<?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>7</volume_number>
		<issue_number>6</issue_number>
		<publication_year>2010</publication_year>
	</journal>
	<doi>10.5194/bg-7-1809-2010</doi>
	<article_url>http://www.biogeosciences.net/7/1809/2010/</article_url>
	<abstract_html>http://www.biogeosciences.net/7/1809/2010/bg-7-1809-2010.html</abstract_html>
	<fulltext_pdf>http://www.biogeosciences.net/7/1809/2010/bg-7-1809-2010.pdf</fulltext_pdf>
	<start_page>1809</start_page>
	<end_page>1832</end_page>
	<publication_date>2010-06-01</publication_date>
	<article_title content_type="html">The leaf-level emission factor of volatile isoprenoids: caveats, model algorithms, response shapes and scaling</article_title>
	<authors>
		<author numeration="1" affiliations="1">
			<name>Ü. Niinemets</name>
			<email>ylo.niinemets@emu.ee</email>
		</author>
		<author numeration="2" affiliations="2">
			<name>R. K. Monson</name>
		</author>
		<author numeration="3" affiliations="3">
			<name>A. Arneth</name>
		</author>
		<author numeration="4" affiliations="4">
			<name>P. Ciccioli</name>
		</author>
		<author numeration="5" affiliations="5">
			<name>J. Kesselmeier</name>
		</author>
		<author numeration="6" affiliations="6">
			<name>U. Kuhn</name>
		</author>
		<author numeration="7" affiliations="1">
			<name>S. M. Noe</name>
		</author>
		<author numeration="8" affiliations="7">
			<name>J. Peñuelas</name>
		</author>
		<author numeration="9" affiliations="8">
			<name>M. Staudt</name>
		</author>
	</authors>
	<affiliations>
		<affiliation numeration="1" content_type="html">Institute of Agricultural and Environmental Sciences, Estonian University of Life Sciences, Kreutzwaldi 1, Tartu, 51014, Estonia</affiliation>
		<affiliation numeration="2" content_type="html">Department of Ecology and Evolutionary Biology and Cooperative Institute for Research in Environmental Sciences, University of Colorado, Boulder, CO 80309-0334, USA</affiliation>
		<affiliation numeration="3" content_type="html">Division of Physical Geography and Ecosystem Analysis, Lund University, Sölvegatan 12, Lund, 22362, Sweden</affiliation>
		<affiliation numeration="4" content_type="html">Istituto di Metodologie Chimiche del CNR, Area della Ricerca di Roma 1, Monterotondo Scalo, 00016, Italy</affiliation>
		<affiliation numeration="5" content_type="html">Max Planck Institute for Chemistry, Biogeochemistry Department, Joh.-J.-Becher Weg 27, Mainz, 55128, Germany</affiliation>
		<affiliation numeration="6" content_type="html">Federal Research Station Agroscope Reckenholz-Taenikon, ART, Zuerich, Switzerland</affiliation>
		<affiliation numeration="7" content_type="html">Global Ecology Unit CSIC-CEAB-CREAF, Facultat de Ciències, Universitat Autònoma de Barcelona, Bellaterra, 08193, Spain</affiliation>
		<affiliation numeration="8" content_type="html">Centre d&apos;Ecologie Fonctionnelle et Evolutive (CEFE-CNRS), 1919, Route de Mende, Montpellier cedex 5, 34293, France</affiliation>
	</affiliations>
	<abstract content_type="html">In models of plant volatile isoprenoid emissions, the instantaneous compound
emission rate typically scales with the plant&apos;s emission potential under
specified environmental conditions, also called as the emission factor,
&lt;i&gt;E&lt;/i&gt;&lt;sub&gt;S&lt;/sub&gt;. In the most widely employed plant isoprenoid emission models, the
algorithms developed by Guenther and colleagues (1991, 1993), instantaneous
variation of the steady-state emission rate is described as the product of
&lt;i&gt;E&lt;/i&gt;&lt;sub&gt;S&lt;/sub&gt; and light and temperature response functions. When these models are
employed in the atmospheric chemistry modeling community, species-specific
&lt;i&gt;E&lt;/i&gt;&lt;sub&gt;S&lt;/sub&gt; values and parameter values defining the instantaneous response
curves are often taken as initially defined. In the current review, we argue
that &lt;i&gt;E&lt;/i&gt;&lt;sub&gt;S&lt;/sub&gt; as a characteristic used in the models importantly depends on
our understanding of which environmental factors affect isoprenoid
emissions, and consequently need standardization during experimental
&lt;i&gt;E&lt;/i&gt;&lt;sub&gt;S&lt;/sub&gt; determinations. In particular, there is now increasing consensus that
in addition to variations in light and temperature, alterations in
atmospheric and/or within-leaf CO&lt;sub&gt;2&lt;/sub&gt; concentrations may need to be
included in the emission models. Furthermore, we demonstrate that for less
volatile isoprenoids, mono- and sesquiterpenes, the emissions are often
jointly controlled by the compound synthesis and volatility. Because of
these combined biochemical and physico-chemical drivers, specification of
&lt;i&gt;E&lt;/i&gt;&lt;sub&gt;S&lt;/sub&gt; as a constant value is incapable of describing instantaneous
emissions within the sole assumptions of fluctuating light and temperature
as used in the standard algorithms. The definition of &lt;i&gt;E&lt;/i&gt;&lt;sub&gt;S&lt;/sub&gt; also varies
depending on the degree of aggregation of &lt;i&gt;E&lt;/i&gt;&lt;sub&gt;S&lt;/sub&gt; values in different
parameterization schemes (leaf- vs. canopy- or region-scale, species vs.
plant functional type levels) and various aggregated &lt;i&gt;E&lt;/i&gt;&lt;sub&gt;S&lt;/sub&gt; schemes are not
compatible for different integration models. The summarized information
collectively emphasizes the need to update model algorithms by including
missing environmental and physico-chemical controls, and always to define
&lt;i&gt;E&lt;/i&gt;&lt;sub&gt;S&lt;/sub&gt; within the proper context of model structure and spatial and temporal
resolution.</abstract>
	<references>
		<reference numeration="1" content_type="text"> Alonso, W. R. and Croteau, R.: Prenyltransferases and cyclases, in: Enzymes of secondary metabolism, edited by: Lea, P. J., Methods in plant biochemistry, 9, Academic Press, London - San Diego - New York - Boston - Sydney - Tokyo - Toronto, 239–260, 1993. </reference>
		<reference numeration="2" content_type="text"> Amthor, J. S.: Scaling CO&lt;sub&gt;2&lt;/sub&gt;-photosynthesis relationships from the leaf to the canopy, Photosynth. Res., 39, 321–350, 1994. </reference>
		<reference numeration="3" content_type="text"> Arimura, G.-I., Matsui, K., and Takabayashi, J.: Chemical and molecular ecology of herbivore-induced plant volatiles: proximate factors and their ultimate functions, Plant Cell Physiol., 50, 911–923, 2009. </reference>
		<reference numeration="4" content_type="text"> Arneth, A., Niinemets, Ü., Pressley, S., Bäck, J., Hari, P., Karl, T., Noe, S., Prentice, I. C., Serça, D., Hickler, T., Wolf, A., and Smith, B.: Process-based estimates of terrestrial ecosystem isoprene emissions: incorporating the effects of a direct CO&lt;sub&gt;2&lt;/sub&gt;-isoprene interaction, Atmos. Chem. Phys., 7, 31–53, doi:10.5194/acp-7-31-2007, 2007. </reference>
		<reference numeration="5" content_type="text"> Arneth, A., Monson, R. K., Schurgers, G., Niinemets, Ü., and Palmer, P. I.: Why are estimates of global terrestrial isoprene emissions so similar (and why is this not so for monoterpenes)?, Atmos. Chem. Phys., 8, 4605–4620, doi:10.5194/acp-8-4605-2008, 2008a. </reference>
		<reference numeration="6" content_type="text"> Arneth, A., Schurgers, G., Hickler, T., and Miller, P. A.: Effects of species composition, land surface cover, CO&lt;sub&gt;2&lt;/sub&gt; concentration and climate on isoprene emissions from European forests, Plant Biol., 10, 150–152, 2008b. </reference>
		<reference numeration="7" content_type="text"> Arneth, A. and Niinemets, Ü.: Induced BVOCs: how to bug our models?, Trends Plant Sci., 15, 118–125, 2010. </reference>
		<reference numeration="8" content_type="text"> Atkinson, R. and Arey, J.: Atmospheric degradation of volatile organic compounds, Chem. Rev., 103, 4605–4638, 2003a. </reference>
		<reference numeration="9" content_type="text"> Atkinson, R. and Arey, J.: Gas-phase tropospheric chemistry of biogenic volatile organic compounds: a review, Atmos. Environ., 37, 197–219, 2003b. </reference>
		<reference numeration="10" content_type="text"> Baldocchi, D.: Measuring and modelling carbon dioxide and water vapour exchange over a temperate broad-leaved forest during the 1995 summer drought, Plant Cell Environ., 20, 1108–1122, 1997. </reference>
