Frost Resistance

Fr1 {1446}. 5AL {1446}. v: Hobbit.
ma: was mapped to the mid-region of 5AL, 2.1 cM distal from Xcdo504-5A and Xwg644-5A and proximal to Xpsr426-5A {419}.

Gametocidal Genes

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-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}.

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 Th. distichum {889,892} is probably the same as that from Th. elongatum {1162}.
 
Sd1 {1647}. 7D {1647}. v: Agatha Sd2 {1647,1161}.
Sd2 {1161}. 7BL {1163}. v: 88M22-149 {1163,1161}.

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.

2. Suppression of Gametocidal Genes

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

igc1. v: Chinese Spring {1483,1488}. Forty wheats are listed in {1483} and/or {1488}.

Gibberellic Acid Response (insensitivity)

Gai1. [GAI1 {565,1246}]. 4B {406}, 4BS{980}. i: See {408}. v: Norin 10 Der. {565}.List in {407}.

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 wheats 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} consider Gai1 and Rht-B1b, Gai2 and Rht-D1b, Gai3 and Rht-B1c to be pleiotropic genes.

Glaucousness (Waxiness/Glossiness)

Glaucousness is the visual expression of waxiness on stems and leaves and is the preferred term. Plants may be either waxy, glaucous (the normal situation), waxy, non-glaucous (e.g., Mentana) or low wax, non-glaucous (e.g., AUS 2598). The expression of glaucousness depends on the arrangement of wax deposits rather than the amount of wax {603}.

This phenotype was originally thought to be controlled by a series of wax-producing genes and an independent series of inhibitors {e.g., 912,1493}, but later work {1432} indicated that the wax-producing genes and the inhibitors were allelic. Hence recessive alleles and dominant alleles may result in non-glaucousness.

Orthologous loci occur in barley chromosome 2HS (gs1, gs6, gs8) {467}, rye chromosome 7RL (wa1) {725} and maize (gl2) {211}.

W1. 2BS {267,1493}. v: Chinese Spring {1493}. itv: LD222*11/T. Turgidum var. pyramidale recognitum {1546}.

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

W2. v: No genetic stock specified.

W2a. dv: Glaucous forms of T. tauschii.
W2b. v: Chinese Spring - weak hypomorph recognized at increased dosage {1432}.
Epistatic inhibitors of glaucousness

W1I. [I1-W {1493}]. 2BS {267,272}. v: Cornell Selection 5075 {595}. tv: Long Kernel {595}; T. dicoccoides {275}.
W2I. [I2-W {1493}]. 2DS {1240,1493,1060}. v: Non-glaucous forms of T. tauschii.
W3I. [{277}]. I3-W {277}. 1BL {277}. tv:T. turgidum var. dicoccoides.

Note: Each inhibitor inhibits both genes for glaucousness.

Glume Colour

1. Red (brown/bronze)

The majority of studies report that a single dominant gene determines 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: Federation 41 {1517}; Highbury {1121}; Red Egyptian {1304}. tv: Ward {792}.

Rg2. Derived from Aegilops squarrosa. 1DL {769,1241}, 1DS. 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.

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 T. tauschii and synthetic hexaploid wheats. v: K-28535 {729}. i: ANK-23 = Novosibirskaya 67*10/ K-28535 {729}.

Rg3 {924,562}. 1AS {924,562}. i: Saratovskaya 29*3 //F2 CS mono1A/Strela {924}.

v: L'govskaya-47 {1405}. Strela Rg1 {924}. Sobko and Sozinov {1405} 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 {624}.
2. Black

Bg. 1A {1304}, {282}. s: CS*7/Indian 1A {1304}. dv: G1777 {282}; G3116 {282}. ma: Bg and Nor9 co-segregated in T. monococcum {282}3.

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 a non-black glumes.

A single factor for blavk gumes 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.

3. Pseudo-black chaff

Pbc. 3B {742}. s: CS*6/Hope 3B {742}; CS*6/Ciano 5B {939}.

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 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.

4. A black-striped-glume phenotype was reported in group dicoccon.v: E4225 {1417}.

5. An inhibitor of glume pigment was reported on chromosome 3A {106}.

6. Chocolate Chaff

cc {719}. 7B {719}. tv: Langdon mutant {719}; PI 349056 {665}. 7BS {665}. dv: CBC-CDd1 {665}.

