Review

Breeding on Durable Leaf Rust Resistant on Wheat

Table of Contents
ABSTRACTS

II

INTRODUCTION

GENE FOR GENE INTERACTION

CONCEPT OF DURABLE RESISTANCE

BREEDING FOR LEAF RUST RESISTANCE

Lr34
Lr34/Yr18

5
6

Lr 46
Lr46/Yr29

6
7

Lr67

CONCLUSION

REFERENCES

Abstracts
Leaf rust caused by Puccinia recondita f.sp. tritici. is an important disease of wheat (Triticum
aestivum L.) in worldwide. Rust diseases continue to cause significant losses to wheat production
worldwide. Genetic resistance is the most effective approach to managing wheat leaf rust. Several
rust resistance genes have been identified and used in breeding for resistance but new variants of
the pathogen (referred to as races) overcome the resistance over a period of time. The semi-dwarf
wheat varieties with race specific resistance could not survive longer due to the evolution of new
rust races. Most plant breeders and pathologists now advocate durable resistance to rusts based on
multiple genes as best source of resistance as breeding for this type of resistance tends to produce
long-lasting solutions. Wheat breeders are now increasingly focusing on the identification and
incorporation of race nonspecific resistance genes that may provide only partial resistance but
when used in combination with other genes can condition highly effective resistance. Genes
expressed in seedling plants have not provided long-lasting effective leaf rust resistance. Adultplant resistance genes Lr34 and Lr46 singly and together have provided the most durable resistance
to leaf rust in wheat throughout the world. Continued efforts to isolate, characterize, and map leaf
rust resistance genes is essential given the ability of the leaf rust fungus to overcome deployed
resistance genes. Molecular markers are becoming available for many genes and their use in
marker-assisted selection will certainly have a remarkable impact in practical breeding.
Keywords: Leaf rust, durable resistance, APR, wheat, Lr genes

[Note: Durable resistance, minor gene resistance, slow rusting resistance, non-specific resistance
and adult plant resistance are terms that have been used interchangeably in the literature.]

Introduction
Wheat is the major cereals crop in the world that is most consumed and traded. In developing
countries, it is the second most important food crop after rice. It is the major source of calorie of
the people all over the world. It is no doubt that increase in population has alarmed the pressure
on performance of wheat production to ensure the food security globally. Demand of wheat has
outstripped its production so its global stocks are constantly under pressure
[http://www.icarda.org]. Some concerned is still require to make more effort on certain factor so
that production could enhance. There are many activities that have been carried out for wheat
performance but in this complex nature, the problems are challenging with their different avatar.
Nature has some surprise up her sleeve- and not all of them are pleasant, wheat rusts included.
Wheat rust, a devastating disease known as polio of agriculture, has spread from Africa to South
and Central Asia, the Middle East and Europe, with calamitous losses for the worlds second most
important grain [www.independent.co.uk].
Rust, cosmopolitan disease, is the disease caused by fungus of Pucciniaceae family of genus
Puccinia. The rust is named after the symptoms caused by the pathogen on plant surface which
resembles with the rust of iron rod. In wheat, there are mainly three types of rust which has
economic importance in agriculture and their concerned field. The yellow rust, also called as stripe
rust is caused by Puccinia stiriformis, stem rust caused by P. graminis f.sp. tritici and brown rust
also called as leaf rust is caused by P. recondita f.sp. tritici. The yellow rust of wheat has
microcylic lifecycle with single host wheat, while leaf rust and stem rust has macrocylic lifecycle.
P. graminis f.sp. tritici have two other alternate host: Barberis spp and Mohonia spp. and P.
recondita has three other alternate host: Thalictrum,. Anchusa and Isopyrum (RA McIntosh et al.,
1995). The causal organism of rust is considered to be an obligate parasite. The fungus produced
yellow uredinospores 20-30 um in diameter. Uredinospore is followed by the teliospore production
in late in the growing season. These uredinospore from the initial infection site gets dispersed with
the help of air by thousands of kilometer if the environment is dry and cool (Khan et al., 2013).
Leaf rust, though causes less loss as compare to other two rusts, occurs more regularly and in more
worldwide region than stem or stripe rust of wheat (Huerta-Espino et al. 2014). Because of its
regular and widespread occurrence, the global leaf rust damages are greater than the other two rust
(Kolmar 1996). According to Roefls (1989) yield loss in wheat from P. triticina infection are
usually the results of decrease in weight per grain and number of grain per head. Upto 40% losses
has been estimated due to the infection of leaf rust on susceptible wheat.
With the upheavals of economic and social condition that is resulted from the yield losses
following with epidemic has driven the attention of most of the crop scientist towards research to
get the economically and financially justifiable solution. Many conventional methods and use of
hazardous chemical pesticides were adopted for several years which were not economically and
ecologically sustainable. After the progressive studies and research, scientists find the disease
biology and relationship with host which was one of the door for resistant breeding. In present,
most of the plant protectionists are oriented towards the resistant breeding. Gene for gene
relationship is the major tool that is used in resistant breeding. However, identifying and utilizing

