Earlier spring leaf unfolding is a frequently observed response of plants to climate warming. Many deciduous tree species require chilling for dormancy release, and warming-related reductions in chilling may counteract the advance of leaf unfolding in response to warming. Empirical evidence for this, however, is limited to saplings or twigs in climate-controlled chambers. Using long-term in situ observations of leaf unfolding for seven dominant European tree species at 1,245 sites, here we show that the apparent response of leaf unfolding to climate warming (ST, expressed in days advance of leaf unfolding per [deg]C warming) has significantly decreased from 1980 to 2013 in all monitored tree species. Averaged across all species and sites, ST decreased by 40% from 4.0 [plusmn] 1.8 days [deg]C-1 during 1980-1994 to 2.3 [plusmn] 1.6 days [deg]C-1 during 1999-2013. The declining ST was also simulated by chilling-based phenology models, albeit with a weaker decline (24-30%) than observed in situ. The reduction in ST is likely to be partly attributable to reduced chilling. Nonetheless, other mechanisms may also have a role, such as /`photoperiod limitation/' mechanisms that may become ultimately limiting when leaf unfolding dates occur too early in the season. Our results provide empirical evidence for a declining ST, but also suggest that the predicted strong winter warming in the future may further reduce ST and therefore result in a slowdown in the advance of tree spring phenology.

Worldwide expansion of agriculture is impacting the earth's climate by altering carbon, water, and energy fluxes, but the climate in turn is impacting crop production. To study this two-way interaction and its impact on seasonal dynamics of carbon, water, and energy fluxes, we implemented dynamic crop growth processes into a land surface model, the Integrated Science Assessment Model (ISAM). In particular, we implemented crop-specific phenology schemes and dynamic carbon allocation schemes. These schemes account for light, water, and nutrient stresses while allocating the assimilated carbon to leaf, root, stem, and grain pools. The dynamic vegetation structure simulation better captured the seasonal variability in leaf area index (LAI), canopy height, and root depth. We further implemented dynamic root distribution processes in soil layers, which better simulated the root response of soil water uptake and transpiration. Observational data for LAI, above- and belowground biomass, and carbon, water, and energy fluxes were compiled from two AmeriFlux sites, Mead, NE, and Bondville, IL, USA, to calibrate and evaluate the model performance. For the purposes of calibration and evaluation, we use a corn–soybean (C4–C3) rotation system over the period 2001–2004. The calibrated model was able to capture the diurnal and seasonal patterns of carbon assimilation and water and energy fluxes for the corn–soybean rotation system at these two sites. Specifically, the calculated gross primary production (GPP), net radiation fluxes at the top of the canopy, and latent heat fluxes compared well with observations. The largest bias in model results was in sensible heat flux (SH) for corn and soybean at both sites. The dynamic crop growth simulation better captured the seasonal variability in carbon and energy fluxes relative to the static simulation implemented in the original version of ISAM. Especially, with dynamic carbon allocation and root distribution processes, the model's simulated GPP and latent heat flux (LH) were in much better agreement with observational data than for the static root distribution simulation. Modeled latent heat based on dynamic growth processes increased by 12–27% during the growing season at both sites, leading to an improvement in modeled GPP by 13–61% compared to the estimates based on the original version of the ISAM.

Summary: In higher plants, photosystems II and I are found in grana stacks and unstacked stroma lamellae, respectively. To connect them, electron carriers negotiate tortuous multi‐media paths and are subject to macromolecular blocking. Why does evolution select an apparently unnecessary, inefficient bipartition? Here we systematically explain this perplexing phenomenon. We propose that grana stacks, acting like bellows in accordions, increase the degree of ultrastructural control on photosynthesis through thylakoid swelling/shrinking induced by osmotic water fluxes. This control coordinates with variations in stomatal conductance and the turgor of guard cells, which act like an accordion's air button. Thylakoid ultrastructural dynamics regulate macromolecular blocking/collision probability, direct diffusional pathlengths, division of function of Cytochrome b₆f complex between linear and cyclic electron transport, luminal pH via osmotic water fluxes, and the separation of pH dynamics between granal and lamellar lumens in response to environmental variations. With the two functionally asymmetrical photosystems located distantly from each other, the ultrastructural control, nonphotochemical quenching, and carbon‐reaction feedbacks maximally cooperate to balance electron transport with gas exchange, provide homeostasis in fluctuating light environments, and protect photosystems in drought. Grana stacks represent a dry/high irradiance adaptation of photosynthetic machinery to improve fitness in challenging land environments. Our theory unifies many well‐known but seemingly unconnected phenomena of thylakoid structure and function in higher plants.