The relationships between the net photosynthetic rate of bean leaves and severity of rust, angular leaf spot and anthracnose were quantified at different temperatures of plant incubation, stages of disease development, and phenological stages of the crop, in two bean cultivars. Several experiments were performed in controlled environment chambers and in the field. The virtual lesion concept was used to quantify the reduction in photosynthetic efficiency of the green leaf tissue surrounding the lesions. The b parameter that expressed the ratio between areas of the virtual and visual lesions was estimated according to the model P x /P 0 (1 2 x) b , where P x was the net photosynthetic rate of a leaf with severity x and P 0 was the average net photosynthetic rate of healthy leaves. The b-value was 2´17^0´02 (R 2 0´88) for rust, 3´81^0´04 (R 2 0´91) for angular leaf spot, and 7´97^0´13 (R 2 0´94) for anthracnose. For each disease, the parameter b was consistent regardless of incubation temperature, stage of disease development, bean phenological stage and bean cultivar. In addition, the relationships between bean yield and the variables of area under the disease progress curve (AUDPC), healthy leaf area index duration (HAD) and healthy leaf area index absorption (HAA), published in the literature for rust, angular leaf spot and anthracnose, were recalculated with the virtual disease severity. Bean yield was more closely related to the new variables photosynthesizing leaf area index duration' (PAD) and`photosynthesizing leaf area index absorption' (PAA) than to those variables previously published, but only for diseases with a large b-value and at high levels of disease severity.
Measurements related to gas exchange and chlorophyll fluorescence emission were taken from healthy and diseased bean leaves with rust, angular leaf spot, and anthracnose during lesion development for each disease. The experiments were performed at different temperatures of plant incubation, and using two bean cultivars. The main effect of temperature of plant incubation was in disease development. There was no significant difference between cultivars in relation to disease development and in magnitude of physiological alterations when disease severity was the same for each cultivar. These diseases reduced the net photosynthetic rate and increased the dark respiration of infected leaves after the appearance of visible symptoms and the differences between healthy and diseased leaves increased with disease development. The transpiration rate and stomatal conductance were stable during the monocycle of rust, however, these two variables decreased in leaves with angular leaf spot and anthracnose beginning with symptom appearance and continuing until lesion development was complete. Carboxylation resistance was probably the main factor related to reduction of photosynthetic rate of the apparently healthy area of leaves with rust and angular leaf spot. Reduction of the intercellular concentration of CO2, due to higher stomatal resistance, was probably the main factor for leaves with anthracnose. Chlorophyll fluorescence assessments suggested that there was no change in electron transport capacity and generation of ATP and NADPH in apparently healthy areas of diseased leaves, but decreases in chlorophyll fluorescence emission occurred on visibly lesioned areas for all diseases. Minimal fluorescence was remarkably reduced in leaves with angular leaf spot. Maximal fluorescence and optimal quantum yield of photosystem II of leaves were reduced for all three diseases. Bean rust, caused by a biotrophic pathogen, induced less damage to the regulation mechanisms of the physiological processes of the remaining green area of diseased leaves than did bean angular leaf spot or anthracnose, caused by hemibiotrophic pathogens. The magnitude of photosynthesis reduction can be related to the host–pathogen trophic relationships.
The effects of rust (caused by Uromyces appendiculatus) and anthracnose (caused by Colletotrichum lindemuthianum) and their interaction on the photosynthetic rates of healthy and diseased bean (Phaseolus vulgaris) leaves were determined by gas-exchange analysis, in plants with each disease, grown under controlled conditions. The equation P(x)/P(0) = (1 - x)() was used to relate relative photosynthetic rate (P(x)/P(0)) to proportional disease severity (x), where beta represents the ratio between virtual and visual lesion. The beta values obtained for rust were near one, indicating that the effect of the pathogen on the remaining green leaf area was minimal. The high values of beta obtained for anthracnose (8.46 and 12.18) indicated that the photosynthesis in the green area beyond the necrotic symptoms of anthracnose was severely impaired. The impact of anthracnose on bean leaf photosynthesis should be considered in assessments of the proportion of healthy tissue in diseased leaves. The accurate assessment of the healthy portion of the leaf could improve the use of concepts such as healthy leaf area duration and healthy leaf area absorption, which are valuable predictors of crop yield. The equation used to analyze the interaction between rust and anthracnose on the same leaf was P(z) = P(0) (1 - x)(x) x (1 - y)(y), where P(z) is the relative photosynthetic rate of any given leaf, P(0) is the maximum relative photosynthetic rate, x is anthracnose severity, y is rust severity, betax is the beta value for anthracnose in the presence of rust, and betay is the beta value for rust in the presence of anthracnose. From the resulting response surface, no interaction of the two diseases was observed. Dark respiration rate increased on diseased leaves compared with control leaves. The remaining green leaf area of leaves with both diseases was not a good source to estimate net photosynthetic rate because the effect of anthracnose extended far beyond the visual lesions, whereas the effect of rust on photosynthesis was essentially limited to the pustule plus halo.
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