Biology Reference
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study is needed to elucidate the biochemical
function of Xa25 protein in rice-
Xoo
interaction.
Genetic studies have revealed that recessive
xa25
gene has the nature of dominance reversal;
it mediates
Xoo
resistance recessively at seeding
stage but dominantly at adult stage (thus it was
named
Xa25(t)
in Chen et al. 2002; Liu et al.
2011). However, transgenic rice plants carrying
both recessive
xa25
and its dominant allele
Xa25
as a transgene are susceptible to
Xoo
at both
seedling and adult stages, confirming that
xa25
is a recessive gene. This dominance reversal of
xa25
-carrying plants is associated with reduction
of
Xoo
-induced expression of dominant
Xa25
at
the adult stage as compared to
Xa25
expression
at seedling stage.
including
Xa2
,
Xa4
,
Xa7
,
Xa10
,
xa24
,
Xa30
,
Xa31(t)
, and
xa34(t)
, have been fine-mapped.
The
Xa2
gene is mapped to an approximately
190-kb region on the long arm of chromosome
4 (He et al. 2006). The
Xa4
gene is defined by
a 47-kb DNA fragment on the long arm of chro-
mosome 11, and the
Xa4
locus is linked or tightly
linked to the
Xa3/Xa26
locus (Sun et al. 2003,
2004).
Xa7
is located in a 118.3-kb region on the
long arm of chromosome 6 (Chen et al. 2008).
Xa10
is mapped to a 74-kb region on the long
arm of chromosome 11 (Gu et al. 2008). The
Xa22(t)
is localized to a 100-kb region of chro-
mosome 11, and this locus is also tightly linked
to
Xa3/Xa26
locus (Wang et al. 2003). The reces-
sive
xa24
is mapped to a 71-kb DNA fragment
on the long arm of chromosome 2 (Wu X. et al.
2008). The
Xa30
is mapped to a 38-kb region
on the long arm of chromosome 4 (Cheema et al.
2008). The
Xa31(t)
is limited to a length of about
100 kb on the long arm of chromosome 11 (Wang
et al. 2009). The recessive
xa34(t)
is defined to
a 204-kb DNA fragment near the centromeric
region of chromosome 1 (Chen et al. 2011). The
fine-mapping information of these genes will
facilitate breeding programs by marker-assisted
selection (MAS). Furthermore, the isolation of
these
MR
genes will deepen understanding of
molecular mechanism underlying BB disease in
general, and the opportunity to develop func-
tional markers for more precise breeding.
Xa27
Xa27
, localized on the long arm of chromosome
6, mediates resistance to diverse strains of
Xoo
,
including Chinese
Xoo
strains and Philippine
Xoo
races 2 to 6. It was isolated from
indica
rice line IRBB27 by map-based cloning (Gu
et al. 2005).
Xa27
encodes an apoplast protein
of 113 amino acids that has no distinguishable
sequence similarity to proteins from organisms
other than rice (Wu L. et al. 2008). The resis-
tant and susceptible alleles of
Xa27
encode an
identical protein, whereas the promoters of this
pair of alleles have crucial sequence differences
that determine the specific recognition of
Xoo
(Gu et al. 2005). The resistance of
Xa27
is dose
dependent. The TAL effector AvrXa27 from
Xoo
induces
Xa27
expression by binding to the UPA
box (upregulated by AvrBs3) of the
Xa27
pro-
moter (Boch et al. 2009). However, the recessive
MR
gene
xa5
can attenuate the
Xa27
-mediated
resistance in rice, which suggests that
Xoo
TA L
effector could not use protein encoded by the
recessive
xa5
as a transcription machinery to
activation of
Xa27
(Gu et al. 2009).
Quantitative Resistance to Xoo
Researchers commonly study quantitative resis-
tance by identifying disease resistance QTLs. At
least 74 QTLs against
Xoo
have been identified
in different rice cultivars interacting with differ-
ent
Xoo
strains (Figure 2.3; Li et al. 1999; Luo
et al. 1998; Yu et al. 2003; Wang et al. 2005;
Li et al. 2006; Yang CD et al. 2006; Hu et al.
2008; Kou et al. 2010; Fu et al. 2011; Deng et al.
2012). These QTLs are distributed on all 12 chro-
mosomes. Several resistance QTLs span a large
segment of a chromosome, indicating the poor
quality of the data. However, resistance QTLs
Fine-MappedMR Genes
In addition to the fine-mapping of the seven
characterized
MR
genes, eight other
MR
genes,
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