Surveys across the world oceans have shown that phytoplankton biomass and production are dominated by small cells (picoplankton) where nutrient concentrations are low, but large cells (microplankton) ...dominate when nutrient-rich deep water is mixed to the surface. I analyzed phytoplankton size structure in samples collected over 25 yr in San Francisco Bay, a nutrient-rich estuary. Biomass was dominated by large cells because their biomass selectively grew during blooms. Large-cell dominance appears to be a characteristic of ecosystems at the land–sea interface, and these places may therefore function as analogs to oceanic upwelling systems. Simulations with a size-structured NPZ model showed that runs of positive net growth rate persisted long enough for biomass of large, but not small, cells to accumulate. Model experiments showed that small cells would dominate in the absence of grazing, at lower nutrient concentrations, and at elevated (+5°C) temperatures. Underlying these results are two fundamental scaling laws: (1) large cells are grazed more slowly than small cells, and (2) grazing rate increases with temperature faster than growth rate. The model experiments suggest testable hypotheses about phytoplankton size structure at the land–sea interface: (1) anthropogenic nutrient enrichment increases cell size; (2) this response varies with temperature and only occurs at mid-high latitudes; (3) large-cell blooms can only develop when temperature is below a critical value, around 15°C; (4) cell size diminishes along temperature gradients from high to low latitudes; and (5) large-cell blooms will diminish or disappear where planetary warming increases temperature beyond their critical threshold.
Phytoplankton blooms are dynamic phenomena of great importance to the functioning of estuarine and coastal ecosystems. We analysed a unique (large) collection of phytoplankton monitoring data ...covering 86 coastal sites distributed over eight regions in North America and Europe, with the aim of investigating common patterns in the seasonal timing and species composition of the blooms. The spring bloom was the most common seasonal pattern across all regions, typically occurring early (February–March) at lower latitudes and later (April–May) at higher latitudes. Bloom frequency, defined as the probability of unusually high biomass, ranged from 5 to 35% between sites and followed no consistent patterns across gradients of latitude, temperature, salinity, water depth, stratification, tidal amplitude or nutrient concentrations. Blooms were mostly dominated by a single species, typically diatoms (58% of the blooms) and dinoflagellates (19%). Diatom-dominated spring blooms were a common feature in most systems, although dinoflagellate spring blooms were also observed in the Baltic Sea. Blooms dominated by chlorophytes and cyanobacteria were only common in low salinity waters and occurred mostly at higher temperatures. Key bloom species across the eight regions included the diatoms Cerataulina pelagica and Dactyliosolen fragilissimus and dinoflagellates Heterocapsa triquetra and Prorocentrum cordatum. Other frequent bloom-forming taxa were diatom genera Chaetoceros, Coscinodiscus, Skeletonema, and Thalassiosira. Our meta-analysis shows that these 86 estuarine-coastal sites function as diatom-producing systems, the timing of that production varies widely, and that bloom frequency is not associated with environmental factors measured in monitoring programs. We end with a perspective on the limitations of conclusions derived from meta-analyses of phytoplankton time series, and the grand challenges remaining to understand the wide range of bloom patterns and processes that select species as bloom dominants in coastal waters.
The annual cycles of phytoplankton biomass Winder, Monika; Cloern, James E.
Philosophical transactions - Royal Society. Biological sciences,
10/2010, Letnik:
365, Številka:
1555
Journal Article
Recenzirano
Odprti dostop
Terrestrial plants are powerful climate sentinels because their annual cycles of growth, reproduction and senescence are finely tuned to the annual climate cycle having a period of one year. ...Consistency in the seasonal phasing of terrestrial plant activity provides a relatively low-noise background from which phenological shifts can be detected and attributed to climate change. Here, we ask whether phytoplankton biomass also fluctuates over a consistent annual cycle in lake, estuarine–coastal and ocean ecosystems and whether there is a characteristic phenology of phytoplankton as a consistent phase and amplitude of variability. We compiled 125 time series of phytoplankton biomass (chlorophyll a concentration) from temperate and subtropical zones and used wavelet analysis to extract their dominant periods of variability and the recurrence strength at those periods. Fewer than half (48%) of the series had a dominant 12-month period of variability, commonly expressed as the canonical spring-bloom pattern. About 20 per cent had a dominant six-month period of variability, commonly expressed as the spring and autumn or winter and summer blooms of temperate lakes and oceans. These annual patterns varied in recurrence strength across sites, and did not persist over the full series duration at some sites. About a third of the series had no component of variability at either the six- or 12-month period, reflecting a series of irregular pulses of biomass. These findings show that there is high variability of annual phytoplankton cycles across ecosystems, and that climate-driven annual cycles can be obscured by other drivers of population variability, including human disturbance, aperiodic weather events and strong trophic coupling between phytoplankton and their consumers. Regulation of phytoplankton biomass by multiple processes operating at multiple time scales adds complexity to the challenge of detecting climate-driven trends in aquatic ecosystems where the noise to signal ratio is high.
