Role of organic acids in the integration of cellular redox metabolism and mediation of redox signalling in photosynthetic tissues of higher plants

Organic acids play a crucial role in numerous metabolic processes accompanied by transfer of electrons and protons and linked to the reduction/ oxidation of major redox couples in plant cells, such as NAD, NADP, glutathione, and ascorbate. Fluxes through the pathways metabolizing organic acids modulate redox states in cell compartments, contribute to generation of reactive oxygen and nitrogen species, and mediate signal transduction processes. Organic acid metabolism not only functions to equilibrate the redox potential in plant cells but also to transfer redox equivalents between cell compartments supporting various metabolic processes. The most important role in this transfer belongs to different forms of malate dehydrogenase interconverting malate and oxaloacetate or forming pyruvate (malic enzymes). During photosynthesis malate serves as a major form of transfer of redox equivalents from chloroplasts to the cytosol and other compartments via the malate valve. On the other hand, mitochondria, via alterations of their redox potential, become a source of citrate that can be transported to the cytosol and support biosynthesis of amino acids. Citrate is also an important retrograde signalling compound that regulates transcription of several genes including those encoding the alternative oxidase. The alternative oxidase, which is activated by increased redox potential and by pyruvate, is, in turn, important for the maintenance of redox potential in mitochondria. The roles of organic acids in establishing redox equilibrium, supporting ionic gradients on membranes, acidification of the extracellular medium, and regulation of production of reactive oxygen and nitrogen species are discussed.