Barley
Genetics Newsletter (2007) 37: 188-301
BGS 1, Brachytic 1, brh1
Stock number: BGS
1
Locus name: Brachytic
1
Locus symbol: brh1
Previous nomenclature and gene symbolization:
Brachytic = br (10, 12).
Breviaristatum-i =
ari-i (5, 8).
Dwarf x = dx1 (6).
Inheritance:
Monofactorial recessive (10, 12).
Located in chromosome 7HS [1S] (3), about 9.3 cM distal from
the fch12 (chlorina seedling 12)
locus (12), 0.8 cM distal from RFLP marker BCD129 (9), about 5.0 cM from AFLP
marker E4134-8 in subgroup 1 of the Proctor/Nudinka map (11), and about 13.6 cM
proximal from SSR marker HVM04 in bin 1H-02 (2).
Description:
Plants have short leaves, culms, spikes, awns, and kernels.
The seedling leaf is about 2/3 normal length. A similar reduction in the size
of other organs is observed, but the awns are less than 1/2 normal length (6).
The mutant phenotype is easy to classify at all stages of growth. The
approximately 20% reduction in kernels size is caused primarily by a reduction
in kernel length. The yields of the brh1
mutants are about 2/3 normal and lodging is greatly reduced in the Bowman brh1 lines (2). Börner (1) reported that
ari-i.38 seedlings are sensitive to
gibberellic acid. Powers (10) states that the assigned gene symbol for this
mutant is br and that L.J. Stadler
selected this symbol.
Origin of mutant:
A spontaneous mutant in
Mutational events:
brh1.a in Himalaya (12); brh1.c
(GSHO 229) in Moravian (PI 539135) (13); ari-i.38
(NGB 115888, GSHO 1657) in Bonus (PI 189763) (8, 14); brh1.e (GSHO 1690) in Aramir (PI 467786) (14); brh1.f (dx1, GSHO 1422)
in Domen (CIho 9562) (6); brh1.t
(OUM136, GSHO 1691) in Akashinriki (PI 467400, OUJ659); brh1.x (7125, DWS1224, GSHO 1692) in Volla (PI 280423); brh1.z (Hja80001) in Aapo; brh1.aa (Hja80051) in a Hja80001 cross
(4, 7); and brh1.ae (FN53) in Steptoe
(CIho 15229) (4).
Mutant used for description and seed stocks:
brh1.a in
References:
1. Börner, A. 1996. GA response in semidwarf barley. Barley
Genet. Newsl. 25:24-26.
2. Dahleen, L.S., L.J. Vander Wal, and J.D. Franckowiak.
2005. Characterization and molecular mapping of genes determining semidwarfism
in barley. J. Hered. 96:654-662.
3. Fedak, G., T. Tsuchiya, and S.B. Helgason. 1972. Use
of monotelotrisomics for linkage mapping in barley.
4. Franckowiak, J.D. 1995. The brachytic class of semidwarf
mutants in barley. Barley Genet. Newsl. 24:56-59.
5. Gustafsson, Å., A. Hagberg, U. Lundqvist, and G. Persson.
1969. A proposed system of symbols for the collection of barley mutants at
Svalöv. Hereditas 62:409-414.
6. Holm, E., and K. Aastveit. 1966. Induction and effects of
the brachytic allele in barley. Adv. Front Plant Sci. 17:81-94.
7. Kivi, E. 1986. (personal communications).
8. Kucera, J., U. Lundqvist, and Å. Gustafsson. 1975.
Inheritance of breviaristatum mutants in barley. Hereditas 80:263-278.
9. Li , M., D. Kudrna, and A. Kleinhofs. 2000. Fine mapping
of a semi-dwarf gene brachytic 1 in barley. p. 72-74. In S. Logue (ed.) Barley Genetics VIII. Volume III, Proc. Eighth
Int. Barley Genet. Symp.,
10. Powers, L. 1936. The nature of the interactions of genes
affecting four quantitative characters in a cross between Hordeum deficiens and vulgare.
Genetics 21:398-420.
11. Pozzi, C., D. di Pietro, G. Halas, C. Roig, and F.
Salamini. 2003. Integration of a barley (Hordeum
vulgare) molecular linkage map with the position of genetic loci hosting 29
developmental mutants. Heredity 90:390-396.
