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	<journal>
		<journal_title>Biogeosciences</journal_title>
		<journal_url>www.biogeosciences.net</journal_url>
		<issn>1726-4170</issn>
		<eissn>1726-4189</eissn>
		<volume_number>4</volume_number>
		<issue_number>1</issue_number>
		<publication_year>2007</publication_year>
	</journal>
	<doi>10.5194/bg-4-27-2007</doi>
	<article_url>http://www.biogeosciences.net/4/27/2007/</article_url>
	<abstract_html>http://www.biogeosciences.net/4/27/2007/bg-4-27-2007.html</abstract_html>
	<fulltext_pdf>http://www.biogeosciences.net/4/27/2007/bg-4-27-2007.pdf</fulltext_pdf>
	<start_page>27</start_page>
	<end_page>36</end_page>
	<publication_date>2007-01-12</publication_date>
	<article_title content_type="html">Effects of climate warming and declining species richness in grassland model ecosystems: acclimation of CO&lt;sub&gt;2&lt;/sub&gt; fluxes</article_title>
	<authors>
		<author numeration="1" affiliations="1">
			<name>S. Vicca</name>
			<email>sara.vicca@ua.ac.be</email>
		</author>
		<author numeration="2" affiliations="2">
			<name>P. Serrano-Ortiz</name>
		</author>
		<author numeration="3" affiliations="1">
			<name>H. J. De Boeck</name>
		</author>
		<author numeration="4" affiliations="1">
			<name>C. M. H. M. Lemmens</name>
		</author>
		<author numeration="5" affiliations="1">
			<name>I. Nijs</name>
		</author>
		<author numeration="6" affiliations="1">
			<name>R. Ceulemans</name>
		</author>
		<author numeration="7" affiliations="2">
			<name>A. S. Kowalski</name>
		</author>
		<author numeration="8" affiliations="1">
			<name>I. A. Janssens</name>
		</author>
	</authors>
	<affiliations>
		<affiliation numeration="1" content_type="html">Research Group of Plant and Vegetation Ecology, Department of Biology, University of Antwerp (Campus Drie Eiken), Universiteitsplein 1, 2610 Wilrijk, Belgium</affiliation>
		<affiliation numeration="2" content_type="html">Department of Applied Physics, University of Granada, C/ Fuentenueva S/N, 18071 Granada, Spain</affiliation>
	</affiliations>
	<abstract content_type="html">To study the effects of warming and declining species richness on the carbon
balance of grassland communities, model ecosystems containing one, three or
nine species were exposed to ambient and elevated (ambient +3&amp;deg;C) air
temperature. In this paper, we analyze measured ecosystem CO&lt;sub&gt;2&lt;/sub&gt; fluxes to
test whether ecosystem photosynthesis and respiration had acclimated to
warming after 28 months of continuous heating, and whether the degree of
acclimation depended on species richness. In order to test whether
acclimation occurred, short term temperature response curves were
established for all communities in both treatments. At similar temperatures,
lower flux rates in the heated communities as compared to the unheated
communities would indicate thermal acclimation. Because plant cover was
significantly higher in the heated treatment, we normalized the data for
plant cover. Subsequently, down-regulation of both photosynthesis and
respiration was observed. Although CO&lt;sub&gt;2&lt;/sub&gt; fluxes were larger in
communities with higher species richness, species richness did not affect
the degree of acclimation to warming. These results imply that models need
to take thermal acclimation into account to simulate photosynthesis and
respiration in a warmer world.</abstract>
	<references>
		<reference numeration="1" content_type="text"> Atkin, O. K., Bruhn, D., Hurry, V. M., and Tjoelker, M. G.: Response of root respiration to changes in temperature and its relevance to global warming, New Phytol., 147, 141&amp;ndash;154, 2000a. </reference>
		<reference numeration="2" content_type="text"> Atkin, O. K., Evans, J. R., Ball, M. C., Lambers, H., and Pons, T. L.: Leaf respiration of snow gum in the light and the dark. Interactions between temperature and irradiance, Plant Physiol., 122, 915&amp;ndash;923, 2000b. </reference>
		<reference numeration="3" content_type="text"> Atkin, O. K., Scheurwater, I., and Pons, T. L.: High thermal acclimation potential of both photosynthesis and respiration in two lowland \textitPlantago species in contrast to an alpine congeneric, Global Change Biol., 12, 500&amp;ndash;515, 2006. </reference>
