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Principles of Plant Genetics and Breeding

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elative to a benchmark. A genotype that is immune to a<br />

disease is completely or totally resistant to it. In practice,<br />

plant breeders arbitrarily set the susceptibility–resistance<br />

scale. A resistant cultivar has less disease than the st<strong>and</strong>ard<br />

cultivar – it could be a little or a lot less diseased.<br />

True disease resistance has a genetic basis <strong>and</strong> hence<br />

is amenable to plant breeding methodologies. It is<br />

manifested in two basic forms: inhibition <strong>of</strong> infection<br />

or inhibition <strong>of</strong> subsequent growth <strong>of</strong> the pathogen,<br />

the former being the more common form <strong>of</strong> resistance.<br />

Resistance can be a qualitative or quantitative trait.<br />

As A. L. Hooker observed, breeding for pest resistance<br />

is an integral part <strong>of</strong> any system <strong>of</strong> crop pest<br />

control. If yield <strong>and</strong> other desirable traits are maintained,<br />

resistance (host resistance) is an ideal method<br />

<strong>of</strong> breeding to control plant pests. The development<br />

<strong>and</strong> use <strong>of</strong> resistant cultivars has several advantages. It<br />

is relatively inexpensive to deploy (versus use <strong>of</strong> other<br />

pest control measures) <strong>and</strong> has no adverse environmental<br />

consequences. Also, as Hooker indicated, it is<br />

convenient for farmers to use, requiring no additional<br />

production costs or decision-making during the production<br />

season. However, when host resistance is not<br />

satisfactory, farmers may need to supplement host resistance<br />

with other pest management practices (e.g., use <strong>of</strong><br />

pesticides).<br />

Concepts <strong>of</strong> pathogen <strong>and</strong> host<br />

The pathogen<br />

The pathogen is a living organism that is capable <strong>of</strong><br />

inflicting a distinct disease or disorder in another organism<br />

(the host). The capacity <strong>of</strong> the pathogen to cause<br />

disease or disorder in a member <strong>of</strong> a host species is<br />

called its pathogenecity, while the extent <strong>of</strong> disorder<br />

or disease development it causes is its virulence. The<br />

pathogenecity <strong>and</strong> virulence <strong>of</strong> a pathogen vary among<br />

pathogen types (races or pathotypes). Races or pathotypes<br />

that fail to cause disease symptoms or successfully<br />

attack a given host are said to be avirulent. It should<br />

be pointed out that the presence <strong>of</strong> a pathogen on a<br />

susceptible host is not enough to cause disease symptoms<br />

to occur. A third factor – favorable environment –<br />

is needed, the trio (pathogen plus susceptible host<br />

plus favorable environment) referred to as the disease<br />

triangle. Pathotypes or races <strong>of</strong> pathogens may also be<br />

described in terms <strong>of</strong> aggressiveness or non-aggressiveness<br />

in relation to the rate at which they produce<br />

disease symptoms.<br />

BREEDING FOR RESISTANCE TO DISEASES AND INSECT PESTS 369<br />

The host<br />

The host (genotype, plant) is the organism in which a<br />

pathogen may produce disease symptoms. A susceptible<br />

host is one in which a pathotype or race can manifest a<br />

disease symptom. A host may employ one <strong>of</strong> several<br />

mechanisms (defense mechanisms) to resist pathogens.<br />

These mechanisms may be pre-existing or induced upon<br />

infection.<br />

1 Pre-existing defense mechanisms. These include<br />

morphological features that pose as barriers to the<br />

penetration <strong>of</strong> the pathogen into the plant (e.g., presence<br />

<strong>of</strong> lignin, cork layer, callose layers), or secondary<br />

metabolites (phenols, alkaloids, glycosides) that have<br />

antimicrobial properties.<br />

2 Infection-induced defense mechanisms. Upon<br />

infection, the host quickly produces chemical products<br />

(e.g., peroxidases, hydrolases, phytoalexins, etc.)<br />

to combat the infection.<br />

Mechanisms <strong>of</strong> defense in plants<br />

against pests<br />

<strong>Plant</strong>s exhibit a wide variety <strong>of</strong> strategies <strong>and</strong> mechanisms<br />

<strong>of</strong> defense against pathogens <strong>and</strong> insects pest that<br />

may be classified into three major groups – avoidance,<br />

resistance, <strong>and</strong> tolerance. The mechanisms are given<br />

different names as pertain to insect pests, as will be<br />

discussed.<br />

Avoidance<br />

Also described as escape, avoidance is a mechanism that<br />

reduces the probability <strong>of</strong> contact between pathogens or<br />

insect pests <strong>and</strong> the plant. That is, it operates prior to the<br />

establishment <strong>of</strong> any intimate host–pest/pathogen contact.<br />

This mechanism is rare in pathogens <strong>and</strong> hence is<br />

primarily applicable to insect pests where entomologists<br />

describe it as antixenosis (or non-preference). A plant<br />

using this mechanism has characteristics (morphological<br />

or chemical) that make it unattractive to insects for feeding,<br />

oviposition, or shelter. In the cabbage, a variant leaf<br />

color makes the plant unattractive to the cabbage aphid<br />

(Brevicoryne brassicae), while pubescence in cotton prevents<br />

oviposition in the cotton boll weevil (Heliothis<br />

zea). Also, in cotton, a repulsive odor deters the boll<br />

weevil from feeding on the plant.<br />

Avoidance or non-preference may not be effective<br />

if there are no alternative hosts. Non-preference

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