Terrestrial gross primary productivity (GPP) varies greatly over time and space. A better understanding of this variability is necessary for more accurate predictions of the future climate-carbon cycle feedback. Recent studies have suggested that variability in GPP is driven by a broad range of biotic and abiotic factors operating mainly through changes in vegetation phenology and physiological processes. However, it is still unclear how plant phenology and physiology can be integrated to explain the spatiotemporal variability of terrestrial GPP. Based on analyses of eddy-covariance and satellite-derived data, we decomposed annual terrestrial GPP into the length of the CO 2 uptake period (CUP) and the seasonal maximal capacity of CO 2 uptake (GPP max ). The product of CUP and GPP max explained >90% of the temporal GPP variability in most areas of North America during 2000-2010 and the spatial GPP variation among globally distributed eddy flux tower sites. It also explained GPP response to the European heatwave in 2003 (r 2 = 0.90) and GPP recovery after a fire disturbance in South Dakota (r 2 = 0.88). Additional analysis of the eddy-covariance flux data shows that the interbiome variation in annual GPP is better explained by that in GPP max than CUP. These findings indicate that terrestrial GPP is jointly controlled by ecosystem-level plant phenology and photosynthetic capacity, and greater understanding of GPP max and CUP responses to environmental and biological variations will, thus, improve predictions of GPP over time and space. ecosystem carbon uptake | growing season length | photosynthetic capacity | spatiotemporal variability | climate extreme L arge variability exists among estimates of terrestrial carbon sequestration, resulting in substantial uncertainty in modeled dynamics of atmospheric CO 2 concentration and predicted future climate change (1). The variability in carbon sequestration is partially caused by variation in terrestrial gross primary productivity (GPP) (2), which is the cumulative rate over time of gross plant Significance Terrestrial gross primary productivity (GPP), the total photosynthetic CO 2 fixation at ecosystem level, fuels all life on land. However, its spatiotemporal variability is poorly understood, because GPP is determined by many processes related to plant phenology and physiological activities. In this study, we find that plant phenological and physiological properties can be integrated in a robust index-the product of the length of CO 2 uptake period and the seasonal maximal photosynthesis-to explain the GPP variability over space and time in response to climate extremes and during recovery after disturbance.

In view of future changes in climate, it is important to better understand how differ-

Grasslands and agroecosystems occupy one-third of the terrestrial area, but their contribution to the global carbon cycle remains uncertain. We used a set of 316 site-years of CO₂ exchange measurements to quantify gross primary productivity, respiration, and light-response parameters of grasslands, shrublands/savanna, wetlands, and cropland ecosystems worldwide. We analyzed data from 72 global flux-tower sites partitioned into gross photosynthesis and ecosystem respiration with the use of the light-response method (Gilmanov, T. G., D. A. Johnson, and N. Z. Saliendra. 2003. Growing season CO₂ fluxes in a sagebrushsteppe ecosystem in Idaho: Bowen ratio/energy balance measurements and modeling. Basic and Applied Ecology 4:167-183) from the RANGEFLUX and WORLDGRASSAGRIFLUX data sets supplemented by 46 sites from the FLUXNET La Thuile data set partitioned with the use of the temperature-response method (Reichstein, M., E. Falge, D. Baldocchi, D. Papale, R. Valentini, M. Aubinet, P. Berbigier, C. Bernhofer, N. Buchmann, M. Falk, T. Gilmanov, A. Granier, T. Grunwald, K. Havrankova, D. Janous, A. Knohl, T. Laurela, A. Lohila, D. Loustau, G. Matteucci, T. Meyers, F. Miglietta, J.M. Ourcival, D. Perrin, J. Pumpanen, S. Rambal, E. Rotenberg, M. Sanz, J. Tenhunen, G. Seufert, F. Vaccari, T. Vesala, and D. Yakir. 2005. On the separation of net ecosystem exchange into assimilation and ecosystem respiration: review and improved algorithm. Global Change Biology 11: 1.424-1439). Maximum values of the quantum yield (alpha = 75 mmol.mol(-1)), photosynthetic capacity (A(max) = 3.4 mg CO₂ . m(-2).s-1), gross photosynthesis (P-g,P-max = 1.16 g CO₂ . m(-2).d(-1)), and ecological light-use efficiency (epsilon(ecol) = 59 mmol . mol(-1)) of managed grasslands and high-production croplands exceeded those of most forest ecosystems, indicating the potential of nonforest ecosystems for uptake of atmospheric CO₂. Maximum values of gross primary production (8 600 g CO₂ . m(-2).yr(-1)), total ecosystem respiration (7 900 g CO₂ . m(-2).yr(-1)), and net CO₂ exchange (2 400 g CO₂ . m(-2).yr(-1)) were observed for intensively managed grasslands and high-yield crops, and are comparable to or higher than those for forest ecosystems, excluding some tropical forests. On average, 80% of the nonforest sites were apparent sinks for atmospheric CO₂, with mean net uptake of 700 g CO₂ . m(-2).yr(-1) for intensive grasslands and 933 g CO₂ . m(-2).d(-1) for croplands. However, part of these apparent sinks is accumulated in crops and forage, which are carbon pools that are harvested, transported, and decomposed off site. Therefore, although agricultural fields may be predominantly sinks for atmospheric CO₂, this does not imply that they are necessarily increasing their carbon stock