Thirst beats hunger – declining hydration during drought prevents carbon starvation in Norway spruce saplings
TL;DR: In a manipulation experiment, the carbon balance of Norway spruce saplings exposed to either drought or carbon starvation (CO2 withdrawal), or both treatments, is manipulated and the dynamics of carbon exchange, allocation and storage in different tissues are compared.
Abstract: Summary � Drought-induced tree mortality results from an interaction of several mechanisms. Plant water and carbon relations are interdependent and assessments of their individual contributions are difficult. Because drought always affects both plant hydration and carbon assimilation, it is challenging to disentangle their concomitant effects on carbon balance and carbon translocation. Here, we report results of a manipulation experiment specifically designed to separate drought effects on carbon and water relations from those on carbon translocation. � In a glasshouse experiment, we manipulated the carbon balance of Norway spruce saplings exposed to either drought or carbon starvation (CO2 withdrawal), or both treatments, and compared the dynamics of carbon exchange, allocation and storage in different tissues. � Drought killed trees much faster than did carbon starvation. Storage C pools were not depleted at death for droughted trees as they were for starved, well-watered trees. Hence drought has a significant detrimental effect on a plant’s ability to utilize stored carbon. � Unless they can be transported to where they are needed, sufficient carbon reserves alone will not assure survival of a drought except under specific conditions, such as moderate drought, or in species that maintain plant water relations required for carbon re-mobilization.
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TL;DR: In this article, the authors identify ten contrasting perspectives that shape the vulnerability debate but have not been discussed collectively and present a set of global vulnerability drivers that are known with high confidence: (1) droughts eventually occur everywhere; (2) warming produces hotter Droughts; (3) atmospheric moisture demand increases nonlinearly with temperature during drought; (4) mortality can occur faster in hotter Drought, consistent with fundamental physiology; (5) shorter Drought can become lethal under warming, increasing the frequency of lethal Drought; and (6) mortality happens rapidly
Abstract: Patterns, mechanisms, projections, and consequences of tree mortality and associated broad-scale forest die-off due to drought accompanied by warmer temperatures—“hotter drought”, an emerging characteristic of the Anthropocene—are the focus of rapidly expanding literature. Despite recent observational, experimental, and modeling studies suggesting increased vulnerability of trees to hotter drought and associated pests and pathogens, substantial debate remains among research, management and policy-making communities regarding future tree mortality risks. We summarize key mortality-relevant findings, differentiating between those implying lesser versus greater levels of vulnerability. Evidence suggesting lesser vulnerability includes forest benefits of elevated [CO2] and increased water-use efficiency; observed and modeled increases in forest growth and canopy greening; widespread increases in woody-plant biomass, density, and extent; compensatory physiological, morphological, and genetic mechanisms; dampening ecological feedbacks; and potential mitigation by forest management. In contrast, recent studies document more rapid mortality under hotter drought due to negative tree physiological responses and accelerated biotic attacks. Additional evidence suggesting greater vulnerability includes rising background mortality rates; projected increases in drought frequency, intensity, and duration; limitations of vegetation models such as inadequately represented mortality processes; warming feedbacks from die-off; and wildfire synergies. Grouping these findings we identify ten contrasting perspectives that shape the vulnerability debate but have not been discussed collectively. We also present a set of global vulnerability drivers that are known with high confidence: (1) droughts eventually occur everywhere; (2) warming produces hotter droughts; (3) atmospheric moisture demand increases nonlinearly with temperature during drought; (4) mortality can occur faster in hotter drought, consistent with fundamental physiology; (5) shorter droughts occur more frequently than longer droughts and can become lethal under warming, increasing the frequency of lethal drought nonlinearly; and (6) mortality happens rapidly relative to growth intervals needed for forest recovery. These high-confidence drivers, in concert with research supporting greater vulnerability perspectives, support an overall viewpoint of greater forest vulnerability globally. We surmise that mortality vulnerability is being discounted in part due to difficulties in predicting threshold responses to extreme climate events. Given the profound ecological and societal implications of underestimating global vulnerability to hotter drought, we highlight urgent challenges for research, management, and policy-making communities.
1,786 citations
Cites background or methods from "Thirst beats hunger – declining hyd..."
...2014), online isotopic measurements (Hartmann et al. 2013), highly controlled chamber systems for experimental manipulations of both [CO2] (e....
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...…temperature changes with changes in canopy cover (Royer et al. 2012), phloem function sensors (Sevanto et al. 2014), online isotopic measurements (Hartmann et al. 2013), highly controlled chamber systems for experimental manipulations of both [CO2] (e.g., Quirk et al. 2013, Duan et al. 2014),…...
