Note: Levy and Feldman {797} studied the inheritance of more than 20 morphological and biochemical traits in crosses of four T. dicoccoides lines and T. durum. Similarly, Kuspira et al. {744} studied 12 qualitative characters in T. monococcum. The symbols applied to the characters examined in these studies are not being reserved and listed in the Catalogue. However, both studies should serve as bases for future work.

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 Sl 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. Squarehead/spelt

Q {881}. [k {1550}]. 5AL {1293}. v: Common wheats. CS.

q {881}. [K {1550}] v: Macha wheats; spelt wheats; vavilovi wheats. s: CS*8/White Spring

Spelt {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}.

2. Club

C {1517}. [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.

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.

3. Sphaerococcum

s1. [sp1{1286}]. 3D {1292}; 3DS {1193,1194}; 3DL {692}. i: S-615*11/T. sphaerococcum

var. rotundatum {1500}. s: CS*7/T. sphaerococcumrubiginosum 3D {1304}. v: Sphaerococcum wheats.

s2. [sp2 {1286}]. Partially dominant{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 factor controlling the sphaerococcum character in tetraploid wheat, Joppa {621} using the same stock found that two recessive genes were necessary to produce this phenotype.

4. Branched spike

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

bh {665}. 2AS {665}. dv: PI 349056 {665}.

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.

P {911}. [Eg {922}]. 7AL {922,1547}, 7A or 7B (based on linkage of 0.2 with a gene for red coleoptile {911}). i: Saratovskaya29*8//Novsibirskaya 67*2/T.polonicum {922}. itv: P-LD222 = LD222*11/T. turgidum var polonicum {1546,1547}.

Accumulation of Abscisic Acid

A QTL was mapped on 5AL between Xpsr575-5A {proximal} and Xpsr426-5A {distal} {1180}.

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}.

Anthocyanin Pigmentation

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}.

2. Purple/Red auricles. Purple leaf base

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

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.

Genes controlling purple leaf base were:
Ra2 {983}. 4B {983}.
Ra3 {983}. 6B {983}.
An5 {983}. 5R {983}.

3. Red/purple coleoptiles.

Rc1. [R {401}]. 7A {769,1293}. s: CS*6/Hope 7A {1293}. v: Hope Rc2.

Rc2. [R2 {401}]. 7B {742},7BS {401,769}. s:CS*6/Hope 7B {769}. v: Hope Rc1.

Rc3. 7D {596}, 7DS {1241,1444}. 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: Rc3 (distal) - 3 cM - Xpsr108-7D {180}.

Tahir & Tsunewaki {1453} reported that T. spelta var. duhamelianum carries genes promoting pigmentation in chromosomes 7A and 7D and genes suppressing pigmentation in 2A, 2B, 2D, 3B and 6A. Sutka {1444} reported a fourth factor in chromosome 6B and suppressors in 2A, 2B, 2D, 4B and 6A.

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. 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 Rc2 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.


1. Dominant Inhibitors

1.1 Hooded

Hd {1551}. 4AS {1195,1293}. i: S-615*11/CS {1500}. v: Chinese Spring B2 {1293}.

hd. s: CS*6/Hope 4A; CS*5/Thatcher 4A; CS*6/Timstein 4A.

1.2 Tipped 1

B1 {1551}. 5AL {1293}. i: S-615*11/Jones Fife {1500}. v: Timstein {741}; Redman {160}.

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.

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.

1.4 Awned.

hd b1 b2. Bearded or fully awned genotype:

Genotypes Hd B2 (e.g., Chinese Spring) and B1 B2 (e.g., Federation) are awnless. Presumably Hd B1 is awnless. Watkins & Ellerton {1551} noted the probability of a third allele "b1a" leading to a half-awned condition, and in discussion they consider the possibility of a similar third allele at the B2 locus. In view of more recent cytogenetic analyses, it seems that the half-awned condition could result from epistatic interactions between the alleles B1 and/or B2 and various promotor genes.

Although hooded, half-awned, tip-awned and awnless variants occur among tetraploid wheats, these are relatively infrequent. It has not been established with certainty that the above inhibitors are involved.

The inhibitor alleles have a pleiotropic effect on glume-beak shape {1348}. Acuminate beak is associated with full beardedness and occurs only in b1 b2 types. B2 reduces beak length producing an acute beak shape. B1 reduces beak length producing an obtuse beak shape. In this effect B1 is epistatic to B2.

2. Promotors

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

3. Smooth Awns

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

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 Bsthe 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 have been 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

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.

Ba1 {643}. [Ba {643}]. Derived from Elytrigia pontica (2n=70). 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}.

For review see {1643}.

Boron Tolerance

Genes controlling tolerance to high concentrations of soil boron act additively.
Bo1 {1111,1113}. 7B {177}. v:  Halberd Bo2Bo3.
Bo2 {1111,1113}.   v:  (W1*MMC). Warigal Bo3. Halberd Bo1 Bo3.
Bo3 {1111,1113}.   v:  Warigal Bo2. HalberdBo1 Bo2.

Very sensitive genotype: Kenya Farmer bo1 bo2 bo3.

Cadmium Uptake

Low Cadmium Uptake
Cdu1 {962}. [Cdu1 {1128}]. dv:  Biodur {1128}; Hercules {1128}; Nile {1128}.
cdu1 {962}. [cdu 1 {1128}]. dv:  Kyle {1128}.

ma: Cdu1 - 4.6cM - OPC-20 {1128}; Cdu1 - 21.2 cM - UBC-180 {1128}.

Chlorophyll Abnormalities

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.

2. Chlorina
Cn-A1    7A {1132};


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. {1545}. (CS type)
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}.

3. Striato-virescens

A mutant of this type was described {376} but has been lost.

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}.

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}.
ad: CS+5R {463}.
su: CS 5R{5D} {463}.
  5BS = T5BS.5RL. v: Sears' stock HN-2 {464}. Backcross derivatives to Warigal and Timgalen {464}.

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}.

Crossability with Rye andHordeum and Aegilops spp.

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 kr1is 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}.

Some genotypes:
kr1 kr2. v: Chinese 446 {790}; Chinese Spring {762,1216,1025}; Martonvársári 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: Martonvársáari 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}.