Time series of environmental measurements are essential for detecting, measuring and understanding changes in the Earth system and its biological communities. Observational series have accumulated ...over the past 2–5 decades from measurements across the world's estuaries, bays, lagoons, inland seas and shelf waters influenced by runoff. We synthesize information contained in these time series to develop a global view of changes occurring in marine systems influenced by connectivity to land. Our review is organized around four themes: (i) human activities as drivers of change; (ii) variability of the climate system as a driver of change; (iii) successes, disappointments and challenges of managing change at the sea‐land interface; and (iv) discoveries made from observations over time. Multidecadal time series reveal that many of the world's estuarine–coastal ecosystems are in a continuing state of change, and the pace of change is faster than we could have imagined a decade ago. Some have been transformed into novel ecosystems with habitats, biogeochemistry and biological communities outside the natural range of variability. Change takes many forms including linear and nonlinear trends, abrupt state changes and oscillations. The challenge of managing change is daunting in the coastal zone where diverse human pressures are concentrated and intersect with different responses to climate variability over land and over ocean basins. The pace of change in estuarine–coastal ecosystems will likely accelerate as the human population and economies continue to grow and as global climate change accelerates. Wise stewardship of the resources upon which we depend is critically dependent upon a continuing flow of information from observations to measure, understand and anticipate future changes along the world's coastlines.
Poised at the interface of rivers, ocean, atmosphere and dense human settlement, estuaries are driven by a large array of natural and anthropogenic forces. San Francisco Bay exemplifies the ...fast‐paced change occurring in many of the world's estuaries, bays, and inland seas in response to these diverse forces. We use observations from this particularly well‐studied estuary to illustrate responses to six drivers that are common agents of change where land and sea meet: water consumption and diversion, human modification of sediment supply, introduction of nonnative species, sewage input, environmental policy, and climate shifts. In San Francisco Bay, responses to these drivers include, respectively, shifts in the timing and extent of freshwater inflow and salinity intrusion, decreasing turbidity, restructuring of plankton communities, nutrient enrichment, elimination of hypoxia and reduced metal contamination of biota, and food web changes that decrease resistance of the estuary to nutrient pollution. Detection of these changes and discovery of their causes through environmental monitoring have been essential for establishing and measuring outcomes of environmental policies that aim to maintain high water quality and sustain services provided by estuarine‐coastal ecosystems. The many time scales of variability and the multiplicity of interacting drivers place heavy demands on estuarine monitoring programs, but the San Francisco Bay case study illustrates why the imperative for monitoring has never been greater.
