Seagrasses and eutrophication

Abstract

This review summarizes the historic, correlative field evidence and experimental research that implicate cultural eutrophication as a major cause of seagrass disappearance. We summarize the underlying physiological responses of seagrass species, the potential utility of various parameters as indicators of nutrient enrichment in seagrasses, the relatively sparse available information about environmental conditions that exacerbate eutrophication effects, and the better known array of indirect stressors imposed by nutrient over-enrichment that influence seagrass growth and survival. Seagrass recovery following nutrient reductions is examined, as well as the status of modeling efforts to predict seagrass response to changing nutrient regimes. The most common mechanism invoked or demonstrated for seagrass decline under nutrient over-enrichment is light reduction through stimulation of high-biomass algal overgrowth as epiphytes and macroalgae in shallow coastal areas, and as phytoplankton in deeper coastal waters. Direct physiological responses such as ammonium toxicity and water-column nitrate inhibition through internal carbon limitation may also contribute. Seagrass decline under nutrient enrichment appears to involve indirect and feedback mechanisms, and is manifested as sudden shifts in seagrass abundance rather than continuous, gradual changes in parallel with rates of increased nutrient additions. Depending on the species, interactions of high salinity, high temperature, and low light have been shown to exacerbate the adverse effects of nutrient over-enrichment. An array of indirect effects of nutrient enrichment can accelerate seagrass disappearance, including sediment re-suspension from seagrass loss, increased system respiration and resulting oxygen stress, depressed advective water exchange from thick macroalgal growth, biogeochemical alterations such as sediment anoxia with increased hydrogen sulfide concentrations, and internal nutrient loading via enhanced nutrient fluxes from sediments to the overlying water. Indirect effects on trophic structure can also be critically important, for example, the loss of herbivores, through increased hypoxia/anoxia and other habitat shifts, that would have acted as “ecological engineers” in promoting seagrass survival by controlling algal overgrowth; and shifts favoring exotic grazers that out-compete seagrasses for space. Evidence suggests that natural seagrass population shifts are disrupted, slowed or indefinitely blocked by cultural eutrophication, and there are relatively few known examples of seagrass meadow recovery following nutrient reductions. Reliable biomarkers as early indicators of nutrient over-enriched seagrass meadows would benefit coastal resource managers in improving protective measures. Seagrasses can be considered as “long-term" integrators (days to weeks) of nutrient availability, especially through analyses of their tissue content, and of activities of enzymes such as nitrate reductase and alkaline phosphatase. The ratio of leaf nitrogen content to leaf mass has also shown promise as a “nutrient pollution indicator” for the seagrass Zostera marina, with potential application to other species. In modeling efforts, seagrass response to nutrient loading has proven difficult to quantify beyond localized areas because long-term data consistent in quality are generally lacking, and high inter-annual variability in abundance and productivity depending upon stochastic meteorological and hydrographic conditions. Efforts to protect remaining seagrass meadows from damage and loss under eutrophication, within countries and across regions, are generally lacking or weak and ineffective. Research needs to further understand about seagrasses and eutrophication should emphasize experimental studies to assess the response of a wider range of species to chronic, low-level as well as acute, pulsed nutrient enrichment. These experiments should be conducted in the field or in large-scale mesocosms following appropriate acclimation, and should emphasize factor interactions (N, P, C; turbidity; temperature; herbivory) to more closely simulate reality in seagrass ecosystems. They should scale up to address processes that occur over larger scales, including food-web dynamics that involve highly mobile predators and herbivores. Without any further research, however, one point is presently very clear: Concerted local and national actions, thus far mostly lacking, are needed worldwide to protect remaining seagrass meadows from accelerating cultural eutrophication in rapidly urbanizing coastal zones.