The distribution of prokaryotic metabolism between maintenance and growth activities has a profound impact on the transformation of carbon substrates to either biomass or CO 2 . Knowledge of key factors influencing prokaryotic maintenance respiration is, however, highly limited. This mesocosm study validated the significance of prokaryotic maintenance respiration by mimicking temperature and nutrients within levels representative of winter and summer conditions. A global range of growth efficiencies (0.05-0.57) and specific growth rates (0.06-2.7 d À1 ) were obtained. The field pattern of cell-specific respiration versus specific growth rate and the global relationship between growth efficiency and growth rate were reproduced. Maintenance respiration accounted for 75% and 15% of prokaryotic respiration corresponding to winter and summer conditions, respectively. Temperature and nutrients showed independent positive effects for all prokaryotic variables except abundance and cell-specific respiration. All treatments resulted in different taxonomic diversity, with specific populations of amplicon sequence variants associated with either maintenance or growth conditions. These results validate a significant relationship between specific growth and respiration rate under productive conditions and show that elevated prokaryotic maintenance respiration can occur under cold and oligotrophic conditions. The experimental design provides a tool for further study of prokaryotic energy metabolism under realistic conditions at the mesocosm scale.
Plankton respiration is a major process removing oxygen from pelagic environments and constitutes one of the largest oxygen transformations in the sea. Where the O2 supplies due to dissolution, advection and oxygenic photosynthesis are not sufficient, hypoxic, or anoxic waters may result. Coastal waters with limited water exchange are especially prone to have low oxygen levels due to eutrophication and climate change. To support marine environmental management in a period of rapid climate change, we investigated the current knowledge of regulating plankton respiration based on field and experimental studies reported in the literature. Models for regulation of plankton respiration was tested on a three-year field data set. Temperature is the most reported predictor positively influencing plankton respiration (mean r2 = 0.50, n=15). The organic carbon supply driven by primary production has a similar coefficient of determination but fewer reported relationships (mean r2 = 0.52, n=6). Riverine discharges of dissolved organic carbon can override the influence of primary production in estuaries precluding effects of nutrient reductions. The median predictions of respiration regulation produced by current models vary by a factor of 2 from the median of observed values and extreme values varied even more. Predictions by models are therefore still too uncertain for application at regional and local scales. Models with temperature as predictor showed best performance but deviated from measured values in some seasons. The combined dependence of plankton respiration on temperature, phytoplankton production and discharge of riverine organic carbon will probably lead to increased oxygen consumption and reduced oxygen levels with projected climate change. This will be especially pronounced where increased precipitation is expected to enhance riverine discharges of carbon compounds. The biologically mediated transfer of carbon for long-term storage in deeper layers will slow down. Implementation of plankton respiration measurements in long-term ecological monitoring programs at water body and basin scales is advocated, which would enable future multivariate analyses and improvements in model precision across aquatic environments.
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