2R#IL{380,389}. i: = Transec/8*Chancellor. v: Transfed{269}; Transec{273}.
Pm8. Derived from Petkus rye - see Yr9, Lr26, Sr31. 1R{1B}.1BL.1RS. v: Corinthian{1531}; Dauntless{1531}; Ambassador{1531}; Disponent{541}; GR876{753}; Halle Stamm{097}; Hammer{098}; Others{1208}; ST1-25{201}; Slejpner{1531}; Stetson{1531}; Stuart{096}. v2: Apollo Pm2 Pm4b{541}; Granada Pm5{541}; Hornet Pm2{1531}; Kristall Pm5{541}; Kronjuwel Pm4b{541}; Sensor Pm5{541}. tv: Cando*2/Veery=KS91WGRC14{381}.
1BS/1RS recombinants 2.9 cM proximal to Gli-B1/GluB3{0084}.i:MA1 and MA2, four-breakpoint double translocation lines 1RS-1BS-1RS-1BS. 1BL in Pavon{0084}.
Crosses between three lines with Pm8 and Helami-105, a 1BL.1RS line with Pm17, indicated that Pm8 and Pm17 were allelic{524}. Earlier, these genes were reported to be genetically independent{1463}. A STS marker distinguished Pm17 from Pm8{0286}.
Pm9{347}. 7A{347}.7AL. v: N14{562}. v2: Anfield Pm1a{1287}; Mephisto Pm1a Pm2{540}; Normandie Pm1a Pm2{347}; Pompe Pm1a{1287}; Ring Pm1a{1287}.
Pm10{1482}. 1D{1482}. v: Norin 4{1482}; Norin 26{1482}; Norin 29{1482}; Penjamo 62{1482}; Shin-chunaga{1482}. v2: T. spelta duhamelianum Pm11{1481}.
Pm10 was detected using a culture derived from a hybrid of B. g. tritici and B. g. agropyri.
Pm11{1481}. 6BS{1481}. v: Chinese Spring{1481}; Salmon{1481}; T. compactum No. 44{1481}. v2: T. spelta duhamelianum Pm10{1481}.
Pm11 was detected using a culture derived from a hybrid of B. g. tritici and B. g. agropyri
Pm12{1017}. Derived from Ae. speltoides.
The earlier location of 6A {1017} was not correct. 6B = 6BS-6SS.6SL{598,572}. v: Wembley*6/Ae. speltoides, #31{1017,598}; 6S1S{598}. al: Ae. speltoides CL214008 = K{1017}. ma: Pm12 was mapped to a translocated segment proximal to Xpsr551-6B{598}.
Pm13. Derived from Ae. longissima
3B{173} = T3BL.3BS-3S1#1S{389}.v:R1A{174}; R1B{0055}; R4A{0055}; R6A{0055}.
3D{173} = T3DL.3DS-3S1#1S{389}.tv:R1D{174}. v: R2A{0055}; R2B{0055}.
3S1#1S.al:Ae. longissima. ma: STS marker Xutv13{0036}; several other markers located in introgressed segments{0036}; Pm13 was mapped to a translocated 3S1S segment distal to Xcdo-460-3B{0036}.
Pm14. 6B{1478}. v2: Akabozu Pm15{1478}; Kokeshikomugi Pm15{1478}; Norin 10 Pm15{1478}.
Pm14 and Pm15 were detected using hybrids between B. g. tritici and B. g. agropyri cultures.
Pm15. 7DS{1478}. v2: Akabozu Pm14{1478}; Chinese Spring Pm11{1478}; Kokeshikomugi Pm14{1478}; Norin 4 Pm10{1478}; Norin 10 Pm14{1478}; Norin 26 Pm10{1478}; Shinchunaga Pm10{1478}; T. macha subletschumicum Pm10{1478}; T. compactum No. 44 Pm11{1478}.
Pm14 and Pm15 were detected using hybrids between B. g. tritici and B. g. agropyri cultures.
Pm16{1201}. 4A{1201}. v: Norman lines with resistance from T. dicoccoides CL1060025{1201}. tv: T. dicoccoides CL1060025{1201}.
Pm17{097,838,544}.
1AS = T1AL.1R#2S{1624,185,389}.v:Amigo{561}; Century{216}; Nekota{0021}; Neobrara{0021}; TAM107{216}; TAM200{216}; TAM201{216}; TAM202{0021}.
1BS = T1BL.1R#2S{561}.v2:Helami 105 Pm5{561}. ma: A STS marker distinguished Pm17 from Pm8{0286}.
Pm8 and Pm17 were reported to be allelic {524}, see note under Pm8.
Pm18. Deleted, see Pm1c.
Pm19{853}. 7D{853}. v: T. durum 'Moroccos 183'/Ae. squarrossa AE 457/78{853}. v2: Synthetic XX186 Pm2{853}. dv: Ae. squarrosa{853}.
Pm20{386}. [M1P6L{543}]. 6BL = T6BS.6R#2L{543,386,389}. v: KS93WGRC28 = PI 583795{386,382}; 6RL. su: 6R{6D}{543}. ad: 6R addition{543}. al: Prolific rye{543}.
Pm21{1177}. 6AS = T6AL.6VS{1177}. v: 9 independent translocations{1177}. ma: RAPD OPH171900 (synonym 'OPH17-1900') was associated with Pm21 and RAPD OPH171000 (synonym OPH17-1000') with its absence{1176}; RAPD OPH171400 and SCAR markers SCAR1400 and SCAR1265 associated with Pm21 are described in{0014}.
Pm22{1134}.
Deleted. See Pm1e.
Pm23{1618}. 5A{1618}. v2: Line 81-7241 Pm8{1618}.
Pm24{571}. 6D{571}.1DS{0150}. v: Chiyacao{571}.
Although Pm24 had previously been located to chromosome 6D {571}, Pm24 was mapped on chromosome arm 1DS in the cross Chinese Spring (susceptible) x Chiyacao (resistant){0150}.
Pm25{1343}. [PmTmb{1343,1344}]. 1A{1343}. v: NC94-3778{1344}. v2: NC96BGTA5 = Saluda*3/PI 427662 Pm3a{1343}. dv: T. monococcum PI 427662{1343}. ma: Linked with 3 RAPDs, the nearest, OPAG04950, at 12.8 +/- 4.0 cM{1343}; Associated with 3 RAPDs{1344}.
Pm26{0001}. Recessive {0001}. 2BS{0001}. s: Bethlehem*8/T. turgidum var. dicoccoides 2BS{0001}. tv: T. turgidum var. dicoccoides TTD140{0001}. ma: Co-segregation with Xwg516-2B{0001}.
Pm27{0002}. 6B (6B-6G){0002}. v: Line 146-155-T{0002}. tv: T. timopheevii var. timopheevii K-38555{0022}. ma: 6BS......Xpsr8/Xpsr964-6B - Pm27 - Xpsr154/Xpsr546-6B ......6BL{0002}; Co-segregation with Xpsr3131-6B{0002}.
Pm28{0022}. 1B{0022}. v: Meri{0022}.
Pm29{0129}. v: Pova{0129}. ma: Location confirmed by co-segregation with molecular markers{0129}.
Pm30{0163}. [MIC20] 5BS{0163}. v: 87-1/C20//2*8866 Seletion{0163}. ma: Pm30 - 5.6cM - Xgwm159-5B{0163}.
Pm31{0301}. v: G-305-M/781//3*Jing411{0301}. tv: T. dicoccoides G-305-M{0301}.

Genotype lists:Chinese wheats{1608,572}; Finnish wheats {0028}; French wheats{1629}; Hungarian wheats {02104}; Western Siberian wheats{1101}
Complex genotypes:
Drabent {heterogeneous} Pm2 Pm4bPm9/Pm1 Pm2 Pm4b Pm9 {1287}; :
Nemares Pm1 Pm2Pm4b Pm6 Pm9 {1287}; :
Planet, Sappo & Walter Pm1 Pm2 Pm4b Pm9 {096,097,540,1287,1428}

76.2. Suppressors of Pm
Some wheats which, on the basis of cytological and rust tests carry 1RS from Petkus rye, do not express resistance due to presence of a suppressor {385}. Zeller & Hsam {1625} located a suppressor of Pm8 and Pm17 in chromosome 7D of Caribo. Mildew resistance was suppressed in Florida, Heinrich, Ikarus, Olymp and Sabina, which are derivatives of Caribo with 1BL.1RS. According to Ren et al. {1209}, SuPm8 does not suppress Pm17. Hanusova et al. {492} listed 16 wheats that carry a suppressor of Pm8; 111 wheats did not carry the suppressor. In contrast, a high frequency of suppression occurred in CIMMYT wheats {108,1208}. Further genotypes are identified in {491}. Although Line 81-7241 carries Pm8 as well as Pm23, evidence was presented to indicate that Pm8 was suppressed in Line 81-7241{1618} and , by inference, indicated that Chinese Spring possessed SuPm8.
SuPm8{1209}. 1AS{1209}. v: Wheats with Gli-A1a{1209}; Lists in{108,491,1208}.
A suppressor of Pm17, as well as of Pm8, was located in chromosome 7D of Caribo {1625}. Gene SuPm8 identified by Ren et al. {1209} did not suppress Pm17.


76.3. Temporarily designated gene for resistance to Blumeria grminis
Ml-Ad{854}. v: Adlungs Alemannen{854}.
Ml-Br{854}. v: Bretonischer Bartweizen{854}.
Mld{096}. 4B{097}. v2: Halle 13471 Pm2{096}; H8810/47 Pm2{096}; Maris Dove Pm2{096}. tv: T. durum line{096}.
Ml-Ga{854}. v: Garnet{854}; many old German cultivars{854}.
mljy{0339}. Recessive, hemizygous-effective {0339} 7B{0339}. v2: Jieyan 94-1-1 Pm8{0339}.
mlsy{0339}. Recessive, hemizygous-effective {0339} 7B{0339}. v: Siyan 94-2-1 {0339}.
Mlre{1220}. 6AL{0142}. v2: RE714 Pm4b{0142,1220}. tv: T. dicoccum 119{1220}.
Mlre showed a residual effect on the quantitative expression of APR in the presence of B. graminis pathotypes considered virulent for Mlre in standard seedling tests{0016}. In addition to Mlre, a QTL for resistance effective at the seedling stage was associated with microsatellite marker Xgwm174-5D {0146}.
Mlxbd{0259}. Recessive and hemizygous-effective {0258} 7B{0259}. v: Xiaobaidong{0258}.



