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<article language="en">
	<journal>
		<journal_title>Biogeosciences</journal_title>
		<journal_url>www.biogeosciences.net</journal_url>
		<issn>1726-4170</issn>
		<eissn>1726-4189</eissn>
		<volume_number>6</volume_number>
		<issue_number>8</issue_number>
		<publication_year>2009</publication_year>
	</journal>
	<doi>10.5194/bg-6-1423-2009</doi>
	<article_url>http://www.biogeosciences.net/6/1423/2009/</article_url>
	<abstract_html>http://www.biogeosciences.net/6/1423/2009/bg-6-1423-2009.html</abstract_html>
	<fulltext_pdf>http://www.biogeosciences.net/6/1423/2009/bg-6-1423-2009.pdf</fulltext_pdf>
	<start_page>1423</start_page>
	<end_page>1444</end_page>
	<publication_date>2009-08-05</publication_date>
	<article_title content_type="html">Improved understanding of drought controls on seasonal variation in  Mediterranean forest canopy CO&lt;sub&gt;2&lt;/sub&gt; and water fluxes through combined in situ measurements and ecosystem modelling</article_title>
	<authors>
		<author numeration="1" affiliations="1">
			<name>T. Keenan</name>
			<email>t.keenan@creaf.uab.es</email>
		</author>
		<author numeration="2" affiliations="2">
			<name>R. GarcÃ­a</name>
		</author>
		<author numeration="3" affiliations="3">
			<name>A. D. Friend</name>
		</author>
		<author numeration="4" affiliations="2">
			<name>S. Zaehle</name>
		</author>
		<author numeration="5" affiliations="1,4">
			<name>C. Gracia</name>
		</author>
		<author numeration="6" affiliations="1,4">
			<name>S. Sabate</name>
		</author>
	</authors>
	<affiliations>
		<affiliation numeration="1" content_type="html">CREAF, Autonomous University of Barcelona (UAB), 08139, Bellaterra, Barcelona, Spain</affiliation>
		<affiliation numeration="2" content_type="html">LSCE, Orme des Merisiers, Gif sur Yvette, Paris, France</affiliation>
		<affiliation numeration="3" content_type="html">Department of Geography, University of Cambridge, Cambridge CB2 3EN, UK</affiliation>
		<affiliation numeration="4" content_type="html">Department of Ecology, University of Barcelona (UB), Diagonal 645, 08028, Spain</affiliation>
	</affiliations>
	<abstract content_type="html">Water stress is a defining characteristic of Mediterranean ecosystems,
and is likely to become more severe in the coming decades.
Simulation models are key tools for making predictions, but
our current understanding of how soil moisture controls ecosystem
functioning is not sufficient to adequately constrain
parameterisations.
&lt;br&gt;&lt;br&gt;
Canopy-scale flux data from four forest ecosystems with
Mediterranean-type climates were used in order to analyse the
physiological controls on carbon and water flues through the
year. Significant non-stomatal limitations on photosynthesis were
detected, along with lesser changes in the conductance-assimilation relationship.
New model parameterisations were derived and implemented in
two contrasting modelling approaches.
&lt;br&gt;&lt;br&gt;
The effectiveness of two models, one a dynamic global vegetation model (&quot;ORCHIDEE&quot;), and the
other a forest growth model particularly developed for Mediterranean
simulations (&quot;GOTILWA+&quot;), was assessed and modelled
canopy responses to seasonal changes in soil moisture were analysed
in comparison with in situ flux measurements.
&lt;br&gt;&lt;br&gt;
In contrast to commonly held assumptions, we find that changing the ratio of
conductance to assimilation under natural,
seasonally-developing, soil moisture stress is not sufficient to reproduce
forest canopy CO&lt;sub&gt;2&lt;/sub&gt; and water fluxes. However, accurate
predictions of both CO&lt;sub&gt;2&lt;/sub&gt; and water fluxes under all soil moisture
levels encountered in the field are obtained if photosynthetic capacity is
assumed to vary with soil moisture. This new parameterisation has
important consequences for simulated responses of carbon and water
fluxes to seasonal soil moisture stress, and should greatly improve our
ability to anticipate future impacts of climate changes on the
functioning of ecosystems in Mediterranean-type climates.</abstract>
	<references>
		<reference numeration="1" content_type="text"> Allen, H D.: Mediterranean Ecogeography, Prentice Hall, London, UK, 2001. </reference>
		<reference numeration="2" content_type="text"> Allard, V., Ourcival, J M., Rambal, S., Joffre, R., and Rocheteau, A.: Seasonal and annual variation of carbon exchange in an evergreen Mediterranean forest in southern France, Glob. Change Biol., 4, 714â€“715, 2008. </reference>
