Louis Legendre

Marine biodiversity, ecosystem functioning, and carbon cycles

Although recent studies suggest that climate change may substantially accelerate the rate of species loss in the biosphere, only a few studies have focused on the potential consequences of a spatial reorganization of biodiversity with global warming. Here, we show a pronounced latitudinal increase in phytoplanktonic and zooplanktonic biodiversity in the extratropical North Atlantic Ocean in recent decades. We also show that this rise in biodiversity paralleled a decrease in the mean size of zooplanktonic copepods and that the reorganization of the planktonic ecosystem toward dominance by smaller organisms may influence the networks in which carbon flows, with negative effects on the downward biological carbon pump and demersal Atlantic cod (Gadus morhua). Our study suggests that, contrary to the usual interpretation of increasing biodiversity being a positive emergent property promoting the stability/resilience of ecosystems, the parallel decrease in sizes of planktonic organisms could be viewed in the North Atlantic as reducing some of the services provided by marine ecosystems to humans.

Global latitudinal variations in marine copepod diversity and environmental factors

Latitudinal gradients in diversity are among the most striking features in ecology. For terrestrial species, climate (i.e. temperature and precipitation) is believed to exert a strong influence on the geographical distributions of diversity through its effects on energy availability. Here, we provide the first global description of geographical variation in the diversity of marine copepods, a key trophic link between phytoplankton and fish, in relation to environmental variables. We found a polar–tropical difference in copepod diversity in the Northern Hemisphere where diversity peaked at subtropical latitudes. In the Southern Hemisphere, diversity showed a tropical plateau into the temperate regions. This asymmetry around the Equator may be explained by climatic conditions, in particular the influence of the Inter-Tropical Convergence Zone, prevailing mainly in the northern tropical region. Ocean temperature was the most important explanatory factor among all environmental variables tested, accounting for 54 per cent of the variation in diversity. Given the strong positive correlation between diversity and temperature, local copepod diversity, especially in extra-tropical regions, is likely to increase with climate change as their large-scale distributions respond to climate warming.

Autotrophic carbon budget in coral tissue: a new 13C-based model of photosynthate translocation

SUMMARY Corals live in symbiosis with dinoflagellates of the genus Symbiodinum. These dinoflagellates translocate a large part of the photosynthetically fixed carbon to the host, which in turn uses it for its own needs. Assessing the carbon budget in coral tissue is a central question in reef studies that still vexes ecophysiologists. The amount of carbon fixed by the symbiotic association can be determined by measuring the rate of photosynthesis, but the amount of carbon translocated by the symbionts to the host and the fate of this carbon are more difficult to assess. In the present study, we propose a novel approach to calculate the budget of autotrophic carbon in the tissue of scleractinian corals, based on a new model and measurements made with the stable isotope 13C. Colonies of the scleractinian coral Stylophora pistillata were incubated in H13CO –3-enriched seawater, after which the fate of 13C was followed in the symbionts, the coral tissue and the released particulate organic carbon (i.e. mucus). Results obtained showed that after 15 min, ca. 60% of the carbon fixed was already translocated to the host, and after 48 h, this value reached 78%. However, ca. 48% of the photosynthetically fixed carbon was respired by the symbiotic association, and 28% was released as dissolved organic carbon. This is different from other coral species, where <1% of the total organic carbon released is from newly fixed carbon. Only 23% of the initially fixed carbon was retained in the symbionts and coral tissue after 48 h. Results show that our 13C-based model could successfully trace the carbon flow from the symbionts to the host, and the photosynthetically acquired carbon lost from the symbiotic association.

A new look at ocean carbon remineralization for estimating deepwater sequestration

The “biological carbon pump” causes carbon sequestration in deep waters by downward transfer of organic matter, mostly as particles. This mechanism depends to a great extent on the uptake of CO2 by marine plankton in surface waters and subsequent sinking of particulate organic carbon (POC) through the water column. Most of the sinking POC is remineralized during its downward transit, and modest changes in remineralization have substantial feedback on atmospheric CO2 concentrations, but little is known about global variability in remineralization. Here we assess this variability based on modern underwater particle imaging combined with field POC flux data and discuss the potential sources of variations. We show a significant relationship between remineralization and the size structure of the phytoplankton assemblage. We obtain the first regionalized estimates of remineralization in biogeochemical provinces, where these estimates range between −50 and +100% of the commonly used globally uniform remineralization value. We apply the regionalized values to satellite-derived estimates of upper ocean POC export to calculate regionalized and ocean-wide deep carbon fluxes and sequestration. The resulting value of global organic carbon sequestration at 2000u2009m is 0.33u2009Pgu2009Cu2009yr−1, and 0.72u2009Pgu2009Cu2009yr−1 at the depth of the top of the permanent pycnocline, which is up to 3 times higher than the value resulting from the commonly used approach based on uniform remineralization and constant sequestration depth. These results stress that variable remineralization and sequestration depth should be used to model ocean carbon sequestration and feedback on the atmosphere.