76.4. QTLs for resistance to Blumeria graminis
QTL: Several QTLs were detected in two RE714/Hardi populations when tested at two growth stages and with different cultures over three years. The most persistent and effective QTL was located in the vicinity of Xgwm174-5D {0272}. Three QTLs, QPm.vt-1B, QPm.vt-2A and QPm.vt-2B, with additive gene action, accounted for 50% of the variation in a population developed from Becker/Massey{0284}. QTLs on chromosomes 1A, 2A, 2B, 3A, 5D, 6A and 7B were detected in a RE714/Festin population in multiple locations and over multiple years. The QTLs on chromosome 5D was detected in all environments and all years and is associated with markers Xgwm639-5D and Xgwm174-5D. Resistance was contributed by RE714. A QTL coinciding with MlRE on 6A was also detected in all environments. The QTL on chromosome 5D and 6A accounted for 45% to 61% of the phenotypic variation {0354}.
QPm.sfr-1A{0051}. 1A{0051}. v: Forno/T. spelta var. Oberkulmer mapping population; the resistance was contributed by Oberkulmer{0051}. ma: Associated with Xpsr1201-1A and Xpsr941-1A{0051}.
QPm.sfr-1B{0051}. 1B{0051}. v: Forno/T. spelta var. Oberkulmer mapping population; the resistance was contributed by Forno{0051}. ma: Associated with Xsfr3(LRR)-1B and Xpsr593-1B{0051}.
QPm.sfr-1D{0051}. 1D{0051}. v: Forno/T. spelta var. Oberkulmer mapping population; the resistance was contributed by Oberkulmer{0051}. ma: Associated with Xpsr168-1D and Xglk558-1D{0051}.
QPm.sfr-2A{0051}. 2A{0051}. v: Forno/T. spelta var. Oberkulmer mapping population; the resistance was contributed by Oberkulmer{0051}. ma: Associated with Xpsr380-2A and Xglk293-2A{0051}.
QPm.sfr-2D{0051}. 2D{0051}. v: Forno/T. spelta var. Oberkulmer mapping population; the resistance was contributed by Oberkulmer{0051}. ma: Associated with Xpsr932-2D and Xpsr331-2D{0051}.
QPm.sfr-3A{0051}. 3A{0051}. v: Forno/T. spelta var. Oberkulmer mapping population; the resistance was contributed by Forno{0051}. ma: Associated with Xpsr598-3A and Xpsr570-3A {0051}.
QPm.sfr-3D{0051}. 3D{0051}. v: Forno/T. spelta var. Oberkulmer mapping population; the resistance was contributed by Oberkulmer{0051}. ma: Associated with Xpsr1196-3D and Xsfr2(Lrk10)-3D{0051}.
QPm.sfr-4A.1{0051}. 4A{0051}. v: Forno/T. spelta var. Oberkulmer mapping population; the resistance was contributed by Forno{0051}. ma: Associated with Xgwm111-4A and Xpsr934-4A {0051}.
QPm.sfr-4A.2{0051}. 4A{0051}. v: Forno/T. spelta var. Oberkulmer mapping population; the resistance was contributed by Forno{0051}. ma: Associated with Xmwg710-4A and Xglk128-4A{0051}.
QPm.sfr-4B{0051}. 4B{0051}. v: Forno/T. spelta var. Oberkulmer mapping population; the resistance was contributed by Forno{0051}. ma: Associated with Xpsr593-4B and Xpsr1112-4B{0051}.
QPm.sfr-4D{0051}. 4D{0051}. v: Forno/T. spelta var. Oberkulmer mapping population; the resistance was contributed by Forno{0051}. ma: Associated with Xglk302-4D and Xpsr1101-4D{0051}.
QPm.sfr-5A.1{0051}. 5A{0051}. v: Forno/T. spelta var. Oberkulmer mapping population; the resistance was contributed by Oberkulmer{0051}. ma: Associated with Xpsr644-5A and Xpsr945-5A{0051}.
QPm.sfr-5A.2{0051}. 5A{0051}. v: Forno/T. spelta var. Oberkulmer mapping population; the resistance was contributed by Oberkulmer{0051}. ma: Associated with Xpsr1194-5A and Xpsr918-5A{0051}.
QPm.sfr-5B{0051}. 5B{0051}. v: Forno/T. spelta var. Oberkulmer mapping population; the resistance was contributed by Oberkulmer{0051}. ma: Associated with Xpsr580-5B and Xpsr143-5B{0051}.
QPm.sfr-6B{0051}. 6B{0051}. v: Forno/T. spelta var. Oberkulmer mapping population; the resistance was contributed by Forno{0051}. ma: Associated with Xpsr167-6B and Xpsr964-6B{0051}.
QPm.sfr-7B.1{0051}. 7B{0051}. v: Forno/T. spelta var. Oberkulmer mapping population; the resistance was contributed by Forno{0051}. ma: Associated with Xpsr593-7B and Xpsr129-7B{0051}.
QPm.sfr-7B.2{0051}. This QTL corresponds to Pm5 {0051}. 7B{0051}. v: Forno/T. spelta var. Oberkulmer mapping population; the resistance was contributed by Forno{0051}. ma: Associated with Xglk750-7B and Xmwg710-7B{0051}.
QPm.ipk-2B{0255}. 2BS{0255}. v: Opata/W-7984 (ITMI) RI mapping population{0255}; Resistance was contributed by Opata{0255}. ma: Associated with Xcdo405 and Xmwg950{0255}.
QPm.ipk-4B{0255}. 4B{0255}. v: Opata/W-7984 (ITMI) RI mapping population{0255}; Resistance was contributed by W-7984{0255}. ma: Associated with Xcdo795 and Xbcd1262{0255}.
QPm.ipk-7D{0255}. 7DS{0255}. v: Opata/W-7984 (ITMI) RI mapping population{0255}; Resistance was contributed by Opata{0255}. ma: Associated with Xwg834 and Xbcd1872{0255}.




77.Reaction to Cochliobolus sativus Ito & Kurib.
Disease: Cochliobolus root rot.
Crr{764}. Recessive. 5BL{764,765}. v: Apex{764}; Cadet{765}.


78.Reaction to Diuraphis noxia (Mordvilko)
Insect pest: Russian aphid, Russian wheat aphid.
Dn1{286}. 7D{1288}.7DS{0211}. i: Betta-Dn1{0004,0211}; Caledon{0004}; Gariep{0004}; Karee-Dn1{0211}; Limpopo-Dn1{0004}; Tugela-Dn1{0004,0211}. v: PI 137739{286}. ma: Xgwm111-7D210 - 3.20 +/- 0.20 cM - Dn1{0211}.
Dn2{286}. 7DL{863}.7DS{0211}. i: Betta-Dn2{0211}; Karee-Dn2{0211}; Tugela-Dn2{0211}. v: PI 262660{286,863}. ma: XksuA1-7D - 9.8 cM - Dn2{863}; Myburg et al.{9968} identified two SCAR markers that mapped 3.3cM proximal to Dn2{9968}; Xgwm111-7D200 - 3.05 +/- 0.18 cM - Dn2{0211}; XksuA1-7D - 9.9 cM - Dn2 - 2.8 cM - Xgwm437-7D{0353}.
According to Saidi & Quick {1250}, Dn1 and Dn2 are probably allelic. Reference stocks with each gene showed allelism with a gene in PI 262605.
Dn3{1086}. Recessive. v: T. tauschii SQ24/T. turgidum TD65{1086}. dv: T. tauschii SQ24{1086}.
Dn4{1250}. 1DL{863}. v: CORWAI{260}; CI 2401{260}; Halt{0209}; PI 151918{260}; PI 372129{1250}. ma: Xabc156-1D - 11.6 cM - Dn4 {863}; Xgwm106-1D - 7.4 cM - Dn4 - 12.9 cM - Xgwm337-1D{0352}.
Dn5{1249}. 7D{259}.7DL{287}.7DS{0211}. i: Betta-DN5{0211}; Palmiet derivative 92RL28{287}; Palmiet DN5{0004}. v: STARS - 9302W-sib{259}; PI 294994{259}. ma: A SCAR marker developed from the RAPD fragment OPF141083 mapped 5.5 cM proximal to Dn5{0172}; Xgwm111-7D220 - less than 3.20 cM - Dn5{0211}.
Dn6{1250}. v: CI 6501{260}; PI 243781{1250,1249}. ma: Dn6 - 3.0 cM - Xgwm111{0352}.
Dn7{9918}. Derived from S. secale cv. Turkey 77 {9918} 1B = 1BL.1RS{9918}.1R{9918}. v: 93M45-14{9918}.
Dn8{0211}. 7DS{0211}. i: Karee-Dn8. v2: PI 294994 Dn5Dn9{0211}. ma: Xgwm635-7D100 - less than 3.20 cM - Dn8{0211}.
Dn9{0211}. 1DL{0211}. i: Betta-DN9. v2: PI 294994 Dn5Dn8{0211}. ma: Xgwm642-7D180 - less than 3.20 cM - Dn9{0211}.
Dnx{0211}. 7DS{0211}. v: PI 220127{0211}. ma: Xgwm111-7D210 - 1.52 +/- 0.15 cM - Dnx{0211}.
Dnx was considered to be located at a locus different from Dn1, Dn2 or Dn5 {0211}, which were likely to be identical or allelic.


79.Reaction to Fusarium graminearum
Disease : Fusarium head scab.
Fhs1{1096}. v: Line A{1096}. v2: Ning 7840 Fhs2{1096}.
Fhs2{1096}. v: Line B{1096}. v2: Ning 7840 Fhs1{1096}.
A major QTL was associated with several linked AFLP markers tentatively located in chromosome 7BL of Ning 7840{0005}.
QFhs.ndsu.2A{9925,0175}. 2AL{9925}. v: Sumai 3/Stoa RI mapping population; the QTL was contributed by Stoa{9925}. ma: Association with RFLP XksuH16-2A (LOD>3){9925,0175}.
QFhs.ndsu.3AS{0372}. 3AS{0372}. tv: T. turgidum var. dicocooides. Recombinant substitution lines LDN and LDN(Dic-3A). The resistant allele was contributed by dicoccoides{0372}. ma: Associated with Xgwm2-3A (explained 37% of the phenotypic variation){0372}.
QFhs.ndsu.3B{9925,0175}. 3BS{9925}. v: Sumai 3/Stoa RI mapping population; the QTL was contributed by Sumai 3{9925,0175}. ma: Association with Xbcd907-3B.2(LOD>3){9925}; and microsatellite markers Xgwm1533-3B and Xgwm493-3B{0175}.
This QTL explained 42% of the variation in Sumai 3/Stoa{0175}.
Two additional QTL for resistance to Fusarium graminearum were identified in the cross Sumai3/Stoa {0175}. The QTL on 4BS was associated with Xwg909-4B and the QTL on 6BS was associated with Xbarc101-6B and Xbcd1383-6B {0175}. The QTL associated with markers Xgwm493/Xgwm533 (explaining 24.8 % of the variation), and Xbarc101/Xbcd1383 were also identified in a RIL population from the cross ND2603/Butte 86 {0175}. In addition, one QTL on chromosome 3AL associated with Xbcd941-3A and one on chromosome 6AS associated with XksuH4-6A were identified in RILs from the cross ND2603/Butte 86 {0175}.
The resistance QTL on chromosome 3BS associated with Xgwm493-3B and Xgwm533-3B was also identified in a DH population of the cross CM-82036 (a Sumai 3 derivative) Remus {0240}. Additional QTL in this cross were detected on chromosome 5A, associated with Xgwm293-5A and Xgwm304-5A, and possibly on 1B, associated with Glu-B1 {0240}.
Two major genes with additive effects were reported in cross between Sumai 3 (resistant) and two susceptible cultivars{0174}. One of the genes was assigned to 5AL based on linkage to the dominant awn suppressor B1 (RF 15.1-21.4%).
For review see{0283}.
Mesterhazy et al.{0006} reported a strong genetic correlation in resistance to different species of Fusarium.
QTLs were located in 3BS, 2BL and 2AS in Ning 7840/Clark. The most effective QTL was probably in interval, flanked by deletions 3BS-3 and -8 and was close to Xgwm533-3B and Xbarc147-3B {0328}.


