<?xml version="1.0" encoding="utf-8" standalone="no"?>
<!DOCTYPE article SYSTEM "http://www.biogeosciences.net/inc/bg/copernicus.dtd">
<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>1</issue_number>
		<publication_year>2009</publication_year>
	</journal>
	<doi>10.5194/bg-6-25-2009</doi>
	<article_url>http://www.biogeosciences.net/6/25/2009/</article_url>
	<abstract_html>http://www.biogeosciences.net/6/25/2009/bg-6-25-2009.html</abstract_html>
	<fulltext_pdf>http://www.biogeosciences.net/6/25/2009/bg-6-25-2009.pdf</fulltext_pdf>
	<start_page>25</start_page>
	<end_page>31</end_page>
	<publication_date>2009-01-07</publication_date>
	<article_title content_type="html">Carbon and oxygen isotope analysis of leaf biomass reveals contrasting photosynthetic responses to elevated CO&lt;sub&gt;2&lt;/sub&gt; near geologic vents in Yellowstone National Park</article_title>
	<authors>
		<author numeration="1" affiliations="1">
			<name>S. Sharma</name>
			<email>shikha@uwyo.edu</email>
		</author>
		<author numeration="2" affiliations="2">
			<name>D. G. Williams</name>
		</author>
	</authors>
	<affiliations>
		<affiliation numeration="1" content_type="html">Department of Renewable Resources, University of Wyoming, Laramie, WY 82071, USA</affiliation>
		<affiliation numeration="2" content_type="html">Departments of Renewable Resources and Botany, University of Wyoming, Laramie, WY 82071, USA</affiliation>
	</affiliations>
	<abstract content_type="html">In this study we explore the use of natural CO&lt;sub&gt;2&lt;/sub&gt; emissions in
Yellowstone National Park (YNP) in Wyoming, USA to study responses of
natural vegetation to elevated CO&lt;sub&gt;2&lt;/sub&gt; levels. Radiocarbon (&lt;sup&gt;14&lt;/sup&gt;C)
analysis of leaf biomass from a conifer (&lt;i&gt;Pinus contortus&lt;/i&gt;; lodgepole pine) and an invasive,
non-native herb (&lt;i&gt;Linaria dalmatica&lt;/i&gt;; Dalmation toadflax) was used to trace the inputs of vent
CO&lt;sub&gt;2&lt;/sub&gt; and quantify assimilation-weighted CO&lt;sub&gt;2&lt;/sub&gt; concentrations
experienced by individual plants near vents and in comparable locations with
no geologic CO&lt;sub&gt;2&lt;/sub&gt; exposure. The carbon and oxygen isotopic composition
and nitrogen percent of leaf biomass from the same plants was used to
investigate photosynthetic responses of these plants to naturally elevated
atmospheric CO&lt;sub&gt;2&lt;/sub&gt; concentrations. The coupled shifts in carbon and oxygen
isotope values suggest that dalmation toadflax responded to elevated
CO&lt;sub&gt;2&lt;/sub&gt; exposure by increasing stomatal conductance with no change in
photosynthetic capacity and lodgepole pine apparently responded by
decreasing stomatal conductance and photosynthetic capacity. Lodgepole pine
saplings exposed to elevated levels of CO&lt;sub&gt;2&lt;/sub&gt; likewise had reduced leaf
nitrogen concentrations compared to plants with no enhanced CO&lt;sub&gt;2&lt;/sub&gt;
exposure, further suggesting widespread and dominant conifer
down-regulated photosynthetic capacity under elevated CO&lt;sub&gt;2&lt;/sub&gt; levels near
geologic vents.</abstract>
	<references>
		<reference numeration="1" content_type="text"> % vor jede Referenz Ainsworth, E. A. and Long, S. P.: What have we learned from fifteen years of Free Air Carbon Dioxide Enrichment (FACE)? A Meta-analytic review of the responses of photosynthesis, canopy properties and plant production to rising CO&lt;sub&gt;2&lt;/sub&gt;, New Phytol., 165, 351–372, 2005. </reference>
		<reference numeration="2" content_type="text"> Ainsworth, E. A. and Rogers, A.: The response of photosynthesis and stomatal conductance to rising [CO&lt;sub&gt;2&lt;/sub&gt;]: mechanisms and environmental interactions, Plant, Cell Env., 30, 258–270, 2007. </reference>
		<reference numeration="3" content_type="text"> Barbour, M. M. and Farquhar, G. D.: Relative humidity- and ABA-induced variation in carbon and oxygen isotope ratios of cotton leaves, Plant, Cell Env., 23, 473–485, 2000. </reference>
		<reference numeration="4" content_type="text"> Barbour, M. M., Andrews, T. J., and Farquhar, G. D.: Correlations between oxygen isotope ratios of wood constituents of Quercus and Pinus samples from around the world, Aust. J. Plant Physiol., 28, 335–348, 2001. </reference>
		<reference numeration="5" content_type="text"> Barbour, M. M.: Stable oxygen isotope composition of plant tissue: a review, Funct. Plant Biol., 34, 83–94, 2007. </reference>
