International Database for Barley Genes and Barley Genetic Stocks

BGS 339, High lysine 3, lys3

BGN  43:127
Stock number: BGS 339
Locus name: High lysine 3
Locus symbol: lys3

Previous nomenclature and gene symbolization:

Shrunken endosperm xenia 3 = sex3 (18, 19).


Monofactorial recessive (3, 5, 18).
Located in 5HL (9, 13, 20); lys3.a is near the msg19 (male sterile genetic 19) locus (18); lys3.a is about 14.5 cM proximal from the fst1 (fragile stem 1) locus; lys3.a is about 20.7 cM proximal from the ddt1 (reaction to DDT 1) locus (9, 11); lys3.a is likely in the cluster of hordein genes in 1H bin 02. A second variant isolated from Risø 1508 is associated with SNP markers 2_0479 to 3_0918 (positions 0.54 to 17.27 cM) in 1H bins 01 to 02 of the Bowman backcross-derived line BW496 (6).


After the hard dough stage, kernels develop a depression in the dorsal side or lemma which becomes progressively more distinct with maturity. The 1000-kernel weight of the mutant is about 75% of normal. The mutant expresses xenia permitting classification of kernels from heterozygous plants as normal or shrunken with an expected 3:1 ratio (3). Field establishment and growth of mutant plants is normal (3, 5). Mutant kernels have a higher lysine content than normal kernels (3) and a pleiotropic the association with shrunken endosperm (13, 19). The endosperm is nearly devoid of B- and C-hordeins (8, 14, 16). The B-hordein genes in developing barley endosperm were highly methylated in the lys3.a mutant compared to wild-type endosperm (17). A deficiency of kernels with shrunken endosperm occurs in some crosses (1, 2, 5, 18) and was caused likely highly distorted male gamete transmission (4). The gamete lethal factor in Risø 1508 was named Gam3 (gametophyte factor 3) and was position in 5HL about 6.5 cM from the lys3 locus (21). Based on SNP mapping information (6), the Bowman backcross-derived line from Risø 1508, BW496, may also contain a shrunken endosperm mutant near the Hor2 locus in the 1HS cluster of hordein genes (15), which is not the lys3.a mutant. The Bowman backcross-derived line BW496 is apparently more sensitive to environmental stress than Bowman. Heading of BW496 is delayed by 6 to 10 days, plants are 2/3 to almost normal in height, and peduncles are 1/2 to normal in length. Kernel weights for BW496 were 30% lower than those for Bowman. Grain yields of BW496 were about half of normal (6).

Origin of mutant:

An induced mutant in Bomi Abed (NGB 5096, PI 43371) (3).

Mutational events:

lys3.a (Risø 1508, GSHO 1785) in Bomi Abed (NGB 5096, PI 43371) (3, 9); lys3.b (Risø 18), lys3.c (Risø 19) in Bomi Abed (9, 12).

Mutant used for description and seed stocks:

lys3.a (GSHO 1785, Risø 1508) in Bomi Abed; lys3.a from Risø 1508 in Bowman (PI 483237) (GSHO 2096); lys3.a in Bowman*6 (BW496, NGB 20724).


1. Ahokas, H. 1979. Further results on the suppression of shrunken endosperm in high lysine mutants. Barley Genet. Newsl. 9:3-7.
2. Ahokas, H. 1988. High-lysine gene segregation distorted in the barley cross Risø 1508 x Crypt CI 1090: Patterns of endosperm proteins by an electrophorectic method. Hereditas 108:129-131.
3. Doll, H. 1973. Inheritance of a high-lysine character of a barley mutant. Hereditas 74:293-294.
4. Doll, H. 1991. A factor on barley chromosome 7 causing distorted male segregation of gene lys3a. Barley Genet. Newsl. 20:23-24.
5. Doll, H., and R. Oram. 1989. Deviating Mendelian segregation of barley gene lys3a. Hereditas 110:97-99.
6. 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. 2011. Genetic dissection of barley morphology and development. Plant Physiol. 155:617-627.
7. Franckowiak, J.D. (Unpublished).
8. Hopp, H.E., S.K. Rasmussen, and A. Brandt. 1983. Organization and transcription of B1 hordein genes in high lysine mutants of barley. Carlsberg Res. Commun. 48:201-216.
9. Jensen, J. 1979. Location of a high-lysine gene and the DDT-resistant gene on barley chromosome 7. Euphytica 28:47-56.
10. Jensen, J. 1979. Chromosomal location of one dominant and four recessive high-lysine genes in barley mutants. Vol. 1. p. 89-96. In Seed Protein Improvement in Cereals and Grain Legumes. Proc. Int. Symp., Neuherberg, Germany, 1978. Int. Atomic Energy Agency, Vienna.
11. Jensen, J. 1981. Construction of a barley chromosome 7 linkage map. p. 927-939. In M.J.C. Asher, R.P. Ellis, A.M. Hayter, and R.N.H. Whitehouse (eds.) Barley Genetics IV. Proc. Fourth Int. Barley Genet. Symp., Edinburgh. Edinburgh Univ. Press, Edinburgh.
12. Jensen, J., and H. Doll. 1979. Gene symbols for barley high-lysine mutants. Barley Genet. Newsl. 9:33-37.
13. Karlsson, K.-E. 1977. Linkage studies in a gene for high lysine content in Risø barley mutant 1508. Barley Genet. Newsl. 7:40-41.
14. Kries, M., P.R. Shewry, B.G. Forde, S. Rahman, M.B. Bahramian, and B.J. Miflin. 1984. Molecular analysis of the effects of the lys3a gene on the expression of Hor loci in developing endosperms of barley (Hordeum vulgare L.). Biochem. Genet. 22:231-255.
15: Pelger, S., T. Säll, and B.O. Bengtsson. 1993. Evolution of hordein gene organization in three Hordeurn species. Hereditas 119:219-231.
16. Shewry, P.R., A.J. Faulks, and B.J. Miflin. 1980. Effect of high-lysine mutations on the protein fractions of barley grains. Biochem. Genet. 18:133-151.
17. Sørensen, M.B. 1992. Methylation of B-hordein genes in barley endosperm is inversely correlated with gene activity and affected by the regulatory gene Lys3. Proc. Natl. Acad. Sci. USA. 89:4119-4123.
18. Ullrich, S.E., and R.F. Eslick. 1977. Inheritance of the shrunken endosperm character, sex3c, of Bomi Risø mutant 1508 and its association with lysine content. Barley Genet. Newsl. 7:66-73.

19. Ullrich, S.E., and R.F. Eslick. 1978. Chromosome location evidence for Risø induced high lysine shrunken endosperm mutants of barley. Barley Genet. Newsl. 8:114-125.

20. Ullrich, S.E., and R.F. Eslick. 1978. Inheritance of the associated kernel characters, high lysine and shrunken endosperm, of the barley mutant Bomi, Risø 1508. Crop Sci. 18:828-831.

21. Yamae, G., and T. Konishi. 1993. Distorted segregation of the lys3a mutant in barley kernels. Barley Genet. Newsl. 22:68-70.


J.D. Franckowiak. 1997. Barley Genet. Newsl. 26:287-288.


J.D. Franckowiak. 2011. Barley Genet. Newsl. 41:138-139.
J.D. Franckowiak. 2013. Barley Genet. Newsl. 43:127-129.