Peroxisomal plant metabolism - An update on nitric oxide, Ca2+ and the NADPH recycling network

Plant peroxisomes are recognized organelles that - with their capacity to generate greater amounts of HO than other subcellular compartments - have a remarkable oxidative metabolism. However, over the last 15 years, new information has shown that plant peroxisomes contain other important molecules and enzymes, including nitric oxide (NO), peroxynitrite, a NADPH-recycling system, Ca and lipid-derived signals, such as jasmonic acid (JA) and nitrofatty acid (NO-FA). This highlights the potential for complex interactions within the peroxisomal nitro-oxidative metabolism, which also affects the status of the cell and consequently its physiological processes. In this review, we provide an update on the peroxisomal interactions between all these molecules. Particular emphasis will be placed on the generation of the free-radical NO, which requires the presence of Ca, calmodulin and NADPH redox power. Peroxisomes possess several NADPH regeneration mechanisms, such as those mediated by glucose-6-phosphate dehydrogenase (G6PDH) and 6-phosphogluconate dehydrogenase (6PGDH) proteins, which are involved in the oxidative phase of the pentose phosphate pathway, as well as that mediated by NADP-isocitrate dehydrogenase (ICDH). The generated NADPH is also an essential cofactor across other peroxisomal pathways, including the antioxidant ascorbate- glutathione cycle and unsaturated fatty acid β-oxidation, the latter being a source of powerful signaling molecules such as JA and NO-FA.Research in our laboratories is supported by an European Regional Development Fund (ERDF) grant co-financed by the Ministerio de Economía y Competitividad (projects AGL2015-65104-P and BIO2015-66390-P) and the Junta de Andalucía (groups BIO192 and BIO286) in Spain.Peer Reviewe