		<reference numeration="6" content_type="text"> Bettarini, I., Calderoni, G., Miglietta, F., Raschi, A., and Ehleringer, J.: Isotopic carbon discrimination and leaf nitrogen content of \textitErica arborea L. along a CO&lt;sub&gt;2&lt;/sub&gt; concentration gradient in a CO&lt;sub&gt;2&lt;/sub&gt; spring in Italy, Tree Physiol., 15, 327–332, 1995. </reference>
		<reference numeration="7" content_type="text"> Dang, Q. L. and Cheng, S.: Effects of soil temperature on ecophysiological traits in seedlings of four boreal tree species, Forest Ecol. Management, 194, 379–387, 2004. </reference>
		<reference numeration="8" content_type="text"> Ehleringer, J. R., Hall, A. E., and Farquhar, G. D.: Stable Isotopes and Plant Carbon – Water Relations, Elsevier, New York, 1993. </reference>
		<reference numeration="9" content_type="text"> Farquhar, G. D., Ehleringer, J. R., and Hubick, K. T.: Carbon isotope discrimination and photosynthesis, Ann. Rev. Plant Physiol. Mol. Biol., 40, 503–537, 1989. </reference>
		<reference numeration="10" content_type="text"> Farquhar, G. D. and Lloyd, J.: Carbon and oxygen isotope effects in the exchange of carbon dioxide between terrestrial plants and the atmosphere, in: Stable Isotopes and Plant Carbon-Water Relations, edited by: Ehleringer, J. R., Hall, A. E., and Farquhar, G. D., Academic Press, San Diego, CA, USA , 47–70, 1993. </reference>
		<reference numeration="11" content_type="text"> Field, C. and Mooney, H. A.: The photosynthesis - nitrogen relationship in wild plants, in: On the Economy of Plant Form and Function, edited by: Givnish T. J., Cambridge University Press, New York, NY, USA, 25–55, 1986. </reference>
		<reference numeration="12" content_type="text"> Field, C. B., Lund, C. P., Chiariello, N. R., and Mortimer, B. E.: CO&lt;sub&gt;2&lt;/sub&gt; effects on the water budget of grassland microcosm communities, Global Change Biol., 3, 197–206, 1997. </reference>
		<reference numeration="13" content_type="text"> Gill, R. A., Polley, H. W., Johnson, H. B., Anderson, L. J., Maherali, H., and Jackson, R. B.: Nonlinear grassland responses to past and future atmospheric CO$_2,$, Nature, 417, 279–282, 2002. </reference>
		<reference numeration="14" content_type="text"> Grams, T. E. E., Kozovitis, A. R., Haberele, K. H., Matyssek, R., and Dawson, T. E.: Combining $\delta ^13$C and $\delta ^18$O analyses to unravel competition, CO&lt;sub&gt;2&lt;/sub&gt; and O&lt;sub&gt;3&lt;/sub&gt; effects on the physiological performance of different-aged trees, Plant Cell Env., 30, 1023–1034, 2007. </reference>
		<reference numeration="15" content_type="text"> Keeling, C. D., Piper, S. C., Bacastow, R. B., Wahlen, M., Whorf, T. P., Heimann, M., and Meijer, H. A.: Exchanges of Atmospheric CO&lt;sub&gt;2&lt;/sub&gt; and $^13$CO&lt;sub&gt;2&lt;/sub&gt; with the Terrestrial Biosphere and Oceans from 1978 to 2000, I. Global Aspects, SIO Reference No. 01–06 (Revised from SIO Reference No. 00–21), 1–28 June 2001. </reference>
		<reference numeration="16" content_type="text"> Knapp, A. K., Hamerlynck, E. P., Ham, J. M., and Owensby, C. E.: Responses in stomatal conductance to elevated CO&lt;sub&gt;2&lt;/sub&gt; in 12 grassland species that differ in growth form, Plant Ecol., 125, 31–41, 1996. </reference>
		<reference numeration="17" content_type="text"> Lockheart, M. J., Van Bergen, P. F., and Evershed, R. P.: Variations in the stable carbon isotope compositions of individual lipids from the leaves of modern angiosperms: implications for the study of higher land plant-derived sedimentary organic matter, Org. Geochem., 26,137–153, 1997. </reference>
		<reference numeration="18" content_type="text"> Luo, Y., Currie, W., Dukes, J., Finzi, A., Hartwig, U., Hungate, B., McMurtrie, R., Oren, R., Parton, W., Pataki, D., Shaw, M., Zak, D., and Field, C.: Progressive nitrogen limitation of ecosystem responses to rising atmospheric carbon dioxide, BioScience, 54, 731–739, 2004. </reference>
		<reference numeration="19" content_type="text"> Miglietta, F., Bettarini, I., Raschi, A., Körner, C., and Vaccari, F. P.: Isotope discrimination and photosynthesis of vegetation growing in the Bossoleto CO&lt;sub&gt;2&lt;/sub&gt; spring, Chemosphere, 36, 771–776, 1998. </reference>
		<reference numeration="20" content_type="text"> Mooney, H. A., Field, C., VasquezYanes, C., and Chu, C.: Environmental controls on stomatal conductance in a shrub of the humid tropics, Proc. Natl. Acad. Sci., 80, 1295–1297, 1983. </reference>