		<reference numeration="11" content_type="text"> Baldocchi, D. and Meyers, T.: On using eco-physiological, micrometeorological and biogeochemical theory to evaluate carbon dioxide, water vapor and trace gas fluxes over vegetation: a perspective, Agr. Forest Meteorol., 90, 1–25, 1998. </reference>
		<reference numeration="12" content_type="text"> Baldocchi, D. D.: Canopy control of trace gas emissions, in: Trace gas emissions by plants, edited by: Sharkey, T. D., Holland, E. A., and Mooney, H. A., Physiological ecology, A series of monographs, texts, and treatises, Academic Press, Inc., San Diego - New York - Boston - London - Sydney - Tokyo - Toronto, 293–333, 1991. </reference>
		<reference numeration="13" content_type="text"> Baldocchi, D. D., Fuentes, J. D., Bowling, D. R., Turnipseed, A. A., and Monson, R. K.: Scaling isoprene fluxes from leaves to canopies: test cases over a boreal aspen and a mixed species temperate forest, J. Appl. Meteorol., 38, 885–898, 1999. </reference>
		<reference numeration="14" content_type="text"> Beauchamp, J., Wisthaler, A., Hansel, A., Kleist, E., Miebach, M., Niinemets, Ü., Schurr, U., and Wildt, J.: Ozone induced emissions of biogenic VOC from tobacco: relations between ozone uptake and emission of LOX products, Plant Cell Environ., 28, 1334–1343, 2005. </reference>
		<reference numeration="15" content_type="text"> Bertin, N. and Staudt, M.: Effect of water stress on monoterpene emissions from young potted holm oak (\textitQuercus ilex L.) trees, Oecologia, 107, 456–462, 1996. </reference>
		<reference numeration="16" content_type="text"> Bertin, N., Staudt, M., Hansen, U., Seufert, G., Ciccioli, P., Foster, P., Fugit, J. L., and Torres, L.: Diurnal and seasonal course of monoterpene emissions from \textitQuercus ilex (L.) under natural conditions – applications of light and temperature algorithms, Atmos. Environ., 31, 135–144, 1997. </reference>
		<reference numeration="17" content_type="text"> Blande, J. D., Tiiva, P., Oksanen, E., and Holopainen, J. K.: Emission of herbivore-induced volatile terpenoids from two hybrid aspen (\textitPopulus tremula~$\times$~\textittremuloides) clones under ambient and elevated ozone concentrations in the field, Global Change Biol., 13, 2538-2550, 2007. </reference>
		<reference numeration="18" content_type="text"> Bowles, E. J.: The chemistry of aromatherapeutic oils, Allen &amp; Unwin, Crows Nest, NSW, Australia, 256~pp., 2003. </reference>
		<reference numeration="19" content_type="text"> Brilli, F., Ciccioli, P., Frattoni, M., Prestininzi, M., Spanedda, A. F., and Loreto, F.: Constitutive and herbivore-induced monoterpenes emitted by \textitPopulus x euroamericana leaves are key volatiles that orient \textitChrysomela populi beetles, Plant Cell Environ., 32, 542–552, 2009. </reference>
		<reference numeration="20" content_type="text"> Caldwell, M. M., Meister, H. P., Tenhunen, J. D., and Lange, O. L.: Canopy structure, light microclimate and leaf gas exchange of \textitQuercus coccifera L in a Portuguese macchia: measurements in different canopy layers and simulations with a canopy model, Trees, 1, 25–41, 1986. </reference>
		<reference numeration="21" content_type="text"> Calogirou, A., Larsen, B. R., and Kotzias, D.: Gas-phase terpene oxidation products: a review, Atmos. Environ., 33, 1423–1439, 1999. </reference>
		<reference numeration="22" content_type="text"> Canard, D., Perru, O., Tauzin, V., Devillard, C., and Bonhoure, J. P.: Terpene composition variations in diverse provenances of \textitCedrus libani (A.) Rich and \textitCedrus atlantica Manet, Trees, 11, 504–510, 1997. </reference>
		<reference numeration="23" content_type="text"> Cardoza, Y. J., Alborn, H. T., and Tumlinson, J. H.: In vivo volatile emissions from peanut plants induced by simultaneous fungal infection and insect damage, J. Chem. Ecol., 28, 161–174, 2002. </reference>
		<reference numeration="24" content_type="text"> Carvalhais, N., Reichstein, M., Seixas, J., Collatz, G. J., Pereira, J. S., Berbigier, P., Carrara, A., Granier, A., Montagnani, L., Papale, D., Rambal, S., Sanz, M. J., and Valentini, R.: Implications of carbon cycle steady state assumptions for biogeochemical modeling performance and inverse parameter retrieval, Global Biogeochem. Cycles, 22, GB2007, doi:10.1029/2007GB003033, 2008. </reference>
		<reference numeration="25" content_type="text"> Cescatti, A. and Niinemets, Ü.: Sunlight capture. Leaf to landscape, in: Photosynthetic adaptation, Chloroplast to landscape, edited by: Smith, W. K., Vogelmann, T. C., and Chritchley, C., Ecological Studies, 178, Springer Verlag, Berlin, 42–85, 2004. </reference>
		<reference numeration="26" content_type="text"> Chen, J. W., Harner, T., Schramm, K.-W., Quan, X., Xue, X. Y., and Kettrup, A.: Quantitative relationships between molecular structures, environmental temperatures and octanol/air partition coefficients of polychlorinated biphenyls, Comput. Biol. Chem., 27, 405–421, 2003. </reference>
		<reference numeration="27" content_type="text"> Ciccioli, P., Fabozzi, C., Brancaleoni, E., Cecinato, A., Frattoni, M., Cieslik, S., Kotzias, D., Seufert, G., Foster, P., and Steinbrecher, R.: Biogenic emission from the Mediterranean pseudosteppe ecosystem present in Castelporziano, Atmos. Environ., 31, 167–175, 1997a. </reference>
		<reference numeration="28" content_type="text"> Ciccioli, P., Fabozzi, C., Brancaleoni, E., Cecinato, A., Frattoni, M., Loreto, F., Kesselmeier, J., Schäfer, L., Bode, K., Torres, L., and Fugit, J.-L.: Use of the isoprene algorithm for predicting the monoterpene emission from the Mediterranean holm oak \textitQuercus ilex L.: performance and limits of this approach, J. Geophys. Res., 102, 23319–23328, 1997b. </reference>
		<reference numeration="29" content_type="text"> Copolovici, L. and Niinemets, Ü.: Salting-in and salting-out effects of ionic and neutral osmotica on limonene and linalool Henry&apos;s law constants and octanol/water partition coefficients, Chemosphere, 69, 621–629, doi:10.1016/j.chemosphere.2007.02.066, 2007. </reference>
		<reference numeration="30" content_type="text"> Copolovici, L. O., Filella, I., Llusià, J., Niinemets, Ü., and Peñuelas, J.: The capacity for thermal protection of photosynthetic electron transport varies for different monoterpenes in \textitQuercus ilex, Plant Physiol., 139, 485–496, 2005. </reference>
		<reference numeration="31" content_type="text"> Copolovici, L. O. and Niinemets, Ü.: Temperature dependencies of Henry&apos;s law constants and octanol/water partition coefficients for key plant volatile monoterpenoids, Chemosphere, 61, 1390–1400, doi:10.1016/j.chemosphere.2005.05.003, 2005. </reference>
		<reference numeration="32" content_type="text"> Corchnoy, S. B., Arey, J., and Atkinson, R.: Hydrocarbon emissions from twelve urban shade trees of Los Angeles, California, air basin, Atmos. Environ., 26, 339–348, 1992. </reference>
		<reference numeration="33" content_type="text"> Dai, Y.-J., Dickinson, R. E., and Wang, Y. P.: A two-big-leaf model for canopy temperature, photosynthesis, and stomatal conductance, J. Climate, 17, 2281–2299, 2004. </reference>
		<reference numeration="34" 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, 537–557, 1997. </reference>
		<reference numeration="35" content_type="text"> Dindorf, T., Kuhn, U., Ganzeveld, L., Schebeske, G., Ciccioli, P., Holzke, C., Köble, R., Seufert, G., and Kesselmeier, J.: Significant light and temperature dependent monoterpene emissions from European beech (\textitFagus sylvatica L.) and their potential impact on the European volatile organic compound budget, J. Geophys. Res.-Atmos., 111, D16305, doi:10.1029/2005JD006751, 2006. </reference>
		<reference numeration="36" content_type="text"> Duhl, T. R., Helmig, D., and Guenther, A.: Sesquiterpene emissions from vegetation: a review, Biogeosciences, 5, 761–777, doi:10.5194/bg-5-761-2008, 2008. </reference>
		<reference numeration="37" content_type="text"> Eder, B. K., Davis, J. M., and Bloomfield, P.: A characterization of the spatiotemporal variability of non-urban ozone concentrations over the eastern United States, Atmos. Environ., 27, 2645–2668, 1993. </reference>