Key Points
Estuarine‐coastal ecosystems are changing at a breathtaking pace
Four decades of study in San Francisco Bay reveal the drivers of those changes
The imperative for estuarine‐coastal monitoring has never been greater
Estuaries are connected to both land and ocean so their physical, chemical, and biological dynamics are influenced by climate patterns over watersheds and ocean basins. We explored climate‐driven ...oceanic variability as a source of estuarine variability by comparing monthly time series of temperature and chlorophyll‐a inside San Francisco Bay with those in adjacent shelf waters of the California Current System (CCS) that are strongly responsive to wind‐driven upwelling. Monthly temperature fluctuations inside and outside the Bay were synchronous, but their correlations weakened with distance from the ocean. These results illustrate how variability of coastal water temperature (and associated properties such as nitrate and oxygen) propagates into estuaries through fast water exchanges that dissipate along the estuary. Unexpectedly, there was no correlation between monthly chlorophyll‐a variability inside and outside the Bay. However, at the annual scale Bay chlorophyll‐a was significantly correlated with the Spring Transition Index (STI) that sets biological production supporting fish recruitment in the CCS. Wind forcing of the CCS shifted in the late 1990s when the STI advanced 40 days. This shift was followed, with lags of 1–3 years, by 3‐ to 19‐fold increased abundances of five ocean‐produced demersal fish and crustaceans and 2.5‐fold increase of summer chlorophyll‐a in the Bay. These changes reflect a slow biological process of estuary–ocean connectivity operating through the immigration of fish and crustaceans that prey on bivalves, reduce their grazing pressure, and allow phytoplankton biomass to build. We identified clear signals of climate‐mediated oceanic variability in this estuary and discovered that the response patterns vary with the process of connectivity and the timescale of ocean variability. This result has important implications for managing nutrient inputs to estuaries connected to upwelling systems, and for assessing their responses to changing patterns of upwelling timing and intensity as the planet continues to warm.
We explored climate‐driven oceanic variability as a source of estuarine variability by comparing monthly temperature and chlorophyll‐a inside San Francisco Bay with those in adjacent shelf waters. We identified clear signals of climate‐mediated oceanic variability in this estuary that depended on the process of connectivity and the time scale of ocean variability. This result has important implications for managing nutrient inputs to estuaries and for assessing their responses to climate change.
The salinity gradient of estuaries plays a unique and fundamental role in structuring spatial patterns of physical properties, biota, and biogeochemical processes. We use variability along the ...salinity gradient of San Francisco Bay to illustrate some lessons about the diversity of spatial structures in estuaries and their variability over time. Spatial patterns of dissolved constituents (e.g., silicate) can be linear or nonlinear, depending on the relative importance of river-ocean mixing and internal sinks (diatom uptake). Particles have different spatial patterns because they accumulate in estuarine turbidity maxima formed by the combination of sinking and estuarine circulation. Some constituents have weak or no mean spatial structure along the salinity gradient, reflecting spatially distributed sources along the estuary (nitrate) or atmospheric exchanges that buffer spatial variability of ecosystem metabolism (dissolved oxygen). The density difference between freshwater and seawater establishes stratification in estuaries stronger than the thermal stratification of lakes and oceans. Stratification is strongest around the center of the salinity gradient and when river discharge is high. Spatial distributions of motile organisms are shaped by species-specific adaptations to different salinity ranges (shrimp) and by behavioral responses to environmental variability (northern anchovy). Estuarine spatial patterns change over time scales of events (intrusions of upwelled ocean water), seasons (river inflow), years (annual weather anomalies), and between eras separated by ecosystem disturbances (a species introduction). Each of these lessons is a piece in the puzzle of how estuarine ecosystems are structured and how they differ from the river and ocean ecosystems they bridge.
Accumulating evidence shows that the planet is warming as a response to human emissions of greenhouse gases. Strategies of adaptation to climate change will require quantitative projections of how ...altered regional patterns of temperature, precipitation and sea level could cascade to provoke local impacts such as modified water supplies, increasing risks of coastal flooding, and growing challenges to sustainability of native species.
We linked a series of models to investigate responses of California's San Francisco Estuary-Watershed (SFEW) system to two contrasting scenarios of climate change. Model outputs for scenarios of fast and moderate warming are presented as 2010-2099 projections of nine indicators of changing climate, hydrology and habitat quality. Trends of these indicators measure rates of: increasing air and water temperatures, salinity and sea level; decreasing precipitation, runoff, snowmelt contribution to runoff, and suspended sediment concentrations; and increasing frequency of extreme environmental conditions such as water temperatures and sea level beyond the ranges of historical observations.