		<reference numeration="3" content_type="text"> Baldocchi, D., Falge, E., Gu, L., Olson, R., Hollinger, D., Running, S., Anthoni, P. Bernhofer, Ch., Davis, K., Fuentes, J., Goldstein, A., Katul, G., Law, B., Lee, X., Malhi, Y., Meyers, T., Munger, J. W., Oechel, W., Pilegaard, K., Schmid, H.P., Valentini, R., Verma, S., Vesala, T., Wilson, K., and Wofsy, S.: FLUXNET: A new tool to study the temporal and spatial variability of ecosystem-scale carbon dioxide, water vapor and energy flux densities, Bulletin of the American Meteorological Society, 82, 2415â€“2435, 2001. </reference>
		<reference numeration="4" content_type="text"> Ball, J T., Woodrow, I E., and Berry, J A.: A~model predicting stomatal conductance and its contribution to the control of photosynthesis under different environmental conditions, in: Progress in Photosynthesis Research, edited by: Biggins, J. and Nijhoff, M., 4, 221â€“224, Martinus-Nijhoff Publishers, Dordrecht, The Netherlands, 1987. </reference>
		<reference numeration="5" content_type="text"> Bernacchi, C J., Singsaas, E L., Pimentel, C., Portis Jr., ~A R., Long, S P.: Improved temperature response functions for models of Rubisco-limited photosynthesis, Plant Cell Environ., 24, 253â€“259, 2001. </reference>
		<reference numeration="6" content_type="text"> Blyth, E M., Dolman, A J., and Noilhan, J.: The effect of forest on mesoscale rainfall â€“ An example from Hapex-Mobilhy, J. Appl. Meteorol., 33, 445â€“454, 1994. </reference>
		<reference numeration="7" content_type="text"> BrÃ©da, N., Huc, R., Granier, A., and Dreyer, E.: Temperate forest trees and stands under severe drought: a review of ecophysiological responses, adaptation processes and long-term consequences, Ann. For. Sci., 63, 625â€“644, 2006. </reference>
		<reference numeration="8" content_type="text"> Buckley, T N., Farquhar, G D., and Mott, K A.: Qualitative effects of patchy stomatal conductance distribution features on gas-exchange calculations, Plant Cell Environ., 20, 867â€“880, 1997. </reference>
		<reference numeration="9" content_type="text"> Byers, F M., Schelling, G T., and Greene, L W.: Development of growth functions to describe energy density of growth in beef cattle, in: Proceedings of Energy Metabolism of Farm Animals, edited by: van-der Honing, Y., and Close, W H., Pudoc, Wageningen, The Netherlands, 43, 195â€“198, 1989. </reference>
		<reference numeration="10" content_type="text"> Chaves, M M., Pereira, J S., Maroco, J., Rodrigues, M L., Ricardo, C P. P., Osorio, M L., Carvalho, I., Faria, T., and Pinheiro, C.: How plants cope with water stress in the field. Photosynthesis and growth, Ann. Bot.-London, 89, 907â€“916, 2002. </reference>
		<reference numeration="11" content_type="text"> Colello, G D., Grivet, C., Sellers, P J., and Berry, J A.: Modeling of energy, water, and \chemCO_2 flux in a temperate grassland ecosystem with SiB2: Mayâ€“October 1987, J. Atmos. Sci., 55, 1141â€“1169, 1998. </reference>
		<reference numeration="12" content_type="text"> Collatz, G J., Ball, J T., Grivet, C., and Berry, J A.: Physiological and environmental regulation of stomatal conductance, photosynthesis and transpiration: a model that includes a laminar boundary layer, Agr. For. Met., 54, 107â€“136, 1991. </reference>
		<reference numeration="13" content_type="text"> Collatz, G J., Ribas-Carbo, M., and Berry, J A.: Coupled Photosynthesis-Stomatal conductance model for leaves of C4 plants, Aust. J. Plant Physiol., 19, 519â€“538, 1992. </reference>
		<reference numeration="14" content_type="text"> Cowan, I R.: Stomatal behaviour and environment, Adv. Bot. Res. 4, 117â€“228, 1977. </reference>
		<reference numeration="15" content_type="text"> Daikoku, K., Hattori, S., Deguchi, A., Aoki, Y., Miyashita, M., Matsumoto, K., Akinyama, J., Iida, S., Toba, T., Fujita, Y., Ohta, T.: Influence of evaporation from the forest floor on evapotranspiration from the dry canopy, Hydrol. Proc., 20, 4083â€“4096, 2008. </reference>
		<reference numeration="16" content_type="text"> Deardoff, J W.: Empirical evidence of the eddy coefficient for heat upon stability above the lowest 50 m, J. Appl. Meteor., 6, 631-643, 1967. </reference>
		<reference numeration="17" content_type="text"> De Pury, D G G., and Farquhar, G D.: Simple scaling of photosynthesis from leaves to canopies without the errors of big-leaf models. Plant Cell Environ., 20, 537â€“557, 1997. </reference>
		<reference numeration="18" content_type="text"> Dore, S., Hymus, G J., Johnson, D P., Hinkle, C R., Valentini, R., and Drake, B.: Cross validation of open-top chamber and eddy covariance measurements of ecosystem CO&lt;sub&gt;2&lt;/sub&gt; exchange in a Florida scrub-oak ecosystem. Glob. Change Biol., 9, 84-95, 2003. </reference>