		<reference numeration="38" content_type="text"> Engelhart, G. J., Asa-Awuku, A., Nenes, A., and Pandis, S. N.: CCN activity and droplet growth kinetics of fresh and aged monoterpene secondary organic aerosol, Atmos. Chem. Phys., 8, 3937–3949, doi:10.5194/acp-8-3937-2008, 2008. </reference>
		<reference numeration="39" content_type="text"> Falge, E., Ryel, R. J., Alsheimer, M., and Tenhunen, J. D.: Effects of stand structure and physiology on forest gas exchange: a simulation study for Norway spruce, Trees, 11, 436–448, 1997. </reference>
		<reference numeration="40" content_type="text"> Fiore, A. M., Jacob, D. J., Mathur, R., and Martin, R. V.: Application of empirical orthogonal functions to evaluate ozone simulations with regional and global models, J. Geophys. Res.-Atmos., 108, 4431, doi:10.1029/2002JD003151, 2003. </reference>
		<reference numeration="41" content_type="text"> Firmage, D. H.: Environmental influences on the monoterpene variation in \textitHedeoma drummondii, Biochem. Syst. Ecol., 9, 53–58, 1981. </reference>
		<reference numeration="42" content_type="text"> Fischbach, R. J., Zimmer, I., Steinbrecher, R., Pfichner, A., and Schnitzler, J.-P.: Monoterpene synthase activities in leaves of \textitPicea abies (L.) Karst and \textitQuercus ilex L., Phytochemistry, 54, 257–265, 2000. </reference>
		<reference numeration="43" content_type="text"> Friend, A. D.: Modelling canopy CO&lt;sub&gt;2&lt;/sub&gt; fluxes: are &quot;big-leaf&quot; simplifications justified?, Global Ecol. Biogeogr., 10, 603–619, 2001. </reference>
		<reference numeration="44" content_type="text"> Fuentes, J. D. and Wang, D.: On the seasonality of isoprene emissions from a mixed temperate forest, Ecol. Appl., 9, 1118–1131, 1999. </reference>
		<reference numeration="45" content_type="text"> Fuentes, J. D., Wang, D., and Gu, L.: Seasonal variations in isoprene emissions from a boreal aspen forest, J. Appl. Meteorol., 38, 855–869, 1999. </reference>
		<reference numeration="46" content_type="text"> Funk, J. L., Giardina, C. P., Knohl, A., and Lerdau, M. T.: Influence of nutrient availability, stand age, and canopy structure on isoprene flux in a \textitEucalyptus saligna experimental forest, J. Geophys. Res.-Biogeo., 111, G02012, doi:10.1029/2005JG000085, 2006. </reference>
		<reference numeration="47" content_type="text"> Graus, M., Hansel, A., Wisthaler, A., Lindinger, C., Forkel, R., Hauff, K., Klauer, M., Pfichner, A., Rappenglück, B., Steigner, D., and Steinbrecher, R.: A relaxed-eddy-accumulation (REA) method using an online gas-chromatographic technique and PTR-MS for the measurement of isoprenoid fluxes, Atmos. Environ., 40, 43–54, 2006. </reference>
		<reference numeration="48" content_type="text"> Gray, D. W., Lerdau, M. T., and Goldstein, A. H.: Influences of temperature history, water stress, and needle age on methylbutenol emissions, Ecology, 84, 765–776, 2003. </reference>
		<reference numeration="49" content_type="text"> Grote, R., Mayrhofer, S., Fischbach, R. J., Steinbrecher, R., Staudt, M., and Schnitzler, J.-P.: Process-based modelling of isoprenoid emissions from evergreen leaves of \textitQuercus ilex L., Atmos. Environ., 40, S152–S165, 2006. </reference>
		<reference numeration="50" content_type="text"> Grote, R.: Sensitivity of volatile monoterpene emission to changes in canopy structure: a model-based exercise with a process-based emission model, New Phytol., 173, 550–561, doi:10.1111/j.1469-8137.2006.01946.x, 2007. </reference>
		<reference numeration="51" content_type="text"> Grote, R. and Niinemets, Ü.: Modeling volatile isoprenoid emission – a story with split ends, Plant Biol., 10, 8–28, doi:10.1055/s-2007-964975, 2008. </reference>
		<reference numeration="52" content_type="text"> Grote, R., Lavoir, A.-V., Rambal, S., Staudt, M., Zimmer, I., and Schnitzler, J.-P.: Modelling the drought impact on monoterpene fluxes from an evergreen Mediterranean forest canopy, Oecologia, 160, 213–223, 2009. </reference>
		<reference numeration="53" content_type="text"> Grote, R., Keenan, T., Lavoir, A.-V., and Staudt, M.: Process-based simulation of seasonality and drought stress in monoterpene emission models, Biogeosciences, 7, 257–274, doi:10.5194/bg-7-257-2010, 2010. </reference>
		<reference numeration="54" content_type="text"> Gu, L., Baldocchi, D. D., Wofsy, S. C., Munger, J. W., Michalsky, J. J., Urbanski, S. P., and Boden, T. A.: Response of a deciduous forest to the Mount Pinatubo eruption: enhanced photosynthesis, Science, 299, 2035–2038, 2003. </reference>
		<reference numeration="55" content_type="text"> Guenther, A., Zimmerman, P. R., and Wildermuth, M.: Natural volatile organic compound emission rates for US~woodland landscapes, Atmos. Environ., 28, 1197–1210, 1994. </reference>
		<reference numeration="56" content_type="text"> Guenther, A., Hewitt, C. N., Erickson, D., Fall, R., Geron, C., Graedel, T., Harley, P., Klinger, L., Lerdau, M., McKay, W. A., Pierce, T., Scholes, B., Steinbrecher, R., Tallamraju, R., Taylor, J., and Zimmerman, P.: A global model of natural volatile compound emissions, J. Geophys. Res., 100, 8873–8892, 1995. </reference>
		<reference numeration="57" content_type="text"> Guenther, A., Baugh, W., Davis, K., Hampton, G., Harley, P., Klinger, L., Vierling, L., Zimmerman, P., Allwine, E., Dilts, S., Lamb, B., Westberg, H., Baldocchi, D., Geron, C., and Pierce, T.: Isoprene fluxes measured by enclosure, relaxed eddy accumulation, surface layer gradient, mixed layer gradient, and mixed layer mass balance techniques, J. Geophys. Res.-Atmos., 101, 18555–18567, 1996a. </reference>
		<reference numeration="58" content_type="text"> Guenther, A., Greenberg, J., Harley, P., Helmig, D., Klinger, L., Vierling, L., Zimmerman, P., and Geron, C.: Leaf, branch, stand and landscape scale measurements of volatile organic compound fluxes from US~woodlands, Tree Physiol., 16, 17–24, 1996b. </reference>
		<reference numeration="59" content_type="text"> Guenther, A., Zimmerman, P., Klinger, L., Greenberg, J., Ennis, C., Davis, K., Pollock, M., Westberg, H., Allwine, G., and Geron, C.: Estimates of regional natural volatile organic compound fluxes from enclosure and ambient measurements, J. Geophys. Res., 101, 1345–1359, 1996c. </reference>
		<reference numeration="60" content_type="text"> Guenther, A.: Seasonal and spatial variations in natural volatile organic compound emissions, Ecol. Appl., 7, 34–45, 1997. </reference>
		<reference numeration="61" content_type="text"> Guenther, A.: Modeling biogenic volatile organic compound emissions to the atmosphere, in: Reactive hydrocarbons in the atmosphere, edited by: Hewitt, C. N., Academic Press, San Diego, 41–94, 1999. </reference>
		<reference numeration="62" content_type="text"> Guenther, A., Baugh, B., Brasseur, G., Greenberg, J., Harley, P., Klinger, L., Serça, D., and Vierling, L.: Isoprene emission estimates and uncertainties for the Central African EXPRESSO study domain, J. Geophys. Res.-Atmos., 104, 30625–30639, 1999. </reference>
		<reference numeration="63" content_type="text"> Guenther, A., Karl, T., Harley, P., Wiedinmyer, C., Palmer, P. I., and Geron, C.: Estimates of global terrestrial isoprene emissions using MEGAN (Model of Emissions of Gases and Aerosols from Nature), Atmos. Chem. Phys., 6, 3181–3210, doi:10.5194/acp-6-3181-2006, 2006. </reference>
		<reference numeration="64" content_type="text"> Guenther, A. B., Monson, R. K., and Fall, R.: Isoprene and monoterpene emission rate variability: observations with \textitEucalyptus and emission rate algorithm development, J. Geophys. Res., 96, 10799–10808, 1991. </reference>
		<reference numeration="65" content_type="text"> Guenther, A. B., Zimmerman, P. R., Harley, P. C., Monson, R. K., and Fall, R.: Isoprene and monoterpene emission rate variability: model evaluations and sensitivity analyses, J. Geophys. Res., 98, 12609–12617, 1993. </reference>
		<reference numeration="66" content_type="text"> Hakola, H., Rinne, J., and Laurila, T.: The hydrocarbon emission rates of tea-leafed willow (\textitSalix phylicifolia), silver birch (\textitBetula pendula) and European aspen (\textitPopulus tremula), Atmos. Environ., 32, 1825–1833, 1998. </reference>
		<reference numeration="67" content_type="text"> Hakola, H., Laurila, T., Lindfors, V., Hellen, H., Gaman, A., and Rinne, J.: Variation of the VOC emission rates of birch species during the growing season, Boreal Environ. Res., 6, 237–249, 2001. </reference>
