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

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120 <strong>Histopathology</strong> <strong>of</strong> <strong>Seed</strong>-<strong>Borne</strong> <strong>Infections</strong>swflflfflssAscembBCFIGURE 5.10 Schematic diagrams to show different pathways <strong>of</strong> development <strong>of</strong> seed-borneinfections <strong>of</strong> smuts and bunts in cereals. A, Ustilago tritici (loose smut) <strong>of</strong> wheat — intraembryalseed inoculum followed by systemic infection. B, Ustilago hordei (covered smut)<strong>of</strong> barley — externally seed-borne inoculum causing systemic infection. C, Tilletia indica(karnal bunt) — externally seed as well as soil-borne inoculum, spores cause blossom infection<strong>of</strong> individual florets. (Abbreviations: mb, embryo; ffl, first foliar leaf; fl, foliar leaves; s, seed;sc, seed coat; ss, soil surface; wfl, weakened foliar leaves.) (Adapted and redrawn fromNeergaard, P. 1979. <strong>Seed</strong> Pathology. Vols. 1 and 2. Macmillan, London.)to form promycelium. They cause secondary infection to developing caryopsis, resultingin embryal infection, which remains dormant in seeds, e.g., Ustilago tritici andUstilago nuda (loose smuts <strong>of</strong> wheat and barley), until germination (Figure 5.10A).The primary inoculum is externally seed-borne or, as in some Tilletia spp., externallyseed- as well as soil-borne, and causes seedling (coleoptile) infection at the time <strong>of</strong>seed germination. Infection is systemic and latent in the vegetative phase andexpresses in ears. All ears <strong>of</strong> tillers <strong>of</strong> a plant and every seed in the infected eardevelop teliospores held by the persistent membrane and pericarp (Figure 5.10B),e.g., Ustilago hordei (covered smut <strong>of</strong> barley). Sometimes, as in the common bunt,individual seeds or ears may remain healthy (Tilletia tritici and Tilletia laevis). Theprimary inoculum is externally seed-borne and soil-borne. Sporidia formed in soilbecomeairborne, cause infection <strong>of</strong> individual florets,and invade ovary and ovuleforming teliospores. Grain infection is either total or partial, covered by pericarp(Figure 5.10C), e.g., Tolyposporium penicillariae (smut <strong>of</strong> pearl millet), Tilletiaindica (Neovossia indica, Karnal bunt).5.5.3.1.1 Type 1 Disease CycleThe embryal infection in wheat grains by Ustilago tritici, causal organism <strong>of</strong> loosesmut <strong>of</strong> wheat, was first reported by Maddox (1896) in Australia and later confirmedby Brefeld (1903) in Germany. Subsequently, many studies have described the course

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