<?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>4</volume_number>
		<issue_number>4</issue_number>
		<publication_year>2007</publication_year>
	</journal>
	<doi>10.5194/bg-4-657-2007</doi>
	<article_url>http://www.biogeosciences.net/4/657/2007/</article_url>
	<abstract_html>http://www.biogeosciences.net/4/657/2007/bg-4-657-2007.html</abstract_html>
	<fulltext_pdf>http://www.biogeosciences.net/4/657/2007/bg-4-657-2007.pdf</fulltext_pdf>
	<start_page>657</start_page>
	<end_page>671</end_page>
	<publication_date>2007-08-14</publication_date>
	<article_title content_type="html">Model analysis of the effects of atmospheric drivers on storage water use in Scots pine</article_title>
	<authors>
		<author numeration="1" affiliations="1">
			<name>H. Verbeeck</name>
		</author>
		<author numeration="2" affiliations="2">
			<name>K. Steppe</name>
		</author>
		<author numeration="3" affiliations="3">
			<name>N. Nadezhdina</name>
		</author>
		<author numeration="4" affiliations="1">
			<name>M. Op De Beeck</name>
		</author>
		<author numeration="5" affiliations="1">
			<name>G. Deckmyn</name>
		</author>
		<author numeration="6" affiliations="4">
			<name>L. Meiresonne</name>
		</author>
		<author numeration="7" affiliations="2">
			<name>R. Lemeur</name>
		</author>
		<author numeration="8" affiliations="3">
			<name>J. Čermák</name>
		</author>
		<author numeration="9" affiliations="1">
			<name>R. Ceulemans</name>
		</author>
		<author numeration="10" 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, Universiteitsplein 1, 2610 Antwerp, Belgium</affiliation>
		<affiliation numeration="2" content_type="html">Laboratory of Plant Ecology, Department of Applied Ecology and Environmental Biology, Ghent University, Coupure Links 653, 9000 Ghent, Belgium</affiliation>
		<affiliation numeration="3" content_type="html">Institute of Forest Ecology, Mendel University of Agriculture and Forestry, Zemědělská 3, 61300 Brno, Czech Republic</affiliation>
		<affiliation numeration="4" content_type="html">Research Institute for Nature and Forest, Ministry of the Flemish Community, Gaverstraat 4, 9500 Geraardsbergen, Belgium</affiliation>
	</affiliations>
	<abstract content_type="html">Storage water use is an indirect consequence of the interplay between
different meteorological drivers through their effect on water flow and
water potential in trees. We studied these microclimatic drivers of storage
water use in Scots pine (&lt;i&gt;Pinus sylvestris L.&lt;/i&gt;) growing in a temperate climate. The storage
water use was modeled using the ANAFORE model, integrating a dynamic water
flow and &amp;ndash; storage model with a process-based transpiration model. The
model was calibrated and validated with sap flow measurements for the
growing season of 2000 (26 May&amp;ndash;18 October).
&lt;br&gt;&lt;br&gt;

Because there was no severe soil drought during the study period, we were
able to study atmospheric effects. Incoming radiation and vapour pressure
deficit (VPD) were the main atmospheric drivers of storage water use. The
general trends of sap flow and storage water use are similar, and follow
more or less the pattern of incoming radiation. Nevertheless, considerable
differences in the day-to-day pattern of sap flow and storage water use were
observed. VPD was determined to be one of the main drivers of these
differences. During dry atmospheric conditions (high VPD) storage water use
was reduced. This reduction was higher than the reduction in measured sap
flow. Our results suggest that the trees did not rely more on storage water
during periods of atmospheric drought, without severe soil drought. The
daily minimum tree water content was lower in periods of high VPD, but the
reserves were not completely depleted after the first day of high VPD, due
to refilling during the night.
