An eco-climatic framework for evaluating the resilience of vegetation to water deficit

The surge in global efforts to understand the causes and consequences of drought on forest ecosystems has tended to focus on specific impacts such as mortality. We propose an ecoclimatic framework that takes a broader view of the ecological relevance of water deficits, linking elements of exposure and resilience to cumulative impacts on a range of ecosystem processes. This ecoclimatic framework is underpinned by two hypotheses: (i) exposure to water deficit can be represented probabilistically and used to estimate exposure thresholds across different vegetation types or ecosystems; and (ii) the cumulative impact of a series of water deficit events is defined by attributes governing the resistance and recovery of the affected processes. We present case studies comprising Pinus edulis and Eucalyptus globulus, tree species with contrasting ecological strategies, which demonstrate how links between exposure and resilience can be examined within our proposed framework. These examples reveal how climatic thresholds can be defined along a continuum of vegetation functional responses to water deficit regimes. The strength of this framework lies in identifying climatic thresholds on vegetation function in the absence of more complete mechanistic understanding, thereby guiding the formulation, application and benchmarking of more detailed modelling

Incorporating non-stomatal limitation improves the performance of leaf and canopy models at high vapour pressure deficit

Vapour pressure deficit (D) is projected to increase in the future as temperature rises. In response to increased D, stomatal conductance (gs) and photosynthesis (A) are reduced, which may result in significant reductions in terrestrial carbon, water and energy fluxes. It is thus important for gas exchange models to capture the observed responses of gs and A with increasing D. We tested a series of coupled A-gs models against leaf gas exchange measurements from the Cumberland Plain Woodland (Australia), where D regularly exceeds 2 kPa and can reach 8 kPa in summer. Two commonly used A-gs models were not able to capture the observed decrease in A and gs with increasing D at the leaf scale. To explain this decrease in A and gs, two alternative hypotheses were tested: hydraulic limitation (i.e., plants reduce gs and/or A due to insufficient water supply) and non-stomatal limitation (i.e., downregulation of photosynthetic capacity). We found that the model that incorporated a non-stomatal limitation captured the observations with high fidelity and required the fewest number of parameters. Whilst the model incorporating hydraulic limitation captured the observed A and gs, it did so via a physical mechanism that is incorrect. We then incorporated a non-stomatal limitation into the stand model, MAESPA, to examine its impact on canopy transpiration and gross primary production. Accounting for a non-stomatal limitation reduced the predicted transpiration by ~19%, improving the correspondence with sap flow measurements, and gross primary production by ~14%. Given the projected global increases in D associated with future warming, these findings suggest that models may need to incorporate non-stomatal limitation to accurately simulate A and gs in the future with high D. Further data on non-stomatal limitation at high D should be a priority, in order to determine the generality of our results and develop a widely applicable model. © The Author(s) 2019. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: [email protected]. was supported by a PhD scholarship from Hawkesbury Institute for the Environment, Western Sydney University. M.G.D.K. acknowledges funding from the Australian Research Council (ARC) Centre of Excellence for Climate Extremes (CE170100023), the ARC Discovery Grant (DP190101823) and support from the NSW Research Attraction and Acceleration Program. EucFACE was built as an initiative of the Australian Government as part of the Nation-building Economic Stimulus Package and is supported by the Australian Commonwealth in collaboration with Western Sydney University. It is also part of a Terrestrial Ecosystem Research Network Super-site facility

The fate of carbon in a mature forest under carbon dioxide enrichment

Atmospheric carbon dioxide enrichment (eCO2) can enhance plant carbon uptake and growth1 5, thereby providing an important negative feedback to climate change by slowing the rate of increase of the atmospheric CO2 concentration6. Although evidence gathered from young aggrading forests has generally indicated a strong CO2 fertilization effect on biomass growth3 5, it is unclear whether mature forests respond to eCO2 in a similar way. In mature trees and forest stands7 10, photosynthetic uptake has been found to increase under eCO2 without any apparent accompanying growth response, leaving the fate of additional carbon fixed under eCO2 unclear4,5,7 11. Here using data from the first ecosystem-scale Free-Air CO2 Enrichment (FACE) experiment in a mature forest, we constructed a comprehensive ecosystem carbon budget to track the fate of carbon as the forest responded to four years of eCO2 exposure. We show that, although the eCO2 treatment of +150 parts per million (+38 per cent) above ambient levels induced a 12 per cent (+247 grams of carbon per square metre per year) increase in carbon uptake through gross primary production, this additional carbon uptake did not lead to increased carbon sequestration at the ecosystem level. Instead, the majority of the extra carbon was emitted back into the atmosphere via several respiratory fluxes, with increased soil respiration alone accounting for half of the total uptake surplus. Our results call into question the predominant thinking that the capacity of forests to act as carbon sinks will be generally enhanced under eCO2, and challenge the efficacy of climate mitigation strategies that rely on ubiquitous CO2 fertilization as a driver of increased carbon sinks in global forests. © 2020, The Author(s), under exclusive licence to Springer Nature Limited

TRY plant trait database – enhanced coverage and open access

Plant traits—the morphological, anatomical, physiological, biochemical and phenological characteristics of plants—determine how plants respond to environmental factors, affect other trophic levels, and influence ecosystem properties and their benefits and detriments to people. Plant trait data thus represent the basis for a vast area of research spanning from evolutionary biology, community and functional ecology, to biodiversity conservation, ecosystem and landscape management, restoration, biogeography and earth system modelling. Since its foundation in 2007, the TRY database of plant traits has grown continuously. It now provides unprecedented data coverage under an open access data policy and is the main plant trait database used by the research community worldwide. Increasingly, the TRY database also supports new frontiers of trait‐based plant research, including the identification of data gaps and the subsequent mobilization or measurement of new data. To support this development, in this article we evaluate the extent of the trait data compiled in TRY and analyse emerging patterns of data coverage and representativeness. Best species coverage is achieved for categorical traits—almost complete coverage for ‘plant growth form’. However, most traits relevant for ecology and vegetation modelling are characterized by continuous intraspecific variation and trait–environmental relationships. These traits have to be measured on individual plants in their respective environment. Despite unprecedented data coverage, we observe a humbling lack of completeness and representativeness of these continuous traits in many aspects. We, therefore, conclude that reducing data gaps and biases in the TRY database remains a key challenge and requires a coordinated approach to data mobilization and trait measurements. This can only be achieved in collaboration with other initiatives

Inputs of organic matter to the ocean

The first objective of this introductory paper is to summarize our present understanding of the quantities of total organic carbon produced in the ocean by photosynthesis and non-biotic photochemical reactions, and the amount entering the ocean from rivers, the atmosphere, and the sediments. In this overview it will become apparent that our knowledge of the primary mechanisms and processes involved with the input of organic matter to the ocean via rivers, the atmosphere, sediments, and in situ photochemical reactions is fragmentary and often completely lacking. This becomes critical when we attempt to estimate the input of certain naturally occurring organic compounds and some synthetic organic compounds, since their primary input to the sea is generally via the atmosphere, rivers, and dumping. Thus the second objective of this paper is to emphasize our need to understand the mechanisms involved in these other input processes and the necessity of developing field programs and mathematical models to evaluate the input of specific organic compounds via these pathways. Polychlorinated biphenyls are used as examples of how necessary it is to understand these other input routes in order to evaluate the cycling of pollutant substances in the ocean

Predicting the decline in daily maximum transpiration rate of two pine stands during drought based on constant minimum leaf water potential and plant hydraulic conductance

International audienceThe effect of drought on forest water use is often estimated with models, but comprehensive models require many parameters, and simple models may not be sufficiently flexible. Many tree species, Pinus species in particular, have been shown to maintain a constant minimum leaf water potential above the critical threshold for xylem embolism during drought. In such cases, prediction of the relative decline in daily maximum transpiration rate with decreasing soil water content is relatively straightforward. We constructed a soil-plant water flow model assuming constant plant conductance and daily minimum leaf water potential, but variable conductance from soil to root. We tested this model against independent data from two sites: automatic shoot chamber data and sap flow measurements from a boreal Scots pine (Pinus sylvestris L.) stand; and sap flow measurements from a maritime pine (Pinus pinaster Ait.) stand. To focus on soil limitations to water uptake, we expressed daily maximum transpiration rate relative to the rate that would be obtained in wet soil with similar environmental variables. The comparison was successful, although the maritime pine stand showed carry-over effects of the drought that we could not explain. For the boreal Scots pine stand, daily maximum transpiration was best predicted by water content of soil deeper than 5 cm. A sensitivity analysis revealed that model predictions were relatively insensitive to the minimum leaf water potential, which can be accounted for by the importance of soil resistance of drying soil. We conclude that a model with constant plant conductance and minimum leaf water potential can accurately predict the decline in daily maximum transpiration rate during drought for these two pine stands, and that including further detail about plant compartments would add little predictive power, except in predicting recovery from severe drought

