Modeling environmental effects on the size-structured energy flow through marine ecosystems. Part 1: The model

This paper presents an original size-structured mathematical model of the energy flow through marine ecosystems, based on established ecological and physiological processes and mass conservation principles. The model is based on a nonlocal partial differential equation which represents the transfer of energy in both time and body weight (size) in marine ecosystems. The processes taken into account include size-based opportunistic trophic interactions, competition for food, allocation of energy between growth and reproduction, somatic and maturity maintenance, predatory and starvation mortality. All the physiological rates are temperature-dependent. The physiological bases of the model are derived from the dynamic energy budget theory. The model outputs the dynamic size-spectrum of marine ecosystems in term of energy content per weight class as well as many other size-dependent diagnostic variables such as growth rate, egg production or predation mortality.In stable environmental conditions and using a reference set of parameters derived from empirical studies, the model converges toward a stationary linear log–log size-spectrum with a slope equal to −1.06, which is consistent with the values reported in empirical studies. In some cases, the distribution of the largest sizes departs from the stationary linear solution and is slightly curved downward. A sensitivity analysis to the parameters is conducted systematically. It shows that the stationary size-spectrum is not very sensitive to the parameters of the model. Numerical simulations of the effects of temperature and primary production variability on marine ecosystems size-spectra are provided in a companion paper [Maury, O., Shin, Y.-J., Faugeras, B., Ben Ari, T., Marsac, F., 2007. Modeling environmental effects on the size-structured energy flow through marine ecosystems. Part 2: simulations. Progress in Oceanography, doi:10.1016/j.pocean.2007.05.001].     

Modeling environmental effects on the size-structured energy flow through marine ecosystems. Part 2: Simulations

Numerical simulations using a physiologically-based model of marine ecosystem size spectrum are conducted to study the influence of primary production and temperature on energy flux through marine ecosystems. In stable environmental conditions, the model converges toward a stationary linear log–log size-spectrum. In very productive ecosystems, the model predicts that small size classes are depleted by predation, leading to a curved size-spectrum.It is shown that the absolute level of primary production does not affect the slope of the stationary size-spectrum but has a nonlinear effect on its intercept and hence on the total biomass of consumer organisms (the carrying capacity). Three domains are distinguished: at low primary production, total biomass is independent from production changes because loss processes dominate dissipative processes (biological work); at high production, ecosystem biomass is proportional to primary production because dissipation dominates losses; an intermediate transition domain characterizes mid-production ecosystems. Our results enlighten the paradox of the very high ecosystem biomass/primary production ratios which are observed in poor oceanic regions. Thus, maximal dissipation (least action and low ecosystem biomass/primary production ratios) is reached at high primary production levels when the ecosystem is efficient in transferring energy from small sizes to large sizes. Conversely, least dissipation (most action and high ecosystem biomass/primary production ratios) characterizes the simulated ecosystem at low primary production levels when it is not efficient in dissipating energy.Increasing temperature causes enhanced predation mortality and decreases the intercept of the stationary size spectrum, i.e., the total ecosystem biomass. Total biomass varies as the inverse of the Arrhenius coefficient in the loss domain. This approximation is no longer true in the dissipation domain where nonlinear dissipation processes dominate over linear loss processes. Our results suggest that in a global warming context, at constant primary production, a 2–4 °C warming would lead to a 20–43% decrease of ecosystem biomass in oligotrophic regions and to a 15–32% decrease of biomass in eutrophic regions.Oscillations of primary production or temperature induce waves which propagate along the size-spectrum and which amplify until a “resonant range” which depends on the period of the environmental oscillations. Small organisms oscillate in phase with producers and are bottom-up controlled by primary production oscillations. In the “resonant range”, prey and predators oscillate out of phase with alternating periods of top-down and bottom-up controls. Large organisms are not influenced by bottom-up effects of high frequency phytoplankton variability or by oscillations of temperature.     

不同温度和盐度对大竹蛏耗氧率和排氨率的影响

在实验室条件下,设置12、17、22、27和32℃五个温度梯度和12、17、22、27和32五个盐度梯度,研究了不同温度和盐度对大竹蛏(Solen grandis)耗氧排氨的影响。结果表明:随着温度的升高耗氧率先降低再逐渐升高,在温度为22℃时为最小值1.571 mg/(g·h);排氨率先降低后升高再降低,在温度17℃时为最小值0.127 mg/(g·h)。随着盐度增加耗氧率先升高后降低,在盐度22时达到最大值3.106 mg/(g·h),排氨率先降低再升高,在盐度为27时为最小值为0.145 mg/(g·h)。在盐度32时,不同温度的O:N范围是5.525~79.037。在温度12℃时,不同盐度下的O:N范围是4.378~16.755。不同温度下耗氧率Q_(10)变化范围为1.054~2.421,排氨率的Q_(10)变化范围是0.748~1.256。研究结果为进一步完善大竹蛏的人工养殖技术提供参考。     

Investigating the effect of ballasting by CaCO3 in Emiliania huxleyi: I. Formation, settling velocities and physical properties of aggregates

To investigate the role of ballasting by biogenic minerals in the export of organic matter in the ocean, a laboratory experiment was conducted comparing aggregate formation and settling velocity of non-calcifying and calcifying strains of the coccolithophore Emiliania huxleyi. Experiments were conducted by making aggregates using a roller table and following aggregate properties during incubation for a period of 40 days. Size, shape, and settling velocities of aggregates were described by image analysis of video pictures recorded during the roller tank incubation. Our results show that biogenic calcite has a strong effect on the formation rate and abundance of aggregates and on aggregate properties such as size, excess density, porosity, and settling velocity. Aggregates of calcifying cells (AGG_CAL) formed faster, were smaller and had higher settling velocities, excess densities, and mass than those of non-calcifying cells (AGG_NCAL). AGG_CAL showed no loss during the duration of the experiment, whereas AGG_NCAL decreased in size after 1 month of incubation. Potential mechanisms that can explain the different patterns in aggregate formation are discussed. Comparison of settling velocities of AGG_CAL and AGG_NCAL with aggregates formed by diatoms furthermore indicated that the ballast effect of calcite is greater than that of opal. Together these results help to better understand why calcite is of major importance for organic matter fluxes to the deep ocean.