Key Laboratory of Agricultural Water Resources, Hebei Key Laboratory of Agricultural Water-Saving, Center for Agricultural Resources Research, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Key Laboratory of Agricultural Wat, Shijiazhuang, Hebei, China
Dr. Arbindra Timilsina
Key Laboratory of Agricultural Water Resources, Hebei Key Laboratory of Agricultural Water-Saving, Center for Agricultural Resources Research, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Key Laboratory of Agricultural Wat, Shijiazhuang, Hebei, China
Oxygen (O2) is the most crucial substrate for numerous biochemical processes in plants. Its deprivation is a critical factor that affects plant growth and may lead to death if it lasts for a long time. However, various biotic and abiotic factors cause O2 deprivation, leading to hypoxia and anoxia in plant tissues. To survive under hypoxia and/or anoxia, plants deploy various mechanisms such as fermentation paths, reactive oxygen species (ROS), reactive nitrogen species (RNS), antioxidant enzymes, aerenchyma, and adventitious root formation, while nitrate (NO3−), nitrite (NO2−), and nitric oxide (NO) have shown numerous beneficial roles through modulating these mechanisms. Therefore, in this review, we highlight the role of reductive pathways of NO formation which lessen the deleterious effects of oxidative damages and increase the adaptation capacity of plants during hypoxia and anoxia. Meanwhile, the overproduction of NO through reductive pathways during hypoxia and anoxia leads to cellular dysfunction and cell death. Thus, its scavenging or inhibition is equally important for plant survival. As plants are also reported to produce a potent greenhouse gas nitrous oxide (N2O) when supplied with NO3− and NO2−, resembling bacterial denitrification, its role during hypoxia and anoxia tolerance is discussed here. We point out that NO reduction to N2O along with the phytoglobin-NO cycle could be the most important NO-scavenging mechanism that would reduce nitro-oxidative stress, thus enhancing plants’ survival during O2-limited conditions. Hence, understanding the molecular mechanisms involved in reducing NO toxicity would not only provide insight into its role in plant physiology, but also address the uncertainties seen in the global N2O budget.
Paddies are a potential source of anthropogenic nitrous oxide (N2O) emission. In paddies, both the soil and the rice plants emit N2O into the atmosphere. The rice plant in the paddy is considered to act as a channel between the soil and the atmosphere for N2O emission. However, recent studies suggest that plants can also produce N2O, while the mechanism of N2O formation in plants is unknown. Consequently, the rice plant is only regarded as a channel for N2O produced by soil microorganisms. The emission of N2O by aseptically grown plants and the distinct dual isotopocule fingerprint of plant-emitted N2O, as reported by various studies, support the production of N2O in plants. Herein, we propose a potential pathway of N2O formation in the rice plant. In rice plants, N2O might be formed in the mitochondria via the nitrate–nitrite–nitric oxide (NO3–NO2–NO) pathway when the cells experience hypoxic or anoxic stress. The pathway is catalyzed by various enzymes, which have been described. So, N2O emitted from paddies might have two origins, namely soil microorganisms and rice plants. So, regarding rice plants only as a medium to transport the microorganism-produced N2O might be misleading in understanding the role of rice plants in the paddy. As rice cultivation is a major agricultural activity worldwide, not understanding the pathway of N2O formation in rice plants would create more uncertainties in the N2O budget.
Plants can produce and emit nitrous oxide (N2O), a potent greenhouse gas, into the atmosphere, and several field-based studies have concluded that this gas is emitted at substantial amounts. However, the exact mechanisms of N2O production in plant cells are unknown. Several studies have hypothesised that plants might act as a medium to transport N2O produced by soil-inhabiting microorganisms. Contrarily, aseptically grown plants and axenic algal cells supplied with nitrate (NO3) are reported to emit N2O, indicating that it is produced inside plant cells by some unknown physiological phenomena. In this study, the possible sites, mechanisms, and enzymes involved in N2O production in plant cells are discussed. Based on the experimental evidence from various studies, we determined that N2O can be produced from nitric oxide (NO) in the mitochondria of plants. NO, a signaling molecule, is produced through oxidative and reductive pathways in eukaryotic cells. During hypoxia and anoxia, NO3 in the cytosol is metabolised to produce nitrite (NO2), which is reduced to form NO via the reductive pathway in the mitochondria. Under low oxygen condition, NO formed in the mitochondria is further reduced to N2O by the reduced form of cytochrome c oxidase (CcO). This pathway is active only when cells experience hypoxia or anoxia, and it may be involved in N2O formation in plants and soil-dwelling animals, as reported previously by several studies. NO can be toxic at a high concentration. Therefore, the reduction of NO to N2O in the mitochondria might protect the integrity of the mitochondria, and thus, protect the cell from the toxicity of NO accumulation under hypoxia and anoxia. As NO3 is a major source of nitrogen for plants and all plants may experience hypoxic and anoxic conditions owing to soil environmental factors, a significant global biogenic source of N2O may be its formation in plants via the proposed pathway.