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 CO 2 with a future increase in ocean temperature. This feedback of marine biota will further aggravate the anthropogenic effects on global warming.global change ͉ metabolic theory ͉ oceanic carbon cycle T he role of the oceans in the CO 2 budget of the biosphere depends largely on the balance between the uptake of carbon by phytoplankton photosynthesis and its remineralization by the respiration of the whole planktonic community (1). For large areas of the epipelagic ocean, planktonic community respiration (CR) exceeds gross primary production (GPP), resulting in net heterotrophy and a source of CO 2 (2-4). The solution of the contentious debate over the extent of such heterotrophic areas (5-7) is hindered by the limited spatiotemporal coverage achievable by traditional incubation methods (8, 9). Here, we tackle this question from a different perspective based on the metabolic theory of ecology (MTE) (10). The flux rates within an ecosystem are the result of the sum of the individual rates of all its constituent organisms (11,12), which, in turn, are governed by the combined effects of body size and temperature (13-15). Although MTE suggests a universal scaling of metabolic rate as the 3͞4 power of body size, it predicts a differential temperaturedependence of heterotrophic processes (driven by ATP synthesis) and autotrophic rates (controlled by Rubisco carboxylation) (12). Following the MTE, the respiration of a heterotrophic planktonic organism B i can be estimated if we know its body size M i and the ambient absolute temperature T:where b 0 is a normalization constant independent of body size and temperature, e ϪEh͞kT is Boltzmann's factor, where E h is the average activation energy for heterotroph respiration (13), and k is Boltzmann's constant (8.62⅐10 Ϫ5 ⅐eV⅐K Ϫ1 ), and ␣ h is the allometric scaling exponent for body size (14,15). For the metabolic rates of marine autotrophs, things are complicated by the dependence of photosyntheti...
Grazing on chlorophyll by microzooplankton (<200 •um) and copepods was measured in the mixed layer of the high-latitude North Atlantic Ocean during May and August 1991. No significant grazing by microzøoplankton occurred in May during a spring bloom dominated by colonial Phaeocystis pouchettii and Nitzschia spp. As the bloom declined, the size distribution of chlorophyll shifted from dominance by the > 20 •um chlorophyll fraction to dominance by the <20 chlorophyll fraction. The impact of grazing by microzooplankton increased as the bloom declined, with microzooplankton consuming 100% of potential daily chlorophyll production following the bloom. In August,.when the phytoplankton was dominated by the <20 #m chlorophyll fraction, m•cr0zooplankton consumed 37-53% (mean = 41%_+11% s.d.)'of potential daily chlorophyll production. Averaged over all experiments, microzooplankton grazing accounted for 81% of daily chlorophyll production. The grazing impact of Calanus finmarchicus stages C4 and C5, which dominated mesozooplankton biomass in the upper euphotic zone in both spring and late summer, was concentrated on chlorophyll > 20/tin in both seasons; C. f in mar c h ic u s did not consume significant amounts of chlorophyll < 20 •um in either season. Compared to the microzooplankton, copepods did not consume a significant fraction of total chlorophyll in either season, accounting for only -1% of daily chlorophyll production. Introduction The objective of this study was to evaluate losses of chlorophyll due to grazing by microzooplankton and mesozooplankton in the mixed layer of the high-latitude North Atlantic Ocean under contrasting biological and hydrographic conditions during spring and summer. The research was conducted on RV Endeavor cruises EN224-3 and EN227 in spring and late summer 1991 under the aegis of the Marine Light Mixed Layers (MLML) Accelerated Research Initiative, a multidisciplinary effort focused on the biological, physical and optical dynamics of the mixed lay_cr. The MLML study site was occupied t'or 8 days •n May 1991 and 9 days in August/Scptcmber 1991. Paper number 94JC00983 0148-0277!95!94JC-00983505.00 tic Front, an area characterized by extreme seasonal forcing of biology and hydrography. Convective cooling of the water column during winter produces adccp mixed layer of at least several hundred meters, which provides new nutrients to the system [Robinson et al., 1979; Plueddemann et al., this issue]. Irradiance and phytoplankton standing stocks are low during winter when phytoplankton growth is believed to be very low. With the onset of stratification, typically in April, standing stocks of chlorophyll increase IWil!iams and Hopkins, 1974; Plueddemann et al., this issue; Stramska et al., this issue], and a strong diatom bloom reaching 2-5/•g L '•. Calanus finmarchicus and other copepod species appear in the euphotic zone at this time, but their grazing appears to have little impact on the bloom [Dam et al., 1993; Morales et al., 1991; Weeks et al., 1993]. Transition to the summer phytoplankto...
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