International Database for Barley Genes and Barley Genetic Stocks
BGS 203, Black lemma and pericarp 1, Blp1
Stock number: BGS 203
Locus name: Black lemma and pericarp 1
Locus symbol: Blp1
Previous nomenclature and gene symbolization:
Black lemma and caryopsis = B (11).
Black pericarp = Bk (1).
Black lemma and pericarp = B (12).
Black pericarp = Bk (1).
Black lemma and pericarp = B (12).
Inheritance:
Monofactorial dominant (1, 8, 11).
Located in chromosome 1HL (5, 9); Blp1.b is about 16.0 cM proximal from the trd1 (third outer glume 1) locus (5); Blp1.b is in 1H bin 13 about 8.8 cM proximal from RFLP marker ABC261 (2); Blp1.b is associated with SNP markers 2_0959 to 2_0603 (positions 173.49 to 199.04 cM) in 1H bins 13 to 15 of the Bowman backcross-derived line BW062 (3); Blp1.g is associated with SNP marker 2_0940 (about position 179 cM) in 1H bin 13 of the Bowman backcross-derived line BW060 (3); Blp1.b in BW062 was located between microsatellites Xgbms0012 and Xgbms0184 (10), Blp1.b is in a 1.66 Mb segment between markers HZSNP63 and HZSNP59 (6); Blp1.b mapped between two Insertion/deletion (Indel) markers MC_1570156 and MC_162350 at 538 Mb (7), in 1H bin 13.
Located in chromosome 1HL (5, 9); Blp1.b is about 16.0 cM proximal from the trd1 (third outer glume 1) locus (5); Blp1.b is in 1H bin 13 about 8.8 cM proximal from RFLP marker ABC261 (2); Blp1.b is associated with SNP markers 2_0959 to 2_0603 (positions 173.49 to 199.04 cM) in 1H bins 13 to 15 of the Bowman backcross-derived line BW062 (3); Blp1.g is associated with SNP marker 2_0940 (about position 179 cM) in 1H bin 13 of the Bowman backcross-derived line BW060 (3); Blp1.b in BW062 was located between microsatellites Xgbms0012 and Xgbms0184 (10), Blp1.b is in a 1.66 Mb segment between markers HZSNP63 and HZSNP59 (6); Blp1.b mapped between two Insertion/deletion (Indel) markers MC_1570156 and MC_162350 at 538 Mb (7), in 1H bin 13.
Description:
Black pigmentation of the lemma and pericarp develops slightly before maturation of the spike. Pigmented organs may include all parts of the spike, awns, the upper portion of the stem, and upper leaves. The intensity of pigmentation associated with each of the dominant alleles at the Blp1 locus is characteristic of that allele, and is relatively stable over environments (12). Black seed is produced by melanin-like pigment in the pericarp (1). Woodward (12) reported that the dominance ranking of alleles at the Blp1 locus is related to the intensity of black pigmentation they confer, with the Blp1.b (B) allele conferring extreme black pigmentation. The Blp1.mb (Bmb) allele is associated with medium black and a reduced distribution pattern; and the Blp1.g (Bg) allele is associated with light black or gray coloration (12, 13). In the Bowman backcross-derived lines for the Blp1 locus, BW060 and BW062, the expression of other morphological traits appeared similar to those of Bowman (4).The flavonoid pigments are not involved in development of the black lemma and pericarp trait (10).
 spike at early maturity to the left compared with normal Bowman'))
 spike showing black colored lemma to the left compared with normal Bowman'))
 grain from the lemma side to the left compared with a normal Bowman grain'))
Origin of mutant:
Natural occurrence in several cultivars (11, 12). ). Genotype-based clustering analyses showed that the black landraces of different geographical background may have evolved from a single origin (7).
Mutational events:
Blp1.b (B) in Hordeum distichon var. nigrinudum No 1 (NSL 3286, GSHO 988) (12); Blp1.mb (Bmb) in CIho 2970 (GSHO 226) (12); Blp1.g (Bg) in Blackhull (CIho 878, GSHO 199) and Black Smyrna (CIho 191, GSHO 222) (12). Since these are naturally occurring variants, the stock numbers are the same for both the mutant and the cultivar.
Mutant used for description and seed stocks:
Blp1.b in Hordeum distichon var. nigrinudum No 1 (GSHO 988); Blp1.b from R.I. Wolfe's Multiple Dominant Marker Stock (GSHO 3450) in Bowman (PI 483237, NGB 22812)*8 (GSHO 2054, BW062, NGB 20470);posibly Blp1.g from a Composite Cross in Bowman*5 (BW060, NGB 20468).
References:
1. Buckley, G.F.H. 1930. Inheritance in barley with special reference to the color of the caryopsis and lemma. Sci. Agric. 10:460-492.
2. Costa, J.M., A. Corey, M. Hayes, C. Jobet, A. Kleinhofs, A. Kopisch-Obusch, S.F. Kramer, D. Kudrna, M. Li, O. Piera-Lizaragu, K. Sato, P. Szues, T. Toojinda, M.I. Vales, and R.I. Wolfe. 2001. Molecular mapping of the Oregon Wolfe Barleys: a phenotypically polymorphic doubled-haploid population. Theor. Appl. Genet. 103:415-424.
3. 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.
4. Franckowiak, J.D. (Unpublished).
5. Ivanova, K.V. 1937. A new character in barley "third outer glume" — Its inheritance and linkage with color of the flowering glumes. Bull. Appl. Bot., Genet., & Pl. Breed. II. 7:339-353.