Chocolate chaff phenotype was suppressed by a gene(s) in chromosome 7D {719}.

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 are determined by three complementary genes designated Bla1, Bla2, Bla3. Bla1 was located in chromosome 1A and linked with Gld 1A (= Gli-A1) and Hg.

Grain Hardness/Endosperm Texture

Grain hardness or endosperm texture signficantly influences flour milling extraction, flour properties and end use. Variation at a single locus was erported by Symes {1452}. Using a Brabender laboratory miller to produce bran samples, Law et al. {777} also showed that grain hardness was controlled by alleles at a single locus. The dominant allele Ha controlling softness was present in Chinese Spring and the allele for hardness, ha was present in Hope. More recently, it was shown that soft wheat possesses greater levels of a 15,000kD starch granule protein, friabilin, than hard wheats {470}.

Ha {777}. 5DS {777}. Soft phenotype. v: Chinese Spring {777}; Cappelle Desprez {470}; Heron {1452,470}.
ha {777}. Hard phenotype. s: CS*6/Cheyenne 5D {915}; CS*6/Hope 5D {777}. Cappelle Desprez*/Besostaya 5D {470}. v: Falcon {1452,470}; Holdfast {470}. 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.

The addition of King II rye chromosome 5R converted Holdfast wheat from hard to soft {470}. A 14,500kD rye analogue was also isolated from 6x triticales which have soft texture {470}. All ryes have soft texture.

Two genes for grain hardness were reported in {055}.

Grain softness proteins (GSPs), or friabilins, are 15-kDa polypeptides that are associated with purified starch granules from soft-grained wheat. Friabilin consists mainly of puroindoline a and puroindoline b and, although both soft and hard wheats possess them, distinction between the two textural types depends upon the manner in which the friabilin binds to starch. See Puroindoline (Proteins 5. 8).

Gsp-A1 {614}. [GSP {614}]. 5A {614}. v: CS {614}.
Gsp-B1 {614}. [GSP {614}]. 5B {614}. v: CS {614}.
Gsp-D1 {614}. [GSP {614}]. 5DS {614}. v: CS {614}. ma: Cosegregation of Gsp-D1 and Ha {614}.

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}. 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}. 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}. v: 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}. v: Cappelle-Desprez D2 {1595}; Cheyenne D2 {1000}. Brevor D2 {1000}.

d1d2d3d4. v: Chinese Spring {534,1000}.

Genotypes 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.

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}.

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}.

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}.

hg1 {1405}. v: Ulyanovkn {1405}; Pionerskaya {715,1405}.

Hairy Leaf

Hl {884}. 4B {884}, 4BL {760}. Weakly hairy. 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}.

hl. v: Chinese Spring {884}.

Kuspira et al. {744} provided evidence for at least three alleles at an Hl locus in T. monococcum. With the reversal of chromosome designations for 4A and 4B, this locus in T. monococcum cannot be allelic with the gene in T. aestivum.

Hairy Neck/Pubescent Peduncle

Translocation from rye chromosome 5R to Chinese Spring.
 
Hp.   4BL {T4B.5R} {274,275}. i: S-615*11/Chinese Spring {1500}.
    5BS {T5B-5R} {1298}. v: HN-2 (CS type) {1298}.
    6D {T6D-5R} {1298}. v: HN-1(CS type) {1298}.

Hairy Node/Pubescent Node

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.

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 the B1 own; own inhibitoe. Observations on 5A trisomics and telosomics of Chinese Spring confirm 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.

Multiple alleles were reported in T. monococcum {744}.

Hairy Leaf Sheath

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}.

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

Height

Ht is the general symbol.

Reduced Height : GA-insensitive

Rht1, see Rht-B1b; Rht2, see RhtD1b; Rht3, see Rht-B1c; Rht10, see Rht-D1c.

Rht-1. {371}.

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

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

Rht-B1b {116}. [Rht1, Sd1 {015}]. Partially recessive {024}, recessive {357}, semi-dominant {408}. i: See {408,414}. v: Frontier {1597}; Guardian {1597}; Selection 14-53/Burt, 5 {015}; Siete Cerros {407}; Wren {1174}; WW15 {407}. 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}.

Rht-B1c {116}. [Rht3, Sd3{565}]. Semi-dominant {1040}. 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}. [Rht1S {1599}]. Semi-dominant {1599,116}. 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.