&lt;br&gt;&lt;br&gt;

Nevertheless, the tree water content deficit was a third important factor
influencing storage water use. When storage compartments were depleted
beyond a threshold, storage water use was limited due to the low water
potential in the storage compartments. The maximum relative contribution of
storage water to daily transpiration was also constrained by an increasing
tree water content deficit.</abstract>
	<references>
		<reference numeration="1" content_type="text"> Baeyens, L., Van Slycken, J., and Stevens, D.: Description of the soil profile in Brasschaat, Internal research paper, Institute for Forestry and Game Management, Geraardsbergen, Belgium, 1993. </reference>
		<reference numeration="2" content_type="text"> Bhaskar, R. and Ackerly, D. D.: Ecological relevance of minimum seasonal water potentials, Physiol. Plant., 127, 353&amp;ndash;359, 2006. </reference>
		<reference numeration="3" content_type="text"> Bunce, J. A.: Does transpiration control stomatal responses to water vapour pressure deficit?, Plant Cell Environ., 20, 131&amp;ndash;135, 1997. </reference>
		<reference numeration="4" content_type="text"> Bunce, J. A.: How do leaf hydraulics limit stomatal conductance at high water vapour pressure deficits?, Plant Cell Environ., 29, 1644&amp;ndash;1650, 2006. </reference>
		<reference numeration="5" content_type="text"> Carrara, A., Kowalski, A. S., Neirynck, J., Janssens, I. A., Curiel Yuste, J., and Ceulemans, R.: Net ecosystem CO&lt;sub&gt;2&lt;/sub&gt; exchange of mixed forest in Belgium over 5 years, Agric. For. Meteorol., 119, 209&amp;ndash;227, 2003. </reference>
		<reference numeration="6" content_type="text"> \vCermák, J., Riguzzi, F., and Ceulemans, R.: Scaling up from the individual trees to the stand level in Scots pine: 1. Needle distribution, overall crown and root geometry, Ann. For. Sci., 55, 63&amp;ndash;88, 1998. </reference>
		<reference numeration="7" content_type="text"> \vCermák, J., Ku&amp;#x010D;era, J., and Nadezhdina, N.: Sap flow measurements with some thermodynamic methods, flow integration within trees and scaling up from sample trees to entire forest stands, Trees, 18, 529&amp;ndash;546, 2004. </reference>
		<reference numeration="8" content_type="text"> \vCermák, J., Ku&amp;#x010D;era, J., Bauerle, W. L., Phillips, N., and Hinckley, T. M.: Tree water storage and its diurnal dynamics related to sap flow and changes in stem volume in old-growth Douglas-fir trees, Tree Physiol., 27, 181&amp;ndash;198, 2007. </reference>
		<reference numeration="9" content_type="text"> Chuang, Y.-L., Oren, R., Bertozzi, A. L., Phillips, N., and Katul, G. G.: The porous media model for the hydraulic system of a conifer tree: Linking sap flux data to transpiration rate, Ecol. Model., 191, 447&amp;ndash;468, 2006. </reference>
		<reference numeration="10" content_type="text"> Curiel Yuste, J., Konôpka, B., Janssens, I. A., Coenen, K., Xiao, C. W. and Ceulemans, R.: Contrasting net primary productivity and carbon distribution between neighboring stand of \textitQuercus robur and \textitPinus sylvestris, Tree Physiol., 25, 701&amp;ndash;712, 2005. </reference>
		<reference numeration="11" content_type="text"> Dewar, R. C.: The Ball-Berry-Leuning and Tardieu-Davies stomatal models: synthesis and extension within a spatially aggregated picture of guard cell function, Plant Cell Environ., 25, 1383&amp;ndash;1398, 2002. </reference>
		<reference numeration="12" content_type="text"> Ewers, B. E., Gower, S. T., Bond-Lamberty, B., and Wang, C. K.: Effects of stand age and tree species on canopy transpiration and average stomatal conductance of boreal forests, Plant Cell Environ., 28, 660&amp;ndash;678, 2005. </reference>
		<reference numeration="13" content_type="text"> Farquhar, G. D., Von Caemmerer, S., and Berry., J.A.: A biochemical model of photosynthetic CO&lt;sub&gt;2&lt;/sub&gt; assimilation in leaves of C3 species, Planta, 149, 78&amp;ndash;90, 1980. %</reference>
		<reference numeration="14" content_type="text"> %Ford, C. R., Goranson, C. E., Mitchell, R. J., Will, R. E., and Teskey, R. %O.: Diurnal and seasonal variability in the radial distribution of sap flow: %predicting total stem flow in Pinus taeda trees, Tree Phsysiol., 24, 951&amp;ndash;960, 2004. </reference>
		<reference numeration="15" content_type="text"> Ford, C. R., Goranson, C. E., Mitchell, R. J., Will, R. E., and Teskey, R. O.: Modeling canopy transpiration using time series analysis: A case study illustrating the effect of soil moisture deficit on Pinus taeda, Agric. For. Meteorol., 130, 163&amp;ndash;175, 2005. </reference>
		<reference numeration="16" content_type="text"> Früh, T. and Kurth, W.: The Hydraulic System of Trees: Theoretical Framework and Numerical Simulation, J. Theor. Biol., 201, 251&amp;ndash;270, 1999. </reference>
		<reference numeration="17" content_type="text"> Gao, Q., Zhao, P., Zeng, X., Cai, X., and Shen, W.: A model of stomatal conductance to quantify the relationship between leaf transpiration, microclimate and soil water stress, Plant Cell Environ., 25, 1373&amp;ndash;1381, 2002. </reference>
		<reference numeration="18" content_type="text"> Goldstein, G., Andrade, J. L., Meinzer, F. C., Holbrook, N. M., Cavelier, J., Jackson, P., and Celis, A.: Stem water storage and diurnal patterns of water use in tropical forest canopy trees, Plant Cell Environ., 21, 397&amp;ndash;406, 1998. </reference>
		<reference numeration="19" content_type="text"> Holtta, T., Vesala, T., Peramaki, M., and Nikinmaa, E.: Relationships between Embolism, Stem Water Tension, and Diameter Changes, J. Theor. Biol., 215, 23&amp;ndash;38, 2002. </reference>
		<reference numeration="20" content_type="text"> Hunt Jr., E. R., Running, S. W., and Federer, C. A.: Extrapolating plant water flow resistances and capacitances to regional scales, Agric. For. Meteorol., 54, 169&amp;ndash;195, 1991. </reference>
		<reference numeration="21" content_type="text"> Irvine, J., Perks, M. P., Magnani, F., and Grace, J.: The response of \textitPinus sylvestris to drought: stomatal control of transpiration and hydraulic conductance, Tree Physiol., 18, 393&amp;ndash;402, 1998. </reference>
		<reference numeration="22" content_type="text"> Janssens, I. A., Sampson, D. A., \vCermák, J., Meiresonne, L., Riguzzi, F., Overloop, S. and Ceulemans, R.: Above- and below-ground phytomass and carbon storage in a Belgian Scots pine stand, Ann. For. Sci., 56, 81&amp;ndash;90, 1999. </reference>
		<reference numeration="23" content_type="text"> Jones, H. G.: Plants and Microclimate, A quantitative approach to environmental plant physiology, second edition, University Press, Cambridge, 428 pp., 1992. </reference>
		<reference numeration="24" content_type="text"> Köstner, B., Falge, E. M., Alsheimer, M., Geyer, R., and Tenhunen, J. D.: Estimating tree canopy water use via xylem sap flow in an old Norway spruce forest and a comparison with simulation-based canopy transpiration estimates, Ann. For. Sci., 55, 125&amp;ndash;139, 1998. </reference>
