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Horticulture Principles and Practices

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2. Instead of excessive growth, the plant may be stunted (as occurs with most viral<br />

infections) or plant organs may be only partially developed. This response is<br />

described as underdevelopment of tissue.<br />

3. To contain the infection, the cells around the point of attack may die so that the<br />

organism cannot spread any further. This response of tissue death is called necrosis<br />

(hypersensitive reaction). Diseases such as leaf spots, blights, cankers (e.g., anthracnose),<br />

<strong>and</strong> decays (e.g., soft rot) are symptoms of diseases involving tissue death.<br />

Dissemination<br />

Agents of dissemination are air, water, insects, other animals, <strong>and</strong> humans. Fungal spores<br />

are usually spread by air currents. Bacteria <strong>and</strong> nematodes are often disseminated by<br />

water. Viral infections are transmitted by vectors. Pathogens are carried on farm tools <strong>and</strong><br />

equipment <strong>and</strong> on the feet or bodies of animals.<br />

Overwintering or Oversummering<br />

Annual plants die after a season, infection or no infection. The pathogen must therefore<br />

be able to survive in the absence of its host. Similarly, pathogens of perennial plants must<br />

survive during the period of dormancy while adverse weather prevails. For example,<br />

nematodes overwinter as eggs in plant debris. Some fungi inhabit the soil.<br />

7.7.2 DEFENSE AGAINST PATHOGENS BY PLANTS<br />

Plants have structural <strong>and</strong> biochemical means of combating an invasion by pathogens.<br />

Structural <strong>and</strong> biochemical defense mechanisms involve the use of physical barriers or<br />

the secretion of chemicals to block entry of pathogens into plant hosts. Plants differ in<br />

the types of mechanisms they possess.<br />

Structural Defense<br />

Preexisting Defense Mechanisms When a pathogen l<strong>and</strong>s on the surface of the plant<br />

<strong>and</strong> attempts to penetrate the tissue, the plant’s first means of defense is its epidermis. Some<br />

plants have deposits of wax in the epidermal cells or a thick cuticle over the epidermal layer,<br />

either of which is impenetrable to pathogens. Some leaves have a waxy, water-repellent surface<br />

or a pubescent surface that makes it difficult for water to settle to provide the moisture<br />

certain pathogens such as fungi <strong>and</strong> bacteria require to grow or multiply. The presence of<br />

such a tough material severely limits direct penetration of pathogens, thus reducing infection<br />

because the remaining avenues for entry are the natural openings (stomata).<br />

Induced Defense Structures If the pathogen gains entrance into the tissue, the<br />

presence of the alien matter may induce the formation of a cork layer for containment of<br />

the invasion. In some cases, especially in stone fruits, an abscission layer may be induced<br />

to form in active leaves. This layer causes a permanent break between the site of infection<br />

<strong>and</strong> the rest of the leaf, consequently forestalling the advancement of the infection.<br />

In some plants, an injury (e.g., resulting from penetration) induces gums to be formed<br />

around the point of infection, thus limiting its spread. Another induced structure is tissue<br />

necrosis, a hypersensitive reaction to invasion. A cell that is penetrated by the hyphae<br />

of fungi, for example, soon experiences degeneration of the nucleus, followed ultimately<br />

by death. Death of tissue means the fungus has no access to nutrients, leading to the<br />

eventual death of an obligate parasite.<br />

Biochemical Defense<br />

Plants under attack by pathogens may secrete certain metabolites to ward off attack.<br />

Preexisting Mechanisms Certain plants under normal conditions exude chemicals<br />

onto the plant surface (stem, root, or leaves). Tomato, for example, is known to release<br />

236 Chapter 7 Biological Enemies of Horticultural Plants

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