BIOCHEMICAL, PHYSIOLOGICAL, AND STRUCTURAL EFFECTS OF EXCESS COPPER IN PLANTS

Heavy metal pollution is one of the most troublesome environmental problems faced by mankind nowadays. Copper, in particular, poses serious problems due to its widespread industrial and agricultural use. Unlike other heavy metals, such as cadmium, lead, and mercury, copper is not readily bioaccumulated and thus its toxicity to man and other mammals is relatively low. On the contrary, plants in general are very sensitive to Cu toxicity, displaying metabolic disturbances and growth inhibition at Cu contents in the tissues only slightly higher than the normal levels. The reduced mobility of Cu in soil and sediments, due to its strong binding to organic and inorganic colloids, constitutes, in a way, a barrier to Cu toxicity in land plants. In aqueous media, however, plants are directly exposed to the harmful effects of Cu and, thus, algae and some species of aquatic higher plants are more easily subjected to Cu toxicity. Excess Cu inhibits a large number of enzymes and interferes with several aspects of plant biochemistry, including photosynthesis, pigment synthesis, and membrane integrity. Perhaps its most important effect is associated with the blocking of photosynthetic electron transport, leading to the production of radicals which start peroxidative chain reactions involving membrane lipids. Copper effects on plant physiology are wide ranging, including interference with fatty acid and protein metabolism and inhibition of respiration and nitrogen fixation processes. At the whole plant level Cu is an effective inhibitor of vegetative growth and induces general symptoms of senescence. The high toxicity of Cu to plants has led to the evolution of several strategies of defense. Among the most important ones is the production of Cu-complexing compounds. Although the nature, structure, and function of these compounds is still controversial, they can be divided into two main groups: metallothionein-like compounds and phytochelatins. The latter appears to constitute the most widespread response of plants to stresses provoked by metals, including Cu.