		<reference numeration="25" content_type="text"> Kowalski, A. S., Overloop, S., and Ceulemans, R.: Eddy fluxes above a Belgian, Campine forest and their relationship with predicting variables, in: Forest Ecosystem Modeling, Upscaling and Remote Sensing, edited by: Ceulemans, R., Veroustraete, F., Gond, V., and Van Rensbergen, J., SPB Academic Publishing, The Hague, 3&amp;ndash;17, 2000. </reference>
		<reference numeration="26" content_type="text"> Lhomme, J. P., Rocheteau, A., Ourcival, J. M., and Rambal, S.: Non-steady-state modelling of water transfer in a Mediterranean evergreen canopy, Agric. For. Meteorol., 108, 67&amp;ndash;83, 2001. </reference>
		<reference numeration="27" content_type="text"> Loustau, D., Berbigier, P., Roumagnac, P., Arruda-Pacheco, C., David, J. S., Ferreira, M. I., Pereira, J. S., and Travares, R.: Transpiration of a 64-year-old maritime pine stand in Portugal. 1. Seasonal course of water flux through maritime pine, Oecologia, 107, 33&amp;ndash;42, 1996. </reference>
		<reference numeration="28" content_type="text"> Martin, T. A., Brown, K. J., \vCermák, J., Ceulemans, R., Ku&amp;#x010D;era, J., Meinzer, F. C., Rombold, J. S., Sprugel, D. G., and Hinckley, T. M.: Crown conductance and tree and stand transpiration in a second-growth Abies amabilis forest, Can. J. For. Res., 27, 797&amp;ndash;808, 1997. </reference>
		<reference numeration="29" content_type="text"> Meinzer, F. C., Hinckley, T. M., and Ceulemans, R.: Apparent responses of stomata to transpiration and humidity in a hybrid poplar canopy, Plant Cell Environ., 20, 1301&amp;ndash;1308, 1997. </reference>
		<reference numeration="30" content_type="text"> Meinzer, F. C., Goldstein, G., Franco, A.C., Bustamante, M., Igler, E., Jackson, P., Caldas, L., and Rundel, P. W.: Atmospheric and hydraulic limitations on transpiration in Brazilian cerrado woody species, Funct. Ecol., 13, 273&amp;ndash;282, 1999. </reference>
		<reference numeration="31" content_type="text"> Meinzer, F. C., Clearwater, M. J., and Goldstein, G.: Water transport in trees: current perspectives, new insights and some controversies, Environ. Exp. Bot., 45, 239&amp;ndash;262, 2001. </reference>
		<reference numeration="32" content_type="text"> Meiresonne, L., Sampson, D. A., Kowalski, A. S., Janssens, I. A., Nadezhdina, N., \vCermák, J., Van Slycken, J., and Ceulemans, R.: Water flux estimates from a Belgian Scots pine stand: a comparison of different approaches, J. Hydrol., 270, 230&amp;ndash;252, 2003. </reference>
		<reference numeration="33" content_type="text"> Monteith, J. L.: Evaporation and environment, in: The state and movement of water in living organisms, edited by: Fogg., G. F., 19th Symposium for Experimental Science, Cambridge University Press, London, 205&amp;ndash;234, 1965. </reference>
		<reference numeration="34" content_type="text"> Monteith, J. L.: A reinterpretation of stomatal responses to humidity, Plant Cell Environ., 18, 357&amp;ndash;64, 1995. </reference>
		<reference numeration="35" content_type="text"> Motzer, T., Munz, N., Küppers, M., Schmitt, D., and Anhuff, D.: Stomatal conductance, transpiration and sap flow of tropical montane rain forest trees in southern Ecuadorian Andes, Tree Physiol., 25, 1283&amp;ndash;1293, 2005. </reference>
		<reference numeration="36" content_type="text"> Nadezhdina, N., \vCermák, J., and Nadezhdin, V.: Heat field deformation method for sap flow measurements, Proceedings Fourth International Workshop on Measuring Sap Flow in Intact Plants. Zidlochoviche, Czech Republic, 3&amp;ndash;5 October, IUFRO Publications. Publishing house of Mendel University of Brno, 72&amp;ndash;92, 1998. </reference>
