Wilts caused by Verticillium species. A cytological survey of vascular alterations in leaves

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Abstract
Plants that are infected with fungi of the species Verticillium frequently develop foliar disease symptoms which may include one or more of the following: flaccidity, drying, chlorosis leading to necrosis, vascular browning, epinasty, and leaf abscission. A number of ultrastructural and chemical alterations occur in the vascular tissues of such leaves: deposition of brown pigments, coating of xylem vessel walls with abnormal material (i.e., lipid-rich coatings or fibrillar coatings), plugging of xylem vessels with gums, gels or tyloses, degeneration of parenchyma cells, and accumulation of abnormal electron dense materials in primary and secondary cell walls. Different host–parasite combinations exhibit different leaf symptoms and different cytological alterations. The purpose of the present survey was to determine whether the extent of any of the possible vascular alterations in leaves could be correlated with the wilting tendency of the host.Chrysanthemums, snapdragons, eggplants, sunflowers, potatoes, sycamore maples and hedge maples were infected with V. dahliae; alfalfa and hops were infected with V. albo-atrum. When leaf symptoms were well advanced, samples were taken from the major lateral leaf veins and were prepared for light (LM) and transmission electron microscopy (TEM) or scanning electron microscopy (SEM). The various types of alterations in the vascular tissues were identified by a correlated LM–TEM method and (or) SEM analysis and for each sample vein the proportion of vessels affected by each type of alteration was calculated. Four leaf samples, each from different plants, were analysed for each host. The visual symptoms, including vascular browning, were estimated subjectively. The degree of leaf flaccidity was correlated positively with the proportion of lipid-coated vessels and inversely with the degree of vascular browning. No other correlations were observed.