rust resistance gene in wheat has been hampered by the continuous and rapid emergence of new
pathogen races.
Breeding for resistance against leaf rust is an economical, efficient and environmentally safe
control measure to reduce wheat production losses. Development of disease resistant varieties is
one of the most economical methods of control of diseases like leaf rust. The virulent gene of
fungus that causes leaf rust disease on plant interact with resistant gene of wheat in a gene-forgene manner. Nevertheless, growing of rust resistant varieties having single gene for resistance
results in rapid evolution of virulent biotypes of the pathogen, thereby making the resistance gene
ineffective and the variety susceptible to rust (Dhillion et al., 2011).
Resistance gene expression is dependent on the genetics of host-parasite interaction, temperature
conditions, plant developmental stage, and interaction between resistance genes with suppressors
or other resistance genes in the wheat genomes. Wheat plant exhibit two types of resistant over the
rust; Seedling resistant and Adult plant resistant (APR). Genes expressed in seedling plants have
not provided long-lasting effective leaf rust resistance. Seedling resistant are more effective than
adult plant resistant for short period. However, pathogen of leaf rust overcomes single-gene
resistance relatively quickly unless it is combined with other genes for resistance (Khan et al.,
2013). In adult plant resistant, also called as slow rusting resistant, genes are considered to be more
durable, and their expression is marked by slow disease development, longer latent period, fewer
and smaller uredinia, and lower spore production (Basnet et al., 2012). Despite of seedling
susceptibility, adult plant resistance is expressed in adult stage of plant. APR develops as the plant
grows: it can start from stem elongation to heading. Adult plant resistance (APR) gene, if present
in single, cannot provide the effective resistant over the high pressure of disease. Research has
revealed that three to five genes combination can provide the effective resistant to the disease as
compare to the highly race specific resistance. Slow-rusting resistance is quantitatively inherited;
therefore, to adopt this kind of resistant in wheat plant, it is important to understand the heritability
of such genes. At present, Shahin and El-Orabey (2015) reported that more than 80 genes and
alleles of leaf rust resistance Lr genes have been identified and described. Among them 33 Lr
genes were transferred from other species into Triticum aestivum L.
Gene for gene interaction
Flor (1946), studied inheritance of pathogenicity in the pathogen and inheritance of resistance in
the host using flax (Linum usitatissimum)-flax rust (Melampsora lini) as a model system. He
concluded, that for every gene that conditions resistance in the plant, there is a corresponding and
complementary gene that conditions avirulence in the pathogen, and only the corresponding
avirulence gene can initiate the hypersensitive reaction (HR) leading to incompatibility. Resistance
and avirulence inherit in most cases in a dominant manner while, susceptibility and virulency in a
recessive manner. This system has the implication that resistance will not remain effective if the
pathogen acquires the corresponding virulence by losing the avirulence alleles that elicits
resistance either by deletion or by genetic change. Gene-for-gene system occurs most clearly in
pathosystems where a biotrophic, highly specialized pathogen is involved such as cereals with
various rusts, smuts and bunts, and with the powdery mildew. There are nevertheless numerous
examples of interactions in which no such relationship has been found. Examples include Flag
2

