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      Tree-ring isotopes suggest atmospheric drying limits temperature–growth responses of treeline bristlecone pine

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          Abstract

          Altitudinally separated bristlecone pine populations in the White Mountains (California, USA) exhibit differential climate–growth responses as temperature and tree-water relations change with altitude. These populations provide a natural experiment to explore the ecophysiological adaptations of this unique tree species to the twentieth century climate variability. We developed absolutely dated annual ring-width chronologies, and cellulose stable carbon and oxygen isotope chronologies from bristlecone pine growing at the treeline (~3500 m) and ~200 m below for the period AD 1710–2010. These chronologies were interpreted in terms of ecophysiological adaptations to climate variability with a dual-isotope model and a leaf gas exchange model. Ring widths show positive tree growth anomalies at treeline and consistent slower growth below treeline in relation to the twentieth century warming and associated atmospheric drying until the 1980s. Growth rates of both populations declined during and after the 1980s when growing-season temperature and atmospheric vapour pressure deficit continued to increase. Our model-based interpretations of the cellulose stable isotopes indicate that positive treeline growth anomalies prior to the 1980s were related to increased stomatal conductance and leaf-level transpiration and photosynthesis. Reduced growth since the 1980s occurred with a shift to more conservative leaf gas exchange in both the treeline and below-treeline populations, whereas leaf-level photosynthesis continued to increase in response to rising atmospheric CO 2 concentrations. Our results suggest that warming-induced atmospheric drying confounds positive growth responses of apparent temperature-limited bristlecone pine populations at treeline. In addition, the observed ecophysiological responses of attitudinally separated bristlecone pine populations illustrate the sensitivity of conifers to climate change.

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          A biochemical model of photosynthetic CO2 assimilation in leaves of C 3 species.

          Various aspects of the biochemistry of photosynthetic carbon assimilation in C3 plants are integrated into a form compatible with studies of gas exchange in leaves. These aspects include the kinetic properties of ribulose bisphosphate carboxylase-oxygenase; the requirements of the photosynthetic carbon reduction and photorespiratory carbon oxidation cycles for reduced pyridine nucleotides; the dependence of electron transport on photon flux and the presence of a temperature dependent upper limit to electron transport. The measurements of gas exchange with which the model outputs may be compared include those of the temperature and partial pressure of CO2(p(CO2)) dependencies of quantum yield, the variation of compensation point with temperature and partial pressure of O2(p(O2)), the dependence of net CO2 assimilation rate on p(CO2) and irradiance, and the influence of p(CO2) and irradiance on the temperature dependence of assimilation rate.
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            Some relationships between the biochemistry of photosynthesis and the gas exchange of leaves.

            A series of experiments is presented investigating short term and long term changes of the nature of the response of rate of CO2 assimilation to intercellular p(CO2). The relationships between CO2 assimilation rate and biochemical components of leaf photosynthesis, such as ribulose-bisphosphate (RuP2) carboxylase-oxygenase activity and electron transport capacity are examined and related to current theory of CO2 assimilation in leaves of C3 species. It was found that the response of the rate of CO2 assimilation to irradiance, partial pressure of O2, p(O2), and temperature was different at low and high intercellular p(CO2), suggesting that CO2 assimilation rate is governed by different processes at low and high intercellular p(CO2). In longer term changes in CO2 assimilation rate, induced by different growth conditions, the initial slope of the response of CO2 assimilation rate to intercellular p(CO2) could be correlated to in vitro measurements of RuP2 carboxylase activity. Also, CO2 assimilation rate at high p(CO2) could be correlated to in vitro measurements of electron transport rate. These results are consistent with the hypothesis that CO2 assimilation rate is limited by the RuP2 saturated rate of the RuP2 carboxylase-oxygenase at low intercellular p(CO2) and by the rate allowed by RuP2 regeneration capacity at high intercellular p(CO2).
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              A world-wide study of high altitude treeline temperatures

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                Author and article information

                Contributors
                Role: Handling Editor
                Journal
                Tree Physiol
                Tree Physiol
                treephys
                Tree Physiology
                Oxford University Press
                0829-318X
                1758-4469
                June 2019
                16 May 2019
                16 May 2019
                : 39
                : 6
                : 983-999
                Affiliations
                [1 ]Department of Environmental Sciences, Utrecht University, Utrecht, The Netherlands
                [2 ]Department of Geography, Swansea University, Swansea, UK
                [3 ]Department of Physical Geography, Utrecht University, Utrecht, The Netherlands
                [4 ]Department of Biosciences, Swansea University, Swansea, UK
                Author notes
                Corresponding author ( H.J.deBoer@ 123456uu.nl )
                Author information
                http://orcid.org/0000-0002-6933-344X
                Article
                tpz018
                10.1093/treephys/tpz018
                6642877
                30976807
                02504c39-0b5f-4f9a-bafe-4f5d1a13607c
                © The Author(s) 2019. Published by Oxford University Press.

                This is an Open Access article distributed under the terms of the Creative Commons Attribution Non-Commercial License ( http://creativecommons.org/licenses/by-nc/4.0/), which permits non-commercial re-use, distribution, and reproduction in any medium, provided the original work is properly cited. For commercial re-use, please contact journals.permissions@ 123456oup.com

                History
                : 22 October 2018
                : 24 April 2019
                Page count
                Pages: 17
                Categories
                Research Paper

                bristlecone pine (pinus longaeva d. k. bailey),cellulose stable isotopes,climate reconstruction,drought stress,tree hydraulics,tree rings,treeline,xylogenesis

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