International Database for Barley Genes and Barley Genetic Stocks
BGS 356, Glossy sheath 6, gsh6
Stock number: BGS 356
Locus name: Glossy sheath 6
Locus symbol: gsh6
Previous nomenclature and gene symbolization:
Glossy sheath 6 = gs6 (17).
Eceriferum-c = cer-c (6).
Eceriferum-cqu = cer-cqu (15, 25).
A member of the gene cluster Cer-cqu (18, 25).
Inheritance:
Monofactorial recessive (6, 17, 20).
Located in chromosome 2HS (20, 21, 22); gsh6.s is in a subterminal position (19); gsh6.s is associated with SNP markers 1_0326 to 2_0563 (positions 16.91 to 21.19 cM) in 2HS bin 02 of the Bowman backcross-derived line BW409 (1) in 2H bin 02; the Cer-cqu gene cluster is near SNP marker 1_0718 and 2.7 cM distal from 1_1059 (19), likely in 2H bins 01 or 02..
Description:
A bright, glossy green color is apparent on the spike, leaf sheath, and stem (17). The gsh6 locus is the cer-c component of the complex locus cer-cqu (2), which controls formation of β-diketone lipids in epicuticular wax layer (23, 25, 27). Most alleles have the wax code - - ++, but a few alleles show delayed development of surface waxes on the sheath (wax code - +/- ++) (6, 16). Since two types of β-diketones and the alkan-2-ol containing esters are absent in the leaf blades (24) and awns (26), mutants in the Cer-cqu cluster of genes do not change the surface wax composition on these organs (24, 26). The only observed morphological difference between Bowman and the backcross-derived line BW409 was an increase in kernel length (3). The Gsh6 (Cer-c) locus is a member of the β-diketone synthase polyketide pathway for synthesis of β-diketone aliphatics, which are part of the complex of epicuticular or surface waxes present in Triticeae species (18). The Cer-c (MLOC_59804) gene is a chalcone synthase-like polyketide synthase, designated diketone synthase (DKS) with two exons and is positioned proximal from the Cer-q (Gsh1) and Cer-u (Gsh8) loci (18). The β-diketone biosynthesis Cer-cqu locus of barley is similar to the wheat W1 locus, which contains a metabolic gene cluster mediating β-diketone biosynthesis (4).
Origin of mutant:
An X-ray induced mutant in Bonus (NGB 14657, PI 189763) (6), a radiation induced mutant in Domen (NGB 2104, CIho 9562) (17). More information on the origin of cer-c mutants is presented in www:nordgen.org/bgs and in Schneider et al. (18).
Mutational events:
cer-c.3 (NGB 101944, GSHO 212. GSHO 424, GSHO 959) in Bonus (NGB 14657, PI 189763) (2, 6); cer-c.7 (NGB 110891), -c.29 (NGB 110913), -c.36 (NGB 110920), -c.44 (NGB 110928), -c.61 (NGB 110945), -c.63 (NGB 110947) in Bonus (5, 6); cer-c.73 (NGB 110957), -c.89 (NGB 110973), -c.95 (NGB 110979), -cu.108 (NGB 110993), -c.115 (NGB 111000), -c.123 (NGB 111008), -cqu.124 (NGB 111009) in Bonus, -c.201 (NGB 111088), -c.206 (NGB 111093), -c.220 (NGB 111107), -c.223 (NGB 111110), -c.234 (NGB 111121), -c.238 (NGB111125), -c.240 (NGB 111127), -c.250 (NGB 111137), -c.270 (NGB 111157), -c.282 (NGB 111169), -c.283 (NGB 111170), -c.284 (NGB 111171), -c.290 (NGB 111177), -c.291 (NGB 111178), -c.293 (NGB 111180). -c.306 (NGB 111193), -c.315 (111202), -c.332 (NGB 111219), -c.343 (NGB 111230), -c.354 (NGB 111241), -c.361 (NGB 111248), -c.364 (NGB 111251), -c.367 (NGB 111254), -c.368 (NGB 111255), -c.374 (NGB 111261), -c.379 (NGB 111266), -c.398h (NGB 111285), -c.398l (NGB 111288), -c.405 (NGB 111293), -c.406 (NGB 111294), -c.413 (NGB 111301), -cqu.416 (NGB 111304), -cqu.420 (NGB 111308), -c.431 (NGB 111319), -c.445 (NGB 111333), -c.455 (NGB 111343), -c.469 (NGB 111357), -c.471 (NGB 111359), -c.502 (NGB 111390) in Foma (NGB 14659, CIho 11333) (7, 15); cer-c.143 (NGB 111029), -c.171 (NGB 111057), -c.175 (NGB 111061) in Bonus, -c.345 (NGB 111232), -c.451 (NGB 111339), -c.459 (NGB 111347), -c.508 (NGB 111396), -c.513 (NGB 111401), -c.514 (NGB 111402), -c.520 (NGB 111408), -c.526 (NGB 111414), -c.544 (NGB 111432), -c.550 (NGB 111438), -c.563 (NGB 111451), -c.596 (NGB 111484), -c.601 (NGB 111489), -c.661 (NGB 111549) in Foma, -c.669 (NGB 111557), -c.673 (NGB 111561), -c.690 (NGB 111578), -c.693 (NGB 111581), -c.698 (NGB 111586), -c.703 (NGB 111591), -c.711 (NGB 111499), -cqu.724 (NGB 111612), -cqu.733 (NGB 111621) in Bonus, -c.1001 (NGB 111889), -c.1004 (NGB 111892), -c.1008 (NGB 111896), -c.1010 (NGB 111898), -c.1015 (NGB 111903), -c.1022 (NGB 111910), -c.1029 (NGB 111917), -c.1034 (NGB 111922), -c.1035 (NGB 111923), -c.1039 (NGB 111927), -c.1062 (NGB 111950), -c.1067 (NGB 111955), -c.1082 (NGB 111970) in Carlsberg II (NGB 5085, CIho 10114) (7); cer-c.589 (NGB 111477) in Foma, -c.748 (NGB 111636), -c.760 (NGB 111648), -c.764 (NGB 111652), -c.767 (NGB 111655), -c.770 (NGB 111658), -c.774 (NGB 111662), -c.786 (NGB 111674), -c.793 (NGB 111681), -c.794 (NGB 111682), -c.841 (NGB 111729), -c.845 (NGB 111733) -, c.847 (NGB 111735), -c.851 (NGB 111739), -c.860 (NGB 111748), -c.870 (NGB 111758), -c.881 (NGB 111769), -c.883 (NGB 111771) in Bonus, -c.1032 (NGB 111920) in Carlsberg II, -c.1098 (NGB 111986), -c.1100 (NGB 111988), -c.1109 (NGB 111997), -c.1117 (NGB 112005), -c.1123 (NGB 112011), -c.1125 (NGB 112013), -c.1135 (NGB 112023), -c.1136 (NGB 112024), -c.1140 (NGB 112028), -c.1147 (NGB 112035) in Kristina (NGB 14661, NGB 1500) (8); cer-c.499 (NGB 111387), -c.609 (NGB 111497) in Foma, -c.630 (NGB 111518), -c.635 (NGB 111523), -c.640 (NGB 111528), -cu.644 (NGB 111532), -cq.645 (NGB 111533), -c.648 (NGB 111536), -c.890 (NGB 111778), -c.897 (NGB 111785), -c.902 (NGB 111790), -c.903 (NGB 111791), -c.904 (NGB 111792), -c.907 (NGB 111795), -c.912 (NGB 111800), -c.915 (NGB 111803), -cqu.925 (NGB 111813), -c.942 (NGB 111830), -cqu.944 (NGB 111832), -c.945 (NGB 111833), -c.980 (NGB 111868) in Bonus, -c.1156 (NGB 112044), -c.1187 (NGB 112075) in Kristina (9); cer-c.144 (NGB 111030), -cu.947 (NGB 111835), -c.987 (NGB 111875) in Bonus, -c.1110 (NGB 111998), -c.1212 (NGB 112100), -c.1229 (NGB 112117), -c.1248 (NGB 112136), -c.1258 (NGB 112146), -c.1264 (NGB 112152), -c.1276 (NGB 112164) in Kristina (10); cer-c.511 (NGB 111399) in Foma, -c.1200 (NGB 112088), -c.1254 (NGB 112271, 1168212), -c.1278 (NGB 116824, 117273), -c.1287 (NGB 112175), -c.1291 (NGB 112179) in Kristina, -c.1319 (NGB 112207), -c.1325 (NGB 112213), -c.1329 (NGB 112217), -c.1334 (NGB 112222), -c.1338 (NGB 112226) in Bonus (11); cer-c.993 (NGB 111881), -c.994 (NGB 111882), -c.995 (NGB 111883), -c.999 (NGB 111887), -c.1348 (NGB 112236), -c.1355 (NGB 112243), -c.1359 (NGB 112247), -c.1365 (NGB 112253), -c.1376 (NGB 112264), -c.1381 (NGB 112269), -c.1382 (NGB 112270), -c.1385 (NGB 112273), -c.1386 (NGB 112274), -c.1389 (NGB 112277), -c.1416 (NGB 112304), -c.1421 (NGB 112309), -c.1439 (NGB 112327), -c.1448 (NGB 112336), -c.1460 (NGB 112348), -c.1462 (NGB 112350), -c.1466 (NGB 112354), -c.1467 (NGB 112355) in Bonus, -c.1501 (NGB 112389), -c.1502 (NGB 112390), -c.1503 (NGB 112391), -c.1505 (NGB 112393), -c.1506 (NGB 112394), -c.1511 (NGB 112399), -c.1515 (NGB 112403) in Nordal (NGB 13680, NGB 4704) (12); cer-c.1491 (112379), -c.1701 (NGB 112514), -c.1702 (NGB 112515), -c.1703 (NGB 112516), -c.1705 (NGB 112518), -c.1707 (NGB 112520), -c.1714 (NGB 112527), -c.1737 (NGB 112550, 117329), -c.1739 (NGB 112552), -c.1749 (NGB 112563), -c.1755 (NGB 112569), -c.1757 (NGB 112572) in Bonus, -c.1516 (NGB 112404), -c.1520 (NGB 112408), -c.1525 (NGB 112413), -c.1526 (NGB 112414), -c.1532 (NGB 112420), -c.1553 (NGB 112441), -c.1555 (NGB 112443), -c.1560 (NGB 112448) in Nordal (13); cer-c.602a (NGB 119348) in Foma (14); gsh6.o in Domen (NGB 2104, CIho 9562), gsh6.p in PI 184877, gsh6.q in Kogane Mugi (OUL010, PI 225020), gsh6.r in Barbless (CIho 5105), gsh6.s (GSHO 740) in Betzes (PI 129430), gsh6.t in Heines Hanna (PI 539131), gsh6.u, gsh6.v, gsh6.w, and gsh6.x in Compana (PI 539111) (15); gsh6.z (OUM017), gsh6.ac (OUM020), gsh6.ai (OUM026) in Akashinriki (PI 467400, OUJ659) (5); gsh6.aq in an Ethiopian introduction (PI 151787, OUE120) (5).
Mutant used for description and seed stocks:
cer-c.3 ( NGB 110887, GSHO 212, GSHO 424, GSHO 959) in Bonus; cer-c.36 (NGB 110920 is used for wax chemistry and wax structure studies (23, 25); gsh6.s (GSHO 740) in Betzes, gsh6.s via R.I. Wolfe’s Multiple Marker Stock for chromosome 2 in Bowman (PI 483237)*7 (GSHO 1869, BW409, NGB 20642).
References:
1. Druka, A., J. Franckowiak, U. Lundqvist, N. Bonar, J. Alexander, K. Houston, S. Radovic, F. Shahinnia, V. Vendramin, M. Morgante, N. Stein, and R. Waugh. 2010. Genetic dissection of barley morphology and development. Plant Physiol. 155:617-627.
2. Eslick, R.F. 1974. Allele tests of gs and cer mutants. Barley Genet. Newsl. 4:9-11.
3. Franckowiak, J.D. (Unpublished).
4. Hen-Avivi, S., O. Savin, R.C. Racovita, W.S. Lee, N. Adamki, S. Malitsky, E. Almekias-Siegl, M. Levy, S. Vautrin, H. Bergès, G. Friedlander, E. Kartvelishvily, G. Ben-Zvi, N. Alkan, C. Uauy, K. Kanyuka, R. Jetter, A. Distelfeld, and A. Aharoni. 2016. A metabolic gene cluster in the wheat W1 and the barley Cer-cqu loci determines β-diketone biosynthesis and glaucousness. Plant Cell 28:1440-1460.
5. Konishi, T. 1973. Genetic analyses of EMS-induced mutants in barley. Barley Genet. Newsl. 3:28-31.
6. Lundqvist, U., and D. von Wettstein. 1962. Induction of eceriferum mutants in barley by ionizing radiations and chemical mutagens. Hereditas 48:342-362.
