Principles of Plant Genetics and Breeding
Principles of Plant Genetics and Breeding
Principles of Plant Genetics and Breeding
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Types <strong>of</strong> genetic resistance<br />
The complexity <strong>of</strong> host–pathogen interaction makes it<br />
difficult to categorize resistance into finite types. A large<br />
number <strong>of</strong> host–pathogen interaction systems occur at<br />
various stages <strong>of</strong> coevolution. Nonetheless, some rough<br />
categorization, albeit simplistic, is widely acceptable for<br />
its convenience <strong>and</strong> practical plant breeding applications.<br />
Resistance reactions may be generally categorized<br />
into two major kinds – vertical or horizontal – based<br />
on their epidemiological status <strong>and</strong> stability <strong>of</strong> resistance.<br />
Vertical (specific) or horizontal (non-specific)<br />
effects are expressed in terms <strong>of</strong> population dynamics,<br />
not in terms <strong>of</strong> genes.<br />
Vertical resistance<br />
Vertical resistance is known by other terms including<br />
hypersensitivity, race- or pathotype-specific resistance,<br />
major gene or oligogenic resistance, nondurable<br />
or qualitative resistance, <strong>and</strong> non-uniform<br />
resistance. This reaction is said to occur when a race <strong>of</strong> a<br />
pathogen produces disease symptoms on some cultivars<br />
<strong>of</strong> a host but fails to do so on others. That is, there is a<br />
complete display <strong>of</strong> resistance in some genotypes while<br />
others are susceptible. Vertical resistance is a differential<br />
resistance, or a host × isolate interaction <strong>of</strong> cultivars<br />
against different races <strong>of</strong> a pathogen.<br />
However, there are examples <strong>of</strong> resistance genes that<br />
the pathogen apparently fails to match (e.g., dominant<br />
resistance in Periconia cincinata, the causal agent <strong>of</strong> the<br />
milo disease <strong>of</strong> sorghum). Major gene resistance is <strong>of</strong>ten<br />
absolute, conferring immunity against the appropriate<br />
pathotype (but not necessarily so). It delays the start <strong>of</strong><br />
an epidemic, <strong>and</strong> disappears when the pathotype virulent<br />
to the vertical resistance gene is prevalent. This type<br />
<strong>of</strong> resistance is relatively easy to breed because the major<br />
genes are easy to identify <strong>and</strong> transfer through simple<br />
crosses. These genes control specific races or genotypes<br />
<strong>of</strong> pests <strong>and</strong> hence do not protect against new races <strong>of</strong><br />
the pests. Cultivars equipped with this narrow-based<br />
genetic protection are <strong>of</strong>ten vulnerable when a genetically<br />
variable population <strong>of</strong> a pathogen occurs in the<br />
environment. They may not succumb immediately to<br />
the new strains <strong>of</strong> the pathogen but under certain cultural<br />
systems (e.g., monoculture without crop rotation),<br />
the populations <strong>of</strong> the new strains could build up high<br />
enough to cause economic injury to the crop. This form<br />
<strong>of</strong> protection is also described as non-durable resistance.<br />
Disease or insect resistance may be conditioned<br />
BREEDING FOR RESISTANCE TO DISEASES AND INSECT PESTS 371<br />
by dominant or recessive genes. Dominant genes occur<br />
in resistance to, for example, rusts, viruses, mildews, <strong>and</strong><br />
nematodes.<br />
Horizontal resistance<br />
Horizontal resistance is also known as slow rusting,<br />
partial resistance, field resistance, race-non-specific<br />
resistance, minor gene reactions, or polygenic inheritance.<br />
There is a lack <strong>of</strong> differential reaction <strong>of</strong> isolates<br />
on different cultivars. The resistance is effective against<br />
all genotypes <strong>of</strong> the parasite species without cultivar ×<br />
isolate interaction (i.e., race-non-specific). Following<br />
the initial establishment <strong>of</strong> the pathogen, the plant may<br />
resist its spread <strong>and</strong> reproduction such that the disease<br />
develops at a slower rate. Unlike vertical resistance,<br />
evolution <strong>of</strong> new races due to selection pressure against<br />
any specific race is absent under horizontal resistance,<br />
making this type <strong>of</strong> resistance more stable.<br />
Horizontal resistance is controlled by polygenes.<br />
Each <strong>of</strong> the genes that condition the disease contributes<br />
toward the level <strong>of</strong> resistance, <strong>and</strong> hence resistance is<br />
also called minor gene resistance. Polygenic resistance<br />
is widespread <strong>and</strong> provides protection against a wide<br />
range <strong>of</strong> races <strong>of</strong> a pest <strong>and</strong> hence is less susceptible to<br />
being overcome by a new race. Polygenic resistance protects<br />
plants by slowing down the spread <strong>of</strong> disease <strong>and</strong><br />
the development <strong>of</strong> epidemics in the field. <strong>Breeding</strong><br />
polygenic resistance is more challenging. The many<br />
minor genes cannot be individually identified <strong>and</strong> consequently<br />
cannot be transferred through crossing in a<br />
predictable fashion. However, the heritability for polygenic<br />
resistance is high.<br />
Durability <strong>of</strong> resistance<br />
An aspect <strong>of</strong> resistance <strong>of</strong> concern to plant breeders<br />
is the durability. Resistance that is durable is one<br />
that remains effective in a widely grown cultivar for a<br />
long time in an environment that favors the parasite.<br />
Horizontal resistance is said to be durable whereas<br />
vertical resistance is not. Adaptations <strong>of</strong> the parasite<br />
to newly introduced resistant cultivars (i.e., breakdown<br />
<strong>of</strong> resistance) occurs frequently in pathogens but less so<br />
in insect pests. Such cultivars lack durable resistance.<br />
Durability <strong>of</strong> resistance is variable even within pathosystems.<br />
For example in one study involving resistance to<br />
stripe rust in wheat, durability ranged from 1 to 18<br />
years, depending on the cultivars.