Organic acids: versatile stress-response roles in plants.

Organic acids (OAs) are central to cellular metabolism. Many plant stress responses involve the exudation of OAs at the root-soil interface, which can improve soil mineral acquisition and toxic metal tolerance. Because of their simple structure, the low-molecular-weight OAs are widely studied. We discuss the conventional roles of OAs, and some newly emerging roles in plant stress tolerance. OAs are more versatile in their role in plant stress tolerance and are more efficient chelating agents than other acids, such as amino acids. Root OA exudation is important in soil carbon sequestration. These functions are key processes in combating climate change and helping with more sustainable food production. We briefly review the mechanisms behind enhanced biosynthesis, secretion, and regulation of these activities under different stresses, and provide an outline of the transgenic approaches targeted towards the enhanced production and secretion of OAs. A recurring theme of OAs in plant biology is their role as 'acids' modifying pH, as 'chelators' binding metals, or as 'carbon sources' for microbes. We argue that these multiple functions are key factors for understanding these molecules' important roles in plant stress biology. Finally, we discuss how the functions of OAs in plant stress responses could be used, and identify the important unanswered questions.