smut disease of wheat. Ellingboe (1982) stated that about 95% of all analyzed disease resistance
operates on gene-forgene pattern with the pathogen.
Concept of durable resistance
The resistance governed by the polygenes are more durable than the resistance governed by few
or single gene (Flor, 1971). However, it is difficult to fix the inheritance of polygene resistance
gene. Durable rust resistance is a mechanism conferring resistance to a cultivar for long period of
time during its widespread cultivation in a favourabe environment for a disease. Many genes were
previously identified for the hypersensitive resistance: governed by few or single genes, which
were more effective for the resistant. But the problem of newly emerged pathogen races has
overcome the resistance. Thus, the identification and use of potential sources of durable resistance
to leaf rust would be significant steps towards control of leaf rust thereby contributing to stability
of production and saving of expenditure on replacement of resistant cultivars. Breeder are giving
their attention towards locating the minor genes that contribute on durable resistance or slow
rusting in combination with other genes. In slow rusting resistance, infection is not completely
stopped but the spread of the disease is delayed. In general, slow rusting wheat has longer latent
periods, fewer uredinia, and smaller uredinia size at 10-14 days after inoculation with leaf rust than
susceptible wheat lines (Messmer et al., 2000). It has been indicated that durable rust resistance is
more likely to be of adult plant type rather than seedling type and is not associated with the genes
conferring hypersensitive reaction (Khan et al., 2013). Many terms and concepts have been
proposed to describe and characterize the types of resistance. Durable type resistance is mainly
explained in terms such as slow rusting, field, intermediate, quantitative, incomplete, general,
partial, horizontal, adult plant, race-non-specific resistance, etc. Each of these terms has a specific
meaning and specific implications, but they all describe a quantitative effect on the epidemic, and
many of them often are taken to imply complex inheritance (Priyamvada et al., 2011). The interest
in these types of resistance is fueled by the hope that they may be more durably effective against
the target pathogen. A general concept of a durable resistance source for a cereal rusts is as follow:

It may be controlled by more than a single gene;

It is more likely to operate at the adult-plant stage rather than at both juvenile or adult stage;
It confers non-hypersensitive response to infection.

Breeding for Leaf Rust resistance

Wheat leaf, stripe and stem rusts have devastating role in reducing crop yield resulting in socioeconomic instability many times across the world. In semi-dwarf wheat varieties rust resistance is
generally based on the race specific resistance gene that could not maintain the longer duration
due to the evolution of new rust races. However, varieties like Lerma Rojo-64, Yaqui-50 and
Lyalpur-73 developed in early part of green revolution retained resistance for longer time due to
presence of adult plant resistance (APR) genes. The durability of the race specific resistance has
created the necessity to search for the more durable type of resistance. Evolution of new rust races
like virulence Yr9 and Yr27 followed by the emergence of stem rust causing race Ug99 and its
mutant from Uganda lead the breeders to revise their breeding strategy (Rehman et al., 2013).
Breeder are now practicing to accumulate the minor genes with major genes for durability of rust
resistance in wheat variety. As durable resistance usually (not always) is governed by several genes
3

rather than by one major gene so breeding to attain durable resistance is more difficult because
several genes need to be transferred at one time, thus requiring large populations for selection, as
well as multiplying the usual problem with linkage drag (undesirable genes that are tightly linked
to the desired ones).In wheat, most of the resistance genes are effective at seedling stage and
remain effective through the adult plant stage. Some of the leaf rust resistance genes express
resistance optimally in adult plants and are known as adult plant resistance (APR) genes, which
depends on the genetics of the host pathogen interaction as well as favored environmental
conditions.
According to Singh et al. (2011), breeding for slow rusting resistance based on minor additive
genes has been challenging and often slow, for several reasons:
(1)
(2)
(3)
(4)

a sufficient number of minor genes may not be present in a single source genotype,
a source genotype may be poorly adapted,
there may be confounding effects from the segregation of both major and minor genes,
crossing and selection schemes and population sizes commonly used by breeding programs
are more suitable for selecting major genes,
(5) reliable molecular markers for several minor genes are unavailable,
(6) high costs associated with identifying and utilizing multiple markers.
Breeding for durable resistance is too much challenging and difficult as the source genotype itself
may be poorly adapted and a sufficient number of minor genes may not be present in a single
source genotype. For checking the presence of minor genes in the source genotypes, reliable
molecular markers are not available. One approach suggested in the literatures is to use recurrent
selection schemes to accumulate several minor genes in a single genetic background. The breeders
select the lines with the lower levels of disease severity and by doing that continuously over the
seasons, the level of durable resistance will increase (Parlevliet & Ommeren, 1988). There is
however one complication that if there is no durable major gene around it has to be taken into
account. Selection for resistance alone will not generate important popular cultivars, unless it
should carry some other traits like good quality and high yield also. The germplasm made of
combination of minor genes could be used in transferring these genes to adapted local cultivars
(Priyamvada et al., 2011). Durable resistance to leaf rust is thought to be more difficult to obtain
than to stem rust (Rubiales and Niks, 1995). But resistance against leaf rust has been identified
that appears more durable than usual.
Adult plant resistance genes express partial rust resistance phenotypes only in adult plants (except
under very specific conditions) and this is characterized by less and slower pathogen growth
without a necrotic response (sometimes referred to as slow rusting). Consequently, APR is
selected by wheat breeders in the field and not in the glasshouse. Reliable field sites for disease
assays that optimize natural infections or induced epidemics for effective selection of resistance
must be available. Although the resistance phenotypes conferred by individual APR genes show
varying levels of partial resistance, it has been reported that when several, mainly undefined APR
genes are combined, near immunity can be achieved in adult field grown plants (Singh et al.,
2014).

