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Summary The partitioning of photosynthate among various forest carbon pools is a key process regulating long‐term carbon sequestration, with allocation to aboveground woody biomass carbon (AGBC) in particular playing an outsized role in the global carbon cycle due to its slow residence time. However, directly estimating the fraction of gross primary productivity (GPP) that goes to AGBC has historically been difficult and time‐consuming, leaving us with persistent uncertainties.We used an extensive dataset of tree‐ring chronologies co‐located at flux towers to assess the coupling between AGBC and GPP, calculate the fraction of fixed carbon that is allocated to AGBC, and understand the drivers of variability in this fraction.We found that annual AGBC and GPP were rarely correlated, and that annual AGBC represented only a small fraction (c. 9%) of fixed carbon. This fraction varied considerably across sites and was driven by differences in stand density and site climate. Annual AGBC was suppressed byc. 30% during drought and remained below average for years afterward.These results imply that assumptions of relatively stationary allocation of GPP to woody biomass and other plant tissues could lead to systematic biases in modeled carbon accumulation in different plant pools and thus in carbon residence time. 

Abstract Forest productivity projections remain highly uncertain, notably because underpinning physiological controls are delicate to disentangle. Transient perturbation of global climate by large volcanic eruptions provides a unique opportunity to retrospectively isolate underlying processes. Here, we use a multi‐proxy dataset of tree‐ring records distributed over the Northern Hemisphere to investigate the effect of eruptions on tree growth and photosynthesis and evaluate CMIP6 models. Tree‐ring isotope records denoted a widespread 2–4 years increase of photosynthesis following eruptions, likely as a result of diffuse light fertilization. We found evidence that enhanced photosynthesis transiently drove ring width, but the latter further exhibited a decadal anomaly that evidenced independent growth and photosynthesis responses. CMIP6 simulations reproduced overall tree growth decline but did not capture observed photosynthesis anomaly, its decoupling from tree growth or the climate sensitivities of either processes, highlighting key disconnects that deserve further attention to improve forest productivity projections under climate change. 

ABSTRACT Stomata control plant water loss and photosynthetic carbon gain. Developing more generalized and accurate stomatal models is essential for earth system models and predicting responses under novel environmental conditions associated with global change. Plant optimality theories offer one promising approach, but most such theories assume that stomatal conductance maximizes photosynthetic net carbon assimilation subject to some cost orconstraintof water. We move beyond this approach by developing a new, generalized optimality theory of stomatal conductance, optimizing any non‐foliar proxy that requires water and carbon reserves, like growth, survival, and reproduction. We overcome two prior limitations. First, we reconcile the computational efficiency ofinstantaneousoptimization with a more biologically meaningfuldynamic feedbackoptimization over plant lifespans. Second, we incorporatenon‐steady‐statephysics in the optimization to account for the temporal changes in the water, carbon, and energy storage within a plant and its environment that occur over the timescales that stomata act, contrary to previous theories. Our optimal stomatal conductance compares well to observations from seedlings, saplings, and mature trees from field and greenhouse experiments. Our model predicts predispositions to mortality during the 2018 European drought and captures realistic responses to environmental cues, including the partial alleviation of heat stress by evaporative cooling and the negative effect of accumulating foliar soluble carbohydrates, promoting closure under elevated CO₂. We advance stomatal optimality theory by incorporating generalized evolutionary fitness proxies and enhance its utility without compromising its realism, offering promise for future models to more realistically and accurately predict global carbon and water fluxes. 

Across forests, photosynthesis and woody growth respond to different climate cues.