Effects of soot deposition on particle dynamics and microbial processes in marine surface waters

Large amounts of soot are continuously deposited on the global ocean. Even though significant concentrations of soot particles are found in marine waters, the effects of these aerosols on ocean ecosystems are currently unknown. Using a combination of in situ and experimental data, and results from an atmospheric transport model, we show that the deposition of soot particles from an oil-fired power plant impacted biogeochemical properties and the functioning of the pelagic ecosystem in tropical oligotrophic oceanic waters off New Caledonia. Deposition was followed by a major increase in the volume concentration of suspended particles, a change in the particle size spectra that resulted from a stimulation of aggregation processes, a 5% decrease in the concentration of dissolved organic carbon (DOC), a decreases of 33 and 23% in viral and free bacterial abundances, respectively, and a factor ~2 increase in the activity of particle-attached bacteria suggesting that soot introduced in the system favored bacterial growth. These patterns were confirmed by experiments with natural seawater conducted with both soot aerosols collected in the study area and standard diesel soot. The data suggest a strong impact of soot deposition on ocean surface particles, DOC, and microbial processes, at least near emission hot spots.

Controlling Effects of Irradiance and Heterotrophy on Carbon Translocation in the Temperate Coral Cladocora caespitosa

Temperate symbiotic corals, such as the Mediterranean species Cladocora caespitosa, live in seasonally changing environments, where irradiance can be ten times higher in summer than winter. These corals shift from autotrophy in summer to heterotrophy in winter in response to light limitation of the symbiont’s photosynthesis. In this study, we determined the autotrophic carbon budget under different conditions of irradiance (20 and 120 µmol photons m−2 s−1) and feeding (fed three times a week with Artemia salina nauplii, and unfed). Corals were incubated in H13CO3 −-enriched seawater, and the fate of 13C was followed in the symbionts and the host tissue. The total amount of carbon fixed by photosynthesis and translocated was significantly higher at high than low irradiance (ca. 13 versus 2.5–4.5 µg cm−2 h−1), because the rates of photosynthesis and carbon fixation were also higher. However, the percent of carbon translocation was similar under the two irradiances, and reached more than 70% of the total fixed carbon. Host feeding induced a decrease in the percentage of carbon translocated under low irradiance (from 70 to 53%), and also a decrease in the rates of carbon translocation per symbiont cell under both irradiances. The fate of autotrophic and heterotrophic carbon differed according to irradiance. At low irradiance, autotrophic carbon was mostly respired by the host and the symbionts, and heterotrophic feeding led to an increase in host biomass. Under high irradiance, autotrophic carbon was both respired and released as particulate and dissolved organic carbon, and heterotrophic feeding led to an increase in host biomass and symbiont concentration. Overall, the maintenance of high symbiont concentration and high percentage of carbon translocation under low irradiance allow this coral species to optimize its autotrophic carbon acquisition, when irradiance conditions are not favourable to photosynthesis.

Estimates of Water-Column Nutrient Concentrations and Carbonate System Parameters in the Global Ocean: A Novel Approach Based on Neural Networks