Lr34
One of the most studied and possibly the most utilized slow rusting leaf rust resistance gene Lr34
located on chromosome arm 7DS has maintained its moderate effectiveness over 60 years of use.
This gene was traced in the Italina var. Mentana using a gene based DNA marker. Lr34 is also
common in Chinese landraces including Chinese spring, and tall var. Frontana and Chris from
Brazil and USA respectively.
In 1930s Boerger and Klein in Uruguay identified a resistant selection from a local landrace, which
was released as Americano-44D in Uruguay and as Universal 2 in Argentina. These cultivars were
widely grown, and since then have been used as source of long term leaf rust resistance. In 1935
Beckman in Brazil made a historian crosses with two landraces Alfredo Chavez 6121 and Polysu,
and selected a number of rust resistance lines (Ginkel & Rajaram, 1993), the most widely used of
which was Frontana (released in 1943). While South America wheats were originally introduced
from Europe, no known past or present European cultivars appear to carry similar level of
resistance (Roelfs, 1988). One of the main source of resistace to leaf rust in CIMMYT has been,
indirectly and directly the Brazilian cultivar Frontana.
Borlaug used the cultivar FKN(Frontana/Kenya58//Newthatch) as the vehicle to infuse this
resistance complex to leaf rust into the joint Mexico/Rockfeller Foundation Wheat Breeding
Program. The resistance in Frontana is based on four additive genes, one of which is Lr34, while
the other three remain unnamed (Singh and Rajaram, 1992). Lr34 alone, however, may allow
relatively high level of disease: upto 60S. The presence of other additive genes along with Lr34
has also been confirmed afterwards (Das 1990).
Relative to a susceptible check; germplasm carrying gene Lr34 displays longer latency periods
(114-149%), fewer uredia(2-58%) a smaller uredia size (22-66%) and a slower disease
development (1-50%), in adult plant stage. Correlations among these four components studied
were quite high and all contributing to the resistance in the field. Hence they appear to be either
tightly linked or pleiotropically controlled. This would allow the breeders to choose; to a certan
extent, the trait that can most easily be introduced into ongoing breeding program (Singh et al.,
1991).
Lr34 found to be the best detectable at lower temperatures (Drijepondt & Pretorius 1989;
Drijepondt et al 1991). Others, however, have warned that it may be unreliable to screen for partial
resistance at the seedling stage (Poyntz and Hyde1987). Dyck(1987) showed that Lr34 may have
effects beyond the leaf rust resistance, in that it could be the non-supressing allele of possible stem
rust resistance suppressor genes on chromosome 7D in the cultivar Thatcher. Thus, Lr34 would
allow otherwise hidden stem rust resistance to be expressed. In another set of CIMMYT
germplasm, 2-4 additive genes were identified, determining slow rusting (Das,1990). Latency
period and uredium size were again highly correlated. Lr34 in combination with other leaf rust
resistance genes have maintained effective resistance for a longer period of time compared with
cultivars that lack Lr34 (Oelke & Kolmer, 2005). The adult-plant genes Lr13 and Lr34 often
interact with seedling resistance genes in seedling plants to produce lower than expected infection
types (Kolmer, 1196). Roelfs (1995) has indicated that Lr13 and Lr34 may be present in many
wheats worldwide that have displayed durable leaf rust resistance. In the present study, cultivars
5

