Sugars play important roles as both nutrients and regulatory molecules throughout plant life. Sugar metabolism and signalling function in an intricate network with numerous hormones and reactive oxygen species (ROS) production, signalling and scavenging systems. Although hexokinase is well known to fulfil a crucial role in glucose sensing processes, a scenario is emerging in which the catalytic activity of mitochondria‐associated hexokinase regulates glucose‐6‐phosphate and ROS levels, stimulating antioxidant defence mechanisms and the synthesis of phenolic compounds. As a new concept, it can be hypothesized that the synergistic interaction of sugars (or sugar‐like compounds) and phenolic compounds forms part of an integrated redox system, quenching ROS and contributing to stress tolerance, especially in tissues or organelles with high soluble sugar concentrations.
Sugars are involved in many metabolic and signalling pathways in plants. Sugar signals may also contribute to immune responses against pathogens and probably function as priming molecules leading to pathogen-associated molecular patterns (PAMP)-triggered immunity and effector-triggered immunity in plants. These putative roles also depend greatly on coordinated relationships with hormones and the light status in an intricate network. Although evidence in favour of sugar-mediated plant immunity is accumulating, more in-depth fundamental research is required to unravel the sugar signalling pathways involved. This might pave the way for the use of biodegradable sugar-(like) compounds to counteract plant diseases as cheaper and safer alternatives for toxic agrochemicals.
Over the past decades, considerable advances have been made in understanding the crucial role and the regulation of sucrose metabolism in plants. Among the various sucrose-catabolizing enzymes, alkaline/neutral invertases (A/N-Invs) have long remained poorly studied. However, recent findings have demonstrated the presence of A/N-Invs in various organelles in addition to the cytosol, and their importance for plant development and stress tolerance. A cytosolic (At-A/N-InvG, At1g35580) and a mitochondrial (At-A/N-InvA, At1g56560) member of the A/N-Invs have been analysed in more detail in Arabidopsis and it was found that At-A/N-InvA knockout plants show an even more severe growth phenotype than At-A/N-InvG knockout plants. The absence of either A/N-Inv was associated with higher oxidative stress defence gene expression, while transient overexpression of At-A/N-InvA and At-A/N-InvG in leaf mesophyll protoplasts down-regulated the oxidative stress-responsive ascorbate peroxidase 2 (APX2) promoter. Moreover, up-regulation of the APX2 promoter by hydrogen peroxide or abscisic acid could be blocked by adding metabolizable sugars or ascorbate. A hypothetical model is proposed in which both mitochondrial and cytosolic A/N-Invs can generate glucose as a substrate for mitochondria-associated hexokinase, contributing to mitochondrial reactive oxygen species homeostasis.
All organisms have an internal timing mechanism, termed the circadian clock, to anticipate the light/dark cycle. The clock, with an oscillating rhythm that approximates 24h, is a rather robust system persisting to a great extent in continuous light and dark. It is widely accepted that plant growth and development are regulated by the clock, hormones, and sugar signals. On the one hand, sugar signalling can affect circadian rhythms by altering the expression pattern of clock-regulated genes. More in particular, the clock seems to be particularly sensitive to sucrose-mediated signalling which is also associated with immunity and abiotic stress responses. Also, hormonal interaction with the clock can contribute to appropriate plant immune responses. Recent data show a prominent role for the clock in growth and stress responses. On the other hand, the clock seems to be essential in controlling the gene expression and activity of an array of carbohydrate-metabolizing enzymes, suggesting a complex reciprocal relationship between the clock and metabolic signalling processes. Therefore, the clock fulfils a crucial role at the heart of cellular networks. The players involved in the complex plant circadian network and their possible contribution to the novel 'sweet immunity' concept are discussed.
Sugars do not only act as source of energy, but they also act as signals in plants. This mini review summarizes the emerging links between sucrose-mediated signaling and the cellular networks involved in flowering time control and defense. Cross-talks with gibberellin and jasmonate signaling pathways are highlighted. The circadian clock fulfills a crucial role at the heart of cellular networks and the bilateral relation between sugar signaling and the clock is discussed. It is proposed that important factors controlling plant growth (DELLAs, PHYTOCHROME INTERACTING FACTORS, invertases, and trehalose-6-phosphate) might fulfill central roles in the transition to flowering as well. The emerging concept of “sweet immunity,” modulated by the clock, might at least partly rely on a sucrose-specific signaling pathway that needs further exploration.
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