The role of nutricline depth in regulating the ocean carbon cycle.

Carbon uptake by marine phytoplankton, and its export as organic matter to the ocean interior (i.e., the "biological pump"), lowers the partial pressure of carbon dioxide (pCO(2)) in the upper ocean and facilitates the diffusive drawdown of atmospheric CO(2). Conversely, precipitation of calcium carbonate by marine planktonic calcifiers such as coccolithophorids increases pCO(2) and promotes its outgassing (i.e., the "alkalinity pump"). Over the past approximately 100 million years, these two carbon fluxes have been modulated by the relative abundance of diatoms and coccolithophores, resulting in biological feedback on atmospheric CO(2) and Earth's climate; yet, the processes determining the relative distribution of these two phytoplankton taxa remain poorly understood. We analyzed phytoplankton community composition in the Atlantic Ocean and show that the distribution of diatoms and coccolithophorids is correlated with the nutricline depth, a proxy of nutrient supply to the upper mixed layer of the ocean. Using this analysis in conjunction with a coupled atmosphere-ocean intermediate complexity model, we predict a dramatic reduction in the nutrient supply to the euphotic layer in the coming century as a result of increased thermal stratification. Our findings indicate that, by altering phytoplankton community composition, this causal relationship may lead to a decreased efficiency of the biological pump in sequestering atmospheric CO(2), implying a positive feedback in the climate system. These results provide a mechanistic basis for understanding the connection between upper ocean dynamics, the calcium carbonate-to-organic C production ratio and atmospheric pCO(2) variations on time scales ranging from seasonal cycles to geological transitions.

Global biodiversity patterns of marine phytoplankton and zooplankton.

Although the oceans cover 70% of the Earth's surface, our knowledge of biodiversity patterns in marine phytoplankton and zooplankton is very limited compared to that of the biodiversity of plants and herbivores in the terrestrial world. Here, we present biodiversity data for marine plankton assemblages from different areas of the world ocean. Similar to terrestrial vegetation, marine phytoplankton diversity is a unimodal function of phytoplankton biomass, with maximum diversity at intermediate levels of phytoplankton biomass and minimum diversity during massive blooms. Contrary to expectation, we did not find a relation between phytoplankton diversity and zooplankton diversity. Zooplankton diversity is a unimodal function of zooplankton biomass. Most strikingly, these marine biodiversity patterns show a worldwide consistency, despite obvious differences in environmental conditions of the various oceanographic regions. These findings may serve as a new benchmark in the search for global biodiversity patterns of plants and herbivores.

Invariant scaling of phytoplankton abundance and cell size in contrasting marine environments.

Scaling relationships such as the variation of population abundance with body size provide links between individual organisms and ecosystem functioning. Previous work, in marine pelagic ecosystems, has focused on the relationship between total phytoplankton abundance and the assemblage mean cell size. However, the relationship between specific population abundance and cell size in marine phytoplankton has received little attention. Here, we show that cell size accounts for a significant amount of variability in the population abundance of phytoplankton species across a cell volume range spanning seven orders of magnitude. The interspecific scaling of population abundance and cell size takes a power exponent near -3/4. Unexpectedly, despite the constraints imposed on large phytoplankton by limited resource acquisition, the size scaling exponent does not differ between contrasting marine environments such as coastal and subtropical regions. These findings highlight the adaptive abilities of individual species to cope with different environmental conditions and suggest that a general rule such as the 'energetic equivalence' constrains the abundance of phytoplankton populations in marine pelagic ecosystems.

Long-term oceanographic and ecological research in the Western English Channel.