selected and investigated in stress conditions had the highest percentage presence of Lr34 and it
seems that this gene was effective in increasing the tolerance of cultivars in environmental stress
conditions (Dadrezaei, 2013).
Lr34/Yr18
Another salient characteristic of Lr34 resistance is that it is genetically tightly linked with Yr18
gene, which confers adult plant resistance (Singh, 1992a, b; McIntosh, 1992). This gene cosegregates with leaf tip necrosis (Ltn1), powdery mildew resistance (Pm38), Barley yellow dwarf
virus (Bydv1) genes (McIntosh, 1992; Singh et al., 1992a, b, Spielmeyer et al., 2005; Liang et al.,
2006). These multi-pathogen resistance traits have made the Lr34/Yr18 locus one of the highly
valuable regions for disease resistance in wheat (Kolmer et al., 2008). If Lr34/Yr18 complex is
present alone the disease level may go high but in combination with other genes it could give
effective control (Ma and Singh, 1996). At low temperature the resistance level conferred by plants
with Lr34 is higher under growth chamber and green house condition.
Because Lr34 and Yr18 work mostly in adult plants and in combination with other rust resistance
genes it is difficult for breeders to determine if they are present when a wheat plant displays
resistance, or if resistance is caused by other resistance genes (Priyamvada et al., 2011). Duo of
Lr34 and Yr18 has long been recognized as carrying the most durable forms of resistance. Other
rust resistance genes may be more effective against specific strains of rust, but their effectiveness
is eventually overcome by new strains. Genes Lr34 and Yr18 confer slow rusting resistance to leaf
and stripe rust, respectively, and are known to be pleiotropic or completely linked to each other
(McIntoch, 1992; Singh, 1992a). Although genes Lr34 and Yr18 may not provide adequate
resistance under high disease pressure when present alone (Ma and Singh, 1996; Singh and Gupta,
1992; Singh and Huerta-Espino, 1997), they could contribute to achieving acceptable levels of
resistance in combination with other slow rusting genes (Singh et al., 2001; Singh and Rajaram,
1992).
A marker associated with csLV34 locus on chromosome 7D was found associated with Lr34/Yr18
gene. Two predominant allelic size variants csLV34a and csLV34b were identified. A strong
association was observed with the presence Lr34/Yr18 gene and csLV34b allele. However, lines
having Lr34/ Yr18 gene and positive for csLv34a allele were rare. The lineage of this gene is
tracked back to varieties Mentana and Ardito developed in Italy during early 1990s (Kolmer et
al., 2008). This gene has been cloned and was shown that Lr34/ Yr18/Pm38/Ltn1 is the same gene
(Kratinger et al., 2009).
Lr 46
Singh et al. (1998) studied adult plant resistance in F2 derived F3 and F5 lines developed through
crossing Pavon 76 with two wheat leaf rust susceptible cultivars, Jupateco 73S and Avocet S. The
offspring lines segregated for resistance and susceptibility in a ratio expected for segregation at
two independent loci. From these results, Singh et al. (1998) concluded that the slow rusting
resistance of Pavon 76 is the result of two genes acting in an additive manner. A slow rusting gene
identified in the cultivar Pavon 76 and was found located on chromosome 1B by crossing with a
monosomic series of adult plant leaf rust susceptible cultivar Lal Bahadur (Singh et al., 1998).
This is the 2nd named minor gene involved in slow rusting. The effect of Lr46 resembled that of
6