12. Swenson, S.P. 1940. Genetic and cytological studies on a
brachytic mutant in barley. J. Agric. Res. 60:687-713.
13. Szarejko,
14. Tsuchiya, T. 1974. Allelic relationships of genes for
short-awned mutants in barley. Barley Genet. Newsl. 4:80-81.
Prepared:
T. Tsuchiya and T.E. Haus. 1971. BGN 1:104.
Revised:
T. Tsuchiya. 1980. BGN 10:100.
J.D. Franckowiak. 1997. BGN 26:44.
J.D. Franckowiak and L. S. Dahleen. 2007. BGN 37:188-189.
BGS 2, Chlorina
seedling 12, fch12
Stock number: BGS
2
Locus name: Chlorina
seedling 12
Locus symbol: fch12
Previous nomenclature and gene symbolization:
Chlorina seedling-c = fc
(3).
Chlorina seedling-fc = clo-fc
(7).
Inheritance:
Monofactorial recessive (3).
Located in chromosome 7HS [1S] (1,
4), about 3.6 cM distal from the gsh3
(glossy sheath 3) locus (6), and about 9.3 cM proximal from the brh1 (brachytic 1) locus (8), in bin
7H-02 about 2.3 cM from RFLP marker KFP027 and co-segregating with markers
BCD130 and ABC327 (5).
Description:
Seedling leaves are yellow with green tips and new leaves
show a yellow base and a green tip. As the plant develops, leaf color changes
to pale green (3). Plants are vigorous, but anthesis is delayed and seed yield
may be low.
Origin of mutant:
A spontaneous mutant in Colsess (CIho 2792) (3).
Mutational events:
fch12.b (fc) in
Colsess (Colsess V) (3); fch12.l
(Trebi chlorina 453, GSHO 155), fch12.m
(Trebi V, GSHO 158), fch12.n
(Trebi IX, GSHO 18), fch12.o (Trebi
XI, GSHO 163) in Trebi (PI 537442) (2); clo-fc.110 in Bonus (PI 189763) (7); fch12.b may be present in the brachytic chlorina stocks (GSHO 124 and GSHO 174) (9).
Mutant used for description and seed stocks:
fch12.b in Colsess (GSHO
36); fch12.b in Bowman (PI 483237)*7 (GSHO 1826).
References:
1. Fedak, G., T. Tsuchiya, and S.B. Helgason. 1972. Use of
monotelotrisomics for linkage mapping in barley.
2. McMullen, M. 1972. Allelism testing of seven chlorina
mutants in Trebi barley. Barley Genet.
Newsl. 2:76-79.
3. Robertson, D.W., and G.W. Deming. 1930. Genetic studies
in barley. J. Hered. 21:283-288.
4. Robertson, D.W., G.W. Deming, and D. Koonce. 1932.
Inheritance in barley. J. Agric. Res. 44:445-466.
5. Schmierer, D., A. Druka, D. Kudrna, and A. Kleinhofs. 2001. Fine mapping
of the fch12 chlorina seedling
mutant. Barley Genet. Newsl. 31:12-13.
6. Shahla, A., and T. Tsuchiya. 1987. Cytogenetic studies in
barley chromosome 1 by means of telotrisomic, acrotrisomic and conventional
analysis. Theor. Appl. Genet. 75:5-12.
7. Simpson, D.J., O. Machold, G. Høyer-Hansen, and D. von
Wettstein. 1985. Chlorina mutants of
barley (Hordeum vulgare L.).
Carlsberg Res. Commun. 50:223-238.
8. Swenson, S.P. 1940. Genetic and cytological studies on a
brachytic mutation in barley. J. Hered. 31:213-214.
9. Wang, S., and T. Tsuchiya. 1991. Genetic analysis of the
relationship between new chlorina mutants in genetic stocks and established f series stocks in barley. Barley Genet.
Newsl. 20:63-65.
Prepared:
T. Tsuchiya and T. E. Haus. 1971. BGN 1:105.
Revised:
T. Tsuchiya. 1980. BGN 10:101.
J.D. Franckowiak and A. Hang. 1997. BGN 26:45.