		<reference numeration="19" content_type="text"> Dufrene, E., Davi, H., Francois, C., le Maire, G., le Dantec, V., and Granier, A.: Modelling carbon and water cycles in a beech forest Part I: Model description and uncertainty analysis on modelled NEE, Ecol. Model., 185, 407â€“436, 2005. </reference>
		<reference numeration="20" content_type="text"> Evans, J R. and von Caemmerer, S.: Carbon dioxide diffusion inside leaves. Plant Physiol., 110, 339-346, 1996. </reference>
		<reference numeration="21" content_type="text"> Evans, J R., Loreto, F.: Acquisition and diffusion of CO&lt;sub&gt;2&lt;/sub&gt; in higher plant leaves, in: Photosynthesis: physiology and metabolism, edited by: Leegood, R C., Sharkey, T D., and Von Caemmerer, S., Kluwer Academic Publishers, Dortrecht, The Netherlands, 321-351, 2000. </reference>
		<reference numeration="22" content_type="text"> Evans, J. R., Terashima, I., Hanba, Y., and Loreto, F.: CO&lt;sub&gt;2&lt;/sub&gt; capture: chloroplast to leaf, in: Photosynthetic adaptation: chloroplast to landscape, edited by: Smith, W. K., Vogelmann, T. C., Critchley C, Springer, New York, USA, 107â€“132, 2005. </reference>
		<reference numeration="23" content_type="text"> Farquhar, G D., Caemmerer, S V., and Berry, J A.: A~Biochemical model of Photosynthetic CO&lt;sub&gt;2&lt;/sub&gt; assimilation in leaves of C3 species, Planta, 149, 78â€“90, 1980. </reference>
		<reference numeration="24" content_type="text"> Field, C B., Randerson, J T., and Malmstrom, C M.: Global Net Primary Production â€“ Combining ecology and remote sensing, Remote Sens. Environ., 51, 74â€“88, 1995. </reference>
		<reference numeration="25" content_type="text"> Flexas, J. and Medrano, H.: Drought inhibition of photosynthesis in C&lt;sub&gt;3&lt;/sub&gt;plants: Stomatal and non-stomatal limitations revisited, Ann. Bot-London, 89, 183â€“189, 2002. </reference>
		<reference numeration="26" content_type="text"> Flexas, J., Bota, J., Loreto, F., Cornic, G., and Sharkey, T D.: Diffusive and metabolic limitations to photosynthesis under drought and salinity in C3 plants, Plant Biol., 6, 269â€“279, 2004. </reference>
		<reference numeration="27" content_type="text"> Flexas, J., Ribas-CarbÃ³, M., Diaz-Espejo, A., GalmÃ©s, J., and Medrano, H.: Mesophyll conductance to \chemCO_2: current knowledge and future prospects, Plant Cell Environ., 31, 602â€“621, 2008. </reference>
		<reference numeration="28" content_type="text"> Friedlingstein, P., Joel, G., Field, C B., and Fung, I Y.: Toward an allocation scheme for global terrestrial carbon models, Glob. Change Biol., 5, 755â€“770, 1999. </reference>
		<reference numeration="29" content_type="text"> Friend, A D., Arneth, A., Kiang, N. Y., Lomas, M., Ogee, J., Rodenbeck, C., Running, S. W., Santaren, J-D, Sitch, S., Viovy, N., Woodwards, F. I., and Zaehle, S.: FLUXNET and modelling the global carbon cycle, Glob. Change Biol., 13, 610â€“633, 2007. </reference>
		<reference numeration="30" content_type="text"> Friend, A D. and Kiang, N Y.: Land Surface Model Development for the GISS GCM: Effects of Improved Canopy Physiology on Simulated Climate, J. Clim., 18, 2883â€“2902, 2005. </reference>
		<reference numeration="31" content_type="text"> Galmes, J., Flexas, J., Keys, A., Cifre, J., Mitchell, R., Madgwick, P., Haslam, R., Medrano, H., and Parry, J.: Rubisco specificity factor tends to be larger in plant species from drier habitats and in species with persistent leaves, Plant Cell Environ., 28, 571â€“579, 2005. </reference>
		<reference numeration="32" content_type="text"> Galmes, J., Medrano, H., and Flexas, J.: Photosynthetic limitations in response to water stress and recovery in Mediterranean plants with different growth forms, New Phytol., 175, 81-93, 2007. </reference>
		<reference numeration="33" content_type="text"> Giorgi, F.: Climate change hot-spots, Geophys. Res. Lett., 33, L08707, doi:10.1029/2006GL025734, 2006. </reference>
		<reference numeration="34" content_type="text"> Giorgi, F., Bi, X., and Pal, J.: Mean, interannual variability and trends in a regional climate change experiment over Europe. II: climate change scenarios (2071â€“2100), Clim. Dynam., 23, 839â€“858, 2004. </reference>
		<reference numeration="35" content_type="text"> Goldstein, A H., Hultman, N E., Fracheboud, J M., Bauer, M R., Panek, J. A., Xu, M., Qi, Y., Guenther, A B., and Baugh W.: Effects of climate variability on the carbon dioxide, water, and sensible heat fluxes above a ponderosa pine plantation in the Sierra Nevada (CA), Agr. For. Meteorol., 101, 113â€“129, 2000. </reference>