		<reference numeration="37" content_type="text"> Nadezhdina, N. and \vCermák, J.: The technique and instrumentation for estimation the sap flow rate in plants (in Czech), Patent No 286438 (PV-1587-98), 2000. </reference>
		<reference numeration="38" content_type="text"> Nadezhdina, N., \vCermák, J., and Ceulemans, R.: Radial patterns of sap flow in woody stems of dominant and understory species: scaling errors associated with positioning of sensors, Tree Physiol., 22, 907&amp;ndash;918, 2002. </reference>
		<reference numeration="39" content_type="text"> Nadezhdina, N., Tributsch, H., and \vCermák, J.: Infra-red images of heat field around a linear heater and sap flow in stems of lime trees under natural and experimental conditions, Ann. For. Sci., 61, 203&amp;ndash;214, 2004. </reference>
		<reference numeration="40" content_type="text"> Nadezhdina, N., \vCermák, J., Gaspárek, J., Nadezhdin, V., and Prax, A.: Vertical and horizontal water redistribution in Norway spruce (\textitPicea abies) roots in the Moravian Upland, Tree Physiol., 26, 1277&amp;ndash;1288, 2006. </reference>
		<reference numeration="41" content_type="text"> Nadezhdina, N., Nadezhdin, V., Ferreira, M. I., and Pitacco, A.: Variability with xylem depth in sap flow in trunks and branches of mature olive trees, Tree Physiol., 27, 105&amp;ndash;113, 2007. </reference>
		<reference numeration="42" content_type="text"> Nelder, J. A. and Mead, R.: A simplex method for function minimization, Comput. J., 7, 308&amp;ndash;313, 1965. </reference>
		<reference numeration="43" content_type="text"> Neirynck, J., Van Ranst, E., Roskams, P., and Lust, N.: Impact of decreasing throughfall depositions on soil solution chemistry at coniferous monitoring sites in northern Belgium, For. Ecol. Manage., 160, 127&amp;ndash;142, 2002. </reference>
		<reference numeration="44" content_type="text"> Op de Beeck, M., Löw, M., Verbeeck, H., and Deckmyn, G.: Suitability of a combined stomatal conductance and photosynthesis model for calculation of leaf level ozone fluxes, Plant Biol., 9, 331&amp;ndash;341, 2007. </reference>
		<reference numeration="45" content_type="text"> Oren, R., Phillips, N., Ewers, B. E., Pataki, D. E., and Megonigal, J. P: Sap-flux-scaled transpiration responses to light, vapour pressure deficit, and leaf area reduction in a flooded \textitTaxodium distichum forest, Tree Physiol., 19, 337&amp;ndash;347, 1999a. </reference>
		<reference numeration="46" content_type="text"> Oren, R., Sperry, J. S., Katul, G. G., Pataki, D. E., Ewers, B. E., Phillips, N., and Schafer, K. V. R.: Survey and synthesis of intra- and interspecific variation in stomatal sensitivity to vapour pressure deficit, Plant Cell Environ., 22, 1515&amp;ndash;1526, 1999b. </reference>
		<reference numeration="47" content_type="text"> Overloop, S. and Meiresonne., L.: Basiskarakteristieken van het proefvlak Brasschaat, domeinbos de Inslag, Communications of the Institute of Forestry and Game Management, no 1999/1, 11&amp;ndash;21, 1999 (in Dutch). </reference>
		<reference numeration="48" content_type="text"> Penman, H. L.: Natural evaporation from open water, bare soil and grass, Proc. R. Soc, A 193, 120&amp;ndash;145, 1948. </reference>
		<reference numeration="49" content_type="text"> Perämäki, M., Vesala, T., and Nikinmaa, E.: Modeling the dynamics of pressure propagation and diameter variation in tree sapwood, Tree Physiol., 25, 1091&amp;ndash;1099, 2005. </reference>
		<reference numeration="50" content_type="text"> Phillips, N., Ryan, M. G., Bond, B. J., McDowell, N. G., Hinckley, T. M., and \vCermák, J.: Reliance on stored water increases with tree size in three species in the Pacific Northwest, Tree Physiol., 23, 237&amp;ndash;245, 2003. </reference>
		<reference numeration="51" content_type="text"> Press, W. H., Teukolsky, S. A., Vetterling, W. T., and Flannery, B. P.: Numerical recipes in fortran 90, The art of scientific computing, Second edition, Vol 2, Cambridge Univerity Press, 921 pp., 1996. </reference>
		<reference numeration="52" content_type="text"> Roskams, P., Sioen, G., and Overloop, S.: Meetnet voor de intensieve monitoring van het bosecosysteem in het Vlaamse Gewest &amp;ndash; resultaten 1991&amp;ndash;1992, Institute for Forestry and Game Management, Ministry of the Flemmish Community, 191 pp., 1997 (in Dutch). </reference>
		<reference numeration="53" content_type="text"> Sampson, D. A., Janssens, I. A., and Ceulemans, R.: Under-story contributions to stand level GPP using the process model SECRETS, Agric. For. Meteorol., 139, 94&amp;ndash;104, 2006. </reference>
		<reference numeration="54" content_type="text"> Scholz, F. G., Bucci, S. J., Goldstein, G., Meinzer, F. C., Franco, A. C., and Miralles-Wilhelm, F.: Biophysical properties and functional significance of stem water storage tissues in Neotropical savanna trees, Plant Cell Environ., 30, 236&amp;ndash;248, 2007. </reference>
		<reference numeration="55" content_type="text"> Sinclair, T. R., Holbrook, N. M., and Zwieniecki, M. A.: Daily transpiration rates of woody species on drying soil, Tree Physiol., 25, 1469&amp;ndash;1472, 2005. </reference>
		<reference numeration="56" content_type="text"> Steppe, K., De Pauw, D. J. W., Lemeur, R., and Vanrolleghem, P. A.: A mathematical model linking tree sap flow dynamics to daily stem diameter fluctuations and radial stem growth, Tree Physiol., 26, 257&amp;ndash;273, 2006. </reference>
		<reference numeration="57" content_type="text"> Stewart, J. B.: Modelling surface conductance of pine forest, Agric. For. Meteorol., 43, 19&amp;ndash;35, 1988. </reference>
		<reference numeration="58" content_type="text"> van Genuchten, M. T.: A closed-form equation for predicting the hydraulic conductivity of the soil, Soil Sci. Soc. Am. J., 44, 892&amp;ndash;898, 1980. </reference>
		<reference numeration="59" content_type="text"> Verbeeck, H., Steppe, K., Nadezhdina, N., Op de Beeck, M., Deckmyn, G., Meiresonne, L., Lemeur, R., \vCermák, J., Ceulemans, R., and Janssens, I. A.: Water storage and transpiration in Scots pine: a modeling analysis using ANAFORE, Tree Physiol., 27, 1671&amp;ndash;1685, 2007. </reference>
		<reference numeration="60" content_type="text"> Whitehead, D.: Regulation of stomatal conductance and transpiration in forest canopies, Tree Physiol., 18, 633&amp;ndash;644, 1998. </reference>
		<reference numeration="61" content_type="text"> Zweifel, R., Item, H., and Häsler, R.: Link between diurnal stem radius changes and tree water relations, Tree Physiol., 21, 869&amp;ndash;877, 2001. </reference>
		<reference numeration="62" content_type="text"> Zweifel, R., Zimmermann, L., and Newbery, D. M.: Modeling tree water deficit from microclimate: an approach to quantifying drought stress, Tree Physiol., 25, 147&amp;ndash;156, 2005. </reference>
	</references>
</article>