		<reference numeration="68" content_type="text"> Hakola, H., Tarvainen, V., Bäck, J., Ranta, H., Bonn, B., Rinne, J., and Kulmala, M.: Seasonal variation of mono- and sesquiterpene emission rates of Scots pine, Biogeosciences, 3, 93–101, doi:10.5194/bg-3-93-2006, 2006. </reference>
		<reference numeration="69" content_type="text"> Hall, G. D. and Langenheim, J. H.: Temporal changes in the leaf monoterpenes of \textitSequoia sempervirens, Biochem. Syst. Ecol., 14, 61–69, 1986. </reference>
		<reference numeration="70" content_type="text"> Hanna, S. R., Russell, A. G., Wilkinson, J. G., Vukovich, J., and Hansen, D. A.: Monte Carlo estimation of uncertainties in BEIS3 emission outputs and their effects on uncertainties in chemical transport model predictions, J. Geophys. Res.-Atmos., 110, D01302, doi:10.1029/2004JD004986, 2005. </reference>
		<reference numeration="71" content_type="text"> Hansen, U. and Seufert, G.: Temperature and light dependence of $\beta $-caryophyllene emission rates, J. Geophys. Res.-Atmos., 108, 4801, doi:10.1029/2003JD003853, 2003. </reference>
		<reference numeration="72" content_type="text"> Harley, P., Guenther, A., and Zimmerman, P.: Effects of light, temperature and canopy position on net photosynthesis and isoprene emission from sweetgum (\textitLiquidambar styraciflua) leaves, Tree Physiol., 16, 25–32, 1996. </reference>
		<reference numeration="73" content_type="text"> Harley, P., Guenther, A., and Zimmerman, P.: Environmental controls over isoprene emission in deciduous oak canopies, Tree Physiol., 17, 705–714, 1997. </reference>
		<reference numeration="74" content_type="text"> Harley, P., Fridd-Stroud, V., Greenberg, J., Guenther, A., and Vasconcellos, P.: Emission of 2-methyl-3-buten-2-ol by pines: a potentially large natural source of reactive carbon to the atmosphere, J. Geophys. Res., 103, 25479–25486, 1998. </reference>
		<reference numeration="75" content_type="text"> Hayashi, N. and Komae, H.: Geographical variation in terpenes from \textitLindera umbellata and \textitLindera sericea, Phytochemistry, 13, 2171–2174, 1974. </reference>
		<reference numeration="76" content_type="text"> He, C., Murray, F., and Lyons, T.: Seasonal variations in monoterpene emissions from \textitEucalyptus species, Chemosphere, Global Change Sci., 2, 65–76, 2000. </reference>
		<reference numeration="77" content_type="text"> Heald, C. L., Wilkinson, M. J., Monson, R. K., Alo, C. A., Wang, G., and Guenther, A.: Response of isoprene emission to ambient CO&lt;sub&gt;2&lt;/sub&gt; changes and implications for global budgets, Global Change Biol., 15, 1127–1140, 2009. </reference>
		<reference numeration="78" content_type="text"> Helmig, D., Revermann, T., Pollmann, J., Kaltschmidt, O., Hernandez, A. J., Bocquet, F., and David, D.: Calibration system and analytical considerations for quantitative sesquiterpene measurements in air, J. Chromatogr. A, 1002, 193–211, 2003. </reference>
		<reference numeration="79" content_type="text"> Herde, M., Gärtner, K., Köllner, T. G., Fode, B., Boland, W., Gershenzon, J., Gatz, C., and Tholl, D.: Identification and regulation of TPS04/GES, an \textitArabidopsis geranyllinalool synthase catalyzing the first step in the formation of the insect-induced volatile C$_16$-homoterpene TMTT, Plant Cell, 20, 1152–1168, 2008. </reference>
		<reference numeration="80" content_type="text"> Holzinger, R., Lee, A., McKay, M., and Goldstein, A. H.: Seasonal variability of monoterpene emission factors for a ponderosa pine plantation in California, Atmos. Chem. Phys., 6, 1267–1274, doi:10.5194/acp-6-1267-2006, 2006. </reference>
		<reference numeration="81" content_type="text"> Huber, D. P. W., Philippe, R. N., Godard, K.-A., Sturrock, R. N., and Böhlmann, J.: Characterization of four terpene synthase \mboxcDNAs from methyl jasmonate-induced Douglas-fir, \textitPseudotsuga menziesii, Phytochemistry, 66, 1427–1439, 2005. </reference>
		<reference numeration="82" content_type="text"> Huber, L., Laville, P., and Fuentes, J. D.: Uncertainties in isoprene emissions from a mixed deciduous forest estimated using a canopy microclimate model, J. Appl. Meteorol., 38, 899–912, 1999. </reference>
		<reference numeration="83" content_type="text"> Janson, R. W.: Monoterpene emissions from Scots pine and Norwegian spruce, J. Geophys. Res., 98, 2839–2850, 1993. </reference>
		<reference numeration="84" content_type="text"> Jones, C. A. and Rasmussen, R. A.: Production of isoprene by leaf tissue, Plant Physiol., 55, 982–987, 1975. </reference>
		<reference numeration="85" content_type="text"> Kahl, J., Hoffmann, T., and Klockow, D.: Differentiation between de novo synthesized and constitutively released terpenoids from \textitFagus sylvatica, Phytochemistry, 51, 383–388, 1999. </reference>
		<reference numeration="86" content_type="text"> Karl, T., Guenther, A., Turnipseed, A., Patton, E. G., and Jardine, K.: Chemical sensing of plant stress at the ecosystem scale, Biogeosciences, 5, 1287–1294, doi:10.5194/bg-5-1287-2008, 2008. </reference>
		<reference numeration="87" content_type="text"> Karl, T. G., Spirig, C., Rinne, J., Stroud, C., Prevost, P., Greenberg, J., Fall, R., and Guenther, A.: Virtual disjunct eddy covariance measurements of organic compound fluxes from a subalpine forest using proton transfer reaction mass spectrometry, Atmos. Chem. Phys., 2, 279–291, doi:10.5194/acp-2-279-2002, 2002. </reference>
		<reference numeration="88" content_type="text"> Karl, T. G., Christian, T. J., Yokelson, R. J., Artaxo, P., Hao, W. M., and Guenther, A.: The Tropical Forest and Fire Emissions Experiment: method evaluation of volatile organic compound emissions measured by PTR-MS, FTIR, and GC from tropical biomass burning, Atmos. Chem. Phys., 7, 5883–5897, doi:10.5194/acp-7-5883-2007, 2007. </reference>
		<reference numeration="89" content_type="text"> Keenan, T., Niinemets, Ü., Sabate, S., Gracia, C., and Peñuelas, J.: Process based inventory of isoprenoid emissions from European forests: model comparisons, current knowledge and uncertainties, Atmos. Chem. Phys., 9, 4053–4076, doi:10.5194/acp-9-4053-2009, 2009. </reference>
		<reference numeration="90" content_type="text"> Kesselmeier, J., Bode, K., Hofmann, U., Müller, H., Schäfer, L., Wolf, A., Ciccioli, P., Brancaleoni, E., Cecinato, A., Frattoni, M., Foster, P., Ferrari, C., Jacob, V., Fugit, J. L., Dutaur, L., Simon, V., and Torres, L.: Emission of short chained organic acids, aldehydes and monoterpenes from \textitQuercus ilex L and \textitPinus pinea L in relation to physiological activities, carbon budget and emission algorithms, Atmos. Environ., 31, 119–133, 1997. </reference>
		<reference numeration="91" content_type="text"> Kim, S., Karl, T., Helmig, D., Daly, R., Rasmussen, R., and Guenther, A.: Measurement of atmospheric sesquiterpenes by proton transfer reaction-mass spectrometry (PTR-MS), Atmos. Meas. Tech., 2, 99–112, doi:10.5194/amt-2-99-2009, 2009. </reference>
		<reference numeration="92" content_type="text"> Komenda, M. and Koppmann, R.: Monoterpene emissions from Scots pine (\textitPinus sylvestris): field studies of emission rate variabilities, J. Geophys. Res., 107, 4161, doi:10.1029/2001JD000691, 2002. </reference>
		<reference numeration="93" content_type="text"> König, G., Brunda, M., Puxbaum, H., Hewitt, C. N., Duckham, S. C., and Rudolph, J.: Relative contribution of oxygenated hydrocarbons to the total biogenic VOC emissions of selected Mid-European agricultural and natural plant species, Atmos. Environ., 29, 861–874, 1995. </reference>
		<reference numeration="94" content_type="text"> Kuhn, U., Rottenberger, S., Biesenthal, T., Wolf, A., Schebeske, G., Ciccioli, P., Brancaleoni, E., Frattoni, M., Tavares, T. M., and Kesselmeier, J.: Isoprene and monoterpene emissions of Amazonian tree species during the wet season: direct and indirect investigations on controlling environmental functions, J. Geophys. Res., D107, 8071, doi:10.1029/2001JD000978, 2002. </reference>
