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

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174 <strong>Histopathology</strong> <strong>of</strong> <strong>Seed</strong>-<strong>Borne</strong> <strong>Infections</strong>for Rhodococcus facians (Corynebacterium fascians) in strawberry and cauliflowerare reported to be essential as vectors <strong>of</strong> these pathogens (Hunger, 1901; Carne,1926; Vasudeva and Hingorani, 1952; Crosse and Pitcher, 1952; Pitcher and Crosse,1958).6.1.2 SPREAD IN PLANT AND COURSE OF ENTRY INTO OVARYAND FRUIT AND OVULE AND SEEDAfter entering the plant tissue, the bacteria multiply in a substomatal cavity(Figure 6.1A, C, E) or space beneath the site <strong>of</strong> entry. Within the host, the bacteriafollow two courses for further spread: they invade xylem elements and move verticallyafter coming in contact with vascular elements and they multiply and spreadin intercellular spaces (Figure 6.1B, D). The latter course usually results in arestricted spread. Vascular bacteria can move over long distances, and they can alsospread laterally after dissolving the walls <strong>of</strong> the vessels (Smith, 1911; Yu, 1933;Pine, Grogan, and Hewitt, 1955) or after escaping through openings (pits) in thewalls <strong>of</strong> vessel elements (Ivan<strong>of</strong>f, 1933; Wolf and Nelson, 1969). Nelson and Dickey(1966) have also suggested lateral movement <strong>of</strong> bacteria through plasmodesmatalconnections in the pit membranes <strong>of</strong> vessels. Association <strong>of</strong> bacteria with phloemis observed, but it probably plays no significant role in their spread (Pine, Grogan,and Hewitt, 1955).Several seed infecting bacteria are systemic vascular pathogens (Zaumeyer,1930; Pine, Grogan, and Hewitt, 1955; Cormack and M<strong>of</strong>fat, 1956; Mukerjee andSingh, 1983; Du Plessis, 1990). Pine, Grogan, and Hewitt (1955) have given adetailed account <strong>of</strong> the spread <strong>of</strong> C. m. subsp. michiganensis, a bacterial cankerpathogen, in young tomato plants and provided evidence <strong>of</strong> longitudinal and lateralmovement <strong>of</strong> the organism in xylem elements (Figure 6.4A to D). Bacterial pocketswere formed in phloem and pith, but they gave no indication <strong>of</strong> vertical movement.During earlier studies, C. m. subsp. michiganensis was described primarily as aphloem parasite (Smith, 1914, 1920). However, Zaumeyer (1930) has also observedinvasion <strong>of</strong> xylem elements in leaf (Figure 6.1F) and stem (Figure 6.1G) by X. a.pv. phaseoli, and its subsequent spread throughout the organism. X. oryzae pv. oryzaehas a similar course in rice (Mukerjee and Singh, 1983).The systemic vascular infection <strong>of</strong>ten enters the reproductive shoot and causesovary and fruit infection (Zaumeyer, 1930; Cormack and M<strong>of</strong>fatt, 1956; Mukerjeeand Singh, 1983; Du Plessis, 1990). Zaumeyer (1930) traced the passage <strong>of</strong> X. a.pv. phaseoli from plant to pods in beans and noted that the bacteria in the xylemvessels <strong>of</strong> the stem travels up and through the vascular elements <strong>of</strong> the pedicel andpasses into the vascular supply <strong>of</strong> the pod. Mukerjee and Singh (1983) observedthat X. o. pv. oryzae reaches the rice husk through vessels <strong>of</strong> the infected rachilla.Skoric (1927) reported that Pseudomonas syringae pv. pisi is largely a parenchymainvader and only occasionally may enter the vessels under field conditions.The bacterium penetrates pods through wounds and spreads in intercellular spaces,forming abundant slime on the inner side <strong>of</strong> the pod. After artificial inoculation byP. s. pv. lachrymans <strong>of</strong> the stem 1.5 m from the fruit, Kritzman and Zutra (1983)observed systemic invasion <strong>of</strong> the cucumber fruits. No histopathological studies were

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