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 E.V. Metakovsky, N.I. Vavilov Institute of General Genetics, Moscow {988}. This nomenclature is reproduced below. Where two cultivars are given as prototypes for an allele, the first named is from the USSR and the second from elsewhere.

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.

A number of novel gliadin alleles were reported in {991}; they will be included in the next supplement to the catalogue.

Note: The catalogue entries reproduced here only refer to alleles in T. aestivum; there is, however, an enormous amount of 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).

The Gli-1 loci may be recognised by probes pcP387 {372} and pTag1436 {065}, and by specific microsatellite primers {252}.
 

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: CS.
Gli-A1b {988}.			v: Bezostaya 1, Mercia.
Gli-A1c {988}.			v: Ukrainka.
Gli-A1d {988}.			v: Dankowska.
Gli-A1e {988}.			v: Falchetto.
Gli-A1f {988}.			v: Mironovskaya 808, Maris Freeman.
Gli-A1g {988}.			v: Gabo.
Gli-A1h {988}.			v: Sadovo I.
Gli-A1i {988}.			v: Saratovskaya 36.
Gli-A1j {988}.			v: Lutescens 62.
Gli-A1k {988}.			v: Skala.
Gli-A1l {988}.			v: Lesostepka 75.
Gli-A1m {988}.			v: Marquis.
Gli-A1n {988}.			v: Intensivnaya.
Gli-A1o {988}.			v: Odesskaya 16, Riband.
Gli-A1p {988}.			v: Pyrotrix 28.
Gli-A1q {988}.			v: Akmolinka 1.
Gli-A1r {988}.			v: Rannaya 73.
Gli-B1 [{1607}], {1125}.	[Gld 1B {1243,1415}, Gld-B1,-B2,-B3,-B4,-B5,-B6 {420}].	1B {1607}, 1BS {150,634}.	s: CS*/Cheyenne {634}. v: CS {1607,150}.
Gli-B1a {988}.			v: CS.
Gli-B1b {988}.			v: Bezostaya1, Gabo.
Gli-B1c {988}.			v: Siete Cerros 66.
Gli-B1d {988}.			v: Dneprovskaya 521, Falchetto.
Gli-B1e {988}.			v: Lutescens 62.
Gli-B1f {988}.			v: Maris Freeman.
Gli-B1g {988}.			v: Galahad.
Gli-B1h {988}.			v: Krasnodonka.
Gli-B1i {988}.			v: Insignia.
Gli-B1j {988}.			v: Cluj 650.
Gli-B1k {988}.			v: Kremena.
Gli-B1l {988}.			v: Dukat.
Gli-B1m {988}.			v: Pyrotrix 28.
Gli-B1n {988}.			v: Intensivnaya.
Gli-B1o {988}.			v: Levent.
Gli-B1p {988}.			v: New Pusa 834.
Gli-B1q (reserved by WJR)
Gli-B1r {995}.			v: Chinook
Gli-D1 [{121}], {1125}.	[Gld 1D {1415}, Gld-D1,-D2,-D3 {420}].	1DS {121,150,634,1334,1607}.	s: CS*/Cheyenne {634}. v: CS {121,150,1334,1607}.
Gli-D1a {988}.			v: CS.
Gli-D1b {988}.			v: Bezostaya 1, Mercia.
Gli-D1c {988}.			v: Skorospelka Uluchshennaya (biotype).
Gli-D1d {988}.			v: Solo.
Gli-D1e {988}.			v: Gerek.
Gli-D1f {988}.			v: Maris Freeman.
Gli-D1g {988}.			v: Mironovskaya 808, Ghurka.
Gli-D1h {988}.			v: Sadovo I.
Gli-D1i {988}.			v: Zelinogradka, Insignia.
Gli-D1j {988}.			v: Promin.
Gli-D1k {988}.			v: Kremena.
Gli-D1l {988}.			v: Longbow.

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 has been 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 are reported in {994}.
 