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Oklahoma State University–Stillwater1, University of Sydney2, Macquarie University3, University of Utah4, Max Planck Society5, University of Alberta6, University of California, Irvine7, University of New Mexico8, University of Nebraska–Lincoln9, United States Geological Survey10, University of Washington11, University of Arizona12, University of Sheffield13, University of Tasmania14, ETH Zurich15, University of California, Davis16, Los Alamos National Laboratory17, Nanchang Institute of Technology18, University of Wyoming19, Swedish University of Agricultural Sciences20, University of Coimbra21, United States Forest Service22, Northern Arizona University23, Swiss Federal Institute for Forest, Snow and Landscape Research24, University of São Paulo25, University of Oxford26, University of California, Santa Cruz27, Humboldt State University28, Fordham University29, Centre national de la recherche scientifique30, University of Edinburgh31, Hobart Corporation32, Spanish National Research Council33, University of Chile34, Idaho State University35, University of Basilicata36, Colorado State University37, United States Department of Agriculture38, University of Montana39, University of Delaware40, Duke University41, University of Western Ontario42, Sonora Institute of Technology43, Pacific Northwest National Laboratory44
TL;DR: It is shown that, across multiple tree species, loss of xylem conductivity above 60% is associated with mortality, while carbon starvation is not universal, indicating that evidence supporting carbon starvation was not universal.
Abstract: Widespread tree mortality associated with drought has been observed on all forested continents and global change is expected to exacerbate vegetation vulnerability. Forest mortality has implications for future biosphere-atmosphere interactions of carbon, water and energy balance, and is poorly represented in dynamic vegetation models. Reducing uncertainty requires improved mortality projections founded on robust physiological processes. However, the proposed mechanisms of drought-induced mortality, including hydraulic failure and carbon starvation, are unresolved. A growing number of empirical studies have investigated these mechanisms, but data have not been consistently analysed across species and biomes using a standardized physiological framework. Here, we show that xylem hydraulic failure was ubiquitous across multiple tree taxa at drought-induced mortality. All species assessed had 60% or higher loss of xylem hydraulic conductivity, consistent with proposed theoretical and modelled survival thresholds. We found diverse responses in non-structural carbohydrate reserves at mortality, indicating that evidence supporting carbon starvation was not universal. Reduced non-structural carbohydrates were more common for gymnosperms than angiosperms, associated with xylem hydraulic vulnerability, and may have a role in reducing hydraulic function. Our finding that hydraulic failure at drought-induced mortality was persistent across species indicates that substantial improvement in vegetation modelling can be achieved using thresholds in hydraulic function.
651 citations
TL;DR: Current knowledge about responses of tree roots to drought supports the view that tree roots are well equipped to withstand drought situations and maintain morphological and physiological functions as long as possible.
Abstract: The ongoing climate change is characterized by increased temperatures and altered precipitation patterns In addition, there has been an increase in both the frequency and intensity of extreme climatic events such as drought Episodes of drought induce a series of interconnected effects, all of which have the potential to alter the carbon balance of forest ecosystems profoundly at different scales of plant organization and ecosystem functioning During recent years, considerable progress has been made in the understanding of how aboveground parts of trees respond to drought and how these responses affect carbon assimilation In contrast, processes of belowground parts are relatively underrepresented in research on climate change In this review, we describe current knowledge about responses of tree roots to drought Tree roots are capable of responding to drought through a variety of strategies that enable them to avoid and tolerate stress Responses include root biomass adjustments, anatomical alterations, and physiological acclimations The molecular mechanisms underlying these responses are characterized to some extent, and involve stress signaling and the induction of numerous genes, leading to the activation of tolerance pathways In addition, mycorrhizas seem to play important protective roles The current knowledge compiled in this review supports the view that tree roots are well equipped to withstand drought situations and maintain morphological and physiological functions as long as possible Further, the reviewed literature demonstrates the important role of tree roots in the functioning of forest ecosystems and highlights the need for more research in this emerging field
501 citations
TL;DR: This work suggests that in most cases, poorly understood processes of tissue formation and cell growth are governing carbon demand, and thus, CO2 uptake, and that this view is not reflecting reality, but emerged from the availability of methods and process understanding at leaf level.
496 citations
Cites background from "Thirst beats hunger – declining hyd..."
...Recently, it has became a hot topic, whether trees die from water shortage because of a depletion of C reserves (C starvation) or because of a failure of the hydraulic system [65,66 ], and several authors prioritized the old paradigm, assuming a dominant role of C starvation (e....
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TL;DR: New isotopic tools allow direct quantification of timescales involved in NSC dynamics, and show that NSC-C fixed years to decades previously is used to support tree functions.
