Abstract
The rise of oxygenic photosynthesis nearly three billion years ago led to the accumulation of free oxygen and to the subsequent diversification of life on Earth; today, nearly half of all oxygen production derives from the photosynthetic activities of marine phytoplankton1. The conclusion that the open sea –– and therefore much of our planet's surface –– is in a net heterotrophic metabolic state2,3,4 is enigmatic and is a first-order question in the global carbon cycle, as discussed by del Giorgio and Duarte5. Our findings suggest that autotrophy in the open sea is episodic and decoupled from the more constant heterotrophic processes. Consequently, the metabolic balance of the open sea depends on proper space and timescale integration to achieve an ecological understanding of life in the sea.
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References
Field, C. B., Behrenfeld, M. J., Randerson, J. T. & Falkowski, P. Science 281, 237–240 (1998).
del Giorgio, P. A., Cole, J. J. & Cimbleris, A. Nature 385, 148–151 (1997).
Duarte, C. M. & Agustí, S. Science 281, 234–236 (1998).
Duarte, C. M., Agustí, S., del Giorgio, P. A. & Cole, J. J. Science 284, 1735 (1999).
del Giorgio, P. A. & Duarte, C. M. Nature 420, 379–384 (2002).
Emerson, S. et al. Nature 389, 951–954 (1997).
Emerson, S., Quay, P. D., Stump, C., Wilbur, D. & Schudlich, R. J. Geophys. Res. 100, 15873–15887 (1995).
Najjar, R. G. & Keeling, R. F. Global Biogeochem. Cycles 14, 573–584 (2000).
Karl, D. M., Bidigare, R. R. & Letelier, R. M. in Phytoplankton Productivity and Carbon Assimilation in Marine and Freshwater Ecosystems (eds Williams, P. J. leB., Thomas, D. R. & Reynolds, C. S.) 222–264 (Blackwell, London, 2002).
McGillicuddy, D. J. Jr et al. Nature 394, 263–266 (1998).
Uz, B. M., Yoder, J. A. & Osychny, V. Nature 409, 597–600 (2001).
Gregg, M. C., Sanford, T. B. & Winkel, D. P. Nature 422, 513–515 (2003).
Young, R. W. et al. Global Biogeochem. Cycles 5, 119–134 (1991).
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Karl, D., Laws, E., Morris, P. et al. Metabolic balance of the open sea. Nature 426, 32 (2003). https://doi.org/10.1038/426032a
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DOI: https://doi.org/10.1038/426032a
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