The pathogen is obligately parasitic on genera within the Vitaceae, including Vitis, Cissus, Parthenocissus and Ampelopsis (Pearson and Gadoury, 1992). The most economically important host is grapevine (Vitis), particularly the European grape, V. vinifera, which is highly susceptible to powdery mildew. Disease symptoms and signs: In the strictest sense, macroscopically visible mildew colonies are signs of the pathogen rather than symptoms resulting from its infection, but, for convenience, we describe the symptoms and signs together as the collective appearance of colonized host tissues. All green tissues of the host may be infected. Ascospore colonies are most commonly found on the lower surface of the first-formed leaves near the bark of the vine, and may be accompanied by a similarly shaped chlorotic spot on the upper surface. Young colonies appear whitish and those that have not yet sporulated show a metallic sheen. They are roughly circular, ranging in size from a few millimetres to a centimetre or more in diameter, and can occur singly or in groups that coalesce to cover much of the leaf. Senescent colonies are greyish, and may bear cleistothecia in various stages of development. Dead epidermal cells often subtend the colonized area, as natural mortality in the mildew colony, the use of fungicides, mycoparasites or resistance responses in the leaf result in the deaths of segments of the mildew colony and infected epidermal cells. Severely affected leaves usually senesce, develop necrotic blotches and fall prematurely. Infection of stems initially produces symptoms similar to those on leaves, but colonies on shoots are eventually killed as periderm forms, producing a dark, web-like scar on the cane (Gadoury et al., 2011). Inflorescences and berries are most susceptible when young, and can become completely coated with whitish mildew. The growth of the berry epidermal tissue stops when severely infected, which may result in splitting as young fruit expand. Berries in a transitional stage between susceptible and resistant (generally between 3 and 4 weeks after anthesis) develop diffuse, nonsporulating mildew colonies only visible under magnification. Diffuse colonies die as berries continue to mature, leaving behind a network of necrotic epidermal cells (Gadoury et al., 2007). Survival over winter as mycelium in buds results in a distinctive foliar symptom. Shoots arising from these buds may be heavily coated with fungal growth, stark white in colour and stand out like white flags in the vine, resulting in the term 'flag shoots'. More commonly, colonization of a flag shoot is less extensive, and infection of a single leaf, or of leaves on one side of the shoot only, is observed (Gadoury et al., 2011).
Berries of Vitis vinifera are reported to be susceptible to infection by Uncinula necator until soluble solids levels (brix) reach 8%, and established colonies are reported to sporulate until brix reach 15%. However, our analysis of disease progress on fruit of selected V. vinifera cultivars indicated that severity became asymptotic several weeks earlier in fruit development. When mildew-free fruit clusters of V. vinifera 'Chardonnay', 'Riesling', 'Gewürztraminer', and 'Pinot Noir' were inoculated at stages ranging from prebloom to 6 weeks postbloom, only fruit inoculated within 2 weeks of bloom developed severe powdery mildew. Substantial ontogenic resistance to infection was expressed in fruit nearly 6 weeks before fruit brix reached 8% and over 2 months before they reached 15%. Rachises of 'Chardonnay' and 'Riesling' fruit clusters developed severe powdery mildew when inoculated at bloom, and disease increased steadily over the next 60 days. The rachis of fruit clusters inoculated 31 days after bloom developed only trace levels of powdery mildew. Berry weight of all four cultivars at harvest was reduced when fruit clusters were inoculated at bloom or 16 days postbloom, primarily by splitting, rotting, and dehydration of mildewed berries, but the weight of later-inoculated berries was not reduced. Inoculation of berries just as ontogenic resistance increased markedly, approximately 3 to 4 weeks postbloom, resulted in the development of inconspicuous, diffuse, non-sporulating mildew colonies on berries, sometimes associated with a network of necrotic epidermal cells. Rather than a protracted and relatively static period of berry susceptibility lasting 3 months, fruit of V. vinifera appear to acquire ontogenic resistance rapidly after fruit set. A refocusing of disease management on this critical period of high fruit susceptibility should greatly improve the efficacy of fungicides directed against powdery mildew.
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