Most of these environmental indicators change substantially over the 21(st) century, and many would present challenges to natural and managed systems. Adaptations to these changes will require flexible planning to cope with growing risks to humans and the challenges of meeting demands for fresh water and sustaining native biota. Programs of ecosystem rehabilitation and biodiversity conservation in coastal landscapes will be most likely to meet their objectives if they are designed from considerations that include: (1) an integrated perspective that river-estuary systems are influenced by effects of climate change operating on both watersheds and oceans; (2) varying sensitivity among environmental indicators to the uncertainty of future climates; (3) inevitability of biological community changes as responses to cumulative effects of climate change and other drivers of habitat transformations; and (4) anticipation and adaptation to the growing probability of ecosystem regime shifts.
Nutrient enrichment has degraded many of the world’s estuaries by amplifying algal production, leading to hypoxia/anoxia, loss of vascular plants and fish/shellfish habitat, and expansion of harmful ...blooms (HABs). Policies to protect coastal waters from the effects of nutrient enrichment require information to determine if a water body is impaired by nutrients and if regulatory actions are required. We compiled information to inform these decisions for San Francisco Bay (SFB), an urban estuary where the best path toward nutrient management is not yet clear. Our results show that SFB has high nutrient loadings, primarily from municipal wastewater; there is potential for high algal production, but that production is not fully realized; and SFB is not impaired by hypoxia or recurrent HABs. However, our assessment includes reasons for concern: nitrogen and phosphorus concentrations higher than those in other estuaries impaired by nutrient pollution, chronic presences of multiple algal toxins, a recent increase of primary production, and projected future hydroclimatic conditions that could increase the magnitude and frequency of algal blooms. Policymakers thus face the challenge of determining the appropriate protective policy for SFB. We identify three crucial next steps for meeting this challenge: (1) new research to determine if algal toxins can be reduced through nutrient management, (2) establish management goals as numeric targets, and (3) determine the magnitude of nutrient load reduction required to meet those targets. Our case study illustrates how scientific information can be acquired and communicated to inform policymakers about the status of nutrient pollution, its risks, and strategies for minimizing those risks.
A primary focus of coastal science during the past 3 decades has been the question: How does anthropogenic nutrient enrichment cause change in the structure or function of nearshore coastal ...ecosystems? This theme of environmental science is recent, so our conceptual model of the coastal eutrophication problem continues to change rapidly. In this review, I suggest that the early (Phase I) conceptual model was strongly influenced by limnologists, who began intense study of lake eutrophication by the 1960s. The Phase I model emphasized changing nutrient input as a signal, and responses to that signal as increased phytoplankton biomass and primary production, decomposition of phytoplankton-derived organic matter, and enhanced depletion of oxygen from bottom waters. Coastal research in recent decades has identified key differences in the responses of lakes and coastal-estuarine ecosystems to nutrient enrichment. The contemporary (Phase II) conceptual model reflects those differences and includes explicit recognition of (1) system-specific attributes that act as a filter to modulate the responses to enrichment (leading to large differences among estuarine-coastal systems in their sensitivity to nutrient enrichment); and (2) a complex suite of direct and indirect responses including linked changes in: water transparency, distribution of vascular plants and biomass of macroalgae, sediment biogeochemistry and nutrient cycling, nutrient ratios and their regulation of phytoplankton community composition, frequency of toxic/harmful algal blooms, habitat quality for metazoans, reproduction/growth/survival of pelagic and benthic invertebrates, and subtle changes such as shifts in the seasonality of ecosystem functions. Each aspect of the Phase II model is illustrated here with examples from coastal ecosystems around the world. In the last section of this review I present one vision of the next (Phase III) stage in the evolution of our conceptual model, organized around 5 questions that will guide coastal science in the early 21st century: (1) How do system-specific attributes constrain or amplify the responses of coastal ecosystems to nutrient enrichment? (2) How does nutrient enrichment interact with other stressors (toxic contaminants, fishing harvest, aquaculture, nonindigenous species, habitat loss, climate change, hydrologic manipulations) to change coastal ecosystems? (3) How are responses to multiple stressors linked? (4) How does human-induced change in the coastal zone impact the Earth system as habitat for humanity and other species? (5) How can a deeper scientific understanding of the coastal eutrophication problem be applied to develop tools for building strategies at ecosystem restoration or rehabilitation?
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