J.D. Franckowiak. 2007. BGN 37:190-191.
BGS 6, Six-rowed spike 1, vrs1
Stock number: BGS
6
Locus name: Six-rowed
spike 1
Locus symbol: vrs1
Previous nomenclature and gene symbolization:
Two-row vs six-row = Zz
(21).
Six-row vs two-row = Aa
(6).
Two-rowed = D
(17).
Six-row vs two-row =
Vv (3).
Six-row vs two-row (distichon)
vs two-row (deficiens) = A, as, af
(8).
Reduced lateral spikelet appendage on the lemma = lr (9).
Allelic series v, Vd, V, and Vt
(22).
Hexastichon mutants = hex-v
(5, 6).
Intermedium spike-d = Int-d
(4).
Reduced lateral spikelet appendage on the lemma = vlr (19).
The vrs1 DNA
sequence identified as HvHox1 (10).
Inheritance:
A multiple allelic series, incomplete dominant allele
interactions based on the size and shape of lateral spikelets (1, 19, 22).
Located in chromosome 2HL (3, 6, 12, 14), about 30.5 cM
distal from the eog1 (elongated outer
glume 1) locus (18), in bin 2H-09 and in a 0.90-cM interval between markers
cMWG699 and MWG865 (11).
Description:
Alleles at this complex locus modify development of the
lateral spikelets and the associated lemma awn. The vrs1.a allele (v gene) is
present in most six-rowed cultivars and produces well-developed lateral
spikelets (6). Based on phylogenetic
analysis of the six-rowed cultivars, the
six-rowed gene originated independently at least three times (vrs1.a1, vrs1.a2, and vrs1.a3)
from different wild type (Vrs1.b)
alleles (10). The lemma awn of lateral spikelets will vary from 3/4 to nearly
as long as those of central spikelets, depending upon alleles present at other
loci. The Vrs1.b allele (V gene, distichon) is present in many two-rowed cultivars and reduces
lateral spikelets to sterile bracts with a rounded tip. The Vrs1.t allele (Vt gene, deficiens)
causes an extreme reduction in the size of lateral spikelets. The lr or vlr (vrs1.c)
gene in Nudihaxtoni and Bozu types will not recombine with the vrs1.a allele (12, 19) and produces
phenotypes similar to the Vrs1.d
allele (Vd gene) of
Svanhals (22). The series of induced mutants in two-rowed barley called hex-v and Int-d mutants differ in the size of lateral spikelets, but they
interact with the vrs1.a allele as
incomplete dominants (5). Many heterozygous combinations with vrs1.a have a pointed tip on the lemma
of sterile lateral spikelets. Alleles at the int-c (intermedium spike-c) locus modify lateral size in the
presence of vrs1.a, Vrs1.b, and Vrs1.d, but not when Vrs1.t is present (22). Multiple origins of vrs1 alleles in six-rowed barley have been confirmed by molecular
analysis (20). Komatsuda et al. (10) found that expression of the Vrs1
gene was strictly localized in the lateral-spikelet primordia of immature
spikes and suggested that the VRS1 protein suppresses development of lateral
spikelets.
Origin of mutant:
Natural occurrence in six-rowed barley and induced
frequently by mutagenic agents (10, 14).
