Interannual variability (IAV, represented by standard deviation) in net ecosystem exchange of CO2 (NEE) is mainly driven by climatic drivers and biotic variations (i.e., the changes in photosynthetic and respiratory responses to climate), the effects of which are referred to as climatic (CE) and biotic effects (BE), respectively. Evaluating the relative contributions of CE and BE to the IAV in carbon (C) fluxes and understanding their controlling mechanisms are critical in projecting ecosystem changes in the future climate. In this study, we applied statistical methods with flux data from 65 sites located in the Northern Hemisphere to address this issue. Our results showed that the relative contribution of BE (CnBE) and CE (CnCE) to the IAV in NEE was 57%±14% and 43%±14%, respectively. The discrepancy in the CnBE among sites could be largely explained by water balance index (WBI). Across water-stressed ecosystems, the CnBE decreased with increasing aridity (slope=0.18%mm−1). In addition, the CnBE tended to increase and the uncertainty reduced as timespan of available data increased from 5 to 15 years. Inter-site variation of the IAV in NEE mainly resulted from the IAV in BE (72%) compared to that in CE (37%). Interestingly, positive correlations between BE and CE occurred in grasslands and dry ecosystems (r>0.45, P<0.05) but not in other ecosystems. These results highlighted the importance of BE in determining the IAV in NEE and the ability of ecosystems to regulate C fluxes under climate change might decline when the ecosystems experience more severe water stress in the future.

Climatic variables not only directly affect the interannual variability (IAV) in net ecosystem exchange of CO2 (NEE) but also indirectly drive it by changing the physiological parameters. Identifying these direct and indirect paths can reveal the underlying mechanisms of carbon (C) dynamics. In this study, we applied a path analysis using flux data from 65 sites to quantify the direct and indirect climatic effects on IAV in NEE and to evaluate the potential relationships among the climatic variables and physiological parameters that represent physiology and phenology of ecosystems. We found that the maximum photosynthetic rate was the most important factor for the IAV in gross primary productivity (GPP), which was mainly induced by the variation in vapour pressure deficit. For ecosystem respiration (RE), the most important drivers were GPP and the reference respiratory rate. The biome type regulated the direct and indirect paths, with distinctive differences between forests and non-forests, evergreen needleleaf forests and deciduous broadleaf forests, and between grasslands and croplands. Different paths were also found among wet, moist and dry ecosystems. However, the climatic variables can only partly explain the IAV in physiological parameters, suggesting that the latter may also result from other biotic and disturbance factors. In addition, the climatic variables related to NEE were not necessarily the same as those related to GPP and RE, indicating the emerging difficulty encountered when studying the IAV in NEE. Overall, our results highlight the contribution of certain physiological parameters to the IAV in C fluxes and the importance of biome type and multi-year water conditions, which should receive more attention in future experimental and modelling research.