Modeling carbon allocation in trees: a search for principles

We review approaches to predicting carbon and nitrogen allocation in forest models in terms of their underlying assumptions and their resulting strengths and limitations. Empirical and allometric methods are easily developed and computationally efficient, but lack the power of evolution-based approaches to explain and predict multifaceted effects of environmental variability and climate change. In evolution-based methods, allocation is usually determined by maximization of a fitness proxy, either in a fixed environment, which we call optimal response (OR) models, or including the feedback of an individual's strategy on its environment (game-theoretical optimization, GTO). Optimal response models can predict allocation in single trees and stands when there is significant competition only for one resource. Game-theoretical optimization can be used to account for additional dimensions of competition, e.g., when strong root competition boosts root allocation at the expense of wood production. However, we demonstrate that an OR model predicts similar allocation to a GTO model under the root-competitive conditions reported in free-air carbon dioxide enrichment (FACE) experiments. The most evolutionarily realistic approach is adaptive dynamics (AD) where the allocation strategy arises from eco-evolutionary dynamics of populations instead of a fitness proxy. We also discuss emerging entropy-based approaches that offer an alternative thermodynamic perspective on allocation, in which fitness proxies are replaced by entropy or entropy production. To help develop allocation models further, the value of wide-ranging datasets, such as FLUXNET, could be greatly enhanced by ancillary measurements of driving variables, such as water and soil nitrogen availability

CO2 Enhancement of Forest Productivity Constrained by Limited Nitrogen Availability

Stimulation of terrestrial productivity by rising CO~2~ concentration is projected to reduce the airborne fraction of anthropogenic CO~2~ emissions; coupled climate-carbon (C) cycle models, including those used in the IPCC Fourth Assessment Report (AR4), are sensitive to this negative feedback on atmospheric CO~2~^1^. The representation of the so-called CO~2~ fertilization effect in the 11 models used in AR4 and subsequent models^2,3^ was broadly consistent with experimental evidence from four free-air CO~2~ enrichment (FACE) experiments, which indicated that net primary productivity (NPP) of forests was increased by 23 +/- 2% in response to atmospheric CO~2~ enrichment to 550 ppm^4^. Substantial uncertainty remains, however, because of the expectation that feedbacks through the nitrogen (N) cycle will reduce the CO~2~ stimulation of NPP^5,6^; these feedbacks were not included in the AR4 models and heretofore have not been confirmed by experiments in forests^7^. Here, we provide new evidence from a FACE experiment in a deciduous Liquidambar styraciflua (sweetgum) forest stand in Tennessee, USA, that N limitation has significantly reduced the stimulation of NPP by elevated atmospheric CO~2~ concentration (eCO~2~). Isotopic evidence and N budget analysis support the premise that N availability in this forest ecosystem has been declining over time, and declining faster in eCO~2~. Model analyses and evidence from leaf- and stand-level observations provide mechanistic evidence that declining N availability constrained the tree response to eCO2. These results provide a strong rationale and process understanding for incorporating N limitation and N feedback effects in ecosystem and global models used in climate change assessments

Plant root distributions and nitrogen uptake predicted by a hypothesis of optimal root foraging

CO2-enrichment experiments consistently show that rooting depth increases when trees are grown at elevated CO2 (eCO2), leading in some experiments to increased capture of available soil nitrogen (N) from deeper soil. However, the link between N uptake an

Tree water uptake enhances nitrogen acquisition in a fertilized boreal forest - but not under nitrogen-poor conditions

Understanding how plant water uptake interacts with acquisition of soil nitrogen (N) and other nutrients is fundamental for predicting plant responses to a changing environment, but it is an area where models disagree. We present a novel isotopic labelling approach which reveals spatial patterns of water and N uptake, and their interaction, by trees. The stable isotopes N-15 and H-2 were applied to a small area of the forest floor in stands with high and low soil N availability. Uptake by surrounding trees was measured. The sensitivity of N acquisition to water uptake was quantified by statistical modelling. Trees in the high-N stand acquired twice as much N-15 as in the low-N stand and around half of their N uptake was dependent on water uptake (H-2 enrichment). By contrast, in the low-N stand there was no positive effect of water uptake on N uptake. We conclude that tree N acquisition was only marginally dependent on water flux toward the root surface under low-N conditions whereas under high-N conditions, the water-associated N uptake was substantial. The results suggest a fundamental shift in N acquisition strategy under high-N conditions