Gli-Agi1.		lAgi {168}.	ad: Vilmorin 27/Ag. intermedium.
Gli-E1 [{781}].		lES {781}.	ad: CS/E. elongata.
Gli-Sl1 {573}.		1Sl {573}.	ad: CS/Ae. longissima.
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}.
Gli-Rm1 [{1340}].		1RmS {1340}.	ad: CS/S. montanum.
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}.
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-A2 [{1334}], {1125}.	[Gld 6A {1415}].	6A {1334}, 6AS {1122}.	v: CS.
Gli-A2a {988}.			v: CS.
Gli-A2b {988}.			v: Bezostaya 1.
Gli-A2c {988}.			v: Siete Cerros 66.
Gli-A2d {988}.			v: Dneprovskaya 521.
Gli-A2e {988}.			v: Sadovo 1.
Gli-A2f {988}.			v: Maris Freeman.
Gli-A2g {988}.			v: Ducat.
Gli-A2h {988}.			v: Hereward.
Gli-A2i {988}.			v: Lesostepka 75.
Gli-A2j {988}.			v: Cluj 650.
Gli-A2k {988}.			v: Skala.
Gli-A2l {988}.			v: Longbow.
Gli-A2m {988}.			v: Marquis.
Gli-A2n {988}.			v: Mironovskaya 808.
Gli-A2o {988}.			v: Dukat.
Gli-A2p {988}.			v: Pliska.
Gli-A2q {988}.			v: Saratovskaya 39.
Gli-A2r {988}.			v: Riband.
Gli-A2s {988}.			v: Saratovskaya 36.
Gli-A2t {988}.			v: Tarasovskaya 2 (biotype).
Gli-A2u {988}.			v: Kirgizskaya Yubileinaya.
Gli-A2v {988}.			v: Kzyl-Bas.
Gli-A2w {988}.			v: Bezenchukskaya 98.
Gli-A2x {988}.			v: Solo.
Gli-B2 [{1607}], {1125}.	[Gld 6B {1415}].	6B {1607}, 6BS {1122}.	v: CS.
Gli-B2a {988}.			v: CS.
Gli-B2b {988}.			v: Bezostaya 1, Kremena.
Gli-B2c {988}.			v: Siete Cerros 66.
Gli-B2d {988}.			v: Tselinnaya 20.
Gli-B2e {988}.			v: Lesostepka 75.
Gli-B2f {988}.			v: Maris Freeman.
Gli-B2g {988}.			v: Galahad.
Gli-B2h {988}.			v: Sadovo 1.
Gli-B2i {988}.			v: Insignia.
Gli-B2j {988}.			v: Cluj 650.
Gli-B2k {988}.			v: Skala.
Gli-B2l {988}.			v: Longbow.
Gli-B2m {988}.			v: Mironovskaya 808.
Gli-B2n {988}.			v: Solo.
Gli-B2o {988}.			v: Odesskaya 16.
Gli-B2p {988}.			v: Pliska.
Gli-B2q {988}.			v: Saratovskaya 39.
Gli-B2r {988}.			v: Omskaya 12.
Gli-B2s {988}.			v: Saratovskaya 36.
Gli-B2t {988}.			v: Tselinogradka.
Gli-B2u {988}.			v: Kirgizskaya Yubileinaya.
Gli-B2v {988}.			v: Poljarka.
Gli-B2w {995}			v: Pembina
Gli-D2 [{1334}], {1125}.	[Gld 6D {1415}].	6D {1334}, 6DS {1122}.	v: CS.
Gli-D2a {988}.			v: CS.
Gli-D2b {988}.			v: Bezostaya 1, Levent (biotype).
Gli-D2c {988}.			v: Siete Cerros 66.
Gli-D2d {988}.			v: Dneprovskaya 521.
Gli-D2e {988}.			v: Mironovskaya 808.
Gli-D2f {988}.			v: Falchetto; Kirgizskaya Yubileinaya.
Gli-D2g {988}.			v: Ghurka.
Gli-D2h {988}.			v: Ducat.
Gli-D2i {988}.			v: Insignia.
Gli-D2j {988}.			v: Promin.
Gli-D2k {988}.			v: Skala.
Gli-D2l {988}.			v: Condor.
Gli-D2m {988}.			v: Marquis.
Gli-D2n {988}.			v: Mercia.
Gli-D2o {988}.			v: Omskaya 12.
Gli-D2p {988}.			v: New Pusa 834.
Gli-D2q {988}.			v: Volshebnitsa (biotype).
Gli-D2r {988}.			v: Kremena.
Gli-D2s {988}.			v: Bezenchukskaya 98.

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}.
 

Gli-Agi2 {374}.		6Agi {374}.	ad: Vilmorin 27/ Ag. intermedium.
Gli-R2 [{781}].	[Sec 2 {1336}].	2R {781,1336}, 2RS {1340}.	ad: CS/Imperial {781,1336,1340}; Holdfast/King II {1340}.
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.

A Gli-3 set of loci coding for omega-type gliadins are located 22 to 31cM proximal to Gli-1 in the short arms of group 1 chromosomes {422,1403,589}.
 

Gli-A3 [{1403}],{1119}].	[Gld-2-1A {1416}].	1AS {1403}.	v: Bezostaya 1.
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}.		S1S {1228}.	ad,su: CS/Ae. longissima

ma: In Ae. longissima 2/Ae. longissima 10, three gliadin loci, one glucose phosphate isomerase, and two glutein 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.

It is not clear how Gli-Sl4 and Gli-Sl5 relate to the Gli-4 and Gli-5 sets described below.

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}. It has yet to be established whether this is one of a series of orthologous loci.
 

Gli-A4 {1205}.

1AS {1205}.

v: Perzivan biotype 2.

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.

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. An estimate for the map distance between Gli-A5 and Gli-A1 was not reported, although evidence was provided that the linkage is of a similar order of magnitude to the Gli-B5 - Gli-B1 distance. Although no orthologous locus was reported for chromosome 1D, the authors cited studies {992,987} reporting a recombination distance of 1 % between two gliadin loci on chromosome 1D, which they considered may have been due to the presence of a locus on 1D orthologous to Gli-B5.
 

Gli-A5 {1147}.		1AS {1147}.	v: Salmone.
Gli-B5 {1147}.		1BS {1147}.	v: Salmone.

The Gli-1 loci may be recognised by pcP387 {372}, pTag1436 {066} and by specific microsaellite primers {252}.

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.