80.Reaction to Heterodera avenaeWoll.
Cereal root eelworm; cereal cyst nematode.
Cre1. [Cre{1388}]. 2B{1388}.2BL{1579,1580}. i: AP = Prins*8/AUS10894{1579}. v: AUS 10894{1056}; Loros CI 3779{1388}. ma: Xglk605-2B - 7.3cM - Cre1 - 8.4cM - Xcdo588-2B/Xabc451-2B{1579}; A PCR-based assay was developed from Xglk605-2B{1580}.
Cre2{238}. Derived from Ae. ventricosa 10 {238,9991}. 6Mv{9991} v2: H-93-8 Cre6{238}.
Although H-93-8 is a double Mv(5A), 7Mv(7D) substitution line, Cre2 was presumed to be located in a separate undetected translocated 6Mv segment{9991}.
Cre3. [CcnD1{329},Ccn-D1{328}]. 2DL{328}. v: Synthetic hexaploids{329}. dv: T. tauschii accessions AUS 18912{328}; AUS 18913{328}; CPI 110809{329}; CPI 110810{328}.
Cre4. [CcnD2{329},Ccn-D2{328}]. 2D{328}. dv: T. tauschii accessions AUS 18914{329}; CPI 110813{328}.
Cre5{0107}. Derived from Aegilops ventricosa {0107,0009}. [CreX{0009,0183}]. 2AS{0107}=2A-2Nv-6Nv. v: VPM1{0107}; Many VPM1 derivatives{0107}; Notable exceptions of lines with Lr37, Sr38 and Yr17, but lacking Cre5 include Trident and Line L22{0107}. su: Moisson 6Nv(6D){0183}. dv: Ae. ventricosa 10{0183}.
Two resistance gene analogues similar to the candidate gene Cre3 were isolated from the Ae. ventricosa segment carrying Cre5
Cre6{0138}. Derived from Aegilops ventricosa {0138}. 5Nv{0138}. ad: Moisson + 5Nv{0138}. v: H-93-35{0138}. v2: H-93-8 Cre2{0138}.
Cre7{0104}. Derived from Aegilops triuncialis {0105}. [CreAet{0105}]. v: TR353 derivatives{0105}.
Cre8{0220}. [CreF{0012,0138}]. 6B{0220}, on basis of linkage with Xbcd1-6B and Xcdo347-6B{0220}. v: Barunga{0220}; Festiguay{0012,0220}; Frame{0138,0220}; Molineaux{0220}. ma: Associated with a unique allele when probed with CDO367 which hybridizes to group 7L{1059}.
CreR{0133,0318}. 6RL{0133}. ad: Wheat + 6R {0318}; Wheat + 6RL{0318}; Various deletion stocks{0318}. su: CS + 6R(6D){0133}. al: Rye accession T701-4-6{0133}; Triticale T-701{0318}. ma: Cent......XksuF37 - 3.7cM - CreR{0133}; Deletion mapping indicated CreR was located near Got-R2{0318}.


81.Reaction to Magnaporthe grisea (Herbert) Barr
M. grisea is a pathogen of blast on many graminaceous species, the best known of which is rice. In Brazil it has become a pathogen of wheat. The wheat pathotype(s) is different from those attacking other species such as rice, oat, millets and weeping lovegrass.
Rmg1{0333}. [Rwt4{0302}]. v: Norin 4{0302}.

A second gene designated Rwt3 {0302} was present in CS and Norin 4. Genes Rwt3 and Rwt4 were detected using hybrids of Triticum- virulent and Avena-virulent pathogen isolates.

82.Reaction to Mayetiola destructor (Say) (Phytophaga destructor) (Say)
Insect pest: Hessian fly.
H1{1087}. i: Dawson/3*Poso, 6179{1087}. v2: Big Club 43 H2{1441}; Dawson H2{166,1087}; Poso 42 H2{1441}.
H2{1087}. i: Dawson/3*Poso, 6232{1087}. v2: Big Club 43 H1{1441}; Dawson H1 {166,1087}; Poso 42 H1{1441}.
H3{156}. Recessive. 5A{425,1105}. i: Carol = Newton-207*5/Larned{1107}. v: Ace{426}; Arthur{426}; Becker{749}; Cardinal{750}; Dual{1273}; Frankenmuth{341}; Georgia 1123{426}; GR855{751}; GR876{753}; Ionia{426}; Larned{824}; Logan{426}; Monon{157}; Norkan{904}; Ottawa{547}; Purdue B 36162 A13-12{156}; PI 468960{1439}; Redcoat{1273}; Reed{1273}; Riley{1273}; Roland{148}; Russell{426}; Shawnee{547}; Titan{747}; Todd{426}; W38{156}. v2: Clara Fay H6{375}. ma: Cosegregation of H3 and a RAPD{296}.
Allan et al.{019} considered that H3 and H4 may be allelic. Also suggested by Shands and Cartwright{1317}. Linkage of 10.5 +/- 2% involving H3 and Pm3a in PI 468960 was attributed to a chromosome 1A/5A translocation{1437}.
H4. Recessive. H4 confered resistance to race A, but not to race B. [h4{1441}]. v: Dixon{1441}; Java{1441}.
H5{1317}. Temperature sensitive {1413}. 1AS{1222}. v: Abe{162}; Arthur 71{162}; Beau{875}; Downy{1223}; Oasis{1109}; Ribeiro{1317}; Sullivan{1110}. tv: Giorgio 331-4{1090}; PI 94567-6{1317}; PI 94571-14{1317}. ma: Cosegregation of H5 and two RAPDs{296}.
H6{019}. 5A{425}. i: Erin = Newton-207*7/Arthur 71{1107}; Flynn = Newton-207*7/Knox 62{1107}. v: Adder{1319}; Benhur{426}; Caldwell{1421}; Compton{1318}; CI 12855{019}; Excel{752}; Fillmore{1106}; Knox 62{426}; Lathrop{426}. v2: Clara Fay H3{375}. tv: Purdue 4835 A4-6{1105}. tv2: PI 94587 H11 H16{019}. ma: Cosegregation with three RAPDs{296}.
H7 & H8{425}. Duplicate factors. H7 is located in chromosome 5D {026}. v: Adena{748}; Seneca{026,425}.
H9{1420}. 5A{162}. i: Iris = Newton-207*7/Ella{1107}. v: Ella{875}; Line 822-34{162}. v2: Elva CI 17714 H10{162}; Line 812-24 H10{1421}; Line 817-2 H10{1421}; Stella H10{875}. ma: Cosegregation with two RAPDs{296}.
H10{1104}. 5A{162}. i: Joy = Newton-207*3/IN76529A5-3-3{1107}. v: IN 76529{875}. v2: Elva CI 17714 H9{162}; Line 817-2H9{162}; Stella H9{875}. ma: Cosegregation with one RAPD and close linkage to another RAPD{296}.
H11{1422}. 1A{1222}. i: Karen = Newton-207*4/IN916-1-3-1-47-1{1107}. v: Kay{875,375}; Line 916{1422}; Line 920{1422}; Line 941{1422}. tv2: T. turgidum PI 94587 H6 H16{1422}. ma: Close linkage with two RAPDs{296}.
H12{1092}. 5A{1098}. i: Lola = Newton-207*4/Luso{1107}. v: Luso{1092}. ma: Cosegregation with one RAPD and close linkage of H12 to another RAPD{296}.
H13{1104}. 6DL{441}. i: Molly = Newton-207*7/3/KU221-19/Eagle/ KS806{1107}. v: KS81H1640HF{441}; T. turgidum var. durum cv. Gulab KU 134/Ae. tauschii KU 2076, KU 221-14{525}; T. turgidum var. persicum straminium KU 138/Ae. tauschii KU 2076, KU221-19{525}. dv: Ae. tauschii KU 2076{525}. ma: Cosegregation with a RAPD{296}.
H14{875}. 5A{875}. tv: IN 81601A2-3-3{875}. tv2: ELS 6404-160 H15{875}. ma: Cosegregation with a RAPD{296}.
H15{875}. 5A{875}. tv: IN 81602C5-3-3{875}. tv2: ELS 6404-160 H14{875}.
H16{1106}. 5A{1098}. tv: IN 80164H5-2-9{1106}; N80164{1097}. tv2: PI 94587 H6 H11{1106}. ma: Cosegregation of H16 and a RAPD{296}.
H17{1090}. 5A{1090}. tv: PI 428435{1090}. ma: Cosegregation of H17 and a RAPD{296}.
H18{1090}. v: Marquillo{426,874}; Shield{198}.
H19{1089}. tv: PI 422297{1089}; This germplasm possesses a second gene which is allelic or closely linked with H16{1089}; IN84702{1097}. tv2: PI422297 H29{1097}.
H20{025}. 2B{025}. tv: Jori{025}.
H21. 2B {383} = 2BS.2R#2L{389}. v: Hamlet = KS89WGRC8{1312}; KSWR 69-2-4-3{383}; KS85HF 011-5{383}. ad: KSWR 297h-1-1-9{383}. al: Chaupon rye{383}. ma: A RAPD amplified by primer OPE-13 was shown to co-segregate with H21{9938}; A STS primer set SJ07 was developed to identify 2RL, and hence H21{0233}.
H22{1199}. 1D{1199}. v: KS86WGRC1{1199}.
H23{1199}. 6D{442}.6DL{1199}. v: KS89WGRC3{442}. ma: H23 - 6.9 cM - XksuH4-6D{861}.
H24{1199}. 3D{442,1199}.6DL{861}. v: KS89WGRC6{442}. ma: H24 - 5.9 cM - Xbcd451-6D/Xcdo482-6D{861}.
H25.
6B{384} = T6BS.6BL-6R#1L{389}.v:88HF16 = WGRC17{384}.
4B{384} = T4BS.4BL-6R#1L{389}.v:88HF79, 88HF80 = WGRC18, 88HF81, 88HF117 = WGRC19{384}.
4A{384} = Ti4AS.4AL-6R#1L-4AL{389}.v:89HF17, 89HF18, 89HF25, 88HF32, 88HF51, 88HF89 = WGRC20{384}.
6R.al:Balbo rye{384}.
H26. 4D{217}. v: KS92WGRC26{217}. dv: Ae. tauschii TA2473{217}.
H27{235}. 4Mv{235}. su: H-93-33{235}. al: Ae. ventricosa No. 10{235}; Ae. ventricosa No. 11{235}.
H28{171}. 5A{171}. tv: PI 59190{171}.
H29{1095}. [H27{171}]. 5A{1097}. tv: PI422297 H19{1097}.
H30{0256}. Derived from Ae. triuncialis {0256}. v: TR-3531{0256}. al: Ae. triuncialis{0256}.
H31{0332}. 5BS{0332}. v: P961696{0332}. tv: CI3984{0332}. ma: STS marker Xupw148-5B - 3 cM - H31{0332}.

A recombination value of 12.0% between leaf-rust reaction {possibly Lr10} and Hessian-fly reaction in Selection 5240 was reported {018}.

83.Reaction to Meloidogyne spp.
Root rot nematode, root knot eelworm
Rkn{632}. dv: Ae. squarrosa G3489. v: Prosquare, a synthetic hexaploid of Produra/A. squarrosa G3489{632}.
Rkn-mn1{1621}. Derived from Ae. variabilis{1621}. 3B{590}. v: X8=CS/Ae. variabilis No. 1//Rescler/3/Lutin{1620}; X35{1620,1621}. ma: Co-segregation with RAPD OpY161065 and close linkage with several markers including Est-B5{0103}.


84.Reaction to Mycosphaerella graminicola (Fuckel) Schroeter
Disease: Septoria tritici blotch
Stb1. [Slb1{1586}]. v: Bulgaria 88{1586}; Oasis{1586}; Sullivan{1586}.
Stb2. [Slb2{1586}]. v: Nova Prata{1586}; Veranopolis{1586}.
Stb3. [Slb3{1586}]. v: Israel 493{1586}.
Stb4{1410}. v: Cleo{1410}; Tadinia{1410}; Tadorna{1410}.
Stb4 segregated independently of Stb1 but its relationship with Stb2 and Stb3 is unknown.
Stb5{0186}. Identified using M. graminicola IPO94269{0186}. Derived from Aegilops tauschii accession 37-1 {0186}. 7DS{0186}. v: Bezostaya{0187}; Hereward{0187}; Sears' Synthetic{0186}; Shafir{0187}; Vivant{0187}. su: CS*8/(Syn7D){0186}. dv: Ae. tauschii 37-1{0186}. ma: Rc3 - 6.6cM - Stb5 - 7.2cM - Xgwm44-7D - Centromere{0186}; Stb6 - 2cM - Xgwm369-3A{0187}.
Stb6{0187}. Confers resistance to M. graminicola isolate IPO323 but not to isolate IPO94269 {0187}. 3AS{0187}. v: Flame{0187}.
Stb7{0311}. 4AL{0311}. v: ST6 = Estanzuela Federal{0310, 0311}. ma: wmc219-4A - 0.8cM - Xwmc-4A - 0.3cM - Stb7{0311}.
Stb8{0326}. 7BL{0326}. v: Synthetic hexaploid W7984 (parent of ITMI population){0326}. ma: Xgwm146-7B - 3.5cM - Stb8 - 5.3cM - Xgwm577-7B{0326}.


