Chan
Francis
Chan
Francis
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ArticleEcological constraints on planktonic nitrogen fixation in saline estuaries. I. Nutrient and trophic controls(Inter-Research, 2006-03-15) Marino, Roxanne ; Chan, Francis ; Howarth, Robert W. ; Pace, Michael L. ; Likens, Gene E.Heterocystous, planktonic cyanobacteria capable of fixing atmospheric N2 into available nitrogen (N) are common and critically important to nutrient cycling in many lakes, yet they are rarely observed in estuaries at salinities >10 ppt, even when strongly N limited. In a series of mesocosm experiments using water from Narragansett Bay (Rhode Island), we manipulated top-down (grazing) and bottom-up (nutrient) factors hypothesized to exclude heterocystous cyanobacteria from estuaries. We previously reported that planktonic, heterocystous cyanobacteria grew and fixed N in the absence of grazers. Here, we focus on responses to phosphorus (P) additions and grazer manipulations. Zooplankton (Acartia sp.) populations typical of temperate zone estuaries suppressed cyanobacteria, and their influence was direct through grazing rather than indirect on nutrient stoichiometry. Cyanobacterial abundance and heterocysts were low in treatments with no external P inputs. Concentrations of dissolved inorganic P comparable to those in Narragansett Bay were obtained only in P-fertilized mesocosms. Unlike previous estuarine mesocosm experiments with P fertilization, planktonic cyanobacteria grew and fixed N in our experimental systems. However, mean cell and heterocyst abundances under the most favorable conditions (high P, low N:P, and low grazers) were much lower than in comparable freshwater experiments, with N limitation maintained. These results support the hypothesis that intrinsic growth of heterocystous cyanobacteria in saline estuaries is slower than in freshwater, and that slower growth is unlikely to be due to systematic differences in P availability. Slow growth, combined with grazing, can severely limit development of planktonic, N-fixing cyanobacterial blooms in estuaries.
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ArticleCoupled biogeochemical cycles : eutrophication and hypoxia in temperate estuaries and coastal marine ecosystems(Ecological Society of America, 2011-02) Howarth, Robert W. ; Chan, Francis ; Conley, Daniel J. ; Garnier, Josette ; Doney, Scott C. ; Marino, Roxanne ; Billen, GillesNutrient fluxes to coastal areas have risen in recent decades, leading to widespread hypoxia and other ecological damage, particularly from nitrogen (N). Several factors make N more limiting in estuaries and coastal waters than in lakes: desorption (release) of phosphorus (P) bound to clay as salinity increases, lack of planktonic N fixation in most coastal ecosystems, and flux of relatively P-rich, N-poor waters from coastal oceans into estuaries. During eutrophication, biogeochemical feedbacks further increase the supply of N and P, but decrease availability of silica – conditions that can favor the formation and persistence of harmful algal blooms. Given sufficient N inputs, estuaries and coastal marine ecosystems can be driven to P limitation. This switch contributes to greater far-field N pollution; that is, the N moves further and contributes to eutrophication at greater distances. The physical oceanography (extent of stratification, residence time, and so forth) of coastal systems determines their sensitivity to hypoxia, and recent changes in physics have made some ecosystems more sensitive to hypoxia. Coastal hypoxia contributes to ocean acidification, which harms calcifying organisms such as mollusks and some crustaceans.
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ArticleRole of external inputs of nutrients to aquatic ecosystems in determining prevalence of nitrogen vs. phosphorus limitation of net primary productivity(Springer, 2021-02-17) Howarth, Robert W. ; Chan, Francis ; Swaney, Dennis P. ; Marino, Roxanne ; Hayn, MelanieWhether net primary productivity in an aquatic ecosystem is limited by nitrogen (N), limited by phosphorus (P), or co-limited by N & P is determined by the relative supply of N and P to phytoplankton compared to their elemental requirements for primary production, often characterized by the “Redfield” ratio. The supply of these essential nutrients is affected by both external inputs and biogeochemical processes within the ecosystem. In this paper, we examine external sources of nutrients to aquatic systems and how the balance of N to P inputs influences nutrient limitation. For ocean subtropical gyres, a relatively balanced input of N and P relative to the Redfield ratio from deep ocean sources often leads to near co-limitation by N and P. For lakes, the external nutrient inputs come largely from watershed sources, and we demonstrate that on average the N:P ratio for these inputs across the United States is well above that needed by phytoplankton, which may contribute to P limitation in those lake that experience this average nutrient loading. Watershed inputs are also important for estuaries and coastal marine ecosystems, but ocean sources of nutrients are also significant contributors to overall nutrient loads. The ocean-nutrient sources of N and P are very often at or below the Redfield ratio of 16:1 molar, and can be substantially so, particularly in areas where the continental shelf is wide. This large input of coastal ocean nutrients with a low N:P ratio is one factor that may make N limitation more likely in many coastal marine ecosystems than in lakes.