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Principles of Plant Genetics and Breeding

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5 The plant or cell overproduces the target compound<br />

for the phytotoxin in large amounts such that it<br />

would take a high concentration <strong>of</strong> the herbicide to<br />

overcome it.<br />

Epidemics <strong>and</strong> plant breeding<br />

Modern agriculture is drastically different to that done<br />

by early plant domesticators. <strong>Plant</strong> breeding, in adapting<br />

wild species for cultivation, <strong>of</strong>ten removes natural<br />

means <strong>of</strong> crop protection needed for survival in the<br />

wild, but undesirable in modern production. Starting<br />

with the early domesticators, mass production <strong>of</strong> a few<br />

desirable genotypes has become the norm <strong>of</strong> crop production.<br />

The monocultures <strong>of</strong> modern agriculture are<br />

extreme versions <strong>of</strong> mass production, in which genetic<br />

diversity is further restricted. It is not difficult to see<br />

how such breeding <strong>and</strong> production practices predispose<br />

crops to widespread disease <strong>and</strong> insect pest damage.<br />

With the arrival <strong>of</strong> a devastating pest attack, breeders<br />

counter with the production <strong>of</strong> resistant cultivars, thus<br />

setting the stage for sequential cycles <strong>of</strong> pest resistance<br />

<strong>and</strong> pest susceptibility <strong>of</strong> crop plants.<br />

The patterns <strong>of</strong> disease incidence vary from one class<br />

<strong>of</strong> pathogen to another. Soil-borne pathogens tend to<br />

be localized <strong>and</strong> persistent in the soil, season after season.<br />

Airborne pathogens have a different biology <strong>and</strong><br />

spread pattern, as well as the potential to become epidemic.<br />

An epidemic may be defined as an outbreak <strong>of</strong><br />

disease characterized by an infection that starts from a<br />

low level <strong>and</strong> then progresses to a high one. Airborne<br />

pathogens tend to have annual epidemic cycles. These<br />

cycles depend on seasonal weather patterns <strong>and</strong> the<br />

presence or absence <strong>of</strong> a susceptible host (crop). Soilborne<br />

pathogens usually stabilize in a new location<br />

after several years, <strong>and</strong> persist in the chronic phase.<br />

Alam, S.N., <strong>and</strong> M.B. Cohen. 1998. Detection <strong>and</strong> analysis <strong>of</strong><br />

QTLs for resistance to the brown planthopper, Nilaparvata<br />

lugens, in a doubled haploid rice population. Theor. Appl.<br />

Genet. 97:1370–1379.<br />

Bogdanove, A.J. 2002. Protein–protein interactions in pathogen<br />

recognition in plants. <strong>Plant</strong> Mol. Biol. 50:981–989.<br />

Flor, H.H. 1956. The complementary genic system in flax <strong>and</strong><br />

flax rust. Adv. Genet. 8:29–54.<br />

Flor, H.H. 1971. Current status <strong>of</strong> the gene-for-gene concept.<br />

Ann. Rev. Phytopathol. 9:275–296.<br />

BREEDING FOR RESISTANCE TO DISEASES AND INSECT PESTS 383<br />

Each type <strong>of</strong> disease pathogen <strong>and</strong> the epidemic it causes<br />

is characterized by its own ecology <strong>and</strong> spatial <strong>and</strong><br />

temporal dynamics. <strong>Plant</strong> fungal diseases are especially<br />

significantly impacted by weather conditions <strong>and</strong> climatic<br />

patterns. Epidemics in agricultural crop production<br />

impact US agriculture by affecting the economic value,<br />

quantity, <strong>and</strong> quality <strong>of</strong> food <strong>and</strong> fiber produced.<br />

J. E. Van der Plank provided the underlying scientific<br />

principles <strong>of</strong> disease epidemics. An epidemic occurs in<br />

phases. It starts with a small inoculum <strong>of</strong> the pathogen<br />

(that may be from overwintering spores, or freshly introduced<br />

into the area from outside). The initial rate <strong>of</strong><br />

infection is proportional to the infection level. As time<br />

proceeds, the infection rate slows down as uninfected<br />

tissue decreases. Van der Plank expressed the initial<br />

infection rate by this mathematical relationship:<br />

dI/dt = rI or I = I 0 exp rt<br />

References <strong>and</strong> suggested reading<br />

where I = infection at time t, I 0 = initial infection, <strong>and</strong><br />

r = multiplication rate <strong>of</strong> the pathogen. A general <strong>and</strong><br />

more realistic expression is:<br />

dI = rI(1 − I)<br />

I = 1/[1 + (exp −rt ) or log e (1/1 − I) = rt + log e (1/C)<br />

where C is a constant that depends upon I 0 but is not<br />

equal to it.<br />

In plant breeding, the breeder’s goal in countering a<br />

disease epidemic is to reduce r (the multiplication rate <strong>of</strong><br />

the pathogen) to a level such that, at the time <strong>of</strong> crop<br />

maturity, the final attack by the pathogen can inflict only<br />

a small <strong>and</strong> acceptable yield loss. It may be best if r = 0<br />

(i.e., immunity), but this is not necessary. In practice,<br />

breeders do not aim for immunity but for reducing<br />

r below that <strong>of</strong> the current commercial cultivar that is<br />

susceptible to the pathogen.<br />

Hill, C.B., Y. Li, <strong>and</strong> G.L. Hartman. 2004. Resistance to the<br />

soybean aphid in soybean germplasm. Crop Sci. 44:98–106.<br />

Hooker, A.L. 1980. <strong>Breeding</strong> to control pests. In: Crop<br />

breeding (Wood, D.R., ed.). American Society <strong>of</strong><br />

Agronomy/Crop Science Society <strong>of</strong> America, Madison,<br />

WI.<br />

Khush, G.S., <strong>and</strong> D.S. Brar. 1991. <strong>Genetics</strong> <strong>of</strong> resistance to<br />

insects in crop plants. Adv. Agron. 45:223–274.<br />

Niks, R.E., P.R. Ellis, <strong>and</strong> J.E. Parlevliet. 1993. Resistance<br />

to parasites. In: <strong>Plant</strong> breeding: <strong>Principles</strong> <strong>and</strong> prospects

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