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Wood is a remarkable material with great cultural, economic, and biogeochemical importance. However, our understanding of its formation is poor. Key properties that have not been explained include the anatomy of growth rings (with consistent transitions from low-density earlywood to high density latewood), strong temperature-dependence of latewood density (used for historical temperature reconstructions), the regulation of cell size, and overall growth-temperature relationships in conifer and ring-porous tree species. We have developed a theoretical framework based on observations on Pinus sylvestris L. in northern Sweden. The observed anatomical properties emerge from our framework as a consequence of interactions in time and space between the production of new cells, the dynamics of developmental zone widths, and the distribution of carbohydrates across the developing wood. Here we find that the diffusion of carbohydrates is critical to determining final ring anatomy, potentially overturning current understanding of how wood formation responds to environmental variability and transforming our interpretation of tree rings as proxies of past climates.

Wood formation determines major long-term carbon (C) accumulation in trees and therefore provides a crucial ecosystem service in mitigating climate change. Nevertheless, we lack understanding of how species with contrasting wood anatomical types differ with respect to phenology and environmental controls on wood formation. In this study, we investigated the seasonality and rates of radial growth and their relationships with climatic factors, and the seasonal variations of stem nonstructural carbohydrates (NSC) in three species with contrasting wood anatomical types (red oak: ring-porous; red maple: diffuse-porous; white pine: coniferous) in a temperate mixed forest during 2017-2019. We found that the high ring width variability observed in both red oak and red maple was caused more by changes in growth duration than growth rate. Seasonal radial growth patterns did not vary following transient environmental factors for all three species. Both angiosperm species showed higher concentrations and lower inter-annual fluctuations of NSC than the coniferous species. Inter-annual variability of ring width varied by species with contrasting wood anatomical types. Due to the high dependence of annual ring width on growth duration, our study highlights the critical importance of xylem formation phenology for understanding and modelling the dynamics of wood formation.

Wood growth constitutes the main process for long-term atmospheric carbon sequestration in vegetation. However, our understanding of the process of wood growth and its response to environmental drivers is limited. Current dynamic global vegetation models (DGVMs) are mainly photosynthesis-driven and thus do not explicitly include a direct environmental effect on tree growth. However, physiological evidence suggests that, to realistically model vegetation carbon allocation under increased climatic stressors, it is crucial to treat growth responses independently from photosynthesis. A plausible growth response function suitable for global simulations in DGVMs has been lacking. Here, we present the first soil water-growth response function and parameter range for deciduous and evergreen conifers. The response curve was calibrated against European larch and Norway spruce in a dry temperate forest in the Swiss Alps. We present a new data-driven approach based on a combination of tree ring width (TRW) records, growing season length and simulated subdaily soil hydrology to parameterize ring width increment simulations. We found that a simple linear response function, with an intercept at zero moisture stress, used in growth simulations reproduced 62.3% and 59.4% of observed TRW variability for larch and spruce respectively and, importantly, the response function slope was much steeper than literature values for soil moisture effects on photosynthesis and stomatal conductance. Specifically, we found stem growth stops at soil moisture potentials of -0.47 MPa for larch and -0.66 MPa for spruce, whereas photosynthesis in trees continues down to -1.2 MPa or lower, depending on species and measurement method. These results are strong evidence that the response functions of source and sink processes are indeed very different in trees, and need to be considered separately to correctly assess vegetation responses to environmental change. The results provide a parameterization for the explicit representation of growth responses to soil water in vegetation models.

Wood formation has received considerable attention across various research fields as a key process to model. Historical and contemporary models of wood formation from various disciplines have encapsulated hypotheses such as the influence of external (e.g., climatic) or internal (e.g., hormonal) factors on the successive stages of wood cell differentiation. This review covers 17 wood formation models from three different disciplines, the earliest from 1968 and the latest from 2020. The described processes, as well as their external and internal drivers and their level of complexity, are discussed. This work is the first systematic cataloging, characterization, and process-focused review of wood formation models. Remaining open questions concerning wood formation processes are identified, and relate to: (1) the extent of hormonal influence on the final tree ring structure; (2) the mechanism underlying the transition from earlywood to latewood in extratropical regions; and (3) the extent to which carbon plays a role as “active” driver or “passive” substrate for growth. We conclude by arguing that wood formation models remain to be fully exploited, with the potential to contribute to studies concerning individual tree carbon sequestration-storage dynamics and regional to global carbon sequestration dynamics in terrestrial vegetation models.

How variations in carbon supply affect wood formation remains poorly understood in particular in mature forest trees. To elucidate how carbon supply affects carbon allocation and wood formation, we attempted to manipulate carbon supply to the cambial region by phloem girdling and compression during the mid- and late-growing season and measured effects on structural development, CO2 efflux and nonstructural carbon reserves in stems of mature white pines. Wood formation and stem CO2 efflux varied with a location relative to treatment (i.e., above or below the restriction). We observed up to twice as many tracheids formed above versus below the treatment after the phloem transport manipulation, whereas the cell-wall area decreased only slightly below the treatments, and cell size did not change relative to the control. Nonstructural carbon reserves in the xylem, needles and roots were largely unaffected by the treatments. Our results suggest that low and high carbon supply affects wood formation, primarily through a strong effect on cell proliferation, and respiration, but local nonstructural carbon concentrations appear to be maintained homeostatically. This contrasts with reports of decoupling of source activity and wood formation at the whole-tree or ecosystem level, highlighting the need to better understand organ-specific responses, within-tree feedbacks, as well as phenological and ontogenetic effects on sink-source dynamics.