		<reference numeration="36" content_type="text"> Gracia C A., Tello, E., Sabate, S., and Bellot, J.: GOTILWA+: An integrated model of water dynamics and forest growth. Ecology of Mediterranean evergreen oak forests, in: Ecology of Mediterranean Evergreen Oak Forests, edited by: RodÃ , F., Retana, J., Bellot, J., and Gracia, C A., Springer, Berlin, Germany, 163â€“178, 1999. </reference>
		<reference numeration="37" content_type="text"> Granier, A., Reichstein, M., BrÃ©da, N., Janssens, I A., Falfe, E., Ciais, P., GrÃ¼nwald, T., Aubinet, M., Berbigier, P., Bernhofer, C., Buchmann, N., Facini, O., Grassi, G., Heinesch, B., Ilvesniemi, H., Keronen, P., Knohl, A., KÃ¶stner, B., Lagergren, F., Lindroth, A., Longdoz, B., Loustau, D., Mateus, J., Montagnani, L., Nys, C., Moors, E., Papale, D., Peiffer, M., Pilegaard, K., Pita, G., Pumpanen, J., Rambal, S., Rebmann, C., Rodrigues, A., Seufert, G., Tenhunen, J., Vesala, T., and Wang, Q.: Evidence for soil water control on carbon and water dynamics in European forests during the extremely dry year, Agr. Forest Meteorol., 143, 123â€“145, 2007. </reference>
		<reference numeration="38" content_type="text"> Grassi, G. and Magnani, F.: Stomatal, mesophyll conductance and biochemical limitations to photosynthesis as affected by drought and leaf ontogeny in ash and oak trees, Plant Cell Environ., 28, 834â€“849, 2005 </reference>
		<reference numeration="39" content_type="text"> Gunasekera, D. and Berkowitz, G A.: Heterogeneous stomatal closure in response to leaf water deficits is not a universal phenomenon, Plant Physiol., 98, 660â€“665, 1992. </reference>
		<reference numeration="40" content_type="text"> Hansen, J., Russell, G., Rind, D., Stone, P., Lacis, A., Lebedeff, S., Ruedy, R., and Travis, L.: Efficient three-dimensional global models for climate studies: Models I and II. Mon. Weather Rev., 111, 609â€“662, 1983. </reference>
		<reference numeration="41" content_type="text"> Harley, P C. and Tenhunen, J D.: Modelling the photosynthetic response of C3 leaves to environmental factors, in: Modeling Crop Photosynthesis â€“ from Biochemistry to Canopy, edited by: Boot, K. J and Loomis, R. S., Am. Soc. Crop Sci., Madison, WI, USA, 19, 17â€“39, 1991. </reference>
		<reference numeration="42" content_type="text"> Hickler, T., Prentice, C., Smith, B., Sykes, M., and Zaehle, S.: Implementing the plant hydraulic architecture within the LPJ Dynamic Global Vegetation Model, Global Ecol. Biogeogr., 15, 567â€“577, 2006. </reference>
		<reference numeration="43" content_type="text"> Hoff, C., Rambal, S., and Joffre, R.: Simulating carbon and water flows and growth in a Mediterranean evergreen \textitQuercus ilex coppice using the FOREST-BGC model, Forest Ecol. Manag., 164, 121â€“136, 2002. </reference>
		<reference numeration="44" content_type="text"> Honeysett, J. L. and Ratkowsky, D. A.: The use of ignition loss to estimate bulk density of forest soils, J. Soil Sc., 40, 299â€“308, 1989. </reference>
		<reference numeration="45" content_type="text"> IPCC: Climate Change 2007: The physical science basis. Contribution of working groups I to the Fourth Assessment Report of the Intergovernamental Panel on Climate Change, edited by: Solomon, S., Qin, D., Manning, M., et al., Cambridge University Press, Cambridge, UK and New York, NY, USA, 2007. </reference>
		<reference numeration="46" content_type="text"> Jones, H G.: Partitioning stomatal and non-stomatal limitations to photosynthesis, Plant Cell Environ., 8, 95â€“104, 1985. </reference>
		<reference numeration="47" content_type="text"> Jump, A., Hunt, J., and Penuelas, J.: Rapid climate change-related growth decline at the southern range edge of \textitFagus sylvatica, Glob. Change Biol., 12, 2163â€“2174, 2006. </reference>
		<reference numeration="48" content_type="text"> Jung, M., Le Maire, G., Zaehle, S., Luyssaert, S., Vetter, M., Churkina, G., Ciais, P., Viovy, N., and Reichstein, M.: Assessing the ability of three land ecosystem models to simulate gross carbon uptake of forests from boreal to Mediterranean climate in Europe, Biogeosciences, 4, 647â€“656, 2007. </reference>
		<reference numeration="49" content_type="text"> Keenan, T., Sabate, S., and Gracia, C.: Forest ecophysiological models and carbon sequestration, in: Managing Forest Ecosystems â€“ The Challange of Climate Change, edited by: Bravo, F., LeMay, V., Jandl, R., and Gadow, K V., Springer, Berlin, ISBN: 978-1-4020-8342-6, 2008. </reference>