		<reference numeration="95" content_type="text"> Kuhn, U., Rottenberger, S., Biesenthal, T., Wolf, A., Schebeske, G., Ciccioli, P., and Kesselmeier, J.: Strong correlation between isoprene emission and gross photosynthetic capacity during leaf phenology of the tropical tree species \textitHymenaea courbaril with fundamental changes in volatile organic compounds emission composition during early leaf development, Plant Cell Environ., 27, 1469–1485, 2004. </reference>
		<reference numeration="96" content_type="text"> Kulmala, M., Suni, T., Lehtinen, K. E. J., Dal Maso, M., Boy, M., Reissell, A., Rannik, Ü., Aalto, P., Keronen, P., Hakola, H., Bäck, J., Hoffmann, T., Vesala, T., and Hari, P.: A new feedback mechanism linking forests, aerosols, and climate, Atmos. Chem. Phys., 4, 557–562, doi:10.5194/acp-4-557-2004, 2004. </reference>
		<reference numeration="97" content_type="text"> Lasslop, G., Reichstein, M., Kattge, J., and Papale, D.: Influences of observation errors in eddy flux data on inverse model parameter estimation, Biogeosciences, 5, 1311–1324, doi:10.5194/bg-5-1311-2008, 2008. </reference>
		<reference numeration="98" content_type="text"> Lehning, A., Zimmer, I., Steinbrecher, R., Brüggemann, N., and Schnitzler, J. P.: Isoprene synthase activity and its relation to isoprene emission in \textitQuercus robur L leaves, Plant Cell Environ., 22, 495–504, 1999. </reference>
		<reference numeration="99" content_type="text"> Lerdau, M. and Throop, H. L.: Sources of variability in isoprene emission and photosynthesis in two species of tropical wet forest trees, Biotropica, 32, 670–676, 2000. </reference>
		<reference numeration="100" content_type="text"> Letchamo, W., Marquard, R., Hölzl, J., and Gosselin, A.: Effects of water supply and light intensity on growth and essential oil of two \textitThymus vulgaris selections, Angew. Bot., 68, 83–88, 1994. </reference>
		<reference numeration="101" content_type="text"> Lichtenthaler, H. K., Schwender, J., Disch, A., and Rohmer, M.: Biosynthesis of isoprenoids in higher plant chloroplasts proceeds via a mevalonate-independent pathway, FEBS Lett., 400, 271–274, 1997. </reference>
		<reference numeration="102" content_type="text"> Lloyd, J., Wong, S. C., Styles, J. M., Batten, D., Priddle, R., Turnbull, C., and McConchie, C. A.: Measuring and modelling whole-tree gas exchange, Aust. J. Plant Physiol., 22, 987–1000, 1995. </reference>
		<reference numeration="103" content_type="text"> Llusià, J. and Peñuelas, J.: Seasonal patterns of terpene content and emission from seven Mediterranean woody species in field conditions, Am. J. Bot., 87, 133–140, 2000. </reference>
		<reference numeration="104" content_type="text"> Logan, J. A.: Ozone in rural areas of the United States, J. Geophys. Res.-Atmos., 94, 8511–8532, 1989. </reference>
		<reference numeration="105" content_type="text"> Loreto, F. and Sharkey, T. D.: A gas-exchange study of photosynthesis and isoprene emission in \textitQuercus rubra L., Planta, 182, 523–531, 1990. </reference>
		<reference numeration="106" content_type="text"> Loreto, F., Ciccioli, P., Brancaleoni, E., Cecinato, A., Frattoni, M., and Sharkey, T. D.: Different sources of reduced carbon contribute to form three classes of terpenoid emitted by \textitQuercus ilex L leaves, P. Natl. Acad. Sci. USA, 93, 9966–9969, 1996a. </reference>
		<reference numeration="107" content_type="text"> Loreto, F., Ciccioli, P., Cecinato, A., Brancaleoni, E., Frattoni, M., Fabozzi, C., and Tricoli, D.: Evidence of the photosynthetic origin of monoterpenes emitted by \textitQuercus ilex L leaves by $^13$C labeling, Plant Physiol., 110, 1317–1322, 1996b. </reference>
		<reference numeration="108" content_type="text"> Loreto, F., Ciccioli, P., Cecinato, A., Brancaleoni, E., Frattoni, M., and Tricoli, D.: Influence of environmental factors and air composition on the emission of α-pinene from \textitQuercus ilex leaves, Plant Physiol., 110, 267–275, 1996c. </reference>
		<reference numeration="109" content_type="text"> Loreto, F., Förster, A., Dürr, M., Csiky, O., and Seufert, G.: On the monoterpene emission under heat stress and on the increased thermotolerance of leaves of \textitQuercus ilex L fumigated with selected monoterpenes, Plant Cell Environ., 21, 101–107, 1998. </reference>
		<reference numeration="110" content_type="text"> Loreto, F., Nascetti, P., Graverini, A., and Mannozzi, M.: Emission and content of monoterpenes in intact and wounded needles of the Mediterranean pine, \textitPinus pinea, Funct. Ecol., 14, 589–595, 2000. </reference>
		<reference numeration="111" content_type="text"> Loreto, F., Fischbach, R. J., Schnitzler, J. P., Ciccioli, P., Brancaleoni, E., Calfapietra, C., and Seufert, G.: Monoterpene emission and monoterpene synthase activities in the Mediterranean evergreen oak \textitQuercus ilex L grown at elevated CO&lt;sub&gt;2&lt;/sub&gt;, Global Change Biol., 7, 709–717, 2001. </reference>
		<reference numeration="112" content_type="text"> Loreto, F., Centritto, M., Barta, C., Calfapietra, C., Fares, S., and Monson, R. K.: The relationship between isoprene emission rate and dark respiration rate in white poplar (\textitPopulus alba L.) leaves, Plant Cell Environ., 30, 662–669, 2007. </reference>
		<reference numeration="113" content_type="text"> Loughner, C. P., Lary, D. J., Sparling, L. C., Cohen, R. C., DeCola, P., and Stockwell, W. R.: A method to determine the spatial resolution required to observe air quality from space, IEEE T. Geosci. Remote, 45, 1308–1314, 2007. </reference>
		<reference numeration="114" content_type="text"> Magel, E., Mayrhofer, S., Müller, A., Zimmer, I., Hampp, R., and Schnitzler, J.-P.: Photosynthesis and substrate supply for isoprene biosynthesis in poplar leaves, Atmos. Environ., 40, S138–S151, 2006. </reference>
		<reference numeration="115" content_type="text"> Martin, D., Gershenzon, J., and Bohlmann, J.: Induction of volatile terpene biosynthesis and diurnal emission by methyl jasmonate in foliage of Norway spruce, Plant Physiol., 132, 1586–1599, 2003. </reference>
		<reference numeration="116" content_type="text"> Martin, M. J., Stirling, C. M., Humphries, S. W., and Long, S. P.: A process-based model to predict the effects of climatic change on leaf isoprene emission rates, Ecol. Model., 131, 161–174, 2000. </reference>
		<reference numeration="117" content_type="text"> Mayrhofer, S., Teuber, M., Zimmer, I., Louis, S., Fischbach, R. J., and Schnitzler, J.-P.: Diurnal and seasonal variation of isoprene biosynthesis-related genes in grey poplar leaves, Plant Physiol., 139, 474–484, 2005. </reference>
		<reference numeration="118" content_type="text"> Mentel, T. F., Wildt, J., Kiendler-Scharr, A., Kleist, E., Tillmann, R., Dal Maso, M., Fisseha, R., Hohaus, T., Spahn, H., Uerlings, R., Wegener, R., Griffiths, P. T., Dinar, E., Rudich, Y., and Wahner, A.: Photochemical production of aerosols from real plant emissions, Atmos. Chem. Phys., 9, 4387–4406, doi:10.5194/acp-9-4387-2009, 2009. </reference>
		<reference numeration="119" content_type="text"> Merk, L., Kloos, M., Schönwitz, R., and Ziegler, H.: Influence of various factors on quantitative composition of leaf monoterpenes of \textitPicea abies (L.) Karst., Trees, 2, 45–51, 1988. </reference>
		<reference numeration="120" content_type="text"> Meylan, W. M. and Howard, P. H.: Estimating octanol-air partition coefficients with octanol-water partition coefficients and Henry&apos;s law constants, Chemosphere, 61, 640–644, 2005. </reference>
		<reference numeration="121" content_type="text"> Mgalobilishvili, M. P., Khetsuriani, N. D., Kalandadze, A. N., and Sanadze, G. A.: Localization of isoprene biosynthesis in poplar leaf chloroplasts, Fiziol. Rast., 25, 1055–1061, 1978. </reference>
		<reference numeration="122" content_type="text"> Monson, R. K. and Fall, R.: Isoprene emission from aspen leaves. Influence of environment and relation to photosynthesis and photorespiration, Plant Physiol., 90, 267–274, 1989. </reference>
		<reference numeration="123" content_type="text"> Monson, R. K., Hills, A. J., Zimmerman, P. R., and Fall, R. R.: Studies of the relationship between isoprene emission rate and CO&lt;sub&gt;2&lt;/sub&gt; or photon-flux density using a real-time isoprene analyser, Plant Cell Environ., 14, 517–523, 1991. </reference>