6. Jia, Q., J. Wang, J. Zhu, W. Hua, Y. Shang, J. Yang, and Z. Liang. 2017. Toward identification of black lemma and pericarp gene Blp1 in barley combining bulked segregant analysis and specific-locus amplified fragment sequencing. Front. Plant Sci. 8:1414.
7. Long, Z., Y. Jia, C. Tan, X.-Q. Zhang, T. Angessa, S. Broughton, S. Westcott, F. Dai, G. Zhang, D Sun, Y. Xu, and C. Li. 2019. Genetic mapping and evolutionary analyses of the black grain trait in barley. Font. Plant Sci. 9:1921.
8. Robertson, D.W. 1929. Linkage studies in barley. Genetics 14:1-36.
9. Robertson, D.W., G.A. Wiebe, R.G. Shands, and A. Hagberg. 1965. A summary of linkage studies in cultivated barley, Hordeum species: Supplement III, 1954-1963. Crop Sci. 5:33-43.
10. Shoeva, O.Y., H.-P. Mock, T.V. Kukoeva, A. Börner, and E.K. Khlestkina. 2016. Regulation of the flavonoid biosynthesis pathway genes in purple and black grains of Hordeum vulgare. PloS ONE 11(10): e0163e782.
11 Tschermak, E. von. 1901. Über Züchtung neuer Getreiderassen mittelst künstlicher Kreuzung. Kritisch-historische Betrachtungen. Zeitschrift für das landwirtschaftliche Versuchswesen Oesterreich 4:1029-1060.
12. Woodward, R.W. 1941. Inheritance of melanin-like pigment in the glumes and caryopses of barley. J. Agric. Res. 63:21-28.
13. Woodward, R.W. 1942. Linkage relationships between the allelomorphic series, B, Bmb, Bg, and Atat factors in barley. J. Amer. Soc. Agron. 34:659-661.
2. Costa, J.M., A. Corey, M. Hayes, C. Jobet, A. Kleinhofs, A. Kopisch-Obusch, S.F. Kramer, D. Kudrna, M. Li, O. Piera-Lizaragu, K. Sato, P. Szues, T. Toojinda, M.I. Vales, and R.I. Wolfe. 2001. Molecular mapping of the Oregon Wolfe Barleys: a phenotypically polymorphic doubled-haploid population. Theor. Appl. Genet. 103:415-424.
3. 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.
4. Franckowiak, J.D. (Unpublished).
5. Ivanova, K.V. 1937. A new character in barley "third outer glume" — Its inheritance and linkage with color of the flowering glumes. Bull. Appl. Bot., Genet., & Pl. Breed. II. 7:339-353.
6. Jia, Q., J. Wang, J. Zhu, W. Hua, Y. Shang, J. Yang, and Z. Liang. 2017. Toward identification of black lemma and pericarp gene Blp1 in barley combining bulked segregant analysis and specific-locus amplified fragment sequencing. Front. Plant Sci. 8:1414.
7. Long, Z., Y. Jia, C. Tan, X.-Q. Zhang, T. Angessa, S. Broughton, S. Westcott, F. Dai, G. Zhang, D Sun, Y. Xu, and C. Li. 2019. Genetic mapping and evolutionary analyses of the black grain trait in barley. Font. Plant Sci. 9:1921.
8. Robertson, D.W. 1929. Linkage studies in barley. Genetics 14:1-36.
9. Robertson, D.W., G.A. Wiebe, R.G. Shands, and A. Hagberg. 1965. A summary of linkage studies in cultivated barley, Hordeum species: Supplement III, 1954-1963. Crop Sci. 5:33-43.
10. Shoeva, O.Y., H.-P. Mock, T.V. Kukoeva, A. Börner, and E.K. Khlestkina. 2016. Regulation of the flavonoid biosynthesis pathway genes in purple and black grains of Hordeum vulgare. PloS ONE 11(10): e0163e782.
11 Tschermak, E. von. 1901. Über Züchtung neuer Getreiderassen mittelst künstlicher Kreuzung. Kritisch-historische Betrachtungen. Zeitschrift für das landwirtschaftliche Versuchswesen Oesterreich 4:1029-1060.
12. Woodward, R.W. 1941. Inheritance of melanin-like pigment in the glumes and caryopses of barley. J. Agric. Res. 63:21-28.
13. Woodward, R.W. 1942. Linkage relationships between the allelomorphic series, B, Bmb, Bg, and Atat factors in barley. J. Amer. Soc. Agron. 34:659-661.
Prepared:
T.E. Haus and T. Tsuchiya. 1971. Barley Genet. Newsl. 1:148.
Revised:
J.D. Franckowiak and U. Lundqvist. 1997. Barley Genet. Newsl. 26:209.
J.D. Franckowiak. 2007: Barley Genet. Newsl. 37:245-246.
J.D. Franckowiak. 2010. Barley Genet. Newsl. 40:69-70.
J.D. Franckowiak. 2016. Barley Genet. Newsl. 46:67-68.
J.D. Franckowiak. 2018. Barley Genet. Newsl. 48:102-103.
J.D. Franckowiak. 2007: Barley Genet. Newsl. 37:245-246.
J.D. Franckowiak. 2010. Barley Genet. Newsl. 40:69-70.
J.D. Franckowiak. 2016. Barley Genet. Newsl. 46:67-68.
J.D. Franckowiak. 2018. Barley Genet. Newsl. 48:102-103.