		<reference numeration="50" content_type="text"> Keenan, T., Sabate, S., and Gracia, C.: The importance of mesophyll conductance in regulating forest ecosystem productivity during drought periods, Glob. Change Biol., in press, 2009. </reference>
		<reference numeration="51" content_type="text"> Kowalski, A S., Loustau, D., Berbigier, P., Manca, G., Tedeschi, V., Borghetti, M., Valentini, R., Kolari, P., Berninger, F., Rannik, U., Hari, P., Rayment, M., Mencuccini, M., Moncrieff, J., and Grace, J.: Paired comparisons of carbon exchange between undisturbed and regenerating stands in four managed forests in Europe, Global Change Biol., 10, 1707â€“1723, 2004. </reference>
		<reference numeration="52" content_type="text"> Kramer, P J.: Water Relations of Plants. Academic Press, New York, USA, 489 pp., 1983. </reference>
		<reference numeration="53" content_type="text"> Kramer, K., Leinonen, I., Baterlink, H. H., Berbigier, P., Borguetti, M., Bernhofer, C., Cienciala, E., Dolman, A J., Froer, O., Gracia, C A., Granier, A., GrÃ¼nwald, T., Hari, P., Jans, W., Kellomak\&quot;&amp;#x0131;, S., Loustau, D., Magnani, F., Markkanen, T., Matteucci, G., Mohren, M J., Moors, E., Nissinen, A., Peltola, H., SabatÃ©, S., Sanchez, A., Sontag, M., Valentini, R., and Vesala, I.: Evaluation of six processes-based forest growth models using eddy-covariance measurements of \chemCO_2 and \chemH_2O fluxes at six forest sites in Europe, Glob. Change Biol., 8, 213â€“230, 2002. </reference>
		<reference numeration="54" content_type="text"> Krinner, G., Viovy de Noblet-DucoudrÃ©, N., OgÃ©e, J., Polcher, J., Friendlingstein, P., Ciais, P., Sitch, S., and Prentice, I C.: A~dynamic global vegetation model for studies of the coupled atmosphere-biosphere system, Global Biogeochem. Cy., 19, GB1015, doi:10.1029/2003GB002199, 2005. </reference>
		<reference numeration="55" content_type="text"> Kubiske, M E. and Adams, M D.: Stomatal and non-stomatal limitations of photosynthesis in 19 temperate tree species on contrasting sites during wet and dry years, Plant Cell Environ., 16, 1123â€“1129, 1993. </reference>
		<reference numeration="56" content_type="text"> Kutbay, H G. and Kilinc, M.: Sclerophylly in \textitQuercus cerris L. var. cerris and \textitPhillyrea larifolia L. and edaphic relations of these species, Plant Ecol., 113, 93â€“97, 1994. </reference>
		<reference numeration="57" content_type="text"> Lawlor, D W.: The effects of water deficit on photosynthesis, in: Environment and Plant Metabolism, edited by: Smirnoff, N., Bios Scientific Publishers, Oxford, UK, 129â€“160, 1995. </reference>
		<reference numeration="58" content_type="text"> Lawlor, D W. and Cornic, G.: Photosynthetic carbon assimilation and associated metabolism in relation to water deficits in higher plants, Plant Cell Environ., 25, 275â€“294, 2002. </reference>
		<reference numeration="59" content_type="text"> Leuning, R., Kelliher, F M., de Pury, D G G., and Schulze, E.-D.: Leaf Nitrogen, Photosynthesis, Conductance and Transpiration: Scaling from leaves to canopies, Plant Cell Environ., 18, 1183â€“1200, 1995. </reference>
		<reference numeration="60" content_type="text"> Leuning R.: Modelling stomatal behaviour and photosynthesis of Eucalyptus grandis, Aust. J. Plant Physiol., 17, 159â€“175, 1990. </reference>
		<reference numeration="61" content_type="text"> Leuning R.: A~critical appraisal of a combined stomatal photosynthesis model for C3 plants, Plant Cell Environ., 18, 339â€“355, 1995. </reference>
		<reference numeration="62" content_type="text"> Loague, K. and Green, R E.: Statistical and graphical methods for evaluating solute transport models: Overview and application, J. Contam. Hydrol., 7, 51â€“73, 1991. </reference>
		<reference numeration="63" content_type="text"> Loreto, F., Harley, P C., Di Marco, G., and Sharkey, T D.: Estimation of mesophyll conductance to \chemCO_2 flux by three different methods, Plant Physiol., 98, 1437â€“1443, 1992. </reference>
		<reference numeration="64" content_type="text"> Loreto, F. and Centritto, M.: Leaf Carbon assimilation in a water-limited world, Plant Biosyst., 142, 154â€“161, 2008. </reference>
		<reference numeration="65" content_type="text"> Maroco, J P., Rodrigues, M L., Lopes, C., and Chaves, M.: Limitations to leaf photosynthesis in field-grown grapevine under drought â€“ metabolic and modelling approaches, Funct. Plant Biol., 29, 451â€“459, 2002. </reference>