		<reference numeration="124" content_type="text"> Monson, R. K., Jaeger, C. H., Adams~III, W. W., Driggers, E. M., Silver, G. M., and Fall, R.: Relationships among isoprene emission rate, photosynthesis, and isoprene synthase activity as influenced by temperature, Plant Physiol., 98, 1175–1180, 1992. </reference>
		<reference numeration="125" content_type="text"> Monson, R. K., Trahan, N., Rosenstiel, T. N., Veres, P., Moore, D., Wilkinson, M., Norby, R. J., Volder, A., Tjoelker, M. G., Briske, D. D., Karnosky, D. F., and Fall, R.: Isoprene emission from terrestrial ecosystems in response to global change: minding the gap between models and observations, Philos. T. Roy. Soc. Lond A, 365, 1677–1695, 2007. </reference>
		<reference numeration="126" content_type="text"> Niinemets, Ü., Tenhunen, J. D., Harley, P. C., and Steinbrecher, R.: A model of isoprene emission based on energetic requirements for isoprene synthesis and leaf photosynthetic properties for \textitLiquidambar and \textitQuercus, Plant Cell Environ., 22, 1319–1336, 1999. </reference>
		<reference numeration="127" content_type="text"> Niinemets, Ü., Hauff, K., Bertin, N., Tenhunen, J. D., Steinbrecher, R., and Seufert, G.: Monoterpene emissions in relation to foliar photosynthetic and structural variables in Mediterranean evergreen \textitQuercus species, New Phytol., 153, 243–256, 2002a. </reference>
		<reference numeration="128" content_type="text"> Niinemets, Ü. and Reichstein, M.: A model analysis of the effects of nonspecific monoterpenoid storage in leaf tissues on emission kinetics and composition in Mediterranean sclerophyllous \textitQuercus species, Global Biogeochem. Cycles, 16, 1110, doi:1110.1029/2002GB001927, 2002. </reference>
		<reference numeration="129" content_type="text"> Niinemets, Ü., Reichstein, M., Staudt, M., Seufert, G., and Tenhunen, J. D.: Stomatal constraints may affect emission of oxygenated monoterpenoids from the foliage of \textitPinus pinea, Plant Physiol., 130, 1371–1385, 2002b. </reference>
		<reference numeration="130" content_type="text"> Niinemets, Ü., Seufert, G., Steinbrecher, R., and Tenhunen, J. D.: A model coupling foliar monoterpene emissions to leaf photosynthetic characteristics in Mediterranean evergreen \textitQuercus species, New Phytol., 153, 257–276, 2002c. </reference>
		<reference numeration="131" content_type="text"> Niinemets, Ü. and Reichstein, M.: Controls on the emission of plant volatiles through stomata: sensitivity or insensitivity of the emission rates to stomatal closure explained, J. Geophys. Res.-Atmos., 108, 4208, doi:4210.1029/2002JD002620, 2003. </reference>
		<reference numeration="132" content_type="text"> Niinemets, Ü.: Costs of production and physiology of emission of volatile leaf isoprenoids, in: Advances in Plant Physiology, edited by: Hemantaranjan, A., Scientific Publishers, Jodhpur, 233–268, 2004. </reference>
		<reference numeration="133" content_type="text"> Niinemets, Ü., Loreto, F., and Reichstein, M.: Physiological and physico-chemical controls on foliar volatile organic compound emissions, Trends Plant Sci., 9, 180–186, 2004. </reference>
		<reference numeration="134" content_type="text"> Niinemets, Ü.: Photosynthesis and resource distribution through plant canopies, Plant Cell Environ., 30, 1052–1071, 2007. </reference>
		<reference numeration="135" content_type="text"> Niinemets, Ü.: Getting hold of terpene emissions from vegetation, ILeaps Newsletter, 5, 40–42, 2008. </reference>
		<reference numeration="136" content_type="text"> Niinemets, Ü. and Anten, N. P. R.: Packing photosynthesis machinery: from leaf to canopy, in: Photosynthesis in silico: understanding complexity from molecules to ecosystems, edited by: Laisk, A., Nedbal, L., and Govindjee, Advances in photosynthesis and respiration, Springer Verlag, 29, Berlin, 363–399, 2009. </reference>
		<reference numeration="137" content_type="text"> Niinemets, Ü.: Mild versus severe stress and BVOCs: thresholds, priming and consequences, Trends Plant Sci., 15, 145–153, 2010. </reference>
		<reference numeration="138" content_type="text"> Niinemets, Ü., Arneth, A., Kuhn, U., Monson, R. K., Peñuelas, J., and Staudt, M.: The emission factor of volatile isoprenoids: stress, acclimation, and developmental responses, Biogeosciences Discuss., 7, 1529–1574, doi:10.5194/bgd-7-1529-2010, 2010. </reference>
		<reference numeration="139" content_type="text"> Noe, S. M., Ciccioli, P., Brancaleoni, E., Loreto, F., and \mboxNiinemets, Ü.: Emissions of monoterpenes linalool and ocimene respond differently to environmental changes due to differences in physico-chemical characteristics, Atmos. Environ., 40, 4649–4662, 2006. </reference>
		<reference numeration="140" content_type="text"> Noe, S. M., Copolovici, L., Niinemets, Ü., and Vaino, E.: Foliar limonene uptake scales positively with leaf lipid content: &quot;non-emitting&quot; species absorb and release monoterpenes, Plant Biol., 10, 129–137, doi:10.1055/s-2007-965239, 2008. </reference>
		<reference numeration="141" content_type="text"> Noe, S. M., Niinemets, Ü., and Schnitzler, J.-P.: Modeling the temporal dynamics of monoterpene emission by isotopic labeling in \textitQuercus ilex leaves, Atmos. Environ., 44, 392–399, 2010. </reference>
		<reference numeration="142" content_type="text"> Ortega, J., Helmig, D., Guenther, A., Harley, P., Pressley, S., and Vogel, C.: Flux estimates and OH reaction potential of reactive biogenic volatile organic compounds (BVOCs) from a mixed northern hardwood forest, Atmos. Environ., 41, 5479–5495, 2007. </reference>
		<reference numeration="143" content_type="text"> Owen, S., Boissard, C., Street, R. A., Duckham, S. C., Csiky, O., and Hewitt, C. N.: Screening of 18~Mediterranean plant species for volatile organic compound emissions, Atmos. Environ., 31, 101–117, 1997. </reference>
		<reference numeration="144" content_type="text"> Owen, S. M., Boissard, C., Hagenlochera, B., and Hewitt, C. N.: Field studies of isoprene emissions from vegetation in the Northwest Mediterranean region, J. Geophys. Res.-Atmos., 103, 25499–25511, 1998. </reference>
		<reference numeration="145" content_type="text"> Owen, S. M. and Hewitt, C. N.: Extrapolating branch enclosure measurements to estimates of regional scale biogenic VOC fluxes in the northwestern Mediterranean basin, J. Geophys. Res.-Atmos., 105, 11573–11583, 2000. </reference>
		<reference numeration="146" content_type="text"> Owen, S. M., Boissard, C., and Hewitt, C. N.: Volatile organic compounds (VOCs) emitted from 40~Mediterranean plant species: VOC speciation and extrapolation to habitat scale, Atmos. Environ., 35, 5393–5409, 2001. </reference>
		<reference numeration="147" content_type="text"> Owen, S. M., Harley, P., Guenther, A., and Hewitt, C. N.: Light dependency of VOC emissions from selected Mediterranean plant species, Atmos. Environ., 36, 3147–3159, 2002. </reference>
		<reference numeration="148" content_type="text"> Owen, S. M., MacKenzie, A. R., Stewart, H., Donovan, R., and Hewitt, C. N.: Biogenic volatile organic compound (VOC) emission estimates from an urban tree canopy, Ecol. Appl., 13, 927–938, 2003. </reference>
		<reference numeration="149" content_type="text"> Paluch, G., Grodnitzky, J., Bartholomay, L., and Coats, J.: Quantitative structure-activity relationship of botanical sesquiterpenes: spatial and contact repellency to the yellow fever mosquito, \textitAedes aegypti, J. Agr. Food Chem., 57, 7618–7625, 2009. </reference>
		<reference numeration="150" content_type="text"> Paré, P. W. and Tumlinson, J. H.: Cotton volatiles synthesized and released distal to the site of insect damage, Phytochemistry, 47, 521–526, 1998. </reference>
		<reference numeration="151" content_type="text"> Paré, P. W. and Tumlinson, J. H.: Plant volatiles as a defense against insect herbivores, Plant Physiol., 121, 325–331, 1999. </reference>
		<reference numeration="152" content_type="text"> Peñuelas, J. and Llusià, J.: Effects of carbon dioxide, water supply, and seasonality on terpene content and emission by \textitRosmarinus officinalis, J. Chem. Ecol., 23, 979–993, 1997. </reference>
		<reference numeration="153" content_type="text"> Peñuelas, J. and Llusià, J.: Seasonal emission of monoterpenes by the Mediterranean tree \textitQuercus ilex in field conditions: relations with photosynthetic rates, temperature and volatility, Physiol. Plant., 105, 641–647, 1999. </reference>
