Proposed gene
symbols for resistance to Barley Mild Mosaic Virus (BaMMV) in barley
T. Konishi
294 Okada,
Mabi-cho Kibi-gun, Okayama 710-1311, Japan
Soil infestation
with barley yellow mosaic virus (BaYMV) is the most serious barley cultivation
problem in East Asia and Europe, and it is gradually dispersing to neighboring
regions together with barley mild mosaic virus (BaMMV). Both viruses are
soil-borne and transmitted by the root-infecting fungus Polymyxa graminis Led.
and are classified in the same genus bymovirus. However, they differ in
their biological and serological properties, RNA sequences, and aggressiveness
to barley cultivars.
The best way to
minimize damage caused by these viruses is to grow the resistant cultivars.
Extensive surveys of barley accessions have identified resistant genotypes and
genetic analyses have elucidated inheritance patterns. Takahashi et al. (1973)
first assigned the gene symbol Ym to a gene conferring resistance to
BaYMV, and subsequently resistance genes have been assigned symbols up to ym11
(Bauer et al. 1997). However, some of these genes confer resistance to BaYMV
while others control resistance to BaMMV. Genes for resistance to BaYMV are
designated as rym1, formerly Ym (Konishi et al. 1997), Ym2
(Takahashi et al. 1973), ym3 (Ukai and Yamashita 1980), rym5
(Konishi et al. 1997), and rym6 (Iida et al. 1999).
A mechanical
inoculation technique for BaMMV tests was developed by Friedt (1983) and was
used to identify other resistance genes in this series. Since mechanical
inoculation procedures for BaMMV are easier and more stable than testing for
BaYMV in the infested field, many genetic analyses in Europe, especially in
Germany, have been conducted using the mechanical inoculation method. The ym4
gene was first identified as a BaYMV resistance gene by this method (Kaiser and
Friedt 1989), but later changed to a BaMMV resistance gene by Kaiser and Friedt
(1992). This problem arose because barley cultivars are frequently evaluated in
Europe in fields infected with both BaYMV and BaMMV. Utilization of mechanical
inoculation has identified several genes conferring resistance to BaMMV, but
they were included in the ym series and designated as ym4
followed by ym7, ym8, ym9, ym10 and ym11.
It should be
pointed out that resistance to BaYMV is different from resistance to BaMMV.
Cultivars with the ym3 gene are resistant to BaYMV and susceptible to
BaMMV, the same situation is observed for the ym5 cultivars (Konishi and
Kaiser-Alexnat 2000). These results strongly support a distinction between
genes for BaMMV resistance and those for BaYMV resistance. The gene symbol mm
was previously recommended for BaMMV resistance genes (Götz and Friedt 1993).
Based on the scheme of three letter symbols for barley genes (Franckowiak et
al. 1997), I propose that genes controlling reaction to BaMMV should be
designated as rmm, the abbreviated for reaction to barley mild
mosaic virus, to distinguish them from rym genes for reaction
to barley yellow mosaic virus.
Since the current
symbols for designation of genes for reaction to BaYMV and BaMMV is progressive,
a new nomenclature system should be considered and applied before serious
confusion occurs among barley researchers. The proposed gene symbol changes for
factors controlling reactions of barley to infection with BaMMV are as follows:
ym4 in Franka
and resistant German cultivars to rmm1,
ym7 in HOR3365 to rmm2,
ym8 in 10247 to rmm3,
ym9 in Bulgarian 347
to rmm4,
ym10 in Hiberna to rmm5,
and
ym11 in Russia 57 to rmm6.
Since the cultivar Franka possesses a gene
for resistance to BaYMV in addition to the rmm1 gene for resistance to
BaMMV (Konishi and Furusho 2000), I proposed that the rym4 gene symbol
be assigned to the BaYMV resistance gene in Franka. As a result, the series of
BaYMV resistance genes will be continuous from rym1 to rym6.
For the
recommended nomenclature scheme, the reaction of new genes to either BaYMV or
BaMMV should be determined first. When tested using mechanical inoculation with
BaMMV, the resistance gene should be designated as a BaMMV resistance gene (rmm).
Next, the number of genes governing reaction to BaYMV or BaMMV should be
investigated and the resistance gene(s) should be mapped by, for example, QTL
analysis using molecular markers. The Chinese barley landrace, Mokusekko 3,
possesses two genes for resistance to BaYMV, rym1 and rym5 in
chromosomes 4HL and 3HL, respectively (Konishi et al. 1997). Further
investigations show that some other resistant accessions also contain two
resistance genes for BaYMV. Some questions might remain regarding this new
system. If a resistance gene reacts to both viruses simultaneously, there is a
question as whether the gene symbol should be rym or rmm. I
suggest that the rym symbol have priority over rmm because BaYMV
is more serious and wide spread than BaMMV. Although genes conferring
resistance to both viruses are possible, none have been identified. Most
landraces and cultivars as Franka that are resistance to BaYMV and BaMMV appear
to possess different genes for reaction to each virus.
References:
Bauer, E., J. Weyen, A. Schiemann, A. Graner, and F.
Ordon. 1997. Molecular mapping of novel resistance genes against barley mild
mosaic virus (BaMMV). Theor. Appl. Genet. 95:1263-1269.
Franckowiak, J.D., U. Lundqvist, and T. Konishi. 1997.
New and revised names for barley genes. BGN 26:4-8.
Friedt, W.
1983. Mechanical transmission of
soil-borne Barley Yellow Mosaic Virus. Phytopath. Z. 106:16-22.
Götz, R., and W. Friedt. 1993. Resistance to the
barley yellow mosaic virus complex. Differential genotypic reactions and
genetics of BaMMV-resistance of barley (Hordeum vulgare L.). Plant
Breed. 111:125-131.
Iida, Y., T. Ban, and T. Konishi. 1999.
Linkage analysis of the rym6 resistance gene to Japanese strain II of
barley yellow mosaic virus (BaYMV-II) in barley. BGN 29 (in press).
(http://wheat.pw.usda.gov/ggpages/bgn/29/a29-07.html)
Kaiser, R., and W. Friedt. 1989. Chromosomal
location of resistance to Barley Yellow Mosaic Virus in German winter-barley
identified by trisomic analysis. Theor. Appl. Genet. 77:241-245.
Kaiser, R., and W. Friedt. 1992. Gene for resistance
to barley mild mosaic virus in German winter barley located on chromosome 3L.
Plant Breed. 108:169-172.
Konishi, T., T. Ban, Y. Iida, and R. Yoshimi.
1997. Genetic analysis of disease
resistance to all strains of BaYMV in a Chinese barley landrace, Mokusekko 3.
Theor. Appl. Genet. 94:871‑877.
Konishi, T., and M. Furusho. 2000. German barley cv. >Franka= possesses two
different resistance genes to barley yellow mosaic virus (BaYMV) and barley
mild mosaic virus. BGN 30 (submitted).
Konishi, T., and R. Kaiser-Alexnat. 2000.
Reaction of barley accessions to BaYMV and BaMMV in Japan, compared with
data in Germany. BGN 30 (submitted).
Takahashi, R., J. Hayashi, T. Inouye, I. Moriya, and
C. Hirao. 1973. Studies on resistance to yellow mosaic
disease in barley. I. Tests for varietal reactions and genetic analysis of
resistance to the disease. Ber. Ohara Inst. landw. Biol., Okayama Univ. 16:1-17
Ukai, Y., and A. Yamashita. 1980. Induced mutation
for resistance to barley yellow mosaic virus. Japan J. Breed. 30:125-130.