Agriculture Reference
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Lr2a were widely grown in this region starting in
the mid-1970s. Isolates with virulence to both
genes increased rapidly and were over 65% in the
early 1990s (Fig. 5.1c). Virulence to Lr16 has
increased from the mid-1990s since many current
spring wheat cultivars have this gene. In Australia
cultivars with either combination of genes Lr13 ,
Lr23 , and Lr34 or genes Lr1 , Lr13 , and Lr23 are
currently resistant to leaf rust (Bariana et al.,
2007). Cultivars with various combinations of
Lr13 , Lr24 , Lr34 , and Lr37 are considered mod-
erately susceptible to leaf rust.
Leaf rust resistance genes up to Lr60 have been
designated (McIntosh et al., 2007). Genes Lr1
(Cloutier et al., 2007), Lr10 from common wheat
(Feuillet et al., 2003), and Lr21 from Ae. tauschii
(Huang et al. 2003) have been sequenced. The
three genes have NBS-LRR regions typical of
resistance genes with isolate specifi city. Genes
Lr1 and Lr10 are widely ineffective, and Lr21 has
provided effective resistance in spring wheat cul-
tivars in the US and Canada. Isolates with viru-
lence to Lr21 would be expected to increase if this
gene was used in a winter wheat cultivar in the
US. Many of the other Lr genes for which viru-
lent isolates of P. triticina have not been found
have also not been widely used in wheat improve-
ment programs.
as Lr34 , and mapped to chromosome 7DS
(Dyck 1987). Singh and Rajaram (1992) deter-
mined that Frontana also carries other genes
besides Lr34 that condition adult-plant leaf rust
resistance. Frontana was used as a leaf rust resis-
tant parent in spring wheat programs in Minne-
sota and also at CIMMYT. The Minnesota
cultivar Chris was derived from crosses with
Frontana and released in 1966 as the fi rst spring
wheat in the US to have Lr34 . The CIMMYT
cultivars Penjamo 62, Lerma Rojo, and Nainari
60 also had Lr34 .
Wheat lines and cultivars with Lr34 optimally
express leaf rust resistance in the adult-plant
stage. Isolates of P. triticina with complete viru-
lence to lines with Lr34 have not been found in
North America (Kolmer et al., 2003, McCallum
and Seto-Goh 2006), despite the presence of
wheat cultivars with Lr34 for over 40 years. In
fi eld plots lines with only Lr34 can have moderate
to high levels of leaf rust severity, although these
can usually be distinguished from completely sus-
ceptible lines if leaf rust readings are made when
known susceptible lines are at near-terminal
severity (Color Plate 9a). Lines with Lr34 can also
express resistance in seedling plants at cooler
temperatures (Singh 1992b; Pretorius et al.,
1994). The presence of Lr34 enhances the
response of other effective resistance genes in
the same wheat genotype (German and Kolmer
1992). The presence of Lr34 is also associated
with a distinctive leaf-tip necrosis (Singh 1992a)
that can vary between genotypes and environ-
ments. Wheat cultivars with other Lr genes com-
bined with Lr34 are often more resistant than
lines with only Lr34 or the other genes singly.
Spring wheat cultivars with combinations of
Lr13 , Lr16 , and Lr34 were highly resistant in
Canada (Samborski and Dyck 1982; Liu and
Kolmer 1997) and the US (Ezzahiri and Roelfs
1989).
Diagnostic molecular markers closely linked to
Lr34 have been developed (Bossolini et al., 2006;
Lagudah et al., 2006) that will greatly simplify
selection of breeding materials with Lr34 . In a
survey of wheat classes in the US using the Lr34
marker csLV34 , the allele associated with the
presence of Lr34 was completely absent in soft
Durable leaf rust resistance in wheat
The development of wheat cultivars with high
levels of effective durable resistance will depend
on genes that confer nonspecifi c resistance or
gene combinations that have proven to be effec-
tive over time. The cultivar Frontana released in
Brazil in 1946 has been a valuable source of
durable nonspecifi c leaf rust resistance. Dyck et
al. (1966) backcrossed leaf rust resistance from
Frontana into Thatcher. Backcross lines with the
adult-plant gene Lr13 were characterized, yet
none of the lines was as resistant as Frontana
because an additional gene was needed to recover
the original resistance in Frontana. Dyck and
Samborski (1982) characterized gene LrT2 in a
group of wheat cultivars that included 'Terenzio'
and Frontana. Later, LrT2 was determined to
be the additional gene in Frontana, designated
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