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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.

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