Concepts of plant biotic stress. Some insights into the stress physiology of virus-infected plants, from the perspective of photosynthesis

The consequences of biotic stress have been poorly understood, partly because its application is difficult to control and partly because its physiological consequences are highly variable. Many plant viruses are recognised on the basis of leaf symptoms that depend on localised changes to chloroplast structure and function. This paper reviews recent progress in understanding early interactions between plant viruses and the photosynthetic apparatus, using chlorophyll fluorescence analysis of novel, defined algal-virus systems and using high resolution imaging of chlorophyll fluorescence and other photosynthetic processes in higher plant systems. We then consider the consequences of viral effects on photosynthetic functioning for whole plants and populations with an emphasis on the potential interactions with other environmental factors. Early responses indicated by increase in both non-photochemical quenching of fluorescence and increased reduction state of the primary electron transport acceptor Q(A) suggest that, not surprisingly, bath photoprotective and photoinhibitory processes contribute to the accelerated local demise of the photosynthetic apparatus and symptom development. In other cases, localised accumulations of carbohydrate and source-sink imbalance following infection may inhibit gene expression, leading to altered levels of chloroplast protein complexes and enzymes of photosynthetic metabolism coincident with symptom development. Recent experiments suggest that much of the variability in plant responses to biotic stress may result from interactions with other environmental factors, such as light intensity and nutrition. Experiments suggest that virus infections may have greater effects on fitness and competitive ability in low N, high light environments than in shaded, high nutrient conditions. Some ecological implications of these observations are discussed.