		<reference numeration="154" content_type="text"> Peñuelas, J., Filella, I., Seco, R., and Llusià, J.: Increase in isoprene and monoterpene emissions after re-watering of droughted \textitQuercus ilex seedlings, Biol. Plant., 53, 351–354, 2009. </reference>
		<reference numeration="155" content_type="text"> Peñuelas, J. and Staudt, M.: BVOCs and global change, Trends Plant Sci., 15, 133–144, 2010. </reference>
		<reference numeration="156" content_type="text"> Pétron, G., Harley, P., Greenberg, J., and Guenther, A.: Seasonal temperature variations influence isoprene emission, Geophys. Res. Lett., 28, 1707–1710, 2001. </reference>
		<reference numeration="157" content_type="text"> Pinto, D. M., Blande, J. D., Nykänen, R., Dong, W.-X., Nerg, A.-M., and Holopainen, J. K.: Ozone degrades common herbivore-induced plant volatiles: does this affect herbivore prey location by predators and parasitoids?, J. Chem. Ecol., 33, 683–694, 2007. </reference>
		<reference numeration="158" content_type="text"> Pio, C. A., Nuñes, T. V., and Brito, S.: Volatile hydrocarbon emissions from common and native species of vegetation in Portugal, in: Proceedings of the joint Workshop of CEC/BIATEX of EUROTRAC, General Assessment of Biogenic Emissions and Deposition of Nitrogen Compounds, Sulfur compounds and Oxidants in Europe, edited by: Slanina, J., Angeletti, G., and Beilke, S., Air Pollution Research Report, 47, EC, Directorate-General for Science, Research and Development Aveiro, Portugal, 291–298, 1993. </reference>
		<reference numeration="159" content_type="text"> Pio, C. A., Silva, P. A., Cerqueira, M. A., and Nuñes, T. V.: Diurnal and seasonal emissions of volatile organic compounds from cork oak (\textitQuercus suber) trees, Atmos. Environ., 39, 1817–1827, 2005. </reference>
		<reference numeration="160" content_type="text"> Possell, M., Hewitt, C. N., and Beerling, D. J.: The effects of glacial atmospheric CO&lt;sub&gt;2&lt;/sub&gt; concentrations and climate on isoprene emissions by vascular plants, Global Change Biol., 11, 60–69, 2005. </reference>
		<reference numeration="161" content_type="text"> Rapparini, F., Baraldi, R., Miglietta, F., and Loreto, F.: Isoprenoid emission in trees of \textitQuercus pubescens and \textitQuercus ilex with lifetime exposure to naturally high CO&lt;sub&gt;2&lt;/sub&gt; environment, Plant Cell Environ., 27, 381–391, 2004. </reference>
		<reference numeration="162" content_type="text"> Rasulov, B., Hüve, K., Välbe, M., Laisk, A., and Niinemets, Ü.: Evidence that light, carbon dioxide and oxygen dependencies of leaf isoprene emission are driven by energy status in hybrid aspen, Plant Physiol., 151, 448–460, 2009. </reference>
		<reference numeration="163" content_type="text"> Rinne, H. J. I., Guenther, A. B., Greenberg, J. P., and Harley, P. C.: Isoprene and monoterpene fluxes measured above Amazonian rainforest and their dependence on light and temperature, Atmos. Environ., 36, 2421–2426, 2002. </reference>
		<reference numeration="164" content_type="text"> Rohloff, J.: Monoterpene composition of essential oil from peppermint (\textitMentha x piperita L.) with regard to leaf position using solid-phase microextraction and gas chromatography/mass spectrometry analysis, J. Agr. Food Chem., 47, 3782–3786, 1999. </reference>
		<reference numeration="165" content_type="text"> Rosenstiel, T. N., Potosnak, M. J., Griffin, K. L., Fall, R., and Monson, R. K.: Increased CO&lt;sub&gt;2&lt;/sub&gt; uncouples growth from isoprene emission in an agriforest ecosystem, Nature, 421, 256–259, 2003. </reference>
		<reference numeration="166" content_type="text"> Rosenstiel, T. N., Ebbets, A. L., Khatri, W. C., Fall, R., and Monson, R. K.: Induction of poplar leaf nitrate reductase: a test of extrachloroplastic control of isoprene emission rate, Plant Biol., 6, 12–21, 2004. </reference>
		<reference numeration="167" content_type="text"> Ruuskanen, T. M., Hakola, H., Kajos, M. K., Hellén, H., Tarvainen, V., and Rinne, J.: Volatile organic compound emissions from Siberian larch, Atmos. Environ., 41, 5807–5812, 2007. </reference>
		<reference numeration="168" content_type="text"> Ryel, R. J.: Light relations in tussock grasses as assessed with a new three-dimensional canopy photosynthesis model. Structure and function of foliage organization of a growth form prevalent in environments characterized by stress, Dr rer Nat Thesis, Julius-Maximilians-Universität Würzburg, 172~pp., 1993. </reference>
		<reference numeration="169" content_type="text"> Sabillón, D. and Cremades, L. V.: Diurnal and seasonal variation of monoterpene emission rates for typical Mediterranean species (\textitPinus pinea and \textitQuercus ilex) from field measurements – relationship with temperature and PAR, Atmos. Environ., 35, 4419–4431, 2001. </reference>
		<reference numeration="170" content_type="text"> Schade, G. W., Goldstein, A. H., Gray, D. W., and Lerdau, M. T.: Canopy and leaf level 2-methyl-3-buten-2-ol fluxes from a ponderosa pine plantation, Atmos. Environ., 34, 3535–3544, 2000. </reference>
		<reference numeration="171" content_type="text"> Schiller, G.: Foliage resin composition of \textitCupressus sempervirens L as affected by environmental factors, Silvae Genet., 42, 297–303, 1993. </reference>
		<reference numeration="172" content_type="text"> Schuh, G., Heiden, A. C., Hoffmann, T., Kahl, J., Rockel, P., Rudolph, J., and Wildt, J.: Emissions of volatile organic compounds from sunflower and beech: dependence on temperature and light intensity, J. Atmos. Chem., 27, 291–318, 1997. </reference>
		<reference numeration="173" content_type="text"> Schurgers, G., Arneth, A., Holzinger, R., and Goldstein, A. H.: Process-based modelling of biogenic monoterpene emissions combining production and release from storage, Atmos. Chem. Phys., 9, 3409–3423, doi:10.5194/acp-9-3409-2009, 2009a. </reference>
		<reference numeration="174" content_type="text"> Schurgers, G., Hickler, T., Miller, P. A., and Arneth, A.: European emissions of isoprene and monoterpenes from the Last Glacial Maximum to present, Biogeosciences, 6, 2779–2797, doi:10.5194/bg-6-2779-2009, 2009b. </reference>
		<reference numeration="175" content_type="text"> Schwender, J., Zeidler, J., Gröner, R., Müller, C., Focke, M., Braun, S., Lichtenthaler, F. W., and Lichtenthaler, H. K.: Incorporation of 1-deoxy-D-xylulose into isoprene and phytol by higher plants and algae, FEBS Lett., 414, 129–134, 1997. </reference>
		<reference numeration="176" content_type="text"> Seufert, G., Bartzis, J., Bombol, T., Ciccioli, P., Cieslik, S., Dlugi, R., Foster, P., Hewitt, C. N., Kesselmeier, J., Kotzias, D., Lenz, R., Manes, F., Perez Pastor, P., Steinbrecher, R., Torres, L., Valentini, R., and Versino, B.: An overview of the Castelporziano experiments, Atmos. Environ., 31, 5–17, 1997. </reference>
		<reference numeration="177" content_type="text"> Shao, M., Czapiewski, K. V., Heiden, A. C., Kobel, K., Komenda, M., Koppmann, R., and Wildt, J.: Volatile organic compound emissions from Scots pine: mechanisms and description by algorithms, J. Geophys. Res., 106, 20483–20491, 2001. </reference>
		<reference numeration="178" content_type="text"> Sharkey, T. D., Loreto, F., and Delwiche, C. F.: High carbon dioxide and sun/shade effects on isoprene emission from oak and aspen tree leaves, Plant Cell Environ., 14, 333–338, 1991. </reference>
		<reference numeration="179" content_type="text"> Sharkey, T. D., Singsaas, E. L., Lerdau, M. T., and Geron, C. D.: Weather effects on isoprene emission capacity and applications in emissions algorithms, Ecol. Appl., 9, 1132–1137, 1999. </reference>
		<reference numeration="180" content_type="text"> Singsaas, E. L., Laporte, M. M., Shi, J.-Z., Monson, R. K., Bowling, D. R., Johnson, K., Lerdau, M., Jasentuliytana, A., and Sharkey, T. D.: Kinetics of leaf temperature fluctuation affect isoprene emission from red oak (\textitQuercus rubra) leaves, Tree Physiol., 19, 917–924, 1999. </reference>
		<reference numeration="181" content_type="text"> Singsaas, E. L. and Sharkey, T. D.: The effects of high temperature on isoprene synthesis in oak leaves, Plant Cell Environ., 23, 751–757, 2000. </reference>
		<reference numeration="182" content_type="text"> Smith, E. L.: The influence of light and carbon dioxide on photosynthesis, J. Gen. Physiol., 20, 807–830, 1937. </reference>
