Alterations of the Biochemical Pathways of Plants by the Air Pollutant Ozone: Which are the True Gauges of Injury?

Plant strategies to survive ozone stress include exclusion or tolerance of ozone. If these processes fail, past observations of ozone injury have indicated many physiological and metabolic changes then occur; most of these changes are likely to have been initiated at the level of gene expression, suggesting signal transduction. In the last decade considerable understanding of the biochemical process within plants has been developed. Currently there are several hypotheses regarding a response of plants to ozone fumigation: [1] membrane dysfunction and alteration of purpose; [2] stress ethylene interactions; [3] impairment of photosynthesis via changes in Rubisco levels and the guard cells so that the stomata do not track correctly the environment; [4] antioxidant protection through metabolites and enzyme systems to reduce the oxidant load; and [5] general impairment or disruption of metabolic pathways. Many believe that free radicals and other oxidative products, formed in plant leaves under ozone exposure, are responsible for much of the spread of the biochemical alterations. There are obvious chemicals that may account for the changes that are observed, such as hydrogen peroxide. Once the ozone enters the tissue, evidence suggests the first line of defense is a range of antioxidants, such as ascorbate, glutathione peroxidase, superoxide dismutase, and catalase. If overwhelmed, subsequent events occur which are highly suggestive of systemic acquired resistance. Furthermore, other defensive indicators, such as salicylic acid and jasmonic acid, tend to increase, but more slowly than ethylene, and spread their signaling effects more widely in the plant. The primary set of metabolic reactions that ozone triggers is thought to be “wounding“ responses with a secondary response of senescence. The dramatic strides in understanding the genetic make-up of plants, gene control, and signal transduction/control over the last few years will only accelerate in the future. We need now to have an understanding of those events that can be translated into more detailed schemes of how ozone alters much of the basic metabolism of plants and how plants counteract or cope with ozone. What is now known about how varied biochemicals and their pathways are changed upon ozone exposure will be discussed.