85.Reaction to Pratylenchus spp.
Root lesion nematode; prats

85.1. Reaction to Pratylenchus neglectus
Rlnn1{0121}. 7AL{0121}. v: Excalibur{0121}; Krickauff{0121}. ma: Mapped between markers Xpsr121-7A and Xgwm344-7A and 9 cM proximal to Lr20{0374}.


85.2. Reaction to Pratylenchus thornei
QTLs were located on chromosomes 2BS and 6DS {0122}.



86.Reaction to Phaeosphaeria nodorum (E. Muller) Hedjaroude (anamorph: Stagonospora nodorum (Berk.) Castellani & E.G. Germano).
Disease: Septoria nodorum blotch.
Snb1{856}. 3AL{856}. v: Red Chief{856}. v2: EE8 Snb2{856}.
Snb2{856}. 2AL{856}. v2: EE8 Snb1{856}.
Snb3{1594}. 5DL{1594}. s: CS*/Synthetic 5D{1594}. v: Synthetic{1594}. dv: Ae. tauschii{1594}.
SnbTM{856,857}. 3A{857}.3AL{856}. tv: T. timopheevii /2*Wakooma{856}; T. timopheevii PI 290518. T. timopheevii derivatives: S3-6{857}; S9-10{857}; S12-1{857}. ma: UBC521650 - 15cM - SnbTM - 13.1cM - RC37510{0212}.
UBC521650 was converted to a SCAR marker{0212}.


87.Reaction to Puccinia graminis Pers.
Disease: Black rust; black stem rust; stem rust.
Note: Some near-isogenic lines are based on Marquis. The genes present in the Marquis background are not listed for those NILs.
Sr1. Deleted - see Sr9d.
Sr2{38,677}. Recessive allele. Adult plant response. 3BS{499}. s: CS*6/Hope 3B{499}. v2: Warigo Sr7b Sr17{499}; Suneca Sr8a Sr17{485}; Hopps Sr9d{499}; Lancer Sr9d Sr17{679}; Scout Sr9d Sr17{679}; See also{499,1040}. ma: Xgwm389-3B - 2.7cM - Sr2 - 1.1cM - Xglk683-3B{0358}.
Sr2 is associated with pseudo-black chaff{742,1102} and seedling chlorosis (see {149}) and occurs very frequently in commercial wheats, especially in germplasm produced and distributed by CIMMYT. Sr2 has probably remained effective since the 1920s.
Sr3 & Sr4{047}. v: Marquillo - based on early data. No stocks available.
Sr5{047}. 6D{939,1308,1626}.6DS{939}. i: I Sr5-Ra{828}; I Sr5-Rb{828}; Sr5/7*LMPG{685}; Thatcher/10*Marquis{686}. s: CS*6/Thatcher 6D{1308}. v: Admonter Fruh{072}; Dacia{979}; Dong-Fang-Hong 2{564}; Dong-Fang-Hong 6{564}; Feng-Kong{563}; Hochzucht{046}; Hybrid 80-3{072}; Jubilejna{068}; Juna{076}; Kanred{1308}; Ke-Fang 1{564}; Stabil{072}; Viginta{071}; Vrakunski{072}. v2: Victor Sr6 Sr8a{979}; Glenlea Sr6 Sr9b{327}; Qing-Chung 5 Sr6 Sr11{564}; An-Hewi II Sr8a{564}; Erythrospermum 974 Sr8a{072}; N.P.789 Sr11{1555}; Jing-Hong Sr17{564}; Jing-Hong 2 Sr17{564}; Amika Sr31{076}; Dong-Xie 3 Sr31{563}; Dong-Xie 4 Sr31{563}; Istra Sr31{076}; Solaris Sr31{076}; Beijing 10 SrTmp{564}.
Sr6{687}. [SrKa1{1167}]. 2D{1293,1308,1577}.2DS{942}. i: I Sr6-Ra{828}; Kenya 58/10*Marquis{468,675}; Sr6/9*LMPG{685}. s: CS*5/Red Egyptian 2D{1308}. v: Africa 43{669}; Eureka{468,844}; Kenya stocks{669,670,673,687,689,1167,1557}; McMurachy{679}; Shield{198}. v2: Bowie Sr8a{1553}; Eurga Sr11{1553}; Fortuna Sr7a{679}; Gamut Sr9b Sr11{1555}; Glenlea heterogeneousSr5 Sr9b{327}; Kentana 52 Sr7a{678,1577}; Kiric 66 Sr7b{979}; Lerma Rojo 64 Sr7b Sr9a{979}; No. 466 Sr9b Sr10{689}; Red Egyptian Sr8a Sr9a{687,1308}; Siete Cerros Sr11{033}; Victor I Sr5 Sr8a{979}.
See also {1553}.
Sr7{830}. 4A{671,830,1293}.4AL{939,1308}.
Sr7a{830}. [Sr7{687}]. i: Egypt Na101/6*Marquis{468}; Kenya 117A/6*Marquis{468}; Sr7a/9*LMPG{685}. s: CS*7/Kenya Farmer 4B{830}; CS*8/Sapporo 4B{830}. v: Egypt Na101{669}; Kenya stocks{669,670,673,687,689}; Sapporo Haru Komugi Ichigo{689}. v2: Fortuna Sr6{679}; Kentana 52 Sr6{689}; French Peace Sr9a Sr13{680}; Egypt Na95 Sr9b Sr10{687}; W3746 Sr12{1371}; Khapstein Sr13 Sr14{674}.
Sr7b{830}. i: I Sr7b-Ra{828}. v2: Warigo Sr2 Sr17{499}; Kiric 66 Sr6{979}; Roussalka Sr8a{979}; Red Bobs Sr10{308}; Nell Sr17{1565}; Spica Sr17{939}; Marquis Sr18 Sr19 Sr20{675,830}.
Sr8. 6A{1293,1308}.6AS{929,1368}.
Sr8a{1368}. [Sr8{687}]. i: I Sr8a-Ra{828}; Red Egyptian/10*Marquis{686}; Sr8a/9*LMPG{685}. s: CS*5/Red Egyptian 6A{1308}. v: Marimp 3{979}; Mentana{844}; Strampelli{979}. v2: An-Hewi II Sr5{564}; E-Gan-Zao Sr17{564}; Erythrospermum 974 Sr5{072}; Frontana Sr9b{689}; Golden Valley Sr17{979}; Hartog Sr2 Sr12{127}; Magnif G Sr9b{689}; Pitic 62 Sr9b{033}; Red Egyptian Sr6 Sr9a{687}; Rio Negro Sr9b{689}; Roussalka Sr7b{979}; Suneca Sr2 Sr17{485}; Victor 1 Sr5 Sr6{979}.
Sr8b{1368}. [SrBB]. v: Barleta Benvenuto{1368}; Klein Titan{1368}. v2: Bezostaya Sr5{979}; Klein Cometa Sr30{1368}.
Sr9{676}. 2B{671,677,828,1308}.2BL{944,946,951,1307,1582}.
Sr9a{676}. [Sr9{687}]. i: I Sr9a-Ra{828}; Red Egyptian/10*Marquis{686}; Sr9a/9*LMPG{685}. s: CS*4/Red Egyptian 2B{1308}. v2: Red Egyptian Sr6 Sr8a{687}; French Peace Sr7a Sr13{680}; Excel Sr8a Sr17{752}.
Sr9b{468}. [Sr9{687},SrKb1{468}]. i: Kenya 117A/10*Marquis{686}; Sr9b/10*LMPG{685}. s: CS*7/Kenya Farmer 2B{939}. v: Gamenya{844}; Kenya stocks{669,670,673,687,689,1557}. v2: Egypt Na95 Sr7a Sr10{636}; Festival Sr15{1553}; Frontana Sr8a{689}; Gamut Sr6 Sr11{1555}; Glenlea Sr5 Sr6 heterogeneous{327}; Kenora Sr15{1553}; Magnif G Sr8a{689}; No. 466 Sr6 Sr10{689}; Pitic 62 Sr8a{033}; Rio Negro Sr8a{689}; Robin Sr11{879}; Veadeira Sr10{687}.
See also {1553}.
Sr9c.Originally reserved for Sr36.
Sr9d{678,831}. [Sr1{047,676,677}]. i: Hope/10*Marquis{677}; H-44/10*Marquis{677}; I Hope 2B-Ra{828}; Sr9d/8*LMPG{685}. v: Hopps Sr2{1040}. v2: Lancer Sr2 Sr17{679}; Scout Sr2 Sr17{679}. tv: Arnautka{939}; Mindum{939}; Spelmar{939}.
Sr9e{951}. [Srd1v{642},Srv{1391}]. v: SST 16{1324}; SST 33{785}; SST 66{785}; SST 3R{1324}; Vernstein{845}. v2: Combination III Sr36{841}; Sunstar Sr8a Sr12{939}. tv: Vernal emmer{1391}; CI 7778{845}; Sr9e occurs in many tetraploid wheats{939,1378}.
Sr9f{826}. v: Chinese Spring{826}; Not present in the near-isogenic I Sr9a-Ra{826}.
Sr9g{965}. s: CS*7/Marquis 2B Sr16{965}; CS*4/Thatcher 2B Sr16{965}. v2: Hochzucht Sr5 Sr12{965}; Lee Sr11 Sr16{965}; Celebration Sr12 Sr16{965}; Eagle Sr26{842}. tv: Acme{965}; Iumillo{965}; Kubanka{965}.
See also {504}.
Sr10{687}. 2B{686,939}. i: Egypt Na95/4*Marquis{468}. v: Federation{939}; Geneva{1412}; Hazen{049}; Kenya stocks{669,670,673,687}. v2: Egypt Na95Sr7a Sr9b{687}; No. 466 Sr6 Sr9b{689}; Red Bobs Sr7b{308}.
Sr11{468}. [Sr11{687},Sr12{687}]. 6B{671,1143,1293,1309}.6BL{1297}. i: I Sr11-Ra{828}; Lee/10*Marquis{686}. s: CS*7/Kenya Farmer 6B{830}; CS*9/Timstein 6B{1308}. v: Charter{844}; Flevina{072}; Gabo{687}; Kenya stocks{670,673,844,1557}; Sonora 64{033}; Sylvia{071}; Timstein{687,1308}; Tobari 66{033}; Yalta{844}. v2: Eurga Sr6{1553}; Gamut Sr6 Sr9b{1555}; Lee Sr9g Sr16{687}; N.P.790 Sr5{1555}; Qing-Chung 5 Sr5 Sr6{564}; Robin Sr9b{879}; Prospect SrWld{197}; See also{1553}.
A resistance gene allelic with Sr11 was found in Chinese Spring {938}, but the P. graminis culture for its detection was lost.
Sr12{1332}. Recessive.
3B{1332,682}.3BS{968}. s: CS*7/Marquis Selection 3B Sr16{1332}; CS*5/Thatcher 3B Sr16{1332}. v: Marquillo{682}; Tincurrin{939}; Windebri{939}. v2: W3746 Sr7a{1371}. tv: Postulated for several durums{1378}.
Sr12 is more widespread and probably more effective in conferring resistance than is usually acknowledged {939}.
Sr13{674}. 6AL{929}. i: Khapstein/10*Marquis{686}; Sr13/9*LMPG{685}. v2: French Peace Sr7a Sr9a{680}; Khapstein Sr7a Sr13 Sr14{674}. tv2: Khapli Sr14{674}.
Sr14{674}. 1BL{933}. i: Khapstein/10*Marquis{686}. v: Line A{933}. v2: Khapstein Sr7a Sr13{674}. tv2: Khapli Sr13{674}.
Sr15{1554}. 7A{1293,1554}.7AL{1305}. v: Present in stocks possessing Pm1 and Lr20{931,1554}; See Reaction to Brumeria graminis and Reaction to P. triticina. ma: Associated with clustered markers{0323}.
Sr16{830}. [Srrl2{1238}]. 2B{830,1308}.2BL{1307}. i: I Sr16-Ra{828}; I Th3B-Ra{832}. s: CS*7/Marquis 2B Sr9g{1581}; CS*4/Thatcher 2B Sr9g{1308}; CS*5/Thatcher 3B Sr12{832}. v2: Thatcher Sr5 Sr9g Sr12{939}; Lee Sr9g Sr11{939}.
Sr17. Recessive. [sr17{964}]. 7B{771}.7BL{964}. s: CS*6/Hope 7B{964}. v2: E-Gan Zeo Sr8a{564}; Golden Valley Sr8a{979}; Jing-Hong 1 Sr5{564}; Jing-Hong 2 Sr5{564}; Lancer Sr2 Sr9d{679}; Nell Sr7b{1565}; Scout Sr2 Sr9d{679}; Suneca Sr2 Sr8a{485}; Present in many stocks possessing Pm5{964}; See Reaction to Blumeria graminis.
Sr18{054}. [SrG2{844},Srrl1{1238},Srmq1{099},SrPs1{1263},SrMn1{1263}]. 1D{054,1308,1582}. i: I Hope 1D-Ra{828}; Sr18/8*LMPG{685}. s: CS*6/Hope 1D{1308}. v: Present in the majority of wheat stocks{828}; Stocks not possessing Sr18: Brevit{054}; Chinese Spring{828}; Eureka{054}; Federation{054}; Gular{054}; Kenya C6042{054}; Koala{054}; Little Club{828}; Morocco{054}; Norka{054}; Prelude{828}; Yalta{054}.
Sr19{029}. [Srmq2{099}]. 2B{029}.2BS{1582}. v: Mq-B{029}. v2: Marquis Sr7b Sr18 Sr20{029}.
Sr20{029}. [Srmq3{1238},Srrl3{1238}]. 2B{029}. v: Mq-C{029}; Rl-C{029}. v2: Reliance Sr5 Sr16 Sr18{029}; Marquis Sr7b Sr18 Sr19{029}.
Sr21{1460}. 2AL{1460,1464}. i: Sr21/8*LMPG{685}. v: Hexaploid derivatives of T. monococcum{939}. tv: Tetraploid derivatives of T. monococcum{939}. dv: Einkorn{1460}; Various monococcum accessions. See also Sr45 which has similar specificity to Sr21.
Sr22{1460}. 7A{649}.7AL{1460}. i: Marquis*4//Stewart*3/T. monococcum{649,1460}; Sr22/9*LMPG{685}; Others{1112}. v: CS/3/Steinwedel*2//Spelmar/T. boeoticum{1460}; Schomburgk{880}; Steinwedel*2//Spelmar/T. boeoticum{1460}; Others{1112}. tv: Spelmar/T. boeoticum{1460}; Stewart*6/T. monococcum RL 5244{649}. dv: Various T. monococcum accessions{649,1460}. ma: Hexaploid derivatives with Sr22 carried 'alien' segments of varying lengths; the shortest segment was distal to Xpsr129-7A{1112}; See also{0158}.
Sr23{950}. The following chromosome locations are consistant with the finding that the first location was based on Rescue monosomics. Rescue differs from CS by a 2B-4B reciprocal translocation {939}. 4B{950}.2BS{939}. v: Exchange{950}; Warden{950}; Sr23 is always associated with Lr16{950}. v2: Etoile de Choisy Sr29{950}.
Sr24{956}. Derived from Thin. elongatum.
3DL.
3DL =T3DS.3DL-3Ae#1L{956,389}.i:Sr24/9*LMPG{685}; Sears' 3D/Ag translocations{956,1300}. v: Agent{956}; Blueboy II{956}; Collin{901}; Cloud{956}; Cody{1284}; Fox{956}; Gamka{785}; Karee{785}; Kinko{785}; Palmiet{785}; Sage{825,1024}; SST 23{1324}; SST 25{785}; SST 44=T4R{1324,785}; SST 102{785}; Torres{128}; Wilga{785}. v2: Siouxland Sr31{1283}. v: List of Australian genotypes{0340}.
1BL.T1BL = 1BS-3Ae#1L{600,389}.tr:Amigo{1463,600,389}; Teewon{600,389}; Note:Amigo also carries a 1AL.1RS translocation with resistance from rye{1463}.
3Ae#1.su:Chinese Spring 3Ag {3D}{1304}; TAP48{389}.
Sr24 is completely linked in coupling with Lr24 {956} and often with red grain colour. See Reaction to P. triticina.
Sr25{956}. Derived from Thin. elongatum.
7DL = T7DS.7DL-7Ae#1L{291,956,388,657}.i:Sears' CS 7D/7Ag translocations{956,1300}; Sr25/9*LMPG{685}. v: Agatha Sr5 Sr9g Sr12 Sr16 {956} = T4{1323}; Mutant 28{388}.
7AL = T7A-7Ae#1L{330}.v:Sears' 7A/7Ae#1L No. 12{330,1304}; Sears' 7D/7Ag#11 carries neither Sr25 nor Lr19{939}.
7Ae#1L.su:Chinese Spring + 7Ae#1L{7D}{1304}; .
See Lr19, reaction to Puccinia triticina.
Sr25/Lr19 often show complete linkage in wheat {956}.