		<reference numeration="66" content_type="text"> Mayer, D G. and Butler, D G.: Statistical validation, Ecol. Model., 68, 21â€“32, 1993. </reference>
		<reference numeration="67" content_type="text"> Medrano, H., Parry, M., Socias, X., and Lawlor, D.: Long term water stress inactivates Rubisco in subterranean clover, Ann. Appl. Biol., 131, 491â€“501, 1997. </reference>
		<reference numeration="68" content_type="text"> Medrano, H., Escalona, J M., Bota, J., Julias, J., and Flexas, J.: Regulation of photosynthesis of C3 plants in response to progressive drought: Stomatal conductance as a reference parameter, Ann. Bot., 89, 895â€“905, 2002. </reference>
		<reference numeration="69" content_type="text"> Misson, L., Panek, J A., and Goldstein, A H.: A~comparison of three approaches to modelling leaf gas exchange in annually drought-stressed ponderosa pine forests, Tree Physiol., 24, 529â€“541, 2004. </reference>
		<reference numeration="70" content_type="text"> Misson, L., Tu, K P., Boniello, R A., and Goldstein, A H.: Seasonality of photosynthetic parameters in a multi-specific and vertically complex forest ecosystem in the Sierra Nevada of California, Tree Physiol., 26, 729â€“741, 2006. </reference>
		<reference numeration="71" content_type="text"> Morales, P., Sykes, M T., Prentice, I C., Smith, P., Smith, B., Bugmann, H., Zierl, B., Friedlingstein, P., Viovy, N., Sabate, S., Sanchez, A., Pla, A., Gracia, C A., Sitch, S., Arneth, A M., and Ogee, J.: Comparing and evaluating process-based ecosystem model predictions of carbon and water fluxes in major European forest biomes, Glob. Change Biol., 11, 2211â€“2233, 2005. </reference>
		<reference numeration="72" content_type="text"> Mott, K A. and Buckley, T N.: Patchy stomatal conductance: emergent collective behavior of stomata, Trends Plant Sci., 5, 258â€“262, 2000. </reference>
		<reference numeration="73" content_type="text"> McMurtrie, R., Rook D., and Kelliher F.: Modelling the yield of pinus radiata on a site limited by water and nitrogen. For. Ecol. Manag., 30, 381â€“413, 1990. </reference>
		<reference numeration="74" content_type="text"> Ni, B R. and Pallardy, S G.: Stomatal and non-stomatal limitations to net photosynthesis in seedlings of woody angiosperms, Plant Physiol., 99, 1502â€“1508, 1992. </reference>
		<reference numeration="75" content_type="text"> Owen, P R. and Thompson, W R.: Heat transfer across rough surfaces, J. Fluid Mech., 15, 321â€“334, 1963. </reference>
		<reference numeration="76" content_type="text"> Papale, D. and Valentini, A.: A new assessment of European forests carbon exchanges by eddy fluxes and artificial neural network spatialization, Glob. Change Biol., 9, 525â€“535, 2003. </reference>
		<reference numeration="77" content_type="text"> Parton, W J., Scurlock, J M O., Ojima, D S., Gilmanov, T G., Scholes, R J., Schimmel, D S., Kirchner, T., Menaut, J C., Seastedt, T., Moya, E. G., Kamnalrut, A., and Kinyamario, J I.: Observations and modelling of biomass and soil organic matter dynamics for the grassland biome worldwide, Global Biogeochem. Cy., 7, 785â€“809, 1993. </reference>
		<reference numeration="78" content_type="text"> Parry, M., Andralojc, P., Khan, S., Lea, P., and Keys, A J.: Rubisco activity: Effects of drought stress, Ann. Bot., 89, 833â€“839, 2002. </reference>
		<reference numeration="79" content_type="text"> Peñuelas, J., and Boada, M.: A global change-induced biome shift in the Montseny mountains (NE Spain), Glob. Change Biol., 9, 131â€“140, 2003. </reference>
		<reference numeration="80" content_type="text"> Qualls, R. and Hopson, T.: Combined use of vegetation density, friction velocity and solar elevation to parameterize the scalar roughness for sensible heat, J. Atmos. Sci., 55, 1198â€“1208. 1998. </reference>
		<reference numeration="81" content_type="text"> Rambal, S., Joffre, R., Ourcival, J M., Cavender-Bares, J., and Rocheteau, A.: The growth respiration component in eddy \chemCO_2 flux from a \textitQuercus Ilex Mediterranean forest, Glob. Change Biol., 10, 1460â€“1469, 2004. </reference>
		<reference numeration="82" content_type="text"> Rambal, S, Ourcival, J M., Joffre, R., Mouillot, F., Nouvellon, Y., Reichstein, M., and Rocheteau, A.: Drought controls over conductance and assimilation of a Mediterranean evergreen ecosystem: scaling from leaf to canopy, Glob. Change Biol., 9, 1813â€“1824, 2003. </reference>
		<reference numeration="83" content_type="text"> Reichstein, M., Tenhunen, J., Roupsard, O., Ourcival, J M., Rambal, S., Miglietta, F., Peressotti, A., Pecchiari, M., Giampiero, T., and Valentini R.: Severe drought effects on ecosystem CO&lt;sub&gt;2&lt;/sub&gt; and H&lt;sub&gt;2&lt;/sub&gt;O fluxes at three Mediterranean evergreen sites: revision of current hypotheses? Glob. Change Biol., 8, 999â€“1017, 2003. </reference>
