SPECIAL FEATURE – ESSAY REVIEW PLANT–SOIL INTERACTIONS AND THE CARBON CYCLE The changing global carbon cycle: linking plant–soil carbon dynamics to global consequences

1. Most current climate–carbon cycle models that include the terrestrial carbon (C) cycle are based on a model developed 40 years ago byWoodwell &Whittaker (1968) and omit advances in biogeochemical understanding since that time. Their model treats net C emissions from ecosystems as the balance between net primary production (NPP) and heterotrophic respiration (HR, i.e. primarily decomposition). 2. Under conditions near steady state, geographic patterns of decomposition closely match those of NPP, and net C emissions are adequately described as a simple balance of NPP and HR (the Woodwell-Whittaker model). This close coupling between NPP and HR occurs largely because of tight coupling between C and N (nitrogen) cycles and because NPP constrains the food available to heterotrophs. 3. Processes in addition to NPP and HR become important to understanding net C emissions from ecosystems under conditions of rapid changes in climate, hydrology, atmospheric CO2, land cover, species composition and ⁄or N deposition. Inclusion of these processes in climate–C cycle models would improve their capacity to simulate recent and future climatic change. 4. Processes that appear critical to soil C dynamics but warrant further research before incorporation into ecosystem models include below-ground C flux and its partitioning among roots, mycorrhizas and exudates; microbial community effects on C sequestration; and the effects of temperature and labile C on decomposition. The controls over and consequences of these processes are still unclear at the ecosystem scale. 5. Carbon fluxes in addition to NPP and HR exert strong influences over the climate system under conditions of rapid change. These fluxes include methane release, wildfire, and lateral transfers of food and fibre among ecosystems. 6. Water and energy exchanges are important complements to C cycle feedbacks to the climate system, particularly under non-steady-state conditions. An integrated understanding of multiple ecosystem–climate feedbacks provides a strong foundation for policies tomitigate climate change. 7. Synthesis. Current climate systems models that include only NPP and HR are inadequate under conditions of rapid change.Many of the recent advances in biogeochemical understanding are sufficiently mature to substantially improve representation of ecosystemC dynamics in these models.