Ascorbic acid, a familiar small molecule intertwined in the response of plants to ozone, pathogens, and the onset of senescence

Ascorbic acid is a well-known antioxidant and cellular reductant with an intimate and complex role in the response of plants to ozone. It is clear from a number of studies that sensitivity to ozone is correlated with total ascorbic acid levels, and that a first line of defence against the reactive oxygen species generated in the apoplastic space by ozone is ascorbic acid. For activity, ascorbic acid must be in the fully reduced state. Therefore, both the rate of ascorbic acid synthesis and recycling via dehydroascorbate and monodehydroascorbate reductases are critical in the maintenance of a high ascorbic acid redox state. Active transport of ascorbic acid across the plasma membrane is necessary to achieve reduction of oxidized ascorbic acid by cytoplasm-localized reductases. It has been known for some time that the chlorotic lesions produced by exposure to ozone are not unlike lesions produced by the hypersensitive response to avirulent pathogen attack. Surprisingly, activation of a defence gene-signalling network by both ozone and pathogens is influenced by the level of ascorbic acid. Indeed, in addition to acting simply as an antioxidant in the apoplastic space, ascorbic acid appears to be involved in a complex phytohormone-mediated signalling network that ties together ozone and pathogen responses and influences the onset of senescence.