Knott {681} obtained two mutants (28 and 235) of Agatha with reduced levels of yellow pigment in the flour. One of these mutants lacked Sr25. Marais {890} reported that a gene very similar to Sr25 was present in the putative Inia 66 x Thinopyrum distichum derivative, Indis. Marais {890,892} also obtained mutants with reduced yellow pigment in Indis derivatives and some of these lacked Sr25.
Sr26{956}. Derived from Thin. elongatum. 6AL{364} = T6AS.6AL-6Ae#1L{388,389}. i: Sr26/9*LMPG{685}. v: Avocet{364}; Flinders{1449}; Harrier{939}; Jabiru{956}; King{1451}; Kite{956}; Knott's 6A-6Ae#1L translocation to Thatcher{672}; Takari{253}. v2: Bass Sr36{1450}; Eagle Sr9g{956}. ma: Detected with several RFLP probes{0138}.
Sr27. Derived from S. cereale.
3A (T3A-3R) = T3AS.3R#1S{003,896,389}.i:Sr27/9*LMPG{685}. v: WRT wheat-rye translocation, available in CS, Thatcher and Pembina backgrounds. Translocated from Imperial rye to Chinese Spring by Acosta{003}; Widespread in triticales{966,1384}.
3B = T3BL.3R#1S{896}.:.
3A = T3A.3RS{896}.v:W964 = 3RS.3AL.1/4* Inia 66{}; W968 = 3RS.3AL.1/5* Condor{}; W970 = 3RS.3AL.88/5*SST3{}.
3B = T3BL.3RS{896}.v:W966 = 3RS.3BL.26/4*Inia 66{}.
Sr28{932}. 2BL{932}. i: Line AD{932}. v2: Kota Sr7b Sr18{932}.
Sr29{313}. [SrEC{955}]. 6DL{313}.6DS{1626}. i: Prelude/8*Marquis//Etoile de Choisy{313}. v: Hana{071}; Hela{076}; Mara{068}; Slavia{076}; Vala{076}. v2: Etoile de Choisy Sr23{955}.
Sr30{688}. [SrW]. 5DL{688}. i: Sr30/7*LMPG - Lines 1, 2, and 3{685}. v: Festiguay{688}; Mediterranean W1728{1369}; Webster{688}. v2: Klein Cometa Sr8b{1368}; Relatively common in Australian and Mexican wheats. Various unnamed accessions{208,1321}.
Sr31. Derived from S. cereale.
1B = T1BL.1RS = T1BL.1R#1S{389} or 1R(1B).i:MA1 and MA2 four-breakpoint double translocation lines 1RS-1BS-1RS.1BL in Pavon{0084}. v: Amika {heterogeneous} Sr5{076}; Cougar{0267}; Derivatives of Petkus rye - see Lr26, Yr9. Feng-Kang 2{563}; Feng-Kang 8{563}; Gamtoos{785}; GR876{753}; Jing-Dan 106{563}; Jan 7770-4{563}; Lu-Mai 1{563}; Rawhide (heterogenous){0267}; Yi 78-4078{563}. v2: Dong Xie 3 Sr5{563}; Dong Xie 4 Sr5{563}; Istra Sr5{076}; Solaris Sr5{076}; Siouxland Sr24{1283}. tv: Cando*2/Veery = KS91WGRC14{381}. ma: 1BS/1RS recombinants 4.4cM proximal to Gli-B1/Glu-B3{0084}; Several markers tightly linked with Sr31 were indentified in{0377}.
Sr32. Derived from Ae. speltoides.
2A {939,1304} = T2AL.2S#1L-2S#1S {389}.v:C95.24{389}.
2B {1304 }= T2BL-2S#1S{389}.v:C82.1 = P80-14.1-2{389}.
2D {1304} = T2DL-2S#1L.2S#1S{389}.v:C82.2 = P80-139.1-4{389,1304}; C82.3 = P80-132.2-2{939,1304}; C82.4 = P80-153.1-2{939,1304}.
Sr33. (linked with Gli-D1). [SrSQ{650}]. 1DL{650}.1DS{620}. v: RL 5405 = Tetra Canthatch/Aegilops squarrosa RL 5288{650}. ma: Xmwg60-1D - 5.8cM - Sr33 - 2.2cM - Xwmg2083-1{0360}.
Sr34{967}. Derived from Ae. comosa.
2A {967} = T2AS-2M#1L.2M#1S{389}.v:CS 2A-2M 4/2{967}.
2D {967} = T2DS-2M#1L.2M#1S{389}.i:Sr34/6*LMPG{685}. v: Compair{967}; CS 2D-2M 3/8{967}; Various addition, substitution and translocation lines with Yr8{967}.
2M{967}.su:Chinese Spring 2M(2A){967}.
Sr35{957}. [SrTm1{1522}]. 3AL{957}. v,tv: Tetraploid and hexaploid derivatives of T. monococcum{957}. dv: T. monococcum C69. 69 Selection{957}; G2919{957}.
Sr36{939}. [SrTt1{949}]. 2BS{939}. i: Sr36/8*LMPG{685}. v: Arthur{939}; Arthur 71{1324}; Flemink{1324}; GK Kincso{0235}; Gouritz{1324}; Idaed 59; Maris Fundin{070}; Mengavi{949}; SST 101{1324}; SST 107{785}; Timvera{949}; T. timopheevii derivatives{949}; Zaragoza{785}; Others{572}. v2: Bass Sr26{1450}; Combination III Sr9e{939}; Timson Sr5 Sr6{939}. tv: T. timopheevii{949}.
Sr37{939}. [SrTt2{949}]. 4BL{939}. v,tv: T. timopheevii and derivatives{484,949}; Line W{949}.
Sr38{062}. Derived from Ae. ventricosa. 2AS{062}.6Mv=2MS-6MS.6ML or 2MS-6ML.6MS{0009}. i: RL 6081 =Thatcher + Lr37. This line will carry additional genes from Thatcher. v: Moisson derivatives Mx12 and Mx22{0213}; VPM1{062}.
Sr38 is linked with Lr37 and Yr17. See Reaction to P. triticina Lr37 and P. striiformis tritici Yr17
Sr39{646}. Derived from Ae. speltoides. 2B{651}. v: RL 5711{646,651}. tv: Amphiploid RL 5347 = Ae. speltoides/T. monococcum{651}. ma: Sr39 is closely linked with Lr35{651}; A SCAR marker was developed{9923}.
Although Sr39 produces similar responses to Sr32, also derived from Ae. speltoides, recombination studies based on three crosses showed independent inheritance {646}. Sr39 segregated independently of Lr13 {651}.
Sr40{302}. Derived from T. araraticum. 2BS {302} = T2BL/2G#2S{389}. i: RL 6087 = RL 6071*7/PGR 6126; RL 6088 = RL 6071*7/PGR 6195{302}. tv: T. araraticum PGR 6126{302}; PGR 6195{302}.
Sr41{1215}. 4D{1215}. v: WDR-B1{1214}. v2: Waldron Sr5 (heterogeneous) Sr11 (heterogeneous).
Sr42{667}. 6DS{938}. v: Norin 40{938}.
Sr43. Derived from Thin. elongatum.
7DL = T7DL-7Ae#2L.7Ae#2S{657,389}.tr:KS10-2{653}.
7D = T7DS.7Ae#2L{657,389}.tr:KS23-9{653}; KS24-1{653}; KS24-2{653}.
Sr44{389}. Derived from Thin. intermedium.
T7DS-7Ai#1L.7Ai#S 7Ai#1L{389}.v:Line 86.187{939}; Several 7A-7Ai#1L translocations{0089}.
7Ai#2, 7Ai#2S.su:Group 7 alien substitution lines with 7Ai#1 and 7Ai#1S{939}. ad: TAF2 = L1{169}.
Sr45{894}. [SrD{934},SrX{1805}]. 1D{897}.1DS{894}. v: 87M66-2-1{894}; 87M66-5- 6{897}; Thatcher + Lr21, RL5406{894,934}; Various backcross derivatives developed at PBI Cobbitty{1461}. dv: Ae. tauschii RL5289{894,934}.
Tests of natural and induced mutants of P. graminis f. sp. tritici indicated that Sr45 has identical specificity to Sr21 {934}.
SrA{323}. v: SW55-1{323}; SW56-1{323}. v2: SW33-5 Sr9a Sr13{323}; SW54-3 Sr9d Sr13{323}.
SrR. ma: Several markers tightly linked with SrR were identified in{0377}.
SrTmp{1230}. v: Bai-Yu-Bao{564}; Beijing 9{564}; Beijing 11{564}; Fertodi 293{977}; Martonvasari 5{977}; Mironovska = Mironovskaya 808{068,977}; Nung-Ta 139{564}; Parker{977}; Trison{1230}; Triumph 64{841,1230,977}; Xuzhou 14{564}; Yen-An 15{564}. v2: Beijing 10 Sr5{564}.
SrWld{1230}. v2: Prospect Sr11{197}.
SrZdar{067}. 1B{067}. v: Zdar{067}.
Additional temporary designations are listed in {1230}.
Genotype lists: {323,970}.