Mutational events:
vrs1.a1 in most six-rowed cultivars (1, 10, 22); vrs1.a2 in Dissa and Valenci (10), vrs1.a3 in Natsudaikon Mugi (OUK735)
(10), Vrs1.b in wild barley (10), Vrs1.b2 in Pamella Blue (OUH630) (10), Vrs1.b3 in Bonus (PI 189763) (10), Vrs1.t in a few two-rowed cultivars (10,
22); vrs1.c or lr in Nudihaxtoni (PI 32368) (12, 19); Vrs1.d in Svanhals (PI 5474) (22); 23 induced mutants from
programs in Belgium, Germany, and Hungary (2); hex-v.3 (NGB 115545), -v.4
(NGB 115546), -v.6 (NGB 115547), -v.7 (NGB115548), -v.8 (NGB 115549), -v.9 (NGB
115550), -v.10 (NGB 115551), -v.11 (NGB 115552), -v.12 (NGB 115553), -v.18 (NGB
115559), -v.44 (NGB 115581), -v.45 (NGB 115582 ), -v.46 (NGB 115583 ), -v.47 (NGB 115584), -v.48 (NGB 115585), in Bonus, -v.13
(NGB 115554), -v.14 (NGB 115555), -v.15 (NGB 115556), -v.16 (NGB 115557), -v.17 (NGB
115558), -v.19 (NGB 115560), -v.21 (NGB 115562), -v.22 (NGB 115563),
-v.23 (NGB 115564), -v.24 (NGB 115565), -v.25 (NGB 115566), -v.26 (NGB
115567), -v.27 (NGB 115568), -v.28 (NGB 115569), -v.29 (NGB 115570), -v.30 (NGB
115571), -v.31 (NGB 115572), -v.35 (NGB 115574) in Foma (CIho
11333), -v.20 (NGB 115561) in Ingrid
(CIho 10083), -v.33 (NGB 115573), -v.36 (NGB 115575), -v.38 (NGB 115576), -v.39 (NGB
115577), -v.41 (NGB 115578), -v.42 (NGB 115579), -v.43 (NGB 115580) in Kristina (NGB 1500) (5, 14); hex-v.49 (NGB 115586) in Bonus, -v.50 (NGB 115587), -v.51 (NGB 115588) in Sv 79353, -v.52 (NGB 119353) in Golf (PI 488529) (13); Int-d.11 (NGB 115429), -d.12 (NGB
115430), -d.22 (NGB 115440), -d.24 (NGB 115442), -d.28 (NGB 115446), -d.36 (NGB
115454) in Foma, -d.40 (NGB 115458), -d.41 (NGB 115459), -d.50 (NGB 115468), -d.57 (NGB
115475), -d.67 (NGB 115485), -d.68 (NGB 115486), -d.69 (NGB 115487) in Kristina (5, 15); Int-d.73 (NGB 115491), -d.80 (NGB
115498), -d.82 (NGB 115500) in Bonus, -d.93 (NGB 115511), -d.94 (NGB 115512), -d.96 (NGB
115514), -d.97 (NGB 115515), -d.100 (NGB 115518) in Hege (NGB 13692)
(13); vrs1.o (v1b) in New Golden (16).
Mutant used for description and seed stock:
vrs1.a in Trebi (PI 537442, GSHO 196); vrs1.a in Bonneville (CIho 7248) (7); vrs1.a from Glenn (CIho 15769) in Bowman (PI 483237)*8 (GSHO 1907);
Int-d.12 in Bowman*7 (GSHO 1910).
References:
1. Biffen, R.H. 1906. Experiments on the hybridization of
barleys. Proc. Camb. Phil. Soc. 13:304-308.
2.
3. Griffee, F. 1925. Correlated inheritance of botanical
characters in barley, and manner of reaction to Helminthosporium sativum. J. Agric. Res. 30:915-935.
4. Gustafsson, Å., A. Hagberg, U. Lundqvist, and G. Persson.
1969. A proposed system of symbols for the collection of barley mutants at
Svalöv. Hereditas 62:409-414.
5. Gustafsson, Å., and U. Lundqvist. 1980. Hexastichon and
intermedium mutants in barley. Hereditas 92:229-236.
6. Harlan, H.V., and H.K. Hayes.
1920. Occurrence of the fixed intermediate,
Hordeum intermedium haxtoni, in crosses between H. vulgare pallidium and H. distichum palmella. J. Agric.
Res.19:575-591.
7. Hockett, E.A. 1985. Registration
of two- and six-rowed isogenic Bonneville barley germplasm. Crop Sci. 25:201.
8. Hor, K.S. 1924. Interrelations of
genetic factors in barley. Genetics 9:151-180.
9. Immer, F.R., and M.T. Henderson.
1943. Linkage studies in barley. Genetics 28:419-440.
10. Komatsuda, T., M. Pourkheirandish, C. He, P. Azhaguvel, H. Kanamori,
D. Perovic, N. Stein, A. Graner, T. Wicker, A. Tagiri, U. Lundqvist, T.
Fujimura, M. Matsuoka, T. Matsumoto, and M. Yano. 2007. Six-rowed barley
originated from a mutation in a homeodomain-leucine zipper I-class homeobox
gene. PNAS 104:1424-1429.
