Overview of the physiological ecology of carbon metabolism in seagrasses.

The small but diverse group of angiosperms known as seagrasses form submersed meadow communities that are among the most productive on earth. Seagrasses are frequently light-limited and, despite access to carbon-rich seawaters, they may also sustain periodic internal carbon limitation. They have been regarded as C3 plants, but many species appear to be C3-C4 intermediates and/or have various carbon-concentrating mechanisms to aid the Rubisco enzyme in carbon acquisition. Photorespiration can occur as a C loss process that may protect photosynthetic electron transport during periods of low CO(2) availability and high light intensity. Seagrasses can also become photoinhibited in high light (generally>1000 µE m(-2) s(-1)) as a protective mechanism that allows excessive light energy to be dissipated as heat. Many photosynthesis-irradiance curves have been developed to assess light levels needed for seagrass growth. However, most available data (e.g. compensation irradiance I(c)) do not account for belowground tissue respiration and, thus, are of limited use in assessing the whole-plant carbon balance across light gradients. Caution is recommended in use of I(k) (saturating irradiance for photosynthesis), since seagrass photosynthesis commonly increases under higher light intensities than I(k); and in estimating seagrass productivity from H(sat) (duration of daily light period when light equals or exceeds I(k)) which varies considerably among species and sites, and which fails to account for light-limited photosynthesis at light levels less than I(k). The dominant storage carbohydrate in seagrasses is sucrose (primarily stored in rhizomes), which generally forms more than 90% of the total soluble carbohydrate pool. Seagrasses with high I(c) levels (suggesting lower efficiency in C acquisition) have relatively low levels of leaf carbohydrates. Sucrose-P synthase (SPS, involved in sucrose synthesis) activity increases with leaf age, consistent with leaf maturation from carbon sink to source. Unlike terrestrial plants, SPS apparently is not light-activated, and is positively influenced by increasing temperature and salinity. This response may indicate an osmotic adjustment in marine angiosperms, analogous to increased SPS activity as a cryoprotectant response in terrestrial non-halophytic plants. Sucrose synthase (SS, involved in sucrose metabolism and degradation in sink tissues) of both above- and belowground tissues decreases with tissue age. In belowground tissues, SS activity increases under low oxygen availability and with increasing temperatures, likely indicating increased metabolic carbohydrate demand. Respiration in seagrasses is primarily influenced by temperature and, in belowground tissues, by oxygen availability. Aboveground tissues (involved in C assimilation and other energy-costly processes) generally have higher respiration rates than belowground (mostly storage) tissues. Respiration rates increase with increasing temperature (in excess of 40 degrees C) and increasing water-column nitrate enrichment (Z. marina), which may help to supply the energy and carbon needed to assimilate and reduce nitrate. Seagrasses translocate oxygen from photosynthesizing leaves to belowground tissues for aerobic respiration. During darkness or extended periods of low light, belowground tissues can sustain extended anerobiosis. Documented alternate fermentation pathways have yielded high alanine, a metabolic 'strategy' that would depress production of the more toxic product ethanol, while conserving carbon skeletons and assimilated nitrogen. In comparison to the wealth of information available for terrestrial plants, little is known about the physiological ecology of seagrasses in carbon acquisition and metabolism. Many aspects of their carbon metabolism - controls by interactive environmental factors; and the role of carbon metabolism in salt tolerance, growth under resource-limited conditions, and survival through periods of dormancy - remain to be resolved as directions in future research. Such research will strengthen the understanding needed to improve management and protection of these environmentally important marine angiosperms.