7. Lundqvist, U., and D. von Wettstein. 1971. Stock list for the eceriferum mutants. Barley Genet. Newsl. 1:97-102.
8. Lundqvist, U., and D. von Wettstein. 1973. Stock list for the eceriferum mutants II. Barley Genet. Newsl. 3:110-112.
9. Lundqvist, U., and D. von Wettstein. 1975. Stock list for the eceriferum mutants III. Barley Genet. Newsl. 5:88-91.
10. Lundqvist, U., and D. von Wettstein. 1977. Stock list for the eceriferum mutants IV. Barley Genet. Newsl. 7:92-96.
11. Lundqvist, U., and D. von Wettstein. 1979. Stock list for the eceriferum mutants V. Barley Genet. Newsl. 9:135-137.
12. Lundqvist, U., and D. von Wettstein. 1982. Stock list for the eceriferum mutants VI. Barley Genet. Newsl. 12:169-172.
13. Lundqvist, U., and D. von Wettstein. 1985. Stock list for the eceriferum mutants VII. Barley Genet. Newsl. 15:89-93.
14. Lundqvist, U., and D. von Wettstein. 1988. Stock list for the eceriferum mutants VIII. Barley Genet. Newsl. 18:88-91.
15. Lundqvist, U., P. von Wettstein-Knowles, and D. von Wettstein. 1968. Induction of eceriferum mutants in barley by ionizing radiations and chemical mutagens. II. Hereditas 59:473-504.
16. McProud, W.L., and R.F. Eslick. 1971. Allelism within a series of glossy sheath mutants. Barley Genet. Newsl. 1:31-34.
17. Rasmusson, D.C., and J.W. Lambert. 1965. Inheritance of the glossy-sheath character in barley, Hordeum vulgare L. Crop Sci. 5:251-253.
18. Schneider, L.M., N.M. Adamski, C.E. Christensen, D.B. Stuart, S. Vautrin, M. Hansson, C. Uauy, and P. von Wettstein-Knowles. 2016. The Cer-cqu gene cluster determines three key players in a β-diketone synthase polyketide pathway synthesizing aliphatics in epicuticular waxes. J. Exp. Bot. 67:2715-2730.
19. Schondelmaier, J., G. Fischbeck, and A. Jahoor. 1993. Linkage studies between morphological and RFLP markers in the barley genome. Barley Genet. Newsl. 22:57-62.
20. Singh, R.J., and T. Tsuchiya. 1974. Further information on telotrisomic analysis in barley. Barley Genet. Newsl. 4:66-69.
21. Takahashi, R., J. Yamamoto, S. Yasuda, and Y. Itano. 1953. Inheritance and linkage studies in barley. Ber. Ohara Inst. landw. Forsch. 10:29-52.
22. Tsuchiya, T. 1972. Cytogenetics of telotrisomics in barley. BGN 2:93-98.
23. Wettstein-Knowles, P. von. 1972. Genetic control of β-diketone and hydro-β-diketone synthesis in epicuticular waxes of barley. Planta (Berl) 106:113-130.
24. Wettstein-Knowles, P. von. 1976. Biosynthetic relationships between β-diketones and esterified alkan-2-ols deduced from epicuticular wax of barley mutants. Mol. Gen. Genet. 144:43-48.
25. Wettstein-Knowles, P. von. 1992. Molecular genetics of the lipid synthesis in barley. p. 753-771. In L. Munck, K. Kirkegaard, and B. Jensen (eds.) Barley Genetics VI, Vol. II. Proc. Sixth Int. Barley Genet. Symp., Helsingborg, Sweden, 1991. Munksgaard International Publishers Ltd., Copenhagen.
26. Wettstein-Knowles, P. von, and A.G. Netting. 1976. Composition of epicuticular waxes on barley spikes. Carlsberg Res. Commun. 41:225-235.
27. Wettstein-Knowles, P. von, and B. Søgaard. 1980. The cer-cqu region in barley: Gene cluster or multifunctional gene. Carlsberg Res. Commun. 45:125-141.
NGB number url references to Nordic Baltic Genebankes Information System (GENBIS):
Prepared:
W. L. McProud and R.F. Eslick. 1971. Barley Genet. Newsl. 1:172-173.
Revised:
U. Lundqvist and J.D. Franckowiak. 1997. Barley Genet. Newsl. 26:302-303.
U. Lundqvist and J.D. Franckowiak. 2010. Barley Genet. Newsl. 40:95-97.
U. Lundqvist and J.D. Franckowiak. 2013. Barley Genet. Newsl. 43:135-138.
U. Lundqvist and J.D. Franckowiak. 2016. Barley Genet. Newsl. 46:101-104.