Konvensyen Myprospec tumpu revolusi industri 4.0

Rising atmospheric concentrations of CO 2 (C a) can reduce stomatal conductance and transpiration rate in trees, but the magnitude of this effect varies considerably among experiments. The theory of optimal stomatal behaviour predicts that the ratio of photosynthesis to transpiration (instantaneous transpiration efficiency, ITE) should increase in proportion to C a. We hypothesized that plants regulate stomatal conductance optimally in response to rising C a. We tested this hypothesis with data from young Eucalyptus saligna Sm. trees grown in 12 climate-controlled whole-tree chambers for 2 years at ambient and elevated C a. Elevated C a was ambient + 240 ppm, 60% higher than ambient C a. Leaf-scale gas exchange was measured throughout the second year of the study and leaf-scale ITE increased by 60% under elevated C a, as predicted. Values of leaf-scale ITE depended strongly on vapour pressure deficit (D) in both CO 2 treatments. Whole-canopy CO 2 and H 2O fluxes were also monitored continuously for each chamber throughout the second year. There were small differences in D between C a treatments, which had important effects on values of canopy-scale ITE. However, when C a treatments were compared at the same D, canopy-scale ITE was consistently increased by 60%, again as predicted. Importantly, leaf and canopy-scale ITE were not significantly different, indicating that ITE was not scale-dependent. Observed changes in transpiration rate could be explained on the basis that ITE increased in proportion to C a. The effect of elevated C a on photosynthesis increased with rising D. At high D, C a had a large effect on photosynthesis and a small effect on transpiration rate. At low D, in contrast, there was a small effect of C a on photosynthesis, but a much larger effect on transpiration rate. If shown to be a general response, the proportionality of ITE with C a will allow us to predict the effects of C a on transpiration rate

New insights into carbon allocation by trees from the hypothesis that annual wood production is maximized

Allocation of carbon (C) between tree components (leaves, fine roots and woody structures) is an important determinant of terrestrial C sequestration. Yet, because the mechanisms underlying C allocation are poorly understood, it is a weak link in curren

Leaf-trait variation explained by the hypothesis that plants maximize their canopy carbon export over the lifespan of leaves

Measured values of four key leaf traits (leaf area per unit mass, nitrogen concentration, photosynthetic capacity, leaf lifespan) co-vary consistently within and among diverse biomes, suggesting convergent evolution across species. The same leaf traits co-vary consistently with the environmental conditions (light intensity, carbon-dioxide concentration, nitrogen supply) prevailing during leaf development. No existing theory satisfactorily explains all of these trends. Here, using a simple model of the carbon-nitrogen economy of trees, we show that global leaf-trait relationships and leaf responses to environmental conditions can be explained by the optimization hypothesis (MAXX) that plants maximize the total amount of carbon exported from their canopies over the lifespan of leaves. Incorporating MAXX into larger-scale vegetation models may improve their consistency with global leaf-trait relationships, and enhance their ability to predict how global terrestrial productivity and carbon sequestration respond to environmental change

Effects of CO₂ on plants at different timescales

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Acclimation of the respiration/photosynthesis ratio to temperature: insights from a model

International audienceBased on short-term experiments, many plant growth models - including those used in global change research - assume that an increase in temperature stimulates plant respiration (R) more than photosynthesis (P), leading to an increase in the R/P ratio. Longer-term experiments, however, have demonstrated that R/P is relatively insensitive to growth temperature. We show that both types of temperature response may be reconciled within a simple substrate-based model of giant acclimation to temperature, in which respiration is effectively limited by the supply of carbohydrates fixed through photosynthesis. The short-term, positive temperature response of R/P reflects the transient dynamics of the nonstructural carbohydrate and protein pools; the insensitivity of R/P to temperature on longer time-scales reflects the steady-state behaviour of these pools. Thus the substrate approach may provide a basis for predicting plant respiration responses to temperature that is more robust than the current modelling paradigm based on the extrapolation of results from short-term experiments. The present model predicts that the acclimated R/P depends mainly on the internal allocation of carbohydrates to protein synthesis, a better understanding of which is therefore required to underpin the wider use of a constant R/P as an alternative modelling paradigm in global change research