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      How do trees die? A test of the hydraulic failure and carbon starvation hypotheses

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          Abstract

          Despite decades of research on plant drought tolerance, the physiological mechanisms by which trees succumb to drought are still under debate. We report results from an experiment designed to separate and test the current leading hypotheses of tree mortality. We show that piñon pine ( Pinus edulis) trees can die of both hydraulic failure and carbon starvation, and that during drought, the loss of conductivity and carbohydrate reserves can also co-occur. Hydraulic constraints on plant carbohydrate use determined survival time: turgor loss in the phloem limited access to carbohydrate reserves, but hydraulic control of respiration prolonged survival. Our data also demonstrate that hydraulic failure may be associated with loss of adequate tissue carbohydrate content required for osmoregulation, which then promotes failure to maintain hydraulic integrity.

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          Most cited references34

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          Mechanisms linking drought, hydraulics, carbon metabolism, and vegetation mortality.

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            Temperature sensitivity of drought-induced tree mortality portends increased regional die-off under global-change-type drought.

            Large-scale biogeographical shifts in vegetation are predicted in response to the altered precipitation and temperature regimes associated with global climate change. Vegetation shifts have profound ecological impacts and are an important climate-ecosystem feedback through their alteration of carbon, water, and energy exchanges of the land surface. Of particular concern is the potential for warmer temperatures to compound the effects of increasingly severe droughts by triggering widespread vegetation shifts via woody plant mortality. The sensitivity of tree mortality to temperature is dependent on which of 2 non-mutually-exclusive mechanisms predominates--temperature-sensitive carbon starvation in response to a period of protracted water stress or temperature-insensitive sudden hydraulic failure under extreme water stress (cavitation). Here we show that experimentally induced warmer temperatures (approximately 4 degrees C) shortened the time to drought-induced mortality in Pinus edulis (piñon shortened pine) trees by nearly a third, with temperature-dependent differences in cumulative respiration costs implicating carbon starvation as the primary mechanism of mortality. Extrapolating this temperature effect to the historic frequency of water deficit in the southwestern United States predicts a 5-fold increase in the frequency of regional-scale tree die-off events for this species due to temperature alone. Projected increases in drought frequency due to changes in precipitation and increases in stress from biotic agents (e.g., bark beetles) would further exacerbate mortality. Our results demonstrate the mechanism by which warmer temperatures have exacerbated recent regional die-off events and background mortality rates. Because of pervasive projected increases in temperature, our results portend widespread increases in the extent and frequency of vegetation die-off.
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              Starch turnover: pathways, regulation and role in growth.

              Many plants store part of their photosynthate as starch during the day and remobilise it to support metabolism and growth at night. Mutants unable to synthesize or degrade starch show strongly impaired growth except in long day conditions. In rapidly growing plants, starch turnover is regulated such that it is almost, but not completely, exhausted at dawn. There is increasing evidence that premature or incomplete exhaustion of starch turnover results in lower rates of plant growth. This review provides an update on the pathways for starch synthesis and degradation. We discuss recent advances in understanding how starch turnover and the use of carbon for growth is regulated during diurnal cycles, with special emphasis on the role of the biological clock. Much of the molecular and genetic research on starch turnover has been performed in the reference system Arabidopsis. This review considers to what extent information gained in this weed species maybe applicable to annual crops and perennial species. Copyright © 2012. Published by Elsevier Ltd.
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                Author and article information

                Journal
                Plant Cell Environ
                Plant Cell Environ
                pce
                Plant, Cell & Environment
                BlackWell Publishing Ltd (Oxford, UK )
                0140-7791
                1365-3040
                January 2014
                30 June 2013
                : 37
                : 1
                : 153-161
                Affiliations
                [1 ]Earth and Environmental Sciences Division, Los Alamos National Laboratory Los Alamos, NM, 87545, USA
                [2 ]Department of Biology, University of New Mexico 219 Yale Blvd., Albuquerque, NM, 87131, USA
                Author notes
                Correspondence: S. Sevanto. e-mail: sanna@ 123456lanl.gov
                Article
                10.1111/pce.12141
                4280888
                23730972
                d061423c-c123-49ef-9cee-542eb8c461b9
                Published 2013. This article is a U.S. Government work and is in the public domain in the USA.

                This is an open access article under the terms of the Creative Commons Attribution-NonCommercial License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited and is not used for commercial purposes.

                History
                : 14 March 2013
                : 20 May 2013
                : 03 June 2013
                Categories
                Original Articles

                Plant science & Botany
                cavitation,forest mortality,hydraulic conductance,non-structural carbohydrates,phloem,xylem

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