In growth room regimes arranged to simulate field conditions which coincide with natural infection of oilseed rape by Leptosphaeria maculans, leaf inoculation resulted in systemic infection. After colonizing intercellular spaces in the spongy mesophyll of the lamina, the fungus reached a vascular strand and spread down the petiole mainly in xylem vessels or between cells of the xylem parenchyma and cortex, eventually invading and killing cells of the stem cortex and causing the stem canker symptom. The intercellular systemic phase of growth, which was biotrophic and virtually sytnptomless, occurred under a wide range of temperatures.

The response of seedlings of the monocot Hordeum vulgare L. cv. Bronze to 0, 25 and 50 #M aluminium in factorial combination with 0, 1.4, 2.0 and 2.8 mM Si was tested in hydroponic culture at pH 4.5. Nutrient solution (500 /_tM calcium nitrate) and A1/Si treatments were designed to avoid the precipitation of AI from solution. Silicon treatments gave significant amelioration of the toxic effects of A1 on root and shoot growth and restored calcium levels in roots and shoots at harvest to levels approaching those of control plants. Aluminium uptake by roots was also significantly diminished in the presence of Si. Silicon alone gave a slight stimulation of growth, insufficient to explain its ameliorative effect on A1 toxicity. The mechanism of the Si effect on A1 toxicity in monocotyledons awaits further investigation.Abbreviations: ICP -inductively coupled plasma.

Natural epidemics of Leptosphaeria maculans in cultivated populations of Brassica napus var. oleifera were predominantly monocyclic, involving a sequence of five phases; latent infection of the lamina, leaf lesion expression, symptomless systemic growth down the petiole, latent infection of the stem, and stem canker development. This sequence potentially terminated in severe stem cankers if initiated from ascospore infection of any leaf between a plastochron index of PI =n+ 0.5 and n+1.0 until plant growth stage 3.1 of the Harper & Berkenkamp scale. At PI < n+ 0.5, infection was rare and at PI > n+ 1.0, the infection sequence was usually terminated by leaf abscission. Differential effects of ambient temperature on the rates of progress of infection and leaf development determined whether abscission occurred before infection reached the stem. Based on these interrelationships, a system for predicting the incidence of severe stem cankers is proposed.

In a series of growth room experiments in which leaves of Brassica napus var. oleifera were inoculated with ascospores or pycnidiospores of Leptosphaeria maculans successful infections progressed through three consecutive phases. Initial establishment in the mesophyll was succeeded by a phase of intercellular exploration, when hyphal proliferation was highly variable and host cell necrosis always ensued, and then by a systemic phase when hyphae were consistently sparse. Host cells associated with the hyphal front were capable of autofluorescence, accumulation of vital stains and plasmolysis, indicating that they were viable and that the pathogen was biotrophic throughout this sequence. During either of the first two phases permanent fungistatic containment, involving the formation of vesicles by disintegration of the hyphae, often occurred. Localization at the first phase was symptomless; at the second it was signified by a lesion with a clearly defined margin. There was a negative correlation between biotrophic potential and necrotrophic potential of three pathogenic isolates, on both the moderately susceptible cultivar Primor and the resistant cultivar Jet Neuf. As leaves aged, a progressively larger proportion of infections failed to become systemic. With increasing inoculum load, symptomless localization of infection diminished, the phase of necrosis extended, and the probability of irreversible systemic development increased.

Development of stem cankers in seedlings of Brassica napus var. oleifera inoculated with Leptosphaeria maculans was studied in growth cabinets. As the pathogen progressed from the petiole into the stem, hyphae invaded parenchyma cells for the first time, initiating a necrotrophic phase which resulted in the formation of a lesion. Ultimate containment of the pathogen was associated with a sequence of host reactions including lignification, cambium formation and callose deposition at the perimeter of the lesion. Two isolates of L. maculans differed in the pattern of stem infection. One was confined to the cortex and the other passed through the leaf gap into the pith. Variation in response between cultivars Rapora, Primor and Jet Neuf modified these two colonization patterns but did not eliminate the main differences. There were qualitative and quantitative differences in host cell reactions. The significance of these findings is discussed in relation to differences in resistance of the cultivars in the field.

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