Complex genotypes:

AC Taber: Sr2, Sr9b, Sr11, Sr12{9905}.
Centurk: Sr5 {979}, Sr6 {979}, Sr8a, Sr9a {979}, Sr17 {979}.
Chris: Sr5 {679,1371}, Sr7a {1371}, Sr9g {1371}, Sr12 {1371}.
Egret: Sr5 {939}, Sr8a {939}, Sr9b {939}, Sr12 {939}.
FKN: Sr2, Sr6, Sr7a, Sr8a {791}, Sr9b {791}.
H-44: Sr2, Sr7b {677}, Sr9d {677}, Sr17.
Hartog: Sr2 {127}, Sr8a, Sr9g, Sr12 {939}.
Hope: Sr2 {677}, Sr7b {677}, Sr9d {677}, Sr17.
Kenya Plume: Sr2 {1370}, Sr5 {1370}, Sr6 {1370}, Sr7a {1370}, Sr9b {1370}, Sr12 {1370} Sr17 {1370}.
Khapstein: Sr2, Sr7a, Sr13 {674}, Sr14 {674}.
Lawrence: Sr2, Sr7b {939}, Sr9d, Sr17.
Lerma Rojo 64: Sr2, Sr6, Sr7b {979}, Sr9a {979}.
Madden: Sr2, Sr9b, Sr11, Sr13 {842}.
Manitou: Sr5 {679}, Sr6 {679}, Sr7a, Sr9g {965}, Sr12 {939}.
Mendos: Sr7a {939}, Sr11 {879}, Sr17, Sr36.
Pasqua: Sr5, Sr6, Sr7a, Sr9b, Sr12. Gene Lr34 acted as an enhancer of APR{9905}.
PI 60599: Sr7a {689}, Sr8a, Sr9b, Sr10.
Selkirk: Sr2 {499}, Sr6 {468}, Sr7b {499}, Sr17, Sr23 {950}.
Redman: Sr2, Sr7b {939}, Sr9d {939}, Sr17.
Reliance: Sr5 {1308}, Sr16 {1238}, Sr18, Sr20.
Renown: Sr2, Sr7b {939}, Sr9d {939}, Sr17.
Roblin: Sr5, Sr7a? Sr11, Sr12.
Timgalen: Sr5 {heterogeneous} {1555}, Sr6 {1555}, Sr8a, Sr36.
Thatcher: Sr5 {1308}, Sr9g {965}, Sr12 {939}, Sr16 {1308}.
WW15 = Anza = Karamu = T4: Sr5 {939}, Sr8a {939}, Sr9b {939}, Sr12 {939}.


88.Reaction to Puccinia striiformis Westend.
Disease: Stripe rust, yellow rust.

