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

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2 <strong>Histopathology</strong> <strong>of</strong> <strong>Seed</strong>-<strong>Borne</strong> <strong>Infections</strong>(pistil). The seed formation takes place in situ through a series <strong>of</strong> integrated sequentialsteps in the life cycle <strong>of</strong> the flowering plant (Maheshwari, 1950, 1963; Johri,1984). After pollination and fertilization, changes in different parts <strong>of</strong> the ovule, i.e.,the embryo sac (zygote and primary endosperm nucleus), nucellus, chalaza, andintegument, lead to the formation <strong>of</strong> seed. The term seed, when used sensu lato,includes one-seeded dry indehiscent fruits that are the dispersal propagative unitsin plants <strong>of</strong> several families such as Poaceae, Asteraceae, Apiaceae, and Chenopodiaceae.In the present treatment, the term seed is used in a loose sense.<strong>Seed</strong> is an autonomous living unit and links successive generations. Structurally,it consists <strong>of</strong> an embryo (new plantlet), a protective covering — seed coat, pericarp,or both — and reserve food material, which may be present in the endosperm,perisperm, or embryo. It has the capacity to withstand desiccation and retain viabilityunder unfavorable environments or until it germinates. These properties <strong>of</strong> seed makeit an important commodity for storage as well as transport to new areas or countries,for planting or for edible purposes. The structure <strong>of</strong> seed is fairly constant in aspecies, but varies in different plant taxa (Netolitzky, 1926; Corner, 1976).<strong>Seed</strong>s, if infected by microorganisms, will act as carriers. If the organisms remainviable they will result in development <strong>of</strong> disease in the new crop. Infected seeds are<strong>of</strong>ten responsible for the spread <strong>of</strong> diseases to new areas. For this reason seed hasbecome an object <strong>of</strong> plant quarantine internationally. Countries also use domesticseed certification, including seed health testing, as a method <strong>of</strong> quality control <strong>of</strong>seed.1.2 MICROORGANISMS IN SEEDFungi, bacteria, viruses, and nematodes are known to be seed-borne (Neergaard,1979; Maude, 1996; Agarwal and Sinclair, 1997). Fungi form a major group <strong>of</strong>pathogens that are seed-borne as well as seed-transmitted. In addition to saprophytesand parasites, fungi are known to form an inherent association with seeds <strong>of</strong> somemembers <strong>of</strong> Cistaceae, Ericaceae, and Orchidaceae. In the Orchidaceae, the seedswill not usually germinate without the presence <strong>of</strong> a mycorhizal fungus (Rayner,1915). The seed coat in seeds <strong>of</strong> Helianthemum chamaecistus (Cistaceae) harbors afungus that seems essential for normal germination <strong>of</strong> seed. In the absence <strong>of</strong> thefungus, the plumule fails to emerge, and roots also are not formed at the time <strong>of</strong>germination (Boursnell, 1950).The list <strong>of</strong> saprophytic and parasitic fungi associated with seeds <strong>of</strong> differentplants is very large (Richardson, 1990), and they belong to all fungal classes. Thefungi that are discussed in this book and for which histopathological information isavailable belong to the division Eumycota, subdivision Mastigomycotina, classOomycetes, subdivisions Ascomycotina, Basidiomycotina, and Deuteromycotina.The members <strong>of</strong> Deuteromycotina dominate and these fungi belong to the classesHyphomycetes and Coelomycetes. The endophytic fungi are discussed separately.A large number <strong>of</strong> viruses, including cryptoviruses and viroids, are known tobe seed-borne, but the information on seeds infected by viruses is limited and mostlyinconclusive. The better-studied viruses are barley stripe mosaic virus (BSMV) andpea seed-borne mosaic virus (PSbMV) due to the studies <strong>of</strong> Carroll and co-workers

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