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      Tree defence and bark beetles in a drying world: carbon partitioning, functioning and modelling

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          Abstract

          Drought has promoted large-scale, insect-induced tree mortality in recent years, with severe consequences for ecosystem function, atmospheric processes, sustainable resources and global biogeochemical cycles. However, the physiological linkages among drought, tree defences, and insect outbreaks are still uncertain, hindering our ability to accurately predict tree mortality under on-going climate change. Here we propose an interdisciplinary research agenda for addressing these crucial knowledge gaps. Our framework includes field manipulations, laboratory experiments, and modelling of insect and vegetation dynamics, and focuses on how drought affects interactions between conifer trees and bark beetles. We build upon existing theory and examine several key assumptions: (1) there is a trade-off in tree carbon investment between primary and secondary metabolites (e.g. growth vs defence); (2) secondary metabolites are one of the main component of tree defence against bark beetles and associated microbes; and (3) implementing conifer-bark beetle interactions in current models improves predictions of forest disturbance in a changing climate. Our framework provides guidance for addressing a major shortcoming in current implementations of large-scale vegetation models, the under-representation of insect-induced tree mortality.

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          Most cited references67

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          Cross-scale Drivers of Natural Disturbances Prone to Anthropogenic Amplification: The Dynamics of Bark Beetle Eruptions

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            Plant defense against herbivores: chemical aspects.

            Plants have evolved a plethora of different chemical defenses covering nearly all classes of (secondary) metabolites that represent a major barrier to herbivory: Some are constitutive; others are induced after attack. Many compounds act directly on the herbivore, whereas others act indirectly via the attraction of organisms from other trophic levels that, in turn, protect the plant. An enormous diversity of plant (bio)chemicals are toxic, repellent, or antinutritive for herbivores of all types. Examples include cyanogenic glycosides, glucosinolates, alkaloids, and terpenoids; others are macromolecules and comprise latex or proteinase inhibitors. Their modes of action include membrane disruption, inhibition of nutrient and ion transport, inhibition of signal transduction processes, inhibition of metabolism, or disruption of the hormonal control of physiological processes. Recognizing the herbivore challenge and precise timing of plant activities as well as the adaptive modulation of the plants' metabolism is important so that metabolites and energy may be efficiently allocated to defensive activities.
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              A multi-species synthesis of physiological mechanisms in drought-induced tree mortality

              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.
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                Author and article information

                Journal
                New Phytologist
                New Phytol
                Wiley
                0028-646X
                1469-8137
                September 27 2019
                January 2020
                October 08 2019
                January 2020
                : 225
                : 1
                : 26-36
                Affiliations
                [1 ]Max Planck Institute for Biogeochemistry Hans‐Knöll‐Str. 10 07745 Jena Germany
                [2 ]Department of Forest Health Forest Research Institute Baden‐Württemberg 79100 Freiburg Germany
                [3 ]Department of Land Resources and Environmental Sciences Montana State University, Bozeman Bozeman MT 59717‐3120 USA
                [4 ]Department of Entomology University of Wisconsin Madison WI 53706 USA
                [5 ]Max Planck Institute for Chemical Ecology Hans‐Knöll‐Str. 8 07745 Jena Germany
                [6 ]Department of Zoology and Entomology, Forestry and Agricultural Biotechnology Institute University of Pretoria Private Bag X20 0028 Pretoria South Africa
                [7 ]Department of Plant Biology, Ecology, and Evolution Oklahoma State University Stillwater OK 74078 USA
                [8 ]Canadian Forest Service Natural Resources Canada Victoria BC V8Z 1M5 Canada
                [9 ]Los Alamos National Laboratory Los Alamos NM 87545 USA
                [10 ]School of the Environment Washington State University Pullman WA 99164‐2812 USA
                [11 ]Institute of Silviculture Department of Forest‐ and Soil Sciences University of Natural Resources and Life Sciences 1190 Vienna Austria
                Article
                10.1111/nph.16173
                31494935
                18318bb0-ee19-48b6-90aa-f43c0574d469
                © 2020

                http://creativecommons.org/licenses/by/4.0/

                http://doi.wiley.com/10.1002/tdm_license_1.1

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