RhtB1e {116}. [RhtKrasnodari 1 {452}, Rht1{B-dw} {1600}]. v: Krasnodari 1 (a spontaneous GA-insensitive offtype of Bezostaya 1 {1600}).

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

Rht-D1 {116}. 4D {411,583,1544}, 4DS {980,1266, see also, 116}. ma: Xpsr1871(Pki)-4D - 4cM - Rht-D1 - 6 cM - Xubc821(PhyA)-4D {410}.

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

Rht-D1b {116}. [Rht2, Sd2 {015}]. Partially recessive {024}, recessive {357}, semi-dominant {408}. 4D {411}, 4DS {980}. i: See {408,414}. 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}.

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

Rht-D1d {116}. [RhtAi-bian 1a {115}]. Semi-dominant {116}. v: Ai-bian 1a (spontaneous mutant of Ai-bian 1) {115}.
Note: There has been no report of a break in the linkage of the Rht-1 and Gai phenotypes.
Further genotype lists in {402,1382,1612,1613}.

Reduced Height : GA-sensitive

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}. Norin 10-Brevor, 14 Rht-B1b Rht-D1b {569}.

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

Rht8. 2D {772,1601,1598}, 2DL. s: Cappelle-Desprez*/ Mara 2D {1601}. v: Novasadska Rana 1 {1604}; Sava {1601,414}, Akakomugi Rht9 {1191}; Mara Rht9 {1191}. ma: Xgwm484-2D (proximal) - 19.9 cM - Rht8 - 0.6 cM - Xgwm261-2D (distal) {727}.

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 the gene B1 and closely linked to b-Amy-A1 {1606}. Xgwm291-5A - 5.4cM - Rht12 {726}.

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}.

Börner et al. {116} found no evidence of orthologous GA-insensitive genes in rye, but reviewed evidence for orthologous GA-insensitive gene

Herbicide Response

1. Difenzoquat Insensitivity

Insensitivity/resistance is dominant.
 
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.

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.

3. Chlortoluron Insensitivity

Insensitivity/resistance is dominant {1402}.

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}.
matv: Nor2 (6BS) - 2.7cM - Su1 {1401} - 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.

Hybrid Weakness

1. Hybrid Necrosis. [Progressive lethal necrosis {155}; Firing {971}].

Complementary dominant genes. Descriptive alleles w (weak), m (medium) and s (strong) were allocated by Hermsen {532}. Phenotype is affected by modifying genes (and/or genetic back-ground) and environment {566}. According to Dhaliwal et al. {257} progressive necrosis is suppressed at 28oC.

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

Ne1w. 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. v: Federation group {532}: Cadia {971}; Cleveland {971}. v: Minister group {532}: Rieti group {532}: Mentana {532}; Mara {532}.

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}; Lutescens 1163 {1264}; Marquillo {155,532,550}; 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.

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 - Atlas 40 {532}; Wheats possessing Lr13 {939}, e. g. Manitou {939}.

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}. [Le2 {155,550,1491}, F {971}]. 2B {1491}, 2BS {1085}. v: Vakka {532}; Varma {532}.

Ne2m?{530}. v: Barleta group {532}: South American wheats - 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}. Noé group {532}: Vilmorin 27 {532}.

Unknown Ne2 allele {532}. v: Harvest Queen {550}. tv: Acme {532};
v: 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}.

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,
1497,1502,1503,1512,1505,1506,1507,1508,1509,1510,1630,1631,1632,1633,1635,1637,1638,
1639}.

Rye line 1R136-2 carries Ner1 {1210} that complements wheat geneNe2 {1289,1210} and rye gene Ner2 {1210} to produce necrosis. Rye lines L155 and L256 carry Ner2 {1210} that complements Ne1 {630,1210} and Ner1 {1210}.

Ner1{1210}. 5RL {1211}. al: S. cereale 1R136-2 {1210}.
Ner2{1210}. 7RL {1211}. al: S. cereale L155, L256 {1210}.

2. Hybrid Chlorosis Type 1 (Red hybrid chlorosis)

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}.

Complementary dominant genes.

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}.

Iron Deficiency

Fe1 {926}. 7DL {927}. v: Saratovskaya 29 {926}.
Fe2 {926}. 7BS {927}. v: CS {927}.

