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Surface waters of the subtropical Sargasso Sea contain dissolved inorganic phosphate (DIP) concentrations of 0.2 to 1.0 nanomolar, which are sufficiently low to result in phosphorus control of primary production. The DIP concentrations in this area (which receives high inputs of iron-rich dust from arid regions of North Africa) are one to two orders of magnitude lower than surface levels in the North Pacific (where eolian iron inputs are much lower and water column denitrification is much more substantial). These data indicate a severe relative phosphorus depletion in the Atlantic. We hypothesize that nitrogen versus phosphorus limitation of primary production in the present-day ocean may be closely linked to iron supply through control of dinitrogen (N2) fixation, an iron-intensive metabolic process. Although the oceanic phosphorus inventory may set the upper limit for the total amount of organic matter produced in the ocean over geological time scales, at any instant in geological time, oceanic primary production may fall below this limit because of a persistent insufficient iron supply. By controlling N2 fixation, iron may control not only nitrogen versus phosphorus limitation but also carbon fixation and export stoichiometry and hence biological sequestration of atmospheric carbon dioxide.
Suspended algae, or phytoplankton, are the prime source of organic matter supporting food webs in freshwater ecosystems. Phytoplankton productivity is reliant on adequate nutrient supplies; however, increasing rates of nutrient supply, much of it manmade, fuels accelerating primary production or eutrophication. An obvious and problematic symptom of eutrophication is rapid growth and accumulations of phytoplankton, leading to discoloration of affected waters. These events are termed blooms. Blooms are a prime agent of water quality deterioration, including foul odors and tastes, deoxygenation of bottom waters (hypoxia and anoxia), toxicity, fish kills, and food web alterations. Toxins produced by blooms can adversely affect animal (including human) health in waters used for recreational and drinking purposes. Numerous freshwater genera within the diverse phyla comprising the phytoplankton are capable of forming blooms; however, the blue-green algae (or cyanobacteria) are the most notorious bloom formers. This is especially true for harmful toxic, surface-dwelling, scum-forming genera (e.g., Anabaena, Aphanizomenon, Nodularia, Microcystis) and some subsurface bloom-formers (Cylindrospermopsis, Oscillatoria) that are adept at exploiting nutrient-enriched conditions. They thrive in highly productive waters by being able to rapidly migrate between radiance-rich surface waters and nutrient-rich bottom waters. Furthermore, many harmful species are tolerant of extreme environmental conditions, including very high light levels, high temperatures, various degrees of desiccation, and periodic nutrient deprivation. Some of the most noxious cyanobacterial bloom genera (e.g., Anabaena, Aphanizomenon, Cylindrospermopsis, Nodularia) are capable of fixing atmospheric nitrogen (N2), enabling them to periodically dominate under nitrogen-limited conditions. Cyanobacteria produce a range of organic compounds, including those that are toxic to higher-ranked consumers, from zooplankton to further up the food chain. Both N2- and non-N2-fixing genera participate in mutualistic and symbiotic associations with microorganisms, higher plants, and animals. These associations appear to be of great benefit to their survival and periodic dominance. In this review, we address the ecological impacts and environmental controls of harmful blooms, with an emphasis on the ecology, physiology, and management of cyanobacterial bloom taxa. Combinations of physical, chemical, and biotic features of natural waters function in a synergistic fashion to determine the sensitivity of water bodies. In waters susceptible to blooms, human activities in water- and airsheds have been linked to the extent and magnitudes of blooms. Control and management of cyanobacterial and other phytoplankton blooms invariably includes nutrient input constraints, most often focused on nitrogen (N) and/or phosphorus (P). The types and amount of nutrient input constraints depend on hydrologic, climatic, geographic, and geologic factors, which interact with anthropogenic and natural nutrient input regimes. While single nutrient input constraints may be effective in some water bodies, dual N and P input reductions are usually required for effective long-term control and management of harmful blooms. In some systems where hydrologic manipulations (i.e., plentiful water supplies) are possible, reducing the water residence time by enhanced flushing and artificial mixing (in conjunction with nutrient input constraints) can be particularly effective alternatives. Implications of various management strategies, based on combined ecophysiological and environmental considerations, are discussed.
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