88.1. Designated genes for resistance to stripe rust
Yr1{851}. [L{1622}]. 2A{877,1610}.2AL{940}. v: Chinese 166{851}; Corin{230}; Dalee{083}; Durin{1459}; E2025{1267}; E7700{1267}; E8594{1267}; Feng-Kang 13{1610}; Heines 110{604}; Maris Ranger{1459}; Maris Templar{1459}; Odra{073}. v2: Argent Yr3a Yr4a Yr6{1067}; Avocet (UK)Yr2 Yr6{1459}; Bounty Yr13{1459}; Fenman Yr2{1459}; Galahad Yr2 {heterogeneous} Yr14{1459}; Galahad Yr14{083}; Ibis Yr2{604}; Longbow Yr2 Yr6{083}; Mardler Yr2 Yr3a Yr4a Yr13{604,1459}; Maris Templar Yr3a Yr4a{604}; Marksman {heterogeneous} Yr2 Yr13{1459}; Mithras Yr2 Yr6{1459}; Nudif TP1 Yr3a{1431}; Nudif TP3 Yr3c{1431}; Nudif TP250 Yr6{1431}; Regina Yr2{073}; Rothwell Perdix Yr2{604}; Sportsman Yr13{1459}; Stetson Yr9{083}; Sylvia Yr2{1430}; Tadorna Yr2{1431}; Virtue Yr13{083,1459}.
A report {1267} that Kalyansona and Nadadores carried Yr1 is not correct.
Yr2{851}. Recessive {1351}. [U{1622}]. 7B{746,186,184}. v: Derius{230}; Flevina{1431}; Hana{51,58}; HD2329{1352}; Kalyansona{1351,1352}; Laketch{050}; Leda{1430}; Manella{1431}; Merlin{1622}; Odra{071}; PBW54{1352}; PBW120{1352}; Slavia{073,071}; Soissonais Desprez{851}; WG377{1352}; WH147{1352}; WL711{1352}; WL1562{1352}. v2: Avocet (U.K.) Yr1 Yr6{1459}; Brigand Yr14{083}; Cleo Yr3c{1457}; Cleo Yr3c Yr14{1431}; Fenman Yr1{1459}; Flamingo Yr6{1430}; Flevina Yr7{1430}; Galahad (heterogeneous) Yr1 Yr14{1459}; Garant Yr7{230}; Hardi Yr7{230}; Heines Kolben Yr6{611}; Heines Peko Yr6Yr25{746}; Heines VII Yr25{851}; Ibis Yr1{604}; Lely Yr7{1430}; Liberator Yr3c{1431}; Longbow Yr1 Yr6{083}; Mardler Yr1 Yr3a Yr4a Yr13{1459}; Maris Beacon Yr3b Yr4b{1459}; Maris Huntsman Yr3a Yr4a Yr13{604}; Maris Nimrod Yr13{1459}; Marksman Yr1 (heterogeneous) Yr13{1459}; Mithras Yr1 Yr6{1459}; Nautica Yr9{1430}; Norman Yr6{083}; Rapier Yr4{083}; Rothwell Perdix Yr1{604}; Sonalika YrA{1352}; Stella Yr3{1430}; Sylvia Yr1{1430}; Tadorna Yr1{1431}; Viginta Yr3a Yr4a{073,071}; Wizard (heterogeneous) Yr14{1459}; Yamhill Yr3a Yr4a{181,182, see also, 184}; Zdar Yr4a{073}.
Yr2 originally referred to a gene in Heines VII conferring resistance to European pathotypes. However, Heines VII possesses an additional resistance gene Yr25 {1351} which can be detected with a geographically wider range of pathogen isolates. Yr2 is present in Kalyansona {1351} and a range of spring wheats distributed by CIMMYT.
Yr3. Undesignated allele. v: Enkoy{050}; Vilmorin 23; Staring{1430}. v2: Minister Yr2{1430}; Stella Yr2{1430}.
Yr3a{851}. 1B{185,184}. v: Bon Fermier{1431}; Nudif TP1{1431}; Stephens{182,184}. v2: Argent Yr1 Yr4a Yr6{1067}; Cappelle-Desprez Yr4a{851}; Druchamp Yr4a{185,182, see also, 184}; Hobbit Yr4a Yr14{604}; Kinsman Yr4a Yr6{604}; Mardler Yr1 Yr2 Yr4a Yr13{1459}; Maris Huntsman Yr2 Yr4a Yr13{604}; Maris Freeman Yr4a Yr6{604}; Maris Ranger Yr4a Yr6{604}; Nord Desprez Yr4a{182,184}; Top Yr4a{230}; Viginta Yr2 Yr4a; Yamhill Yr2 Yr4a{182}; Zdar Yr4a{073,071}.
Yr3b{851}. v: Chen & Line {182} found that a second gene in Hybrid 46 - presumably this gene - was not located at the Yr3 locus. v2: Hybrid 46 Yr4b{851}.
Yr3c{851}. 1B{184}. v: Minister{851,182,184}. v2: Cleo Yr2{1430}; Maris Beacon Yr2 Yr4b{1459}.
Yr4. Undesignated allele. v: Kenya Kubangu{050}. v2: Avalon Yr14{083}; Rapier Yr2 Yr14{083}.
Yr4a{851}. 6B{185,184}. v: Vilmorin23{184}. v2: Argent Yr1 Yr3a Yr6{1067}; Cappelle-Desprez Yr3a{851}; Druchamp Yr3a{182}; Hobbit Yr3a Yr14{604}; Huntsman Yr2 Yr3a Yr13{604}; Kinsman Yr3a Yr6{604}; Maris Ranger Yr3a Yr6{604}; Maris Freeman Yr3a Yr6{604}; Mardler Yr1 Yr2 Yr3a Yr13{1459}; Nord Desprez Yr3a{182}; Top Yr3a{230}; Viginta Yr2 Yr3a{073,071}; Yamhill Yr2 Yr3a{182,185, see also, 184}; Zdar Yr3c{073,071}.
Yr4b{851}. 6B{184}. v: Avalon{1160}; Opal{1431}; Staring{1430}. v2: Hybrid 46 Yr3b{851,182, see also, 184}; Maris Beacon Yr2 Yr3b{1160,1459}; Nudif TP12 Yr3c{1431}; Stella Yr2{1430}.
Yr5{877}. 2BL{034}. v: By 33{03102}; E5557{1267}; E8510{1267}; T. spelta album{877}; Seven spelt accessions from Europe and Iran{640}. ma: Yr5 - 10.5 & 13.3cM - Xgwm501-2B{03102}.
Yr6{877}. [B{1622}]. 7B{746}.7BS{331}. v: Austerlitz{230}; Fielder{181}; Heines Kolben{1622}; Koga II{746}; Maris Dove{604}; Recital{230}; Takari{368}. v2: Argent Yr1 Yr3a Yr4a{1067}; Avocet (UK) Yr1 Yr2{1459}; Flamingo Yr2{1430}; Heines Peko Yr2{746,877}; Kinsman Yr3a Yr4a{604}; Kolben Yr2{611}; Longbow Yr1 Yr2{083,1459}; Maris Freeman Yr3a Yr4a{604}; Maris Ranger Yr3a Yr4a{604}; Mithras Yr1 Yr2{1459}; Norman Yr2{083,1459}; Nudif TP241 Yr7{1431}; Nudif TP250 Yr1{1431}; Orca Yr3c{1431}; Pavon 76 Yr7{284}; Penjamo 62 (heterogeneous) Yr18{1562}. tv: Duilio{192}; Latino{192}; Norba{192}; Quadruro{192}; Rodeo (heterogeneous){192}.
Yr7{877}. 2B{612,1429}.2BL{965}. v: Present in many hexaploid wheats with Sr9g - see{965}; Brock{083}; Lee; Nudif TP257{1431}; PBW12{1352}; Prinqual{230}; Renard{083}; Talent{230}; Tango{230}; Tommy{083}; WL2265{1352}. v2: Donata Yr9{1430}; Flevina Yr2{1431}; Garant Yr2{230}; Hardi Yr2{230}; Lely Yr2{1430}; Nudif TP241 Yr6{1431}; Pakistan 81 = Veery#5 Yr9{284}; Pavon 76 Yr6{284}; Reichersberg 2S Yr25{0010}. tv: Iumillo{965}; but not present Acme and Kubanka which also carry Sr9g{965}.
Yr8{1217,1218}. Derived from Ae. comosa.
2D = T2D-2M {1218} = T2DS-2M#1L.2M#1S{389}.tr:Chromosome 2D-2M translocations in Hobbit Sib and Maris Widgeon{1016}; Compair{1217,1218}; CS 3D/2M 3/8{967}; See also Sr34 and{967}.
2A = 2A-2M = T2AS-2M#1L.2M#1S{389}.tr:CA 2A/2M 4/2{967}.
2M-1.su:CS 2M#1(2A){967}.
Yr9{878}. Derived from S. cereale.
1B = 1RS.1BL.v:Almus{998}; Aurora{1623}; Benno{998}; Bezostaya II{998}; Clement{1430,1532}; Cougar{0267}; GR876{753}; Kavkaz{1623}; Lovrin 10{998}; Lovrin 13{998}; Mildress{1027}; Perseus{998}; Predgornaya{998}; Rawhide (heterogeneous){0267}; Salmon{998}; Sarhad 82{284}; Skorospelka 35{998}; Veery{986}; Weique{1627}; Winnetou{998}; Riebesel 47/51{878,1623}; St 2153/63{997}; Weihenstephan 1007/53{1623}. v2: Pakistan 81 = Veery#5 Yr7{284}. tv: Cando*2/Veery, KS91WGRC14{381}.
1R(1B){997,1623}.su:Burgas 2{998}; Clement{1160}; Lovrin 13{998}; Mildress{998}; Mironovskaja 10{998}; Neuzucht{1623}; Orlando{1623}; Roseana{068}; Saladin{997}; Salzmunder Bartweizen{1623}; St 14/44{998}; Weique{1627}; Wentzel W{1623}; Winnetou{1027}; Zorba{998}; See also{050}.
Chromosome status not specified.v:Baron{083}; Branka{071}; Danubia{068}; Hammer{083}; Iris{068}; Kromerzhizhskaya{1149}; Lyutestsens 15{1149}; Roxana{068}; Sabina{068}; Selekta{068}; Shtorm{1149}; Solaris{068}; Stuart{083}. v2: Stetson Yr1{083}; Nautica Yr2{1430}; Agra Yr3{068,071}; Sparta Yr3{071}; Donata Yr7{1430}. ma: Several markers tightly linked with Yr9 were identified in{0377}.
Yr10{878}. [YrVav{0262}]. 1B{641}.1BS{1002}. v: Moro{878}; PI 178383{878}; QLD709 = Janz*2/T. vavilovii{0262}; T. spelta 415{641}; T. vavilovii AUS 22498{0262}. ma: A SCAR marker was described in{0261}; QLD709 and T. spelta415, both with white glumes, failed to amplify the SCAR sequence, but both carried unique alleles at the Gli-B1 and Xpsp3000-1B loci{0262}. These differed from the Moro source of Yr10. Yr10 - 1.5cM - Gli-B1 - 1.1cM - Xpsp3000-1B{0262}.; Yr10 - 1.2cM - Xpsp3000-1B - 4.0cM - Gli-B1{0321}; Cosegregation between a RGA marker RgaYr10a and Yr10 was reported in {0376}.
Yr11. Adult plant resistance. [R11{1157}]. v: Joss Cambier{606}. v2: Heines VII Yr2 Yr25 see{970}.
Yr12. Adult plant resistance. [R12{1157}]. v: Fleurus{1158}; Frontier{1159}; Pride{1157}. v2: Armada Yr3a Yr4a{1160,081}; Mega Yr3a Yr4a{1157,1160}. v: Waggoner Yr3a Yr4a Yr6{1158}.
Yr13. Adult plant resistance. [R13{1157}]. v2: Hustler Yr1 Yr2 Yr3a Yr4a{083,1459}; Mardler Yr1 Yr2 Yr3a Yr4a{1459}; Marksman Yr1 {heterogeneous} Yr2 Yr3a Yr4a{1459}; Bounty Yr1 Yr3a Yr4a{1459}; Sportsman Yr1 Yr3a Yr4a{1459}; Virtue Yr1 Yr3a Yr4a{083,1158,1459}; Guardian Yr2{082}; Maris Huntsman Yr2 Yr3a Yr4a{083,604,1459}; Pageant Yr2 Yr3a Yr4a{082}; Professor Marchal Yr2 Yr3a Yr4a{607}; Maris Nimrod Yr2 Yr3a Yr4a{607,1157,1459}; Brigand Yr2 Yr3a Yr4a Yr14{609}; Gawain Yr2 Yr3a Yr4a Yr14{081}; Copain Yr3a Yr4a{1158}; Kinsman Yr3a Yr4a Yr6{1459}.
Yr14. Adult plant resistance. [R14{1157}]. v: Kador{1158}; Score{1157}; Wembley{610}. v2: Avalon Yr3b Yr4b{083,1459}; Brigand Yr2 Yr3a Yr4a Yr13{083,609,1459}; Galahad Yr1 Yr2 (heterogeneous) Yr3a Yr4a{083,1459}; Gawain Yr2 Yr3a Yr4a Yr13{081}; Hobbit Yr3a Yr4a{1459,1157}; Maris Bilbo Yr3a Yr4a{1157,1459}; Moulin Yr6{083}; Rapier Yr2 Yr3b Yr4b{083}; Wizard Yr2 (heterogeneous) Yr3b Yr4b{083,1459}.
Yr15{432,969}. 1BS{939,969}. v: Agrestis{0330}; Boson{0330}; Hexaploid derivatives of T. dicoccoides G-25{432,466}; V763-2312{969}; V763-254{969}. tv: T. dicoccoides G-25{432,431,466}; D447 derivatives B1, B2, B9, B10{1434}. ma: Xgwm33-1B - 5cM - Yr15{9904}; Xgwm33-1B - 4.5cM - Yr15 - 5.6cM - UBC199200 - 5.6cM Nor-B1{0110}. ma,tv: OPB131420 - 27.1 cM - Yr15 - 11.0 cM - Nor-B1{1434}.
Yr16{1598}. Adult plant resistance. 2D{1598}. v: Bersee{1604}; Cappelle-Desprez{1598}.
Yr17{062}. 2AS{062}.2AS-6Mv.6Mv = 2MS-6MS.6ML or 2MS-6ML.6MS{0009}. v: See Lr37 (Reaction to Puccinia triticina) and Sr38 (Reaction to Puccina graminis); Arche{0044}; Balthazar{0044}; Brigadier{0044}; Cordial{0044}; Eureka{0044}; Hussar{0044}; Lynx{0044}; Pernel{0044}; Renan{0044}; RL 6081{939}; Genotype list in{02105}. ma: Yr17 was closely linked to the SCAR marker SC-Y15, developed from RAPD marker OP-Y15580, and to Xpsr150-2Nv{0044}.
Yr18{1362}. 7D{1362}.7DS{324}. i: Thatcher near-isogenic lines with Lr34 including the 13 2-gene combinations reported in{434,937}. v: Jupateco 73R; Lerma Rojo 64{1375}; Nacazari 76{1375}; Tesia F 79{1375}; Tonichi S 81{1375}; Wheaton{1375}. v2: Penjamo 62 Yr6 (heterogeneous){1375}; Wheats with Lr34 (See Lr34); Others{1376}. ma: Complete linkage with Lr34{937,1362}; Ltn{1361}; and Bdv1{1363}.
Yr19{183}. [YrCom{183}]. 5B{183}. v2: Compair Yr8{183}.
Yr20{183}. [YrFie{181}]. 6D{183}. v2: Fielder Yr6{183}.
Yr21{183}. [YrLem{181}]. 1B{183}. v: Lemhi{183}.
Yr22{183}. [YrLe1{183}]. 4D{183}. v2: Lee Yr7 Yr23{183}.
Yr23{183}. [YrLe2{183}]. 6D{183}. v2: Lee Yr7 Yr22{183}.
Yr24{952}. 1BS{952}. v: Meering*3/K733/T. tauschii AUS18911{952}. tv: K733{952}.
Yr25{158}. 1D{158}. v: Carina{0010}; Hugenout{0010}; TP1295{158}; TP981{158}; Tugela{0314}; Tugela-DN{0010,0314}. v2: Heines Peko Yr2 Yr26{0010}; Reichersberg 42 Yr7{0010}.
Yr25 was predicted to be present in Strubes Dickkopf, Heines VII Yr2, Heines Peko Yr2 Yr6, Reichersberg 42 Yr7 and Clement Yr9 {158}. This prediction was confirmed for Heines VII, Heines Peko and Reichersberg 42 {0010} but the pathogen culture used in {0010} was not virulent on Clement(Yr9) or on Strubes Dickkopf where another, or a different gene, must be present.
Yr26{617}. 1BS{0285}.The earlier location of 6AS (6AL.6VS){617} is not corect.. v: Lines R43, R55, R64 and R77{0285}. tv: T. turgidum Gamma 80-1. tr: Yangmai-5{617}. ma: Yr26 - 1.9cM - Xgwm11-1B/Xgwm18-1B{0285}.
Yr27{953}. [YrSk{928}]. 2BS{953}. v: Ciano 79{953}; Selkirk{953}; Webster{953}.
Many CIMMYT wheat lines {953}. Note: Yr27 is closely linked with Lr13 (repulsion).
Yr28{1377}. Yr22 was also reported for chromosome 4D, but in the absence of an appropriate single gene stock and the unavailability of avirulent cultures in most laboratories, tests of linkage with Yr28 are unlikely in the foreseeable future. 4DS{1377}. v: Synthetic = Altar 84/T. tauschii W-219. Synthetic/Opata 85 SSD population. Genotype lists: {1325,970}. dv: T. tauschii W-219{1377}. ma: Close association with Xmwg634-4DS{1377}.
Yr29{0119}. Adult plant resistance {0119}. 1BL{0119}. v2: Pavon F76 Yr6 Yr7 Yr30{0119}; Yr29 is completely linked with Lr46. See Lr46{0119}.
Yr30{0120}. Adult plant resistance {0120}. 3BS{0120}. v: Opata 85{0120}; Parula{0120}. v2: Inia 66 YrA{0120}; Pavon F76 Yr6 Yr7 Yr29{0120}; Yr30 is closely linked with Sr2 and Lr27{0120}.
Yr31{0325}. 2BS. v: Pastor{0325}. ma: Recombination values: Yr31 - Yr27 0.148; Yr31 - Lr23 0.295; Yr27 - Lr23 0.131{0325}.
Yr32. [YrCv{939}]. 2BS{939}. i: Avocet S*4/Carstens V{970}; Cook*6/Carstens V{970}. v: Caribo{1430}; Cyrano{1430}; Okapi{1430}. v2: Felix Yr3{1430}; Zdar Yr3a Yr4a{067}.
Sources of additional genes for seedling (designated "12") and adult resistances ("13", "14", "15") are listed in {1430}.


