The illusion of balance in the history of the biosphere.

Earth's surface has been irreversibly altered by the activity of organisms, a process that has accelerated as the power of the biosphere (the rate at which life extracts and deploys energy) has increased over time. This trend is incompatible with the expectation that the inputs to Earth's surface of life's materials from the crust and mantle be matched by export from Earth's surface to long-term reservoirs. Here, I suggest that the collective activity of organisms has always violated this balance. The biosphere's ability to extract, retain, recycle, and accumulate materials has allowed living biomass to increase and for exports to decrease over very long timescales. This collective metabolism implies a net transfer of materials from the planet's interior to its surface. The combination of metabolic innovations, competition, adaptive evolution, and the establishment of collaborative economic feedback in ecosystems created dynamic ecological stability despite great spatial and temporal heterogeneity in physical and biological inputs and export of nutrients into and out of the biosphere. Models of geochemical cycling must take the fundamental role of living organisms and the evolutionary changes in these roles into account to explain past and future conditions.