Long-term research in the western English Channel, undertaken by the marine laboratories in Plymouth, is described and details of survey methods, sites, and time series given in this chapter. Major findings are summarized and their limitations outlined. Current research, with recent reestablishment and expansion of many sampling programmes, is presented, and possible future approaches are indicated. These unique long-term data sets provide an environmental baseline for predicting complex ecological responses to local, regional, and global environmental change. Between 1888 and the present, investigations have been carried out into the physical, chemical, and biological components (ranging from plankton and fish to benthic and intertidal assemblages) of the western English Channel ecosystem. The Marine Biological Association of the United Kingdom has performed the main body of these observations. More recent contributions come from the Continuous Plankton Recorder Survey, now the Sir Alister Hardy Foundation for Ocean Science, dating from 1957; the Institute for Marine Environmental Research, from 1974 to 1987; and the Plymouth Marine Laboratory, which was formed by amalgamation of the Institute for Marine Environmental Research and part of the Marine Biological Association, from 1988. Together, these contributions constitute a unique data series-one of the longest and most comprehensive samplings of environmental and marine biological variables in the world. Since the termination of many of these time series in 1987-1988 during a reorganisation of UK marine research, there has been a resurgence of interest in long-term environmental change. Many programmes have been restarted and expanded with support from several agencies. The observations span significant periods of warming (1921-1961; 1985-present) and cooling (1962-1980). During these periods of change, the abundance of key species underwent dramatic shifts. The first period of warming saw changes in zooplankton, pelagic fish, and larval fish, including the collapse of an important herring fishery. During later periods of change, shifts in species abundances have been reflected in other assemblages, such as the intertidal zone and the benthic fauna. Many of these changes appear to be related to climate, manifested as temperature changes, acting directly or indirectly. The hypothesis that climate is a forcing factor is widely supported today and has been reinforced by recent studies that show responses of marine organisms to climatic attributes such as the strength of the North Atlantic Oscillation. The long-term data also yield important insights into the effects of anthropogenic disturbances such as fisheries exploitation and pollution. Comparison of demersal fish hauls over time highlights fisheries effects not only on commercially important species but also on the entire demersal community. The effects of acute ("Torrey Canyon" oil spill) and chronic (tributyltin [TBT] antifoulants) pollution are clearly seen in the intertidal records. Significant advances in diverse scientific disciplines have been generated from research undertaken alongside the long-term data series. Many concepts in marine biological textbooks have originated in part from this work (e.g. the seasonal cycle of plankton, the cycling of nutrients, the pelagic food web trophic interactions, and the influence of hydrography on pelagic communities). Associated projects currently range from studies of marine viruses and bacterial ecology to zooplankton feeding dynamics and validation of ocean colour satellite sensors. Recent advances in technology mean these long-term programmes are more valuable than ever before. New technology collects data on finer temporal and spatial scales and can be used to capture processes that operate on multiple scales and help determine their influence in the marine environment. The MBA has been in the forefront of environmental modelling of shelf seas since the early 1970s. Future directions being pursued include the continued development of coupled physical-ecosystem models using western English Channel time-series data. These models will include both the recent high-resolution data and the long-term time-series information to predict effects of future climate change scenarios. It would be beneficial to provide more spatial and high-resolution temporal context to these data, which are fundamental for capturing processes that operate at multiple scales and understanding how they operate within the marine environment. This is being achieved through employment of technologies such as satellite-derived information and advanced telemetry instruments that provide real-time in situ profile data from the water column.

Scaling the metabolic balance of the oceans.

Oceanic communities are sources or sinks of CO2, depending on the balance between primary production and community respiration. The prediction of how global climate change will modify this metabolic balance of the oceans is limited by the lack of a comprehensive underlying theory. Here, we show that the balance between production and respiration is profoundly affected by environmental temperature. We extend the general metabolic theory of ecology to the production and respiration of oceanic communities and show that ecosystem rates can be reliably scaled from theoretical knowledge of organism physiology and measurement of population abundance. Our theory predicts that the differential temperature-dependence of respiration and photosynthesis at the organism level determines the response of the metabolic balance of the epipelagic ocean to changes in ambient temperature, a prediction that we support with empirical data over the global ocean. Furthermore, our model predicts that there will be a negative feedback of ocean communities to climate warming because they will capture less CO2 with a future increase in ocean temperature. This feedback of marine biota will further aggravate the anthropogenic effects on global warming.

Copepod hatching success in marine ecosystems with high diatom concentrations.

Diatoms dominate spring bloom phytoplankton assemblages in temperate waters and coastal upwelling regions of the global ocean. Copepods usually dominate the zooplankton in these regions and are the prey of many larval fish species. Recent laboratory studies suggest that diatoms may have a deleterious effect on the success of copepod egg hatching. These findings challenge the classical view of marine food-web energy flow from diatoms to fish by means of copepods. Egg mortality is an important factor in copepod population dynamics, thus, if diatoms have a deleterious in situ effect, paradoxically, high diatom abundance could limit secondary production. Therefore, the current understanding of energy transfer from primary production to fisheries in some of the most productive and economically important marine ecosystems may be seriously flawed. Here we present in situ estimates of copepod egg hatching success from twelve globally distributed areas, where diatoms dominate the phytoplankton assemblage. We did not observe a negative relationship between copepod egg hatching success and either diatom biomass or dominance in the microplankton in any of these regions. The classical model for diatom-dominated system remains valid.