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Histopathology of Seed-Borne Infections - Applied Research Center ...

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82 <strong>Histopathology</strong> <strong>of</strong> <strong>Seed</strong>-<strong>Borne</strong> <strong>Infections</strong><strong>Seed</strong>s after harvest and threshing or after natural dissemination are devoid <strong>of</strong>the above surroundings and are directly exposed to the environment that they cometo occupy such as storage areas and soil. <strong>Seed</strong>s have a well-formed cuticule, surfacewax deposits (when present), seed coat, or seed coat and pericarp (one-seeded dryindehiscent fruits) with protective layers, hilum, pits or micropores, and cracks. Themicropyle is narrow or open to various extents. Exposed hilum, micropyle, and raphe(when present) are features <strong>of</strong> true seeds. The latter two do not occur on the surface<strong>of</strong> one-seeded fruits, and the scar left upon separation <strong>of</strong> the fruit is at best analogousto the hilum. Since seeds are stored under dry conditions, no free moisture isavailable. The chief determinants for the development <strong>of</strong> fungi in and on seeds duringstorage are temperature and available moisture (water) <strong>of</strong> the grain. Storage fungican grow with restricted water availability (Christensen and Kaufman, 1969; Jainet al., 1994). The soil environments <strong>of</strong> seeds are highly variable, ranging from dryto water-logged soils.4.2 NATURE OF THE PATHOGENThe seed-borne fungi may be parasitic or saprophytic and, according to Dickinsonand Lucas (1977), may be biotrophs or necrotrophs. Biotrophs cause minimal damageto the host, including seed tissues, and are in fair harmony with the host.Biotrophs have a narrow host range and are usually obligate parasites. Necrotrophscause apparent damage to the host cells and have a wide host range. They secreteenzymes and bring about the disintegration <strong>of</strong> cell components, resulting in celldeath. The released cell contents are used by such pathogens for their growth.Basically, the mode <strong>of</strong> nutrition is like that <strong>of</strong> saprophytes. The necrotrophic fungi,depending upon time <strong>of</strong> infection and humidity, cause superficial or deep infection,whereas the biotrophs generally establish in deeper tissues including the embryo.The majority <strong>of</strong> seed-borne fungi are known to be necrotrophs. The obligate parasitesthat belong to Peronosporaceae, Albuginaceae, Erysiphales, Ustilaginales, and Uredinalesare biotrophs. Many intermediate conditions occur between the true necrotrophsand biotrophs. Maude (1996) believes that necrotrophs, which degrade tissuesas they spread, are rarely transmitted to the embryo through the mother plant.Another quality <strong>of</strong> the pathogen that may determine its passage during infection<strong>of</strong> the ovule and seed in the field is the nature <strong>of</strong> disease in the plant and themechanism <strong>of</strong> transmission for becoming seed-borne. Neergaard (1979) has listedeight disease cycles for seed-borne pathogens taking into consideration the location<strong>of</strong> primary inoculum in seed, course <strong>of</strong> disease development, and reinfection <strong>of</strong> ovuleand seed. The infection may be systemic, local, or organospecific. The systemicinfection may follow a vascular or a nonvascular course predominantly.4.3 INFECTION IN DEVELOPING SEEDSSince the ovules and developing seeds are present inside the ovary, the passages fortheir invasion need to be recognized at two levels: (1) routes leading to internalovary infection and (2) ovary to ovule and seed infection. Infection passages <strong>of</strong> a

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