Seagrass-salinity interactions: Physiological mechanisms used by submersed marine angiosperms for a life at sea

Abstract Due to the nature of coastal and estuarine systems, seagrasses must be able to tolerate short-term salinity fluctuations including both hyposaline and hypersaline conditions. Salt tolerance can be achieved, in part, through vacuolar ion sequestering (mostly Na+, K+, and Cl−) and cytosolic osmolyte accumulation (K+ and organic osmolytes), with differences in cellular ion levels attributed to selective ion flux and ion partitioning between the cytoplasm and vacuole (with lower cytoplasmic-to-vacuolar ratios favoring higher cellular Na+ concentrations). The hydrophilic nature of organic compounds such as organic acids, soluble carbohydrates, and free amino acids allow them to serve as osmoprotectants and low-molecular-weight chaperones which diminishes the inhibitory effects of potentially harmful ions on metabolic processes. Nevertheless, some carbohydrate studies on seagrasses have shown decreased soluble sugar content with increased salinities. During salt stress, carbohydrates are likely converted to other organic compounds that would better facilitate osmotic adjustment in these plants. This is further supported by observed decreases in sucrose-P synthase (a key enzyme involved in sucrose synthesis) activities in seagrass exposed to higher salinities. While modifications in ion flux and organic solute levels often follow changes in environmental salinities, these adjustments are relatively slow (hours to days). Therefore, the initial response to sudden salinity change will include rapid alterations in turgor pressure driven by water flux in the direction of the osmotic gradient. The rate of water movement depends largely on the hydraulic conductivity of the plasmalemma and the elastic properties of the cell wall (bulk elastic modulus; Є). Observations on cell wall elasticity indicate that some seagrasses maintain fairly rigid walls (high Є values), thereby limiting the amount of water influx during hypoosmotic stress. Although high Є would be beneficial to open-water coastal plants living in relatively stable saline environments, in estuaries where salinities fluctuate considerably over shorter intervals, high Є could promote flaccid cells with no turgor pressure during hyperosmotic conditions. Hypo- and hyperosmotic conditions also inhibit photosynthesis in seagrasses. Decreases in photosynthesis have been attributed to declines in chlorophyll content, changes in chloroplast ultrastructure, disruptions of electron flow through photosystems, and inhibitions of key photosynthetic enzymes. The uptake of nutrients can also be strongly influenced by salinity. High affinity Na+-dependent nutrient transport systems (for NO3−, H2PO4−, and HPO4−2) which benefit from the inwardly driving force for Na+ have been observed in seagrasses. Nitrate reductase, the key enzyme involved in nitrate reduction/ assimilation, also has elevated activities at higher salinities which would agree with Na+-dependent NO3− transport. While our basic understanding of how seagrasses survive in saline environments is increasing, it still lags well behind marine algae and terrestrial halophytes. It is likely that further investigations will reveal unique physiological adaptations that have not been observed in other plants.