During the Last Glacial Maximum (LGM), atmospheric CO2 was around 90 ppmv lower than during the pre-industrial period. The reasons for this decrease are most often elucidated through factorial experiments testing the impact of individual mechanisms. Due to uncertainty in our understanding of the real system, however, the different models used to conduct the experiments inevitably take on different parameter values and different structures. In this paper, the objective is therefore to take an uncertainty-based approach to investigating the LGM CO2 drop by simulating it with a large ensemble of parameter sets, designed to allow for a wide range of large-scale feedback response strengths. Our aim is not to definitely explain the causes of the CO2 drop but rather explore the range of possible responses. We find that the LGM CO2 decrease tends to predominantly be associated with decreasing sea surface temperatures (SSTs), increasing sea ice area, a weakening of the Atlantic Meridional Overturning Circulation (AMOC), a strengthening of the Antarctic Bottom Water (AABW) cell in the Atlantic Ocean, a decreasing ocean biological productivity, an increasing CaCO3 weathering flux and an increasing deep-sea CaCO3 burial flux. The majority of our simulations also predict an increase in terrestrial carbon, coupled with a decrease in ocean and increase in lithospheric carbon. We attribute the increase in terrestrial carbon to a slower soil respiration rate, as well as the preservation rather than destruction of carbon by the LGM ice sheets. An initial comparison of these dominant changes with observations and paleoproxies other than carbon isotope and oxygen data (not evaluated directly in this study) suggests broad agreement. However, we advise more detailed comparisons in the future, and also note that, conceptually at least, our results can only be reconciled with carbon isotope and oxygen data if additional processes not included in our model are brought into play.

American Geophysical Union Fall Meeting, 11-15 December 2017, New Orleans, Increased occurrence of extreme climate or weather events is one of the most damaging consequences of global climate change today and in the future. Estimating the impacts of such extreme events across different human and natural systems is crucial for quantifying overall risks from climate change. Are current models fit for this task? Here we use the 2003 European heat wave and drought (EHW) as a historical analogue for comparable events in the future, and evaluate how accurately its impacts are reproduced by a multi-sectoral >super-ensemble> of state-of-the-art impacts models. Our study combines, for the first time, impacts on agriculture, freshwater resources, terrestrial and marine ecosystems, energy, and human health in a consistent multi-model framework. We identify key impacts of the 2003 EHW reported in the literature and/or recorded in publicly available databases, and examine how closely the models reproduce those impacts, applying the same measure of impact magnitude across different sectors. Preliminary results are mixed: While the EHW's impacts on water resources (streamflow) are reproduced well by most global hydrological models, not all crop and natural vegetation models reproduce the magnitude of impacts on agriculture and ecosystem productivity, respectively, and their performance varies by country or region. A hydropower capacity model matches reported hydropower generation anomalies only in some countries, and estimates of heat-related excess mortality from a set of statistical models are consistent with literature reports only for some of the cities investigated. We present a synthesis of simulated and observed impacts across sectors, and reflect on potential improvements in modeling and analyzing cross-sectoral impacts

Turnover concepts in state-of-the-art global vegetation models (GVMs) account for various processes, but are often highly simplified and may not include an adequate representation of the dominant processes that shape vegetation carbon turnover rates in real forest ecosystems at a large spatial scale. Here we evaluate vegetation carbon turnover processes in GVMs participating in the Inter-Sectoral Impact Model Intercomparison Project (ISI-MIP; including HYBRID4, JeDi, JULES, LPJml, ORCHIDEE, SDGVM, and VISIT) using estimates of vegetation carbon turnover rate (k) derived from a combination of remote sensing based products of biomass and net primary production (NPP). We find that current model limitations lead to considerable biases in the simulated biomass and in k (severe underestimations by all models except JeDi and VISIT compared to observation-based average k), likely contributing to underestimation of positive feedbacks of the northern forest carbon balance to climate change caused by changes in forest mortality. A need for improved turnover concepts related to frost damage, drought and insect outbreaks in order to better reproduce observation-based spatial patterns in k is identified. Since direct frost damage effects on mortality are usually not accounted for in these GVMs, simulated relationships between k and winter length in boreal forests are not consistent between different regions and strongly biased compared to the observation-based relationships. Some models show a response of k to drought in temperate forests as a result of impacts of water availability on NPP, growth efficiency or carbon balance dependent mortality as well as soil or litter moisture effects on leaf turnover or fire. However, further direct drought effects like carbon starvation (only in HYBRID4) or hydraulic failure are usually not taken into account by the investigated GVMs. While they are considered dominant large-scale mortality agents, mortality mechanisms related to insects and pathogens are not explicitly treated in these models. This article is protected by copyright. All rights reserved.