Abstract: Contents 386 I. 386 II. 388 III. 392 IV. 392 V. 396 VI. 399 399 References 399 SUMMARY: Carbohydrates provide the building blocks for plant structures as well as versatile resources for metabolic processes. The nonstructural carbohydrates (NSC), mainly sugars and starch, fulfil distinct functional roles, including transport, energy metabolism and osmoregulation, and provide substrates for the synthesis of defence compounds or exchange with symbionts involved in nutrient acquisition or defence. At the whole-plant level, NSC storage buffers the asynchrony of supply and demand on diel, seasonal or decadal temporal scales and across plant organs. Despite its central role in plant function and in stand-level carbon cycling, our understanding of storage dynamics, its controls and response to environmental stresses is very limited, even after a century of research. This reflects the fact that often storage is defined by what we can measure, that is, NSC concentrations, and the interpretation of these as a proxy for a single function, storage, rather than the outcome of a range of NSC source and sink functions. New isotopic tools allow direct quantification of timescales involved in NSC dynamics, and show that NSC-C fixed years to decades previously is used to support tree functions. Here we review recent advances, with emphasis on the context of the interactions between NSC, drought and tree mortality.
441 citations
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TL;DR: Copyright (©) 1999–2012 R Foundation for Statistical Computing; permission is granted to make and distribute verbatim copies of this manual provided the copyright notice and permission notice are preserved on all copies.
Abstract: Copyright (©) 1999–2012 R Foundation for Statistical Computing. Permission is granted to make and distribute verbatim copies of this manual provided the copyright notice and this permission notice are preserved on all copies. Permission is granted to copy and distribute modified versions of this manual under the conditions for verbatim copying, provided that the entire resulting derived work is distributed under the terms of a permission notice identical to this one. Permission is granted to copy and distribute translations of this manual into another language, under the above conditions for modified versions, except that this permission notice may be stated in a translation approved by the R Core Team.
272,030 citations
TL;DR: A hydraulically based theory considering carbon balance and insect resistance that allowed development and examination of hypotheses regarding survival and mortality was developed, and incorporating this hydraulic framework may be effective for modeling plant survival andortality under future climate conditions.
Abstract: Summary Severe droughts have been associated with regional-scale forest mortality worldwide. Climate change is expected to exacerbate regional mortality events; however, pre- diction remains difficult because the physiological mechanisms underlying drought survival and mortality are poorly understood. We developed a hydraulically based theory considering carbon balance and insect resistance that allowed development and examination of hypotheses regarding survival and mortality. Multiple mechanisms may cause mortality during drought. A common mechanism for plants with isohydric
3,302 citations
TL;DR: In this paper, alternative formulations of Levene's test statistic for equality of variances are found to be robust under nonnormality, using more robust estimators of central location in place of the mean.
Abstract: Alternative formulations of Levene's test statistic for equality of variances are found to be robust under nonnormality. These statistics use more robust estimators of central location in place of the mean. They are compared with the unmodified Levene's statistic, a jackknife procedure, and a χ2 test suggested by Layard which are all found to be less robust under nonnormality.
2,559 citations
TL;DR: The role du couvert dans les echanges avec l'atmosphere is rappele puis integre dans l'analyse des reductions de bilan d'eau and de carbone in 2003 dus a regulation stomatique as discussed by the authors.
Abstract: La secheresse exceptionnelle de 2003 a ete l'occasion de faire le point de nos connaissances sur les mecanismes ecophysiologiques permettant aux arbres de traverser un tel evenement climatique extreme. L'analyse a ete conduite a l'echelle de l'arbre et du peuplement, tandis que l'intensite de la secheresse a ete quantifiee a l'aide d'un calcul de bilan hydrique sur neuf sites forestiers europeens contrastes du reseau CARBOEUROPE. Le role du couvert dans les echanges avec l'atmosphere est rappele puis integre dans l'analyse des reductions de bilan d'eau et de carbone en 2003 dus a la regulation stomatique. Les caracteristiques du complexe sol-racine, important a la fois pour l'acces a la ressource et a l'efficience de son absorption, constituent un des premiers traits d'adaptation a la secheresse. La reponse et les adaptations des especes ont surtout ete analysees en termes de diversite inter-specifique de fonctionnement hydraulique et du couplage entre proprietes hydrauliques et regulation stomatique. Enfin, nous discutons l'hypothese selon la quelle les dysfonctionnements hydrauliques ou les deficits de mise en reserve sont impliques dans les reactions differees de croissance, de developpement, d'induction de deperissement. Par exemple, des mesures de reserves glucidiques dans les troncs de chenes menees en fin d'ete 2003 ont permis de predire l'etat des couronnes des arbres au printemps 2004. Les faibles taux d'amidon etaient associes a une forte mortalite de branches et de jeunes pousses.
1,553 citations
980 citations