		<reference numeration="4" content_type="text"> Atkin, O. K. and Tjoelker, M. G.: Thermal acclimation and the dynamic response of plant respiration to temperature, Trends in Plant Sci., 8, 343&amp;ndash;351, 2003. </reference>
		<reference numeration="5" content_type="text"> Berry, J. A. and Björkman, O.: Photosynthetic response and adaptation to temperature in higher plants, Ann. Rev. Plant Physiol., 31, 491&amp;ndash;543, 1980. </reference>
		<reference numeration="6" content_type="text"> Bouma, T. J., De Visser, R., Van Leeuwen, P. H., De Kock, M. J., and Lambers, H.: The respiratory energy requierments involved in nocturnal carbohydrate export from starch-storing mature source leaves and their contribution to leaf dark respiration, J. Exp. Bot., 46, 1185&amp;ndash;1194, 1995. </reference>
		<reference numeration="7" content_type="text"> Bryla, D. R., Bouma, T. J., and Eissenstat, D. M.: Root respiration in citrus acclimates to temperature and slows during drought, Plant Cell Environ., 20, 1411&amp;ndash;1420, 1997. </reference>
		<reference numeration="8" content_type="text"> Bryla, D. R., Bouma, T. J., Hartmond, U., and Eissenstat, D. M.: Influence of temperature and soil drying on respiration of individual roots in citrus: integrating greenhouse observations into a predictive model for the field, Plant Cell Environ., 24, 781&amp;ndash;790, 2001. </reference>
		<reference numeration="9" content_type="text"> Burton, A. and Pregitzer, K. S.: Field measurements of root respiration indicate little to no seasonal temperature acclimation for sugar maple and red pine, Tree Physiol., 23, 273&amp;ndash;280, 2003. </reference>
		<reference numeration="10" content_type="text"> Caldwell, M. M., Dawson, T. E., and Richards, J. H.: Hydraulic lift: consequences of water efflux from the roots of plants, Oecologia, 113, 151&amp;ndash;161, 1998. </reference>
		<reference numeration="11" content_type="text"> Davidson, E. A. and Janssens, I. A.: Temperature sensitivity of soil carbon decomposition and feedbacks to climate change, Nature, 440, 165&amp;ndash;173, 2006. </reference>
		<reference numeration="12" content_type="text"> De Boeck, H. J., Lemmens, C M H M., Bossuyt, H., Malchair, S., Carnol, M., Merckx, R., Nijs, I., and Ceulemans, R.: How do climate warming and plant species richness affect water use in experimental grasslands?, Plant Soil, 288, 249&amp;ndash;261, 2006. </reference>
		<reference numeration="13" content_type="text"> Eliasson, P. E., McMurtrie, R. E., Pepper, D. A., Strömgren, M., Linder, S., and Agren, G. I.: The response of heterotrophic CO&lt;sub&gt;2&lt;/sub&gt; flux to soil warming, Global Change Biol., 11, 167&amp;ndash;181, 2005. </reference>
		<reference numeration="14" content_type="text"> Fridley, D.: The influence of species diversity on ecosystem productivity: how, where and why?, Oikos, 93, 514&amp;ndash;526, 2001. </reference>
		<reference numeration="15" content_type="text"> Gifford, G. M.: The global carbon cycle: A viewpoint on the missing sink, Austr. J. Plant Physiol., 21, 1&amp;ndash;15, 1994. </reference>
		<reference numeration="16" content_type="text"> Gifford, G. M.: Whole plant respiration and photosynthesis of wheat under increased CO&lt;sub&gt;2&lt;/sub&gt; concentration and temperature: long-term vs. short-term distinctions for modeling, Global Change Biol., 1, 385&amp;ndash;396, 1995. </reference>
		<reference numeration="17" content_type="text"> Hector, A., Schmid, B., Beierkuhnlein, C., Caldeira, M. C., Diemer, M., Dimitrakopoulos, P. G., Finn, J. A., Freitas, H., Gilles, P. S., Good, J., Harris, R., Högberg, P., Huss-Danell, K., Joshi, J., Jumponnen, A., Körner, C., Leadley, P. W., Loreau, M., Minns, A., Mulde, C. P. H., O&apos;Donovan, G., Otway, S. J., Pereira, J. S., Prinz, A., Read, D. J., Scherer-Lorenzen, M., Schulze, E.-D., Siamantziouras, A.-S. D., Spehn, E. M., Terry, A. C., Troumbis, A. Y., Woodward, F. I., Yachi, S., and Lawton, J. H.: Plant diversity and productivity experiments in European grasslands, Science, 286, 1123&amp;ndash;1127, 1999. </reference>
		<reference numeration="18" content_type="text"> Hoefnagel, M. H. N., Atkin, O. K., and Wiskich, J. T.: Interdependence between chloroplasts and mitochondria in the light and the dark, Biochim. Bioph. Acta &amp;ndash; Bioenerg., 1366, 235&amp;ndash;255, 1998. </reference>
		<reference numeration="19" content_type="text"> King, A. W., Gunderson, C. A., Post, W. M., Weston, D. J., and Wullschleger, S. D: Plant respiration in a warmer world, Science, 312, 536&amp;ndash;537, 2006. </reference>
		<reference numeration="20" content_type="text"> Kirschbaum, M. U.: Soil respiration under prolonged soil warming: are rate reductions caused by acclimation or substrate loss?, Global Change Biol., 10, 1870&amp;ndash;1877, 2004. </reference>
		<reference numeration="21" content_type="text"> Körner, C. and Larcher, W.: Plant Life in Cold Environments, edited by: Long, S. F. and Woodward, F. I., Symposium of the Society of Experimental Biologists, The Company of Biologists Limited, Cambridge, 1988. </reference>
		<reference numeration="22" content_type="text"> Krömer, S.: Respiration during photosynthesis, Ann. Rev. Plant Physiol. Mol. Biol., 46, 45&amp;ndash;70, 1995. </reference>
		<reference numeration="23" content_type="text"> Lambers, H., Chapin, F. S., and Pons, T. L.: Plant Physiological Ecology, Springer-Verlag, New York, 1998. </reference>
		<reference numeration="24" content_type="text"> Larcher, W.: Physiological Plant Ecology (fourth edition), Springer-Verlag, Berlin, Heidelberg, New York, 2003. </reference>
		<reference numeration="25" content_type="text"> Larigauderie, A. and Körner, C.: Acclimation of leaf dark respiration to temperature in alpine and lowland plant species, Ann. Bot., 76, 245&amp;ndash;252, 1995. </reference>
		<reference numeration="26" content_type="text"> Lemmens, C. M. H. M., De Boeck, H. J., Gielen, B., Bossuyt, H., Malchair, S., Carnol, M., Merckx, R., Nijs, I., and Ceulemans, R.: End-of-season effects of elevated temperature on ecophysiological processes of grassland species at different species richness levels, Environ. Exp. Bot., 56, 245&amp;ndash;254, 2006. </reference>
		<reference numeration="27" content_type="text"> Loveys, B. R., Atkinson, L. J., Sherlock, D. J., Roberts, R. L., Fitter, A. H., and Atkin, O. K.: Thermal acclimation of leaf and root respiration: an investigation comparing inherently fast- and slow-growing plant species, Global Change Biol., 9, 895&amp;ndash;910, 2003. </reference>
		<reference numeration="28" content_type="text"> Loveys, B. R., Scheurwater, I., Pons, T. L., Fitter, A. H., and Atkin, O. K.: Growth temperature influences the underlying components of relative growth rate: an investigation using inherently fast- and slow-growing plant species, Plant Cell Environ., 25, 975&amp;ndash;987, 2002. </reference>
		<reference numeration="29" content_type="text"> Luo, Y. Q., Wan, S. Q., Hui, D. F., and Wallace, L. L.: Acclimatization of soil respiration to warming in a tall grass prairie, Nature, 413, 622&amp;ndash;625, 2001. </reference>
		<reference numeration="30" content_type="text"> Melillo, J. M., Steudler, P. A., Aber, J. D., Newkirk, K., Lux, H., Bowles, F. P., Catricala, C., Magill, A., Ahrens, T., and Morrisseau, S.: Soil warming and carbon cycle feedbacks to the climate system, Science, 298, 2173&amp;ndash;2175, 2002. </reference>
		<reference numeration="31" content_type="text"> Pollock, C. J.: The response of plants to temperature change, J. Agricult. Sci., 115, 1&amp;ndash;5, 1990. </reference>
		<reference numeration="32" content_type="text"> Read, D.: Mycorrhizal fungi &amp;ndash; The ties that bind, Nature, 388, 517&amp;ndash;518, 1997. </reference>
		<reference numeration="33" content_type="text"> Rook, D.: The influence of growing temperature on photosynthesis and respiration of \textitPinus radiate seedlings, New Zealand J. Bot., 7, 43&amp;ndash;55, 1969. </reference>
		<reference numeration="34" content_type="text"> Roy, J.: Biodiversity Control of Primary Production, in: Terrestrial Global Productivity, edited by: Roy, J., Saugier, B., and Mooney, H. A., Academic Press, San Diego, USA, 169&amp;ndash;181, 2001. </reference>