		<reference numeration="84" content_type="text"> Reichstein, M., Tenhunen, J., Roupsard, O., Ourcival, J M., Rambal, S., Miglietta, F., Peressotti, A., Pecchiari, M., Giampiero, T., and Valentini, R.: Inverse modeling of seasonal drought effects on canopy CO&lt;sub&gt;2&lt;/sub&gt;/H2O exchange in threee Mediterranean ecosystems, J. Geophys Res., 108(D23), 4726, doi:10.1029/2003JD003430, 2003. </reference>
		<reference numeration="85" content_type="text"> Reichstein, M., Falge, E., Baldocchi, D., Papale, D., Aubinet, M., Berbigier, P., Bernhofer, C., Buchmann, N., Gilmanov, T., Granier, A., Grunwald, T., Havrankova, K., Ilvesmiemi, H., Janous, D., Knohl, A., Laurila, T., Lohila, A., Loustau, D., Matteucci, G., Meyers, T., Miglietta, F., Ourcival, J M., Pumpanen, J., Rambal, S., Rotenberg, E., Sanz, M., Tenhunen, J., Seufert, G., Vaccari, F., Vesala, T., Yakir, D., and Valentini, R.: On the separation of net ecosystem exchange into assimilation and ecosystem respiration: review and improved algorithm, Glob. Change Biol., 11, 1424â€“1439, 2005. </reference>
		<reference numeration="86" content_type="text"> Reichstein, M., Papale, D., Valentini, R., Aubinet, M., Bernhofer, C., Knohl, A., Laurila, T., Lindroth, A., Moors, E., Pilegaard, K., and Seufert, G.: Determinants of terrestrial ecosystem carbon balance inferred from European eddy covariance flux sites, Geophys. Res. Lett., 34, L01402, doi:10.1029/2006GL027880, 2007. </reference>
		<reference numeration="87" content_type="text"> Sala, A. and Tenhunen J D.: Site-specific water relations and stomatal response of \textitQuercus ilex L. in a Mediterranean watershed, Tree Physiol. 14, 601â€“617, 1994. </reference>
		<reference numeration="88" content_type="text"> Santaren, D., Peylin, P., Viovy, N., and Ciais, P.: Optimizing a process-based ecosystem model with eddy-covariance flux measurements: a pine forest in southern France, Global Biogeochem. Cy., 21, GB2013, doi:10.1029/2006GB002834, 2007. </reference>
		<reference numeration="89" content_type="text"> SchrÃ¶ter, D., Cramer, W., Leemans, R., Prentice, I C., Ara\&apos;ujo, M. B., Arnell, N W., Bondeau, A., Bugmann, H., Carter, T R., Gracia, C A., de la Vega-Leinert, A C., Erhard, M., Ewert, F., Glendining, M., House, J. I., KankaanpÃ¤Ã¤, S., Klein, R J T., Lavorel, S., Lindner, M., Metzger, M J., Meyer, J., Mitchell, T D., Reginster, I., Rounsevell, M., SabatÃ©, S., Sitch, S., Smith, B., Smith, J., Smith, P., Sykes, M T., Thonicke, K., Thuiller, W., Tuck, G., Zaehle,~S., and Zierl,~B.: Ecosystem Service Supply and Vulnerability to Global Change in Europe, Science, 310, 1333â€“1337, 2005. </reference>
		<reference numeration="90" content_type="text"> Sellers, P J., Randall, D A., Collatz, G J., Berry, J A., Field, C B., Dazlich, D A., Zhang, C., Collelo, G D., and Bounoua, L.: A~revised land surface parameterization (SiB2) for atmospheric GCMs. Part I: Model formulation, J. Climate, 9, 676â€“705, 1996b. </reference>
		<reference numeration="91" content_type="text"> Sitch, S., Smith, B., Prentice, I C., Arneth, A., Bondeau, A., Cramer, W., Kaplan, J O., Levis, S., Lucht, W., Sykes, M T., Thonicke, K., and Venevsky, S.: Evaluation of ecosystem dynamics, plant geography and terrestrial carbon cycling in the LPJ dynamic global vegetation model, Glob. Change Biol., 9, 161â€“185, 2003. </reference>
		<reference numeration="92" content_type="text"> Slatyer, R O.: Plant-water Relationships, Academic Press, London, UK, 1967. </reference>
		<reference numeration="93" content_type="text"> Smirnoff, N. and Stewart, G R.: Nitrate assimilation and translocation by higher plants â€“ Comparative physiology and ecological consequences, Physiol. Plantarum, 64, 133â€“140, 1985. </reference>
		<reference numeration="94" content_type="text"> Syvertsen, J P., Lloyd, J., McConchie, C., Kriedemann, P E., and Farquhar, G D.: On the relationship between leaf anatomy and \chemCO_2 diffusion through the mesophyll of hypostomatous leaves, Plant Cell Environ., 18, 149â€“157, 2005. </reference>
		<reference numeration="95" content_type="text"> Tenhunen, J D., Serra, A S., Harley, P C., Dougherty, R., and Reynolds, J.: Factors influencing carbon fixation and water-use by Mediterranean sclerophyll shrubs during summer drought, Oecologia, 82, 381â€“393, 1990. </reference>