the Lr34 in adult plants. Both prolong the latency period and cause a higher percentage of abortion,
a reduced colony size and a lower disease severity, but in seedling stage its effect is weaker than
that of Lr34 (Martinez et al., 2001). The presence of the single slow-rusting gene Lr46 does not
give enough protection under high leaf rust pressure.
Mateos-Hernandez et al. (2006) developed fourteen new markers that potentially link to Lr46.
They narrowed down the physical location of Lr46 to a submicroscopic region between the
breakpoints of deletion lines 1BL-13 and 1BL-10. A substitution line of wheat cultivar Lalbahadur,
carrying Lr46 from Pavon was used. The leaf rust score differed over the years and environment.
Substitution lines carrying Lr46 displayed leaf rust severity of 20-30%, while the susceptible check
Lalbahadur showed a susceptibility of 80-100%.
Lr46/Yr29
Rosewarne et al. (2008) identified chromosomal regions associated with leaf rust and stripe rust
resistance in a cross between Attila and Avocet-S. They found a continuous distribution of
variation for stripe rust and leaf rust resistance among the lines derived from the population. Attila,
a resistant parent, scored very low for both leaf and stripe rust, while Avocet-S showed high
susceptibility to both rusts. Genetic analysis of the population indicated the involvement of two
additive genes in resistance, and the Lr46/Yr29 locus was the main contributor to resistance. They
also identified an epistatic interaction for stripe rust resistance between Lr46/Yr29 locus and
another unmapped region.
Rosewarne et al. (2006) identified the presence of the Lr46/Yr29 locus in a population developed
through the Avocet-Yr-A x Attila cross using AFLP markers. They observed that the population
segregated for leaf tip necrosis (LTN), a trait previously associated with Lr34. Single chromosome
recombinant lines were used to confirm the association of LTN with Lr46. The results of their
study concluded that LTN is also pleiotropic or closely linked to Lr46 and suggested that a new
LTN gene designation should be given to this locus. They suggested that LTN is a good phenotypic
marker when Lr46 and Lr34 are individually used in combination with other leaf rust resistance
genes. In crosses containing both of these genes, the use of molecular markers will help to identify
lines carrying both genes.
Lr67
Lr67 is the new and promising APR gene, located on chromosome arm 4DL, was identified in
Canadian wheat line RL6077 (Basnet et al, 2012). Lr67 is nearly similar in response with Lr34,
but it is not so effective as Lr34 for leaf rust resistance. Lr67 along cannot confer the stem rust
resistance. An important difference of Lr67 to Lr34 is that it lacks seedling resistance to leaf rust
in RL6077 when grown under low temperature condition. In field condition Lr67 confers the
resistance to leaf, stem and stripe rust of wheat. The slow-rusting gene(s) Lr67 and Yr46 can be
utilized in combination with other slow-rusting genes to develop high levels of durable APR to
leaf rust and stripe rust in wheat (Herrera et al., 2011).

Table 1 Leaf rust resistance genes in old wheat varieties (Khan et al., 2013)
S No
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15

Varieties
Lerma Rojo
Champingo 53
Penjamo 62
Pitic 62
Sonora 64
Mexipak 65, Kalyansona
Sonalika, Bluesilver
Lyalpur 73
Bluebird, Yecora 70
Ciano 79
Arz
Pavon F 76, Dollarbird
Parula
Punjab 81
Era

Year
1964
1953
1962
1962
1964
1965
1967, 1971
1973
1970
1979
1973
1983, 1987
1981
1981
1970

Country/Region
Mexico
Mexico
Mexico
Mexico
Mexico
India, Pakistan
India, Pakistan
Pakistan
Mexico, Pakistan
Mexico
Lebnon
Mexico, Australia
CIMMYT
Pakistan
North America

Genes
Lr13, Lr17
Lr34
Lr14a, Lr34
Lr14a
Lr1
Lr14a
Lr13, Lr14a
Lr1, Lr13, Lr34
Lr1, Lr13, Lr34
Lr16
Lr17
Lr1, Lr10, Lr13, Lr46+
Lr34 & Lr46+
Lr10, Lr13, Lr34
Lr10, Lr13, Lr34

Conclusion
The ever changing nature of wheat leaf rusts poses a serious threat to future wheat production.
Breeding for disease resistance is one of the most economical and environmental friendly methods
to control leaf rust (Puccinia recondita f.sp. tritici) in wheat (Triticum aestivum L.). Learning from
wheat breeding history and epidemic losses by wheat rusts, breeders devised the strategy of
pyramiding APR/minor genes. Genes that are effective at the seedling stage (major genes) have
been vulnerable to new races of leaf rust. Race specific resistance under qualitative genetic control
can be rendered ineffective by a sudden shift in the pathogen race, whereas race non-specific
resistance due to its complex genetic nature, makes it very difficult for the pathogen to evolve into
new races that can overcome the mechanism of partial resistance. The resistances that have been
shown to be durable are almost inevitably conditioned by adult-plant resistance genes, often Lr34,
Lr46, Lr67, etc. These genes along are not effective for the durable resistant if plant is subjected
to the high disease pressure. This is why pyramiding the APR/minor genes are crucial for durable
leaf rust resistant. Correlations among the resistant genes and the other performance attributing
genes should be quite high. Lr34/Yr29 is the most significant combinations of APR gene for the
resistant of leaf rust as well as stripe rust in wheat.

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