		<reference numeration="35" content_type="text"> Ryan, M. G., Hunt, E. R., McMurtrie, R. E., Agren, G. I., Aber, J. D., Friend, A. D., Rastetter, E. B., Pulliam, W. M., Raison, R. J., and Linder, S.: Comparing Models of Ecosystem Function for Temperate Conifer Forests. I. Model Description and Validation, in: Global Change: Effects on Coniferous Forests and Grasslands, edited by: Breymeyer, A. I., Hall, D. O., Melillo, J. M., and Agren, G. I., SCOPE 56, Scientific Committee on Problems of the Encironment/ Wiley, Chichester, UK, 313&amp;ndash;362, 1996. </reference>
		<reference numeration="36" content_type="text"> Saxe, H., Cannell, M. G. R., Johnsen, O., Ryan, M. G., and Vourlitis, G.: Tree and forest functioning in response to global warming, New Phytol., 149, 369&amp;ndash;400, 2001. </reference>
		<reference numeration="37" content_type="text"> Sharkey T.D.: Effects of moderate heat stress on photosynthesis: importance of thylakoid reactions, rubisco deactivation, reactive oxygen species, and thermotolerance provided by isoprene, Plant Cell Environ., 28, 269-277, 2005. </reference>
		<reference numeration="38" content_type="text"> Sowell, J. B. and Spomer, G. G.: Ecotypic variation in root respiration rate among elevational populations of Abies lasiocarpa and Picea engelmannii, Oecologia, 68, 375&amp;ndash;379, 1986. </reference>
		<reference numeration="39" content_type="text"> Spehn, E. M., Joshi, J., Schmid, B., Diemer, M., and Körner, C.: Above-ground resource use increases with plant species richness in experimental grassland ecosystems, Funct. Ecol., 14, 326&amp;ndash;337, 2000. </reference>
		<reference numeration="40" content_type="text"> Symstad, A. J., Tilman, D., Willson, J., and Knops, J. M. H.: Species loss and ecosystem functioning: effects of species identity and community composition, Oikos, 81, 389&amp;ndash;397, 1998. </reference>
		<reference numeration="41" content_type="text"> Tillman, D., Wedin, D., and Knops, J.: Productivity and sustainability influenced by biodiversity in grassland ecosystems, Nature, 379, 718&amp;ndash;720, 1996. </reference>
		<reference numeration="42" content_type="text"> Tjoelker, M. G., Oleksyn, J., and Reich, P. B.: Seedlings of five boreal tree species differ in acclimation of net photosynthesis to elevated CO&lt;sub&gt;2&lt;/sub&gt; and temperature, Tree Physiol., 18, 715&amp;ndash;726, 1998. </reference>
		<reference numeration="43" content_type="text"> Turnbull, M. H., Whitehead, D., Tissue, D. T., Schuster, W. S. F., Brown, K. J., and Griffin, K. L.: Responses of leaf respiration to temperature and leaf characteristics in three deciduous tree species vary with site water availability, Tree Physiol., 21, 571&amp;ndash;578, 2001. </reference>
		<reference numeration="44" content_type="text"> Van Ruijven, J. and Berendse, F.: Diversity-productivity relationship: Initial effects, long-term patterns, and underlying mechanisms, Proc. Natl. Acad. Sci. USA, 102, 695&amp;ndash;700, 2005 </reference>
		<reference numeration="45" content_type="text"> Vandermeer, J. H.: The Ecology of Intercropping, Cambridge University Press, 1989. </reference>
		<reference numeration="46" content_type="text"> Weger, H. G. and Guy, R. D.: Cytochrome and alternative pathway respiration in white spruce (\textitPicea glauca) roots. Effects of growth and measurement temperature, Physiol. Plantarum, 83, 675&amp;ndash;681, 1991. </reference>
		<reference numeration="47" content_type="text"> Woodwell, G. M.: The effects of global warming, in: Global Warming: the Greenpeace Report, edited by: Leggett, J., Oxford University Press, Oxford, UK, 116&amp;ndash;132, 1990. </reference>
		<reference numeration="48" content_type="text"> Ziska, L. H. and Bunce, J. A.: The influence of increasing growth temperature and CO&lt;sub&gt;2&lt;/sub&gt; concentration on the ratio of respiration to photosynthesis in soybean seedlings, Global Change Biol., 4, 637&amp;ndash;643, 1998. </reference>
		<reference numeration="49" content_type="text"> Zogg, G. P., Zak, D. R., Burton, A. J., and Pregitzer, K. S.: Fine root respiration in northern hardwood forests in relation to temperature and nitrogen availability, Tree Physiol., 16, 719&amp;ndash;725, 1996. </reference>
	</references>
</article>

