Copper: uptake, toxicity and tolerance in plants and management of Cu-contaminated soil.

Copper (Cu) is an essential mineral nutrient for the proper growth and development of plants; it is involved in myriad morphological, physiological, and biochemical processes. Copper acts as a cofactor in various enzymes and performs essential roles in photosynthesis, respiration and the electron transport chain, and is a structural component of defense genes. Excess Cu, however, imparts negative effects on plant growth and productivity. Many studies have summarized the adverse effects of excess Cu on germination, growth, photosynthesis, and antioxidant response in agricultural crops. Its inhibitory influence on mineral nutrition, chlorophyll biosynthesis, and antioxidant enzyme activity has been verified. The current review focuses on the availability and uptake of Cu by plants. The toxic effects of excess Cu on seed germination, plant growth and development, photosynthesis, and antioxidant response in plants are discussed. Plant tolerance mechanisms against Cu stress, and management of Cu-contaminated soils are presented.