		<reference numeration="21" content_type="text"> Nowak, R. S., Ellsworth, D. S., and Smith, S. D.: Functional responses of plants to elevated atmospheric – do photosynthetic and productivity data from FACE experiments support early predictions?, New Phytol., 162, 253–280, 2004. </reference>
		<reference numeration="22" content_type="text"> Scheidegger, Y., Saurer, M., Bahn, M., and Siegwolf, R.: Linking stable oxygen and carbon isotopes with stomatal conductance and photosynthetic capacity: a conceptual model, Oecologia, 125, 350–357, 2000. </reference>
		<reference numeration="23" content_type="text"> Schomp, J. M.: Rising atmospheric CO&lt;sub&gt;2&lt;/sub&gt; concentrations and plant invasion in the northern mixed-grass prairie: an ecophysiological perspective, M.S. thesis, University of Wyoming, Laramie, 2007. </reference>
		<reference numeration="24" content_type="text"> Schwarz, P. A., Fahey, T. J., and Dawson, T. E.: Seasonal soil temperature effects on photosynthesis in red spruce (Picea rubens) saplings, Tree Physiol., 17, 187–194, 1997. </reference>
		<reference numeration="25" content_type="text"> Shangguan, Z. P., Shao, M. A., and Dyckmans, J.: Nitrogen nutrition and water stress effects on leaf photosynthetic gas exchange and water use efficiency in winter wheat, Env. Exp. Botany, 44, 141–149, 2000. </reference>
		<reference numeration="26" content_type="text"> Sinclair, T. R., Pinter Jr. P. J., Kimball, B. A., Adamsen, F. J., LaMorte, R. L., Wall, G. W., Hunsaker, D. J., Adam, N., Brooks, T. J., Garcia, R. L., Thompson, T., Leavitt, S., and Matthias, A.: Leaf nitrogen concentration of wheat subjected to elevated [CO&lt;sub&gt;2&lt;/sub&gt;] and either water or N deficits, Agriculture Ecosystem and Environment, 79, 53–60, 2000. </reference>
		<reference numeration="27" content_type="text"> Stitt, M. and Krapp, A.: The interaction between elevated carbon dioxide and nitrogen nutrition: the physiological and molecular background, Plant, Cell Env., 22, 583–621, 1999. </reference>
		<reference numeration="28" content_type="text"> Stuiver, M. and Polach, H. A.: Reporting of $^14$C data, Radiocarbon, 19, 355–363, 1977. </reference>
		<reference numeration="29" content_type="text"> Tereck, M.: Plant communities growing under elevated CO&lt;sub&gt;2&lt;/sub&gt; concentrations in Yellowstone National Park, Field survey unpublished report, 2004. </reference>
		<reference numeration="30" content_type="text"> Tognetti, R., Minnocci, A., Peñuelas, J., Raschi, A., and Jones, M. B.: Comparative field water relations of three Mediterranean shrub species co-occurring at a natural CO&lt;sub&gt;2&lt;/sub&gt; vent, J. Exp. Botany, 51, 1135–1146, 2000. </reference>
		<reference numeration="31" content_type="text"> van Gardingen, P. R., Grace, J., Harkness, D. D., Miglietta, F., and Raschi, A.: Carbon dioxide emissions at an Italian mineral spring: measurements of average CO&lt;sub&gt;2&lt;/sub&gt; concentration and air temperature, Agriculture&amp; Forest Meteorology, 73, 17–27, 1995. </reference>
		<reference numeration="32" content_type="text"> Warren, C. R. and Adams, M. A.: Water availability and branch length determine $\delta ^13$C in foliage of Pinus pinaster, Tree Physiol., 20, 637–643, 2000. </reference>
		<reference numeration="33" content_type="text"> Werner, C. and Brantley, S.: CO&lt;sub&gt;2&lt;/sub&gt; emissions from the Yellowstone volcanic systems, Geochem. Geophys. Geosys., 4, 1–27, 2003. </reference>
		<reference numeration="34" content_type="text"> Wullschleger, S. D., Tschaplinski, T. J., and Norby, R. J.: Plant Water Relations at Elevated CO&lt;sub&gt;2&lt;/sub&gt; – Implications for Water-Limited Environments, Plant Cell Env., 25, 319–331, 2002. </reference>
		<reference numeration="35" content_type="text"> Zak, D. R., Pregitzer, K. S., King, J. S., and Holmes, W. E.: Elevated atmospheric CO&lt;sub&gt;2&lt;/sub&gt;, fine roots and the response of soil microorganisms: A review and hypothesis, New Phytol., 147, 201–222, 2000. </reference>
		<reference numeration="36" content_type="text"> Zhang, S. and Dang, Q. L.: Effects of soil temperature and elevated atmospheric CO&lt;sub&gt;2&lt;/sub&gt; concentration on gas exchange, in vivo carboxylation and chlorophyll fluorescence in jack pine and white birch seedlings, Tree Physiol., 25, 523–531, 2005. </reference>
	</references>
</article>