11. Komatsuda, T., and K. Tanno. 2004. Comparative high resolution map of
the six-rowed locus 1 (vrs1) in
several populations of barley, Hordeum
vulgare L. Hereditas 141:68-73.
12.
Leonard, W.H. 1942. Inheritance of reduced lateral spikelet appendages in the
Nudihaxtoni variety of barley. J. Am. Soc. Agron. 34:211-221.
13. Lundqvist, U. (unpublished).
14. Lundqvist, U., and A. Lundqvist.
1987. Barley mutants - diversity and genetics. p. 251-257. In S. Yasuda and T. Konishi (eds.) Barley Genetics V. Proc. Fifth
Int. Barley Genet. Symp.,
15. Lundqvist, U., and A. Lundqvist. 1988. Induced intermedium
mutants in barley: origin, morphology and inheritance. Hereditas 108:13-26.
16. Makino, T., M. Furusho, and T. Fukuoka. 1995. A mutant
having six-rowed gene allelic to v
locus. Barley Genet. Newsl. 24:122.
17. Miyake, K., and Y. Imai. 1922. [Genetic studies in
barley. 1.] Bot. Mag.,
18. Swenson, S.P., and D.G. Wells. 1944. The linkage
relation of four genes in chromosome 1 of barley. J. Am. Soc. Agron.
36:429-435.
19. Takahashi, R., J. Hayashi,
20. Tanno, K., S.
Taketa, K. Takeda, and T. Komatsuda. 2002.
A DNA marker closely linked to the vrs1
locus (row-type gene) indicates multiple origins of six-rowed barley (Hordeum vulgare L.) Theor. Appl.
Genet. 104:54-60.
21. Ubisch, G.
von. 1916. Beitrag zu einer Faktorenanalyse von Gerste. Z. Indukt. Abstammungs. Vererbungsl. 17:120-152.
22. Woodward, R.W. 1949. The inheritance of fertility in the
lateral florets of the four barley groups. Agron. J. 41:317-322.
Prepared:
T.E. Haus. 1975. BGN 5:106.
Revised:
J.D. Franckowiak and U. Lundqvist. 1997. BGN 26:49-50.
U. Lundqvist and J.D. Franckowiak. 2007. BGN 37:192-194.
BGS 7, Naked caryopsis 1, nud1
Stock number: BGS
7
Locus name: Naked
caryopsis 1
Locus symbol: nud1
Previous nomenclature and gene symbolization:
Naked caryopsis = k
(14).
Naked caryopsis = s
(21).
Naked caryopsis = n
(6, 9).
Hulless = h (10).
Inheritance:
Monofactorial recessive (6, 14, 19).
Located in chromosome 7HL [1L] (3, 11, 12, 14, 20), near the
centromere (3, 11), about 9.6 cM proximal from the lks2 (short awn 2) locus (15), about 10.5 cM proximal from the dsp1 (dense spike 1) locus (15, 16), in
bin 7H-07 about 13.1 cM distal from RFLP marker MWG808 (2), co-segregating with
AFLP markers KT3 and KT7 and SCAR marker sKT7 (7), about 0.06 cM distal from
SCAR marker sTK3 and the same distance proximal from sTK9 (17).
Description:
The lemma and palea do not adhere to the caryopsis and the
grain will thresh free of the hull at maturity. The naked caryopsis trait is
expressed in all environments (16). The naked lines fail to produce a cementing
substance present in covered lines (4). The nud1.a
mutant depressed the expression by 10 to 20% of other traits such as plant
height, seed weight (1, 8) and altered malt quality parameters (8). The nud1.a gene is often associated with the
dsp1.a (dense spike 1) gene in
Japanese cultivars (16). Allele IV of
the marker sKT7 near the nud1 locus
was the only one found in naked barley cultivars (18); however, the geographic
distribution for haplotypes of allele IV
suggest migration of naked types toward eastern
Origin of mutant:
In an unknown cultivar, b