Sugar-mediated regulation and the role of HXK1, SnRK1, TOR kinases in Arabidopsis thaliana

Sugars play an important role in plant metabolism. They are not only a source of energy, but also an integral part of the intracellular signalling network. Signals involving sugars regulate a variety of metabolic reactions, e.g. carbon and nitrogen metabolism, and produce plant response to biotic and abiotic stresses. This paper provides a brief review of modern publications about sugars acting as metabolites and signaling molecules. The well-known and recently discovered pathways for the transmission of sugar-mediated signals are described, along with their role in the regulation of plant metabolism and expression of nuclear genes. Hexokinase 1 (an enzyme for glucose synthesis), which was one of the first studied participants in the sugar-mediated signalling network, regulates the expression of many nuclear genes in response to changes in the glucose level. Hexokinase 1 should be considered as a broad-spectrum regulator due to its participation in the transmission of mitochondrial-nuclear signals and regulation of aging processes in plants, which functions lay beyond the regulation of metabolic reactions. Of particular interest is the participation of sugar-mediated signals in the switching of metabolic reactions in response to changes in carbohydrate concentrations. Sugar deficiency deactivates most energy-intensive processes, at the same time as stimulating photosynthesis or (in its absence) enhancing catabolic reactions due to activation of the SnRK1 kinase. Under sufficient sugar levels, the SnRK1 kinase is suppressed and the TOR kinase is activated, which stimulates anabolic reactions and growth. The role of competing SnRK1 and TOR kinases as the main regulators of such a process is considered, along with the function of signals mediated by trehalose-6-phosphate highly similar to the TOR signalling pathway. All these regulatory mechanisms enable plants to adapt to environmental changes and maintain homeostasis. It is possible that additional information will be obtained in the nearest future on the interaction of energy and stress-dependent signals in plants through the antagonism of TOR and SnRK1.

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