Sulfur assimilation in plants and weed control: Potential targets for novel herbicides and action sites of certain safeners

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Sulfur is an indispensable element for plants. It is found in sulfur‐containing amino acids, cysteine and methionine, and in various other important biochemical components and processes. Inhibitors of sulfur assimilation, or cysteine and methionine synthesis, could be potential herbicides. In the present paper, the sulfur assimilation pathway in plants is described, followed by the introduction of several compounds (inhibitors and safeners) acting on this pathway. Uptake of inorganic sulfate through the roots is the first step of sulfur assimilation in plants. Sulfate is reduced mainly in chloroplasts to sulfide by a multistep process, and sulfide is then incorporated into cysteine. Cysteine is converted to cystathionine, homocysteine and methionine. Cysteine is incorporated into glutathione (GSH) by γ‐glutamylcysteine synthetase and GSH synthetase. Three enzymes involved in cysteine and methionine biosynthesis, cysteine synthase, cystathionine γ‐synthase and cystathionine β‐lyase, have been investigated as target sites for herbicides. Several inhibitors of these enzymes (e.g. rhizobitoxine and propargylglycine) were also phytotoxic, suggesting that the synthetic pathway of sulfur‐containing amino acids could be a new target site for herbicides. Some safeners for herbicides were found to act on the sulfur assimilation pathway and on GSH synthesis to increase GSH, which can be involved in herbicide metabolism and detoxification. Several safeners elevate GSH levels by increasing the activities of enzymes involved in sulfur assimilation and GSH synthesis. Further studies on plant sulfur metabolism may lead to the discovery of new herbicides and to the comprehensive understanding of the mode of action of safeners.