A neural network-based method (CANYON: CArbonate system and Nutrients concentration from hYdrological properties and Oxygen using a Neural-network) was developed to estimate water-column biogeochemically relevant variables in the Global Ocean. These are the concentrations of 3 nutrients [nitrate (NO3−), phosphate (PO43−) and silicate (Si(OH)4)] and 4 carbonate system parameters [total alkalinity (AT), dissolved inorganic carbon (CT), pH (pHT) and partial pressure of CO2 (pCO2)], which are estimated from concurrent in situ measurements of temperature, salinity, hydrostatic pressure and oxygen (O2) together with sampling latitude, longitude and date. Seven neural-networks were developed using the GLODAPv2 database, which is largely representative of the diversity of open-ocean conditions, hence making CANYON potentially applicable to most oceanic environments. For each variable, CANYON was trained using 80 % randomly chosen data from the whole database (after eight 10° x 10° zones removed providing an “independent data-set” for additional validation), the remaining 20 % data were used for the neural-network test of validation. Overall, CANYON retrieved the variables with high accuracies (RMSE): 0.93 uf06dmol kg-1 (NO3−), 0.07 uf06dmol kg-1 (PO43-), 3.0 uf06dmol kg-1 (Si(OH)4), 0.019 (pHT), 7 uf06dmol kg-1 (AT), 10 uf06dmol kg-1 (CT) and 28 uf06datm (pCO2). This was confirmed for the 8 independent zones not included in the training process. CANYON was also applied to the Hawaiian Time Series site to produce a 22-years long simulated time series for the above 7 variables. Comparison of modeled and measured data was also very satisfactory (RMSE in the order of magnitude of RMSE from validation test). CANYON is thus a promising method to derive distributions of key biogeochemical variables. It could be used for a variety of global and regional applications ranging from data quality control to the production of datasets of variables required for initialization and validation of biogeochemical models but difficult to obtain. In particular, combining the increased coverage of the global Biogeochemical-Argo program, where O2 is one of the core variables now very accurately measured, with the CANYON approach offers the fascinating perspective of obtaining large-scale estimates of key biogeochemical variables with unprecedented spatial and temporal resolutions.

Comprehensive Model of Annual Plankton Succession Based on the Whole-Plankton Time Series Approach

Ecological succession provides a widely accepted description of seasonal changes in phytoplankton and mesozooplankton assemblages in the natural environment, but concurrent changes in smaller (i.e. microbes) and larger (i.e. macroplankton) organisms are not included in the model because plankton ranging from bacteria to jellies are seldom sampled and analyzed simultaneously. Here we studied, for the first time in the aquatic literature, the succession of marine plankton in the whole-plankton assemblage that spanned 5 orders of magnitude in size from microbes to macroplankton predators (not including fish or fish larvae, for which no consistent data were available). Samples were collected in the northwestern Mediterranean Sea (Bay of Villefranche) weekly during 10 months. Simultaneously collected samples were analyzed by flow cytometry, inverse microscopy, FlowCam, and ZooScan. The whole-plankton assemblage underwent sharp reorganizations that corresponded to bottom-up events of vertical mixing in the water-column, and its development was top-down controlled by large gelatinous filter feeders and predators. Based on the results provided by our novel whole-plankton assemblage approach, we propose a new comprehensive conceptual model of the annual plankton succession (i.e. whole plankton model) characterized by both stepwise stacking of four broad trophic communities from early spring through summer, which is a new concept, and progressive replacement of ecological plankton categories within the different trophic communities, as recognised traditionally.

Marine copepod diversity patterns and the metabolic theory of ecology.

Temperature is a powerful correlate of large-scale terrestrial and marine diversity patterns but the mechanistic links remain unclear. Whilst many explanations have been proposed, quantitative predictions that allow them to be tested statistically are often lacking. As an important exception, the metabolic theory of ecology (MTE) provides a rather robust technique using the relationship between diversity, temperature and metabolic rate in order to elucidate the ultimate underlying mechanisms driving large-scale diversity patterns. We tested if the MTE could explain geographic variations in marine copepod diversity on both ocean-wide and regional scales (East Japan Sea and North East Atlantic). The values of the regression slopes of diversity (ln taxonomic richness) over temperature (1/kT) across all spatial scales were lower than the range predicted by the metabolic scaling law for species richness (i.e. −0.60 to −0.70).We therefore conclude that the MTE in its present form is not suitable for predicting marine copepod diversity patterns. These results further question the applicability of the MTE for explaining diversity patterns and, despite the relative lack of comparable studies in the marine environment, the generality of the MTE across systems.

Water Column Biogeochemistry below the Euphotic Zone

The main focus of the International JGOFS research inititiatives was on the cycling of carbon and of associated elements within the surface layer, and their downward export from the upper ocean. Relatively few coordinated measurements and experiments were made below the photic zone so our understanding and modeling of the biogeochemistry of the ocean’s interior is still in its infancy. However from the numerous data acquired in the 1990s during JGOFS and JGOFS-like process studies it is possible to extract sufficient information to make preliminary statements about the biogeochemistry of the water column below the euphotic zone. An important preliminary result of these studies is that we now are beginning to realize that the biogeochemistry of the surface ocean, of the ocean’s interior, and of the surface sediments appears to be more coupled than was thought fifteen years ago.