Predicting the decline in daily maximum transpiration rate of two pine stands during drought based on constant minimum leaf water potential and plant hydraulic conductance

International audienceThe effect of drought on forest water use is often estimated with models, but comprehensive models require many parameters, and simple models may not be sufficiently flexible. Many tree species, Pinus species in particular, have been shown to maintain a constant minimum leaf water potential above the critical threshold for xylem embolism during drought. In such cases, prediction of the relative decline in daily maximum transpiration rate with decreasing soil water content is relatively straightforward. We constructed a soil-plant water flow model assuming constant plant conductance and daily minimum leaf water potential, but variable conductance from soil to root. We tested this model against independent data from two sites: automatic shoot chamber data and sap flow measurements from a boreal Scots pine (Pinus sylvestris L.) stand; and sap flow measurements from a maritime pine (Pinus pinaster Ait.) stand. To focus on soil limitations to water uptake, we expressed daily maximum transpiration rate relative to the rate that would be obtained in wet soil with similar environmental variables. The comparison was successful, although the maritime pine stand showed carry-over effects of the drought that we could not explain. For the boreal Scots pine stand, daily maximum transpiration was best predicted by water content of soil deeper than 5 cm. A sensitivity analysis revealed that model predictions were relatively insensitive to the minimum leaf water potential, which can be accounted for by the importance of soil resistance of drying soil. We conclude that a model with constant plant conductance and minimum leaf water potential can accurately predict the decline in daily maximum transpiration rate during drought for these two pine stands, and that including further detail about plant compartments would add little predictive power, except in predicting recovery from severe drought

Low sensitivity of gross primary production to elevated CO2 in a mature eucalypt woodland

The response of mature forest ecosystems to a rising atmospheric carbon dioxide concentration (span classCombining double low line"inline-formula"iC/ia/span) is a major uncertainty in projecting the future trajectory of the Earth's climate. Although leaf-level net photosynthesis is typically stimulated by exposure to elevated span classCombining double low line"inline-formula"iC/ia/span (espan classCombining double low line"inline-formula"iC/ia/span), it is unclear how this stimulation translates into carbon cycle responses at the ecosystem scale. Here we estimate a key component of the carbon cycle, the gross primary productivity (GPP), of a mature native eucalypt forest exposed to free-air span classCombining double low line"inline-formula"CO2/span enrichment (the EucFACE experiment). In this experiment, light-saturated leaf photosynthesis increased by 19andthinsp;% in response to a 38andthinsp;% increase in span classCombining double low line"inline-formula"iC/ia/span. We used the process-based forest canopy model, MAESPA, to upscale these leaf-level measurements of photosynthesis with canopy structure to estimate the GPP and its response to espan classCombining double low line"inline-formula"iC/ia/span. We assessed the direct impact of espan classCombining double low line"inline-formula"iC/ia/span, as well as the indirect effect of photosynthetic acclimation to espan classCombining double low line"inline-formula"iC/ia/span and variability among treatment plots using different model scenarios./p At the canopy scale, MAESPA estimated a GPP of 1574andthinsp;gandthinsp;Candthinsp;mspan classCombining double low line"inline-formula"-2/spanandthinsp;yrspan classCombining double low line"inline-formula"-1/span under ambient conditions across 4 years and a direct increase in the GPP of span classCombining double low line"inline-formula"+/span11andthinsp;% in response to espan classCombining double low line"inline-formula"iC/ia/span. The smaller canopy-scale response simulated by the model, as compared with the leaf-level response, could be attributed to the prevalence of RuBP regeneration limitation of leaf photosynthesis within the canopy. Photosynthetic acclimation reduced this estimated response to 10andthinsp;%. After taking the baseline variability in the leaf area index across plots in account, we estimated a field GPP response to espan classCombining double low line"inline-formula"iC/ia/span of 6andthinsp;% with a 95andthinsp;% confidence interval (span classCombining double low line"inline-formula"-/span2andthinsp;%, 14andthinsp;%). These findings highlight that the GPP response of mature forests to espan classCombining double low line"inline-formula"iC/ia/span is likely to be considerably lower than the response of light-saturated leaf photosynthesis. Our results provide an important context for interpreting the espan classCombining double low line"inline-formula"iC/ia/span responses of other components of the ecosystem carbon cycle. © Author(s) 2020.Martin G. De Kauwe was supported by the NSW Research Attraction and Acceleration Program (RAAP). Euc-FACE was built as an initiative of the Australian Government as part of the Nation Building Economic Stimulus Plan and is supported by the Australian Commonwealth in collaboration with Western Sydney University