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<article language="en">
	<journal>
		<journal_title>Biogeosciences</journal_title>
		<journal_url>www.biogeosciences.net</journal_url>
		<issn>1726-4170</issn>
		<eissn>1726-4189</eissn>
		<volume_number>6</volume_number>
		<issue_number>1</issue_number>
		<publication_year>2009</publication_year>
	</journal>
	<doi>10.5194/bg-6-59-2009</doi>
	<article_url>http://www.biogeosciences.net/6/59/2009/</article_url>
	<abstract_html>http://www.biogeosciences.net/6/59/2009/bg-6-59-2009.html</abstract_html>
	<fulltext_pdf>http://www.biogeosciences.net/6/59/2009/bg-6-59-2009.pdf</fulltext_pdf>
	<start_page>59</start_page>
	<end_page>66</end_page>
	<publication_date>2009-01-09</publication_date>
	<article_title content_type="html">Contribution of different grass species to plant-atmosphere ammonia exchange in intensively managed grassland</article_title>
	<authors>
		<author numeration="1" affiliations="1,4">
			<name>M. Mattsson</name>
		</author>
		<author numeration="2" affiliations="2">
			<name>B. Herrmann</name>
		</author>
		<author numeration="3" affiliations="2,3">
			<name>S. Jones</name>
		</author>
		<author numeration="4" affiliations="2">
			<name>A. Neftel</name>
		</author>
		<author numeration="5" affiliations="3">
			<name>M. A. Sutton</name>
		</author>
		<author numeration="6" affiliations="1">
			<name>J. K. Schjoerring</name>
			<email>jks@life.ku.dk</email>
		</author>
	</authors>
	<affiliations>
		<affiliation numeration="1" content_type="html">Plant and Soil Science Laboratory, University of Copenhagen, Faculty of Life Sciences, Thorvaldsensvej 40, 1871 Frederiksberg C, Copenhagen, Denmark</affiliation>
		<affiliation numeration="2" content_type="html">Agroscope Reckenholz-Tänikon Research Station ART, Reckenholzstrasse 191, 8046 Zürich, Switzerland</affiliation>
		<affiliation numeration="3" content_type="html">Natural Environmental Research Council, Centre for Ecology and Hydrology, Edinburgh Research Station, Penicuik EH26 0QB, Midlothian Scotland</affiliation>
		<affiliation numeration="4" content_type="html">now at: Section for Economy and Technology, Halmstad University, Halmstad, 30118 Sweden</affiliation>
	</affiliations>
	<abstract content_type="html">Species diversity in grasslands usually declines with increasing input of
nitrogen from fertilizers or atmospheric deposition. Conversely,
species diversity may also impact the build-up of soil and plant nitrogen
pools. One important pool is NH&lt;sub&gt;3&lt;/sub&gt;/NH&lt;sub&gt;4&lt;/sub&gt;&lt;sup&gt;+&lt;/sup&gt; which also can be
exchanged between plant leaves and the atmosphere. Limited information is
available on how plant-atmosphere ammonia exchange is related to species
diversity in grasslands. We have here investigated grass species abundance
and different foliar nitrogen pools in 4-year-old intensively managed
grassland. Apoplastic pH and NH&lt;sub&gt;4&lt;/sub&gt;&lt;sup&gt;+&lt;/sup&gt; concentrations of the 8 most
abundant species (&lt;i&gt;Lolium perenne, Phleum pratense, Festuca pratensis,
Lolium multiflorum, Poa pratensis, Dactylis glomerata, Holcus lanatus, Bromus mollis&lt;/i&gt;)
were used to calculate stomatal NH&lt;sub&gt;3&lt;/sub&gt; compensation
points. Apoplastic NH&lt;sub&gt;4&lt;/sub&gt;&lt;sup&gt;+&lt;/sup&gt; concentrations differed considerably among
the species, ranging from 13 to 117 μM, with highest values in &lt;i&gt;Festuca pratensis&lt;/i&gt;. Also
apoplastic pH values varied, from pH 6.0 in &lt;i&gt;Phleum pratense&lt;/i&gt;
to 6.9 in &lt;i&gt;Dactylis glomerata&lt;/i&gt;. The observed
differences in apoplastic NH&lt;sub&gt;4&lt;/sub&gt;&lt;sup&gt;+&lt;/sup&gt; and pH resulted in a large span of
predicted values for the stomatal NH&lt;sub&gt;3&lt;/sub&gt; compensation point which ranged