		<reference numeration="183" content_type="text"> Spanke, J., Rannik, U., Forkel, R., Nigge, W., and Hoffmann, T.: Emission fluxes and atmospheric degradation of monoterpenes above a boreal forest: field measurements and modelling, Tellus, 53, 406–422, 2001. </reference>
		<reference numeration="184" content_type="text"> Spirig, C., Neftel, A., Ammann, C., Dommen, J., Grabmer, W., Thielmann, A., Schaub, A., Beauchamp, J., Wisthaler, A., and Hansel, A.: Eddy covariance flux measurements of biogenic VOCs during ECHO~2003 using proton transfer reaction mass spectrometry, Atmos. Chem. Phys., 5, 465–481, doi:10.5194/acp-5-465-2005, 2005. </reference>
		<reference numeration="185" content_type="text"> Spracklen, D. V., Bonn, B., and Carslaw, K. S.: Boreal forests, aerosols and the impacts on clouds and climate, Philos. Trans. Royal Soc. London~A, 366, 4613–4626, 2008. </reference>
		<reference numeration="186" content_type="text"> Staudinger, J. and Roberts, P. V.: A critical compilation of Henry&apos;s law constant temperature dependence relations for organic compounds in dilute aqueous solutions, Chemosphere, 44, 561–576, 2001. </reference>
		<reference numeration="187" content_type="text"> Staudt, M. and Seufert, G.: Light-dependent emission of monoterpenes by holm oak (\textitQuercus ilex L.), Naturwissenschaften, 82, 89–92, 1995. </reference>
		<reference numeration="188" content_type="text"> Staudt, M., Bertin, N., Hansen, U., Seufert, G., Ciccioli, P., Foster, P., Frenzel, B., and Fugit, J. L.: Seasonal and diurnal patterns of monoterpene emissions from \textitPinus pinea (L.) under field conditions, Atmos. Environ., 31, 145–156, 1997. </reference>
		<reference numeration="189" content_type="text"> Staudt, M. and Bertin, N.: Light and temperature dependence of the emission of cyclic and acyclic monoterpenes from holm oak (\textitQuercus ilex L.) leaves, Plant Cell Environ., 21, 385–395, 1998. </reference>
		<reference numeration="190" content_type="text"> Staudt, M., Bertin, N., Frenzel, B., and Seufert, G.: Seasonal variation in amount and composition of monoterpenes emitted by young \textitPinus pinea trees – implications for emission modeling, J. Atmos. Chem., 35, 77–99, 2000. </reference>
		<reference numeration="191" content_type="text"> Staudt, M., Joffre, R., Rambal, S., and Kesselmeier, J.: Effect of elevated CO&lt;sub&gt;2&lt;/sub&gt; on monoterpene emission of young \textitQuercus ilex trees and its relation to structural and ecophysiological parameters, Tree Physiol., 21, 437–445, 2001a. </reference>
		<reference numeration="192" content_type="text"> Staudt, M., Mandl, N., Joffre, R., and Rambal, S.: Intraspecific variability of monoterpene composition emitted by \textitQuercus ilex leaves, Can. J. Forest Res., 31, 174–180, 2001b. </reference>
		<reference numeration="193" content_type="text"> Staudt, M., Joffre, R., and Rambal, S.: How growth conditions affect the capacity of \textitQuercus ilex leaves to emit monoterpenes, New Phytol., 158, 61–73, 2003. </reference>
		<reference numeration="194" content_type="text"> Staudt, M., Mir, C., Joffre, R., Rambal, S., Bonin, A., Landais, D., and Lumaret, R.: Isoprenoid emissions of \textitQuercus spp (\textitQ suber and \textitQ ilex) in mixed stands contrasting in interspecific genetic introgression, New Phytol., 163, 573–584, 2004. </reference>
		<reference numeration="195" content_type="text"> Staudt, M. and Lhoutellier, L.: Volatile organic compound emission from holm oak infested by gypsy moth larvae: evidence for distinct responses in damaged and undamaged leaves, Tree Physiol., 27, 1433–1440, 2007. </reference>
		<reference numeration="196" content_type="text"> Steinbrecher, R., Hauff, K., Rabong, R., and Steinbrecher, J.: Isoprenoid emission of oak species typical for the Mediterranean area: source strength and controlling variables, Atmos. Environ., 31, 79–88, 1997. </reference>
		<reference numeration="197" content_type="text"> Steinbrecher, R., Hauff, K., Hakola, H., and Rössler, J.: A revised parameterisation for emission modelling of isoprenoids for boreal plants, in: Biogenic VOC emissions and photochemistry in the boreal regions of Europe: Biphorep, Final report, Contract No~ENV4-CT95-0022, Air Pollution research report No~70, edited by: Laurila, T. and Lindfors, V., Office for Official Publications of the European Communities, Luxembourg, 29–44, 1999. </reference>
		<reference numeration="198" content_type="text"> Street, R. A., Owen, S., Duckham, S. C., Boissard, C., and Hewitt, C. N.: Effect of habitat and age on variations in volatile organic compound (VOC) emissions from \textitQuercus ilex and \textitPinus pinea, Atmos. Environ., 31, 89–100, 1997. </reference>
		<reference numeration="199" content_type="text"> Tarvainen, V., Hakola, H., Hellén, H., Bäck, J., Hari, P., and Kulmala, M.: Temperature and light dependence of the VOC emissions of Scots pine, Atmos. Chem. Phys., 5, 989–998, doi:10.5194/acp-5-989-2005, 2005. </reference>
		<reference numeration="200" content_type="text"> Tenhunen, J. D., Yocum, C. S., and Gates, D. M.: Development of a photosynthesis model with an emphasis on ecological applications. I Theory, Oecologia, 26, 89–100, 1976. </reference>
		<reference numeration="201" content_type="text"> Tingey, D. T., Manning, M., Grothaus, L. C., and Burns, W. F.: Influence of light and temperature on monoterpene emission rates from slash pine, Plant Physiol., 65, 797–801, 1980. </reference>
		<reference numeration="202" content_type="text"> Tobolski, J. J. and Hanover, J. W.: Genetic variation in the monoterpenes of Scots pine, For. Sci., 17, 293–299, 1971. </reference>
		<reference numeration="203" content_type="text"> van Roon, A., Parsons, J. R., and Govers, H. A. J.: Gas chromatographic determination of vapour pressure and related thermodynamic properties of monoterpenes and biogenically related compounds, J. Chromatogr A, 955, 105–115, 2002. </reference>
		<reference numeration="204" content_type="text"> Voirin, B., Brun, N., and Rayet, C.: Effects of daylength on the monoterpene composition of leaves of \textitMentha~$\times$~\textitpiperita, Phytochemistry, 29, 749–755, 1990. </reference>
		<reference numeration="205" content_type="text"> Vuorinen, T., Nerg, A.-M., Syrjälä, L., Peltonen, P., and Holopainen, J. K.: \textitEpirrita autumnata induced VOC emission of silver birch differ from emission induced by leaf fungal pathogen, Arthropod-Plant Interact., 1, 159–165, 2007. </reference>
		<reference numeration="206" content_type="text"> Wiberley, A. E., Linskey, A. R., Falbel, T. G., and Sharkey, T. D.: Development of the capacity for isoprene emission in kudzu, Plant Cell Environ., 28, 898–905, 2005. </reference>
		<reference numeration="207" content_type="text"> Wilkinson, M. J., Monson, R. K., Trahan, N., Lee, S., Brown, E., Jackson, R. B., Polley, H. W., Fay, P. A., and Fall, R.: Leaf isoprene emission rate as a function of atmospheric CO&lt;sub&gt;2&lt;/sub&gt; concentration, Global Change Biol., 15, 1189–1200, 2009. </reference>
		<reference numeration="208" content_type="text"> Winer, A. M., Fitz, D. R., and Miller, P. R.: Investigation of the role of natural hydrocarbons in photochemical smog formation in California, Contract No AO-056-32, prepared for the California Air Resources Board, Statewide Air Pollution Research Center, Riverside, California, USA, 1983. </reference>
		<reference numeration="209" content_type="text"> Wu, J. and Baldwin, I. T.: Herbivory-induced signalling in plants: perception and action, Plant Cell Environ., 32, 1161–1174, 2009. </reference>
		<reference numeration="210" content_type="text"> Young, P. J., Arneth, A., Schurgers, G., Zeng, G., and Pyle, J. A.: The CO&lt;sub&gt;2&lt;/sub&gt; inhibition of terrestrial isoprene emission significantly affects future ozone projections, Atmos. Chem. Phys., 9, 2793–2803, doi:10.5194/acp-9-2793-2009, 2009. </reference>
		<reference numeration="211" content_type="text"> Zimmer, W., Brüggemann, N., Emeis, S., Giersch, C., Lehning, A., Steinbrecher, R., and Schnitzler, J. P.: Process-based modelling of isoprene emission by oak leaves, Plant Cell Environ., 23, 585–595, 2000. </reference>
	</references>
</article>

