Aluminum (Al) toxicity is a major limiting factor for plant growth and productivity in acidic soil. At pH lower than 5.0 (pH < 5.0), the soluble and toxic form of Al (Al 3+ ions) enters root cells and inhibits root growth and uptake of water and nutrients. The organic acids malate, citrate, and oxalate are secreted by the roots and chelate Al 3+ to form a non-toxic Al-OA complex, which decreases the entry of Al 3+ into the root cells.When Al 3+ enters, it leads to the production of reactive oxygen species (ROS) in cells, which are toxic and cause damage to biomolecules like lipids, carbohydrates, proteins, and nucleic acids. When ROS levels rise beyond the threshold, plants activate an antioxidant defense system that comprises of superoxide dismutase (SOD), catalase (CAT), ascorbate peroxidase (APX), glutathione S-transferase (GST), ascorbic acid (ASA), phenolics and alkaloids etc., which protect plant cells from oxidative damage by scavenging and neutralizing ROS. Besides, ROS also play an important role in signal transduction and influence many molecular and cellular process like hormone signaling, gene expression, cell wall modification, cell cycle, programed cell death (PCD), and development. In the present review, the mechanisms of Al-induced ROS generation, ROS signaling, and crosstalk with other signaling pathways helping to combat Al toxicity have been summarized, which will help researchers to understand the intricacies of Al-induced plant response at cellular level and plan research for developing Al-toxicity tolerant crops for sustainable agriculture in acid soil-affected regions of the world.
| INTRODUCTIONApproximately 50% of the world's arable soils are acidic (Kochian et al., 2015). The major limitations for crop production on acid soils are the plant available form of aluminum (Al +3 ) and suboptimal level of phosphorus. Al is the most abundant metal in the earth's crust. It remains unavailable for plant uptake at near-neutral or alkaline pH as its stays in its mineral form. However, at pH < 5, its most abundant form is trivalent Al ion (Al 3+ ), which is soluble and highly toxic to the plant. Al toxicity inhibits root growth, which impairs the absorption of water and nutrients (Kochian et al., 2004(Kochian et al., , 2015. At cellular level, Al toxicity adversely affects the integrity and functioning of membranes, DNA synthesis, cell elongation, and metabolism (Bojórquez-Quintal et al., 2017).Though Al cannot catalyze redox reactions itself, a rise in oxidative stress under Al toxicity has been reported in many plant species.Previous studies based on gene expression have also reported induction of antioxidant-related genes, like peroxidase (POX), catalase (CAT), superoxide dismutase (SOD), glutathione-S-transferase (GSTs), etc. (Singh et al., 2017).Alok Ranjan and Ragini Sinha contributed equally to this study.