between 0.20 and 6.57 nmol mol&lt;sup&gt;&amp;minus;1&lt;/sup&gt;. Three species (&lt;i&gt;Lolium perenne,
Festuca pratensis&lt;/i&gt; and &lt;i&gt;Dactylis glomerata&lt;/i&gt;) had
sufficiently high NH&lt;sub&gt;3&lt;/sub&gt; compensation point and abundance to contribute to
the bi-directional NH&lt;sub&gt;3&lt;/sub&gt; fluxes recorded over the whole field. The other
5 grass species had NH&lt;sub&gt;3&lt;/sub&gt; compensation points considerably below the
atmospheric NH&lt;sub&gt;3&lt;/sub&gt; concentration and were thus not likely to contribute to
NH&lt;sub&gt;3&lt;/sub&gt; emission but only to NH&lt;sub&gt;3&lt;/sub&gt; uptake from the atmosphere. Evaluated
across species, leaf bulk-tissue NH&lt;sub&gt;4&lt;/sub&gt;&lt;sup&gt;+&lt;/sup&gt; concentrations correlated
well (&lt;i&gt;r&lt;/i&gt;&lt;sup&gt;2&lt;/sup&gt;=0.902) with stomatal NH&lt;sub&gt;3&lt;/sub&gt; compensation points calculated
on the basis of the apoplastic bioassay. This suggests that leaf tissue
NH&lt;sub&gt;4&lt;/sub&gt;&lt;sup&gt;+&lt;/sup&gt; concentrations combined with data for the frequency
distribution of the corresponding species can be used for predicting the
NH&lt;sub&gt;3&lt;/sub&gt; exchange potential of a mixed grass sward.</abstract>
	<references>
		<reference numeration="1" content_type="text"> % vor jede Referenz Bullock, J. M., Pywell, R. F., and Walker, K. J.: Long-term enhancement of agricultural production by restoration of biodiversity, J. Appl. Ecol., 44, 6–12, 2007. </reference>
		<reference numeration="2" content_type="text"> % vor jede Referenz Clark, C. M., Cleland, E. E , Collins, S. L., Fargione, J. E., Gough, L., Gross, K. L., Pennings, S. C., Suding, K. N., and Grace, J. B.: Environmental and plant community determinants of species loss following nitrogen enrichment, Ecol. Lett., 10, 596–607, 2007. </reference>
		<reference numeration="3" content_type="text"> % vor jede Referenz Cleland, E. E., Chiariello, N. R., Loarie, S. R., Mooney, H. A., and Field, C. B.: Diverse responses of phenology to global changes in a grassland ecosystem, Proc. Natl. Acad. Sci. USA, 103, 13740–13744, 2006. </reference>
		<reference numeration="4" content_type="text"> % vor jede Referenz Critchley, C. N. R, Chambers, B. J., Fowbert, J. A., Sanderson, R. A., Bhogal, A., and Rose, S. C.: Association between lowland grassland plant communities and soil properties, Biol. Conserv., 105, 199–215, 2002. </reference>
		<reference numeration="5" content_type="text"> % vor jede Referenz Critchley, C. N. R., Fowbert, J. A., and Wright, B.: Dynamics of species-rich upland hay meadows over 15 years and their relation with agricultural management practices, Appl. Veg. Sci., 10, 307–314, 2007. </reference>
		<reference numeration="6" content_type="text"> % vor jede Referenz David, M., Loubet, B., Cellier, P., Mattsson, M., Schjoerring, J. K., Nemitz, E., Roche, R., Riedo, M., and Sutton, M.A.: Ammonia sources and sinks in an intensively managed grassland using dynamic chambers, Biogeosciences Discuss., accepted, 2008. </reference>
		<reference numeration="7" content_type="text"> % vor jede Referenz Ellenberg, H., Weber H. E., Düll, R., Wirth, V., Werner, W., and Paulissen, D.: Indicator values of plants in Central Europe, Scripta Geobotanica, 18, 1, 1–248, 1991. </reference>
		<reference numeration="8" content_type="text"> % vor jede Referenz Genfa, Z., Dasgupta, P. K., and Dong, S.: Measurement of atmospheric ammonia, Environ. Sci. Technol., 23, 1467–1474, 1989. </reference>
		<reference numeration="9" content_type="text"> % vor jede Referenz Hanstein, S., Mattsson, M., Jaeger, H.-J., and Schjoerring, J. K.: Uptake and utilization of atmospheric ammonia in three native \textitPoaceae species: Leaf conductances, composition of apoplastic solution and interactions with nitrogen supply, New Phytol., 141, 71–83, 1999. </reference>