88.2. Temporarily designated gene for resistance to stripe rust
YrA. Refers to a phenotype specificity that appears to be controlled by complementary genes {1563}. v: Avocet* {*= heterogeneous}; Anza = Karamu = Mexicani =T4 = WW15; Banks*; Condor*; Cocamba; Egret*; Inia 66; Lerma Rojo 64; Lerma Rojo 64A; Nainari 60; Nuri 70; Sanda 73; Sonalika; Zaminder 80. v2: Condor selection P44 Yr6*; Pari 73 Yr6; Saric 70 Yr6; Yecora 70 Yr6{1563}.
YrCle{186}. 4B{186}. v2: Clement Yr9{186}.
YrD{185}. 6A{185}. v: Druchamp{185,185}.
YrDa1{186}. 1A{186}. v2: Daws YrDa2{186}.
YrDa2{186}. 5D{186}. v2: Daws YrDa1{186}.
YrDru{184,185}. 5B{184}.6B{185}. v: Druchamp{184,185}.
YrDru2{184}.Druchamp {184}. 6A{184}.
YrH46. Hybrid 46 {184}.
Not the same gene as YrDru2 {184}. 6A{184}.
YrH52{0003}. 1BS{0003}. tv: T. dicocoides H52{0003}. ma: distal ...Yr15 - 9.6 cM - YrH52 - 1.4cM - Nor-B1 - 0.8cM - Xgwm264a - 0.6cM - Xgwm18{0003}; Xgmw273a - 2.7cM - YrH52 - 1.3cM - Xgwm413/Nor1...centromere{0108}.
YrHVII{186}. 4A{186}. v2: Heines VII Yr2{186}.
YrMin. Minister {184}. 4A{184}.
YrMor{186}. 4B{186}. v2: Moro Yr10{186}. ma: The development of an STS marker, derived from an AFLP fragment, that co-segregates with YrMor was reported in {0357}.
YrND. Nord Desprez {184}.
May be the same as YrMin {184}. 4A{184}.
YrS{185}. 3B{185}. v: Stephens{185}.
YrSte{184}. 2B{184}. v: Stephens{184}.
YrSte2.Stephens {184}. 3B{184}.
YrTye{186}. 6D{186}. v: Tyee{186}.
YrTr1{186}. 6D{186}. v2: Tres YrTr2{186}.
YrTr2{186}. 3A{186}. v2: Tres YrTr1{186}.
YrYam{184,185}. 4B{185}. v: Yamhill{185}.
YrV23. Vilmorin {184}.
Allelic but not the same as YrSte {184}. 2B{184}.
Yrns-B1{0033}. 3BS{0033}. v: Lgst.79-74{0033}. ma: Xgwm493 (distal) - 21 cM - Yrns-B1{0033}.
North American workers {181,186,184} allocated a number of temporary designations to uncatalogued genes detected with North American P. striiformis accessions. Druchamp, Yamhill and Stephens were reported to carry "Yr3a or Yr4a" because these genes could not be distinguished with the cultures that were used.


88.3. Stripe rust QTLs
QTL: In the ITMI mapping population, QTLs were found in 2BS, 7DS, and possibly 5A, 3D and 6D {0287}. The 2BS gene is likely to be Yr27. In Camp Remy/Michigan Amber, QTLs were found in 2AL and 2BL {0287}.

Two QTLs in Camp Remy/Michigan Amber were located on chromosome 2BL (QYR1, LOD score 12) and 2AL (QYR2, LOD 2.2) {0304}. Four QTLs were scored in the ITMI population. The most effective (QYR3, LOD 7.4) on chromosome 2BS was probably Yr27, the others were located in 7DS (QYR4, LOD 3.4). 5A (QYR5, LOD 2.8), 3D(QYR6, LOD 2.8) and 6DL(QYR7, LOD 2.4) {0304}.




89.Reaction to Puccinia triticina
Disease: Brown rust, leaf rust.

89.1. Genes for resistance
Lr1{047}. 1B{1409}.5D{954}.5DL{945}. i: Centenario/6*Thatcher{317}; Malakoff/6*Prelude{317}; Wichita*4/Malakoff{613}. v: Centenario{317}; Chicora 'S'{143}; Daws (heterogeneous){1019}; Dirkwin{1019}; Glenlea{1255,976}; Halle 9H37{074}; Hyslop{1019}; Luke {heterogeneous}{1019}; Malakoff{047}; McDermid{1019}; Mexico 120{933}; Newton{143,1024,1023}; Norco{1019}; Shabati Sonora{842}; Sonora 64{842}; Tarsa{842}; Uruguay{954}; Walliday{1019}. v2: Blueboy Lr10{143}; Blueboy II Lr10 Lr24{143}; Erythrospermum 142 and 953 Lr3{074}; Laura Lr10 Lr34{712}; Norka Lr20{1552}; Plainsman V Lr3{1024}; Suneca Lr13{485}. ma: Co-seg. with Xpsr567-5D and Xglk621-5D in a Frisal/Lr1 resistant line. pTAG621 was converted to a diagnostic STS{354}.
Lr2. 1B{1409}.2DS{843,942}.
Lr2a{320}. [Lr2{047}]. i: Prelude*6/Webster{320}; Red Bobs*6/Webster{320}; Webster/6*Thatcher{306}; Wichita*4/Webster{613}. v: Eureka CI 17738{143}; Festiguay{843}; Webster{047}. v2: Alex Lr10{976}; Guard Lr10{976}; James Lr10{976}; Len Lr10{976}; Marshall Lr10{976}; Mediterranean W1728 Lr3{1369}; Shield Lr3 Lr10{198}; Waldron Lr10{143}.
Lr2b{320}. [Lr22{1409}]. i: Prelude*6/Carina{320}; Red Bobs*6/Carina{320}; Thatcher*6/Carina{320}; Wichita/4*Carina{613}. v: Carina{613}.
Lr2c{320}. [Lr23{1409}]. i: Prelude*5/Brevit{320}; Prelude*6/Loros{320}; Red Bobs*6/Brevit{320}; Red Bobs*6/Loros{320}; Thatcher*4/Brevit{320}; Thatcher*6/Loros{320}; Wichita*4/Brevit{613}; Wichita*4/Loros{613}. v: Brevit{613}; Loros{317,1257}.
Lr3{047}. Because Lr3 appears to be a complex locus {486} Democrat and Democrat/6* Thatcher should be accepted as standards. There is evidence to suggest that the allele in Mentana, and therefore many derivatives, is Lr3b {939}. If this is correct, many genotypes listed under Lr3a are likely to be Lr3b
Lr3a. [Lr3{047}]. 6B{549}.6BL{939}. i: Democrat/6*Thatcher{318}; Wichita*4/ Mediterranean{613}. v: Belocerkovskaja 289{074}; Bennett{1024}; Democrat{047}; Fertodi 293{074}; Gage{1024}; Hana{068}; Homestead{1024}; Ilyitchovka{075}; Juna{075}; Jubilejne{068}; Kawvale{143}; Lancota{1024}; Mara{068}; Mediterranean{047}; Mediterranean W3732{1369}; Mentana{842}; Mironovskaya 264 & 808{074}; Odra{075}; Osetinskaya{074}; Ottawa{143}; Pawnee{1408}; Ponca{143}; Rannaja 12{074}; Shawnee{143}; Shirahada{842}; Skorospelka 3b{074}; Sledkovicova K1004{074}; Viginta{068}; Warrior{143,1024}; Yubileynaya{075}. v2: Amika Lr26{076}; Bezostaya 1 Lr34{074}; Bowie Lr14b{319}; Erythrospermum 142 & 953 Lr1{074}; Istra Lr26{076}; Mediterranean W1728 Lr2a{1369}; Plainsman V Lr1{1024}; Shield Lr2a Lr10{198}; Solaris Lr26{076}; See also{069}. ma: Co-segregation with Xmwg798-6B{9921}.
Lr3b{486}. [Lr3bg{486}]. i: Thatcher*6/Bage; RL6094 = Tc*6/T6{307}. v: Bage{486}. v2: T6 Lr16{307}.
Lr3c{486}. [Lr3ka{486}]. i: Tc*6/Klein Aniversario. v: Klein Aniversario{486}.
Lr4, Lr5, Lr6, Lr7, Lr8{365}. v: Purdue Selection 3369-61-1-10 = Waban{365}; Not available as separate single-gene lines.
Lr9{1408}. Derived from Ae. umbellulata. 6B{954,1296,1299}.6BL = T6BS.6BL-6U#1L{389}. i: T47 = Transfer = CS + Lr9{1408}; Thatcher*6/Transfer; Wichita*4/ Transfer{613}. v: Abe{143}; Arthur 71{1320,1024}; Clemson 201{465}; McNair 701 & 2203{143}; PI 468940{1439}; Riley 67{1320,1024}; Sullivan{1110}. v2: Oasis Lr11{1109}. ma: Co-seg with XksuD27-6B{048}; co-seg with Xmwg684-6B and STS Xsfr1{1272}; Lr9 - 8 cM - Xpsr546-6B{1272}; The structures of additional translocations are given in{389}.
Lr10{199}. [LrL{031}]. 1A{312,546}.1AS{939}. i: Exchange/6*Thatcher{306}; Gabo/6*Thatcher{306}; Lee/6*Thatcher{306}; Selkirk/6*Thatcher{306}; Timstein/6*Thatcher{306}. s: CS*5/Timstein 1A{939}; CS/7*Kenya Farmer 1A{939}. v: Centurk{1024}; Centurk 78{1024}; Concho{143}; Federation{939}; Mayo 52{031}; Mayo 54{031}; Parker{546,1024}; Rocky{1024}; Scout 66{02101}; Sinton{1256}; Tascosa{143}; TAM-105{055}; Unknown accessions{208}; See also{0337}. v2: Blueboy Lr1{143}; Blueboy II Lr1 Lr24{143}; Laura Lr1 Lr34{712}; Alex Lr2a{976}; Guard Lr2a{976}; James Lr2a{976}; Kenya Farmer Lr23{939}; Len Lr2a{976}; Marshall Lr2a{976}; Waldron Lr2a{143}; Shield Lr2a Lr3{198}; Exchange Lr12 Lr16{031}; Era Lr13{143}; Selkirk Lr14a Lr16{031,199}; Warden Lr16{031}; Gabo Lr23{031}; Lee Lr23{031}; Timstein Lr23{031}; Parker 76 Lr24{143}. ma: Xcdo426-1A - 5.1cM - Lr10{1058}; Lr10 - 8cM - Glu-A3{355}; cosegregation with Xsfr1(Lrk10-1A) and Xsfrp1(Lrk10-1A){1270}; complete linkage with Xsfr1(Lrk10-1A), which encodes a protein kinase{639}.
Lrk10. A receptor-like kinase. The locus Xsfr1(Lrk10)-1A, detected by the probe Lrk10, is completely linked with Lr10 in chromosome 1AS {356}. The gene encodes a receptor-like kinase with extracellular and kinase domains {0297}. Using probe pLrk10-A, developed from the extracellular domain, 6 homologues were found in chromosomes 1A (1), 1B (3) and 1D (2) as well as group 1 chromosomes of T. monococcum, Ae. tauschii and barley {0296,0294}. Probes based on the kinase domain identified further homologues in chromosomes 3AS and 3BS as well as the corresponding regions in rice and maize {0294}. Both orthologous and paralogous evolution were suggested.
Lr11{1409}. 2A{1409}. i: Thatcher*6/Hussar{306}; Wichita*4/Hussar{613}. v: Bulgaria 88{142}; Hart{1024}; Hazen{049}; Hussar{1409}. v2: Karl 92 Lr3 Lr10{02101}; Oasis Lr9{143}.
Lr12{326}. Adult pl