3. Endosperm Storage Proteins

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.

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}. 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.
 
Glu-A1 [{780}], {1125}.  [Glt-A1, 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} T. dicoccum. tv: PI 352359, Germany,
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,
T. durum {1526}.
Glu-A1o [{1526,125}].
[Glu A1-V {1526,125}, Glu-A1n {959}]. 2** {125}. tv: Aric 581/1 {125}; PI
61189, T. durum {1525}; USSR.
Glu-A1p [{1146}].
3* {1146}. v: David 1.
Glu-A1q [{125}].
[Glu A1VI {125}]. 2*** {125}. tv: Melianopus 1528,
T. durum.
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, W 3879, W31169.

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}] has been changed to n, because the m allele in {1069} has precedence. Allele n [{1526}] has been changed to o. An earlier reference to an allele designated Glu-A1d {1411} has been 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.
 
Glu-B1 [{107}], {1125}. [Glt-B1,-B2,-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}. 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}. v: Dunav (rare).
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: Mariñar.
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.

The alleles formerly designated t to x in {959} have been renamed x to ab because allele t in {847} and alleles u,v and w in {1069} have 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 varieties has also been observed {1069}; possible low gene expression at Glu-B1 has been noted for Glu-B1w, where subunits 6*+8* stain very faintly {1146}. One of the Glu-B1af subunits has been 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.
 
Glu-D1 [{1100}], {1125}. [Glt-D1, 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}.
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.
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: T. tauschii accession TA2450/2*.
Glu-D1u [{836}].
2+10' {836}. v: Coker 68-15.
Glu-D1v [{755}].
2.1+10.1 {755}. dv: T. tauschii.
Glu-D1w [{755}].
2+T1+T2 {755}. dv: T. tauschii.
Glu-D1x [{755}].
2+T2 {755}. dv: T. tauschii.
Glu-D1y [{755}].
3+T2 {755}. dv: T. tauschii.
Glu-D1z [{755}].
3+10 {755}. dv: T. tauschii.
Glu-D1aa [{755}].
3+10.3 {755}. dv: T. tauschii.
Glu-D1ab [{755}].
4.1+10 {755}. dv: T. tauschii.
Glu-D1ac [{755}].
4+10 {755}. dv: T. tauschii.
Glu-D1ad [{755}].
5.1+10.2 {755}. dv: T. tauschii.
Glu-D1ae [{1578}].
2.1+T1+T2 {1578}. dv: T. tauschii.
Glu-D1af [{1578}].
3+T1+T2 {1578}. dv: T. tauschii.
Glu-D1ag [{1578}].
1.5+T1+T2 {1578}. dv: T. tauschii.
Glu-D1ah [{1578}].
1.5+10 {1578}. dv: T. tauschii.
Glu-D1ai [{1578}].
2.1+10.5 {1578}. dv: T. tauschii.
Glu-D1aj [{1578}].
1.5+12 {1578}. dv: T. tauschii.
Glu-D1ak [{1578}].
3+10.5 {1578}. dv: T. tauschii.

Glu-D1k [{421}] appears to have arisen 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}.

Alleles and subunits at Glu-1: Glu-1-1 encodes x-type glutenins and Glu-1-2 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.
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, T: durum.
Glu-A1-1m
    tv: Nugget Biotype 1, Canada, T. durum.
Glu-A1-1n.
  1". tv: Corado, Portugal, T. durum.
Glu-A1-1o.
  2**. tv: PI 61189, USSR, T. durum; Aric 581/1.
Glu-A1-1p.
  3*. v: David 1.
Glu-A1-1q.
  2***. tv: Melianopus 1528, T. durum.
Glu-A1-1r
  39. i: T. thaoudar IPSR1020006/6* 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-A1-2.      
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.
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. v: Dunav.
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, T. turgidum
Glu-B1-1r.
    tv: Tremez Mollez, Portugal, T. durum.
Glu-B1-1s.
    tv: Quaduro, Italy, T. durum.
Glu-B1-1t.
    tv: Athena, Italy, T. durum.
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: Mariñar.
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-B1-2.
     
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, T. turgidum.
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: Mariñar.
Glu-B1-2x.
  33. v: BG-1943.
Glu-B1-2y.
  35. v: Jeja Almendros.
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.
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: T. tauschii accession TA2450/2*.
Glu-D1-1k {755}.
  4.1 {755}. dv: T. tauschii.
Glu-D1-1j {755}.
  5.1 {755}. dv: T. tauschii.
Glu-D1-1l {1578}.
  1.5 {1578}. dv: T. tauschii.
Glu-D1-2.      
Glu-D1-2a.
  12. v: CS.
Glu-D1-2b.
  10. v: Hope.
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: T. tauschii TA2450/2*
Glu-D1-2j [{836}].
  10' {836}. v: Coker 68-15.
Glu-D1-2k {755}.
  T1 {755}. dv: T. tauschii.
Glu-D1-2l {755}.
  T2 {755}. dv: T. tauschii.
Glu-D1-2m {755}.
  10.1 {755}. dv: T. tauschii.
Glu-D1-2n {755}.
  10.2 {755}. dv: T. tauschii.
Glu-D1-2o {755}.
  10.3 {755}. dv: T. tauschii.
Glu-D1-2p {1578}.
  10.5 {1578}. dv: T. tauschii.
     
Glu-Agi1 {374}.   1Agi {374}. ad: Vilmorin 27/Ag. 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-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-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 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.
 
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 : 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.
Glu-Ht1 {1037}.   1HtL {1037}. ad: CS/E. trachycaulum.

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.

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}.
 
Glu-B2 {819,277}. [XGlu-B2 {277}]. 1BS. s: CS*/Cheyenne 1B {277}. 
      stv: Langdon*/T.turgidum var. dicoccodis 1B {277}.

Gli-B3 was designated Glu-B2 {589} until the name of the locus was changed in {1119}.

The Glu-3 loci are defined as the cluster of LMW glutenin genes previously considered a component of the compound Gli-1 loci.

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 now 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}.
 
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-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}. 
    tv: Duramba-B, Duramba-D, Langdon.
Glu-B3k {476}. 
    tv: ALP-153, Dural, Durati, Edmore.
Glu-B3l {476}.
    tv: Gionp-1954.
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.

Multiple alleles at each of the three Glu-3 wheat loci were observed {479} and effects of allelic differences on bread making quality noted {483}.
 
Glu-E3 [{480}].   1ES {480}. su: CS/E. elongata.
Glu-Sl3 [{480}],[{1228}]   1Sl {480},1S1S {1228}. su: CS/T. longissimam {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/T. umbellulatum.

The Glu-3 loci can be recognised with pTag544 {49} and pTdUCD1 {167} and by specific microsatellite primers {252}.