Mechanisms and regulation of aluminum-induced secretion of organic acid anions from plant roots

Aluminum(Al)is the most abundant metal element in the earth’s crust.On acid soils,at pH 5.5 or lower,part of insoluble Al-containing minerals become solubilized into soil solution,with resultant highly toxic effects on plant growth and development.Nevertheless,some plants have developed Al-tolerance mechanisms that enable them to counteract this Al toxicity.One such well-documented mechanism is the Al-induced secretion of organic acid anions,including citrate,malate,and oxalate,from plant roots.Once secreted,these anions chelate external Al ions,thus protecting the secreting plant from Al toxicity.Genes encoding the citrate and malate transporters responsible for secretion have been identified and characterized,and accumulating evidence indicates that regulation of the expression of these transporter genes is critical for plant Al tolerance.In this review,we outline the recent history of research into plant Al-tolerance mechanisms,with special emphasis on the physiology of Al-induced secretion of organic acid anions from plant roots.In particular,we summarize the identification of genes encoding organic acid transporters and review current understanding of genes regulating organic acid secretion.We also discuss the possible signaling pathways regulating the expression of organic acid transporter genes.

Organic acid anions:An effective defensive weapon for plants against aluminum toxicity and phosphorus deficiency in acidic soils

Aluminum （AI） toxicity and phosphorous （P） deficiency are two major limiting factors for plant growth on acidic soils. Thus, the physiological mechanisms for AJ tolerance and P acquisition have been intensively studied. A commonly observed trait is that plants have developed the ability to utilize organic acid anions （OAs; mainly malate, citrate and oxalate） to combat AI toxicity and P deficiency. OAs secreted by roots into the rhizosphere can externally chelate Al^3＋ and mobilize phosphate （Pi）, while OAs synthesized in the cell can internally sequester Al^3＋ into the vacuole and release free Pi for metabolism. Molecular mechanisms involved in OA synthesis and transport have been described in detail. Ensuing genetic improvement for AI tolerance and P efficiency through increased OA exudation and/or synthesis in crops has been achieved by transgenic and marker-assisted breeding. This review mainly elucidates the crucial roles of OAs in plant Al tolerance and P efficiency through summarizing associated physiological mechanisms, molecular traits and genetic manipulation of crops.