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 the liguleless Chinese Spring derivative {939}.

lg1 {047}. 2B {942}. v: Eligulate W1342 lg2 {942}. Partial backcross derivative of Chinese Spring {939}. tv: Liguleless durum selection {939}.
lg2. 2D {942}. v: Eligulate W1342 lg1 {942}. Partial backcross derivative of Chinese Spring {939}.

Because diploid wheats are liguled, polyploid wheats presumably carry a third recessive factor in chromosome 2A.

Leaf Tip Necrosis

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

Male Sterility

Chromosomal

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

ms1a {268}. v: Briggle's Chancellor Derivative {268}; Pugsley's Male Sterile {268}.
ms1b {268}. 4B {268}, 4BS {064}. v: Probus mutant {268}.
ms1c {064}. v: Cornerstone {064}.
ms2 {806}. [Ta1 {240}]. Dominant allele for sterility. 4DS {806}. v: Taigu = Line 223 {240,807,806}. ms2 confers sterility when present in octaploid triticale {597}.

ms3 {872}. 5AS {872}. Dominant allele for sterility. i: Chris derivative {872}; KS87UP9 {219}.

Megasporogenesis

Control of Megasporogenesis

Msg {625}. 7AS {625}. tv: Langdon {625}.

Meiotic Characters

1. Low-Temperature Pairing

ltp {527}. v: Chinese Spring.

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.

2. Pairing homoeologous

Ph1 {1537}. 5BL {1301}.

ph1a - Not applicable - see  ph2b {1303}.
ph1b {1301}. v: Sears' high pairing mutant {1301}.
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}. Ph1 was physically mapped in 5BL to fraction length 0.55, bracketed by deletions 5BL-1 and ph1b {446}.

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}.
3. Inhibitor of pairing homoeologous

Ph1I al: Aegilops speltoides {1218,439}.

Nitrate Reductase Activity
 
Nra {424} v: UC44-111 {424}.
nra {424} v: Anza {424}.

Nuclear-Cytoplasmic Compatability Enhancers

scs {869}. Derived from T. timopheevi {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.

Nucleolus Organizer Regions

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
 
Nor1a and Nor2a v: Maris is Huntsman {1399}.
Nor1b and Nor2b v: Bezostaya 1 {1399}.
Nor1c and Nor2c v: Cappelle-Desprez, Maris Ranger {1399}.

may or may not be equivalent to those described below.
 
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.
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.
Nor-H1. [Nor-I1 {794}]. 1HS {794}. dv: Sultan barley.
Nor-R1.   1RS {039}. ad: CS/Imperial.
Nor-S1.   1SS {294}. al: Ae. speltoides.
Nor-U1.   1U {906}. su: CS/Ae. umbellulata.
Nor-V1 {241}.    1V {241}. ad: CS/D. villosum.
Nor-B2. [Nor2 {1120}]. 6BS{1041,221,366,835}. v: CS.
Nor-B2a. {918}
   [CS6B {288}]. v: Chinese Spring {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.
Nor-B2h {918}.
     v: Yaktana.
Nor-B2i {918}.
     v: Maris Huntsman; Thatcher.
Nor-E2.6ES {294}.     ad: CS/E. elongata.
Nor-G2.   6G {578}. tv: T. timopheevi IPSR {PBI} No. 1.
Nor-H2. [Rnr1 {1248}]. 6H {1070,039,1248}, 6HS {794}. al: Clipper barley {039}; Sultan barley {794}.
Nor-S2.   6SS {294}. al: Ae. speltoides.
       
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.
Nor-E3.   5ES {294}. ad: CS/E. elongata.
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.
Nor-D4 {1042}.
 7DL {1042}.
7DL {1042}.		 v: CS.
dv: Ae squarrosa.
Nor-H4. [Nor-I4 {794}].  7HS {794,793}. al: Sultan barley.
Nor-H5. [Nor-I5 {794}].  2HS {794,793}. al: Sultan barley.
Nor-B6 {601}.
 1BL {601}. v: CS; Cheyenne, Wichita.
tv: Langdon.
Nor-A7 {601}.
 5AL {601}. v: CS; Cheyenne, Wichita.
tv: Langdon.
Nor-D8 {601}.
   3DS {601}. v: Witchita.

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.

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}. High osmoregulation. 7A{1030}, 7AS {1031}. v: Chinese Spring, Condor, Songlen, Takari {1030}.
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}.

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.