		<reference numeration="96" content_type="text"> Terashima, I., Araya, T., Miyazawa, S., Sone, K., and Yano, S.: Construction and maintenance of the optimal photosynthetic systems of the leaf, herbaceous plant and tree: an eco-developmental treatise, Annal. Bot., 95, 507-519, 2000 </reference>
		<reference numeration="97" content_type="text"> Terashima, I., Hanba, Y T., Tazoe, Y., Vyas, P., and Yano, S.: Irradiance and phenotype: comparative eco-development of sun and shade leaves in relation to photosynthetic CO&lt;sub&gt;2&lt;/sub&gt; diffusion, J. Exp. Bot., 57, 343-354, 2006. </reference>
		<reference numeration="98" content_type="text"> Tezara, W V J., Mitchell, S P., Driscoll, S P., and Lawlor, D W.: Water stress inhibits plant photosynthesis by decreasing coupling factor and ATP, Nature, 401, 914â€“917, 1999. </reference>
		<reference numeration="99" content_type="text"> Tuzet, A., Perrier, A., and Leuning, R.: A~coupled model of stomatal conductance, photosynthesis and transpiration, Plant Cell Environ., 26, 1097â€“1116, 2003. </reference>
		<reference numeration="100" content_type="text"> Valentini, R., DeAngelis, P., Matteucci, G., Monaco, R., Dore, S., and Scarascia-Mugnozza, G E.: Seasonal net carbon exchange of a beech forest with the atmosphere, Glob. Change Biol., 2, 199â€“207, 1996.   </reference>
		<reference numeration="101" content_type="text"> Vassey, T L. and Sharkey, T D.: Mild water-stress of Phaseolus-vulgaris plants leads to reduced starch synthesis and extractable sucrose phosphate synthase activity, Plant Physiol., 89, 1066â€“1070, 1989. </reference>
		<reference numeration="102" content_type="text"> Vetter, M., Churkina, G., Jung, M., Reichstein, M., Zaehle, S., Bondeau, A., Chen, Y., Ciais, P., Feser, F., Freibauer, A., Geyer, R., Jones, C., Papale, D., Tenhunen, J., Tomelleri, E., Trusilova, K., Viovy, N., and Heimann, M.: Analyzing the causes and spatial pattern of the European 2003 carbon flux anomaly in Europe using seven models, Biogeosciences, 5, 561â€“583, 2008. </reference>
		<reference numeration="103" content_type="text"> Von Caemmerer, S. and Farquhar, G D.: Some relationships between the biochemistry of photosynthesis and gas exchange of leaves, Planta, 153, 376â€“387, 1981. </reference>
		<reference numeration="104" content_type="text"> Wang, Y P. and Leuning, R.: A~two-leaf model for canopy conductance, photosynthesis and partitioning of available energy I: Model description and comparison with a multi-layered model, Agr. Forest Meteorol., 19, 89â€“111, 1998. </reference>
		<reference numeration="105" content_type="text"> Warren C R.: Stand aside stomata, another actor deserves centre stage: the forgotten role of the internal conductance to \chemCO_2 transfer, J. Exp. Bot., 59, 1475â€“1487, 2008. </reference>
		<reference numeration="106" content_type="text"> Wong, S., Cowan, I., and Farquhar, G.: Stomatal conductance correlates with photosynthetic capacity, Nature, 282, 424â€“426, 1979. </reference>
		<reference numeration="107" content_type="text"> Wilson, K B., Baldocchi, D D., and Hanson, P J.: Quantifying stomatal and non-stomatal limitations to carbon assimilation resulting from leaf aging and drought in mature deciduous tree species, Tree Physiol., 20, 787â€“797, 2000. </reference>
		<reference numeration="108" content_type="text"> Wilson, K B., Goldstein, A., Falge, E., Aubinet, M., Baldocchi, D., Berbigier, P., Bernhofer, C., Ceulemans, R., Dolmanh, H., Field, C., Grelle, A., Ibrom, A., Lawl, B. E., Kowalski, A., Meyers, T., Moncrieffm, J., Monson, R., Oechel, W., Tenhunen, J., Valentini, R., and Verma, S.: Energy balance closure at FLUXNET sites, Agr. Forest Meteorol., 113, 233â€“243, 2002. </reference>
		<reference numeration="109" content_type="text"> Wohlfahrt, G., Haslwanter, A., Hortnagl, L., Jasoni, R L., Fenstermaker, L F., Arnone, J A., and Hammerle, A.: On the consequences of the energy imbalance for calculating surface conductance to water vapour. Agric. Forest Meteorol., 149, 1556â€“1559,, 2009. </reference>
		<reference numeration="110" content_type="text"> Wofsy, S C., Goulden, M L., Munger, J W., Fan, S M., Bakwin, P S., Daube, B C., Bassow, S L., and Bazzaz, F A.: Net Exchange of \chemCO_2 in a mid-latitude forest, Science, 260, 1314â€“1317, 1993. </reference>
		<reference numeration="111" content_type="text"> Xu, L. and Baldocchi, D D.: Seasonal trends in photosynthetic parameters and stomatal conductance of blue oak (\textitQuercus douglasii) under prolonged summer drought and high temperature, Tree Physiol., 23, 865â€“877, 2003. </reference>
	</references>
</article>