		<reference numeration="10" content_type="text"> % vor jede Referenz Harpole, W. S., Potts, D. L., and Suding, K. N.: Ecosystem responses to water and nitrogen amendment in a California grassland, Global Change Biol., 13, 2341–2348, 2007. </reference>
		<reference numeration="11" content_type="text"> % vor jede Referenz Herrmann, B., Jones, S. K., Fuhrer, J., Feller, U., and Neftel, A.: N budget and NH&lt;sub&gt;3&lt;/sub&gt; exchange of a grass/clover crop at two levels of N application, Plant Soil, 235, 243–252, 2001. </reference>
		<reference numeration="12" content_type="text"> % vor jede Referenz Hill, J. O., Simpson, R. J., Wood, J. T., Moore, A. D., and Chapman, D. F.: The phosphorus and nitrogen requirements of temperate pasture species and their influence on grassland botanical composition, Aust. J. Agric. Res., 56, 1027–1039, 2005. </reference>
		<reference numeration="13" content_type="text"> % vor jede Referenz Hill, M. O. and Carey, P. D.: Prediction of yield in the Rothamsted Park Grass Experiment by Ellenberg indicator values, J. Veg. Sci., 8, 579–586, 1997. </reference>
		<reference numeration="14" content_type="text"> % vor jede Referenz Hill, P. W., Raven, J. A., Loubet, B., Fowler, D., and Sutton, M. A.: Comparison of gas exchange and bioassay determinations of the ammonia compensation point in \textitLuzula sylvatica (Huds.) Gaud., Plant Physiol., 125, 476–487, 2001. </reference>
		<reference numeration="15" content_type="text"> % vor jede Referenz Hopkins, A.: Botanical composition of permanent grassland in England and wales in relation to soil, environment and management factors, Grass Forage Sci., 41, 237–246, 1986. </reference>
		<reference numeration="16" content_type="text"> % vor jede Referenz Husted, S. and Schjoerring, J. K.: Apoplastic pH and ammonium concentration in leaves of \textitBrassica napus L., Plant Physiol., 109, 1453–1460, 1995. </reference>
		<reference numeration="17" content_type="text"> % vor jede Referenz Husted, S. and Schjoerring, J. K.: Ammonia flux between oilseed rape plants and the atmosphere in respons to changes in leaf temperature, light intensity, and air humidity, Plant Physiol., 112, 67–74, 1996. </reference>
		<reference numeration="18" content_type="text"> % vor jede Referenz Husted, S., Schjoerring, J. K., Nielsen, K. H., Nemitz, E., and Sutton, M. A.: Stomatal compensation points for ammonia in oilseed rape plants under field conditions, Agr. Forest Meteorol., 105, 371–383, 2000a. </reference>
		<reference numeration="19" content_type="text"> % vor jede Referenz Husted, S., Hebbern, C. A., Mattsson, M., and Schjoerring, J. K.: Determination of ammonium, low molecular weight amines and amides in plant tissue, Physiol. Plant., 109, 167–179, 2000b. </reference>
		<reference numeration="20" content_type="text"> % vor jede Referenz Klimek, S., Kemmermann, A. R. G., Hofmann, M., and Isselstein, J.: Plant species richness and composition in managed grasslands: The relative importance of field management and environmental factors, Biol. Conserv., 134, 559–570, 2007. </reference>
		<reference numeration="21" content_type="text"> % vor jede Referenz Knapp, R. (Ed.): Sampling methods in taxon analysis in vegetation science, Handbook of Vegetation Science 1. Part 4, Junk, Hague, The Netherlands, 1984. </reference>
		<reference numeration="22" content_type="text"> % vor jede Referenz Lohaus, G., Pennewiss, K., Sattelmacher, B., Hussmann, M., and Muehling, K. H.: Is the infiltration-centrifugation technique appripriate for isolation of apoplastic fluid? A critical evaluation with different plant species, Physiol. Plant., 111, 457–465, 2001. </reference>
		<reference numeration="23" content_type="text"> % vor jede Referenz Loubet, B., Milford, C., Hill, P. W., Tang, Y. S., Cellier, P., and Sutton, M. S.: Seasonal variability of apoplastic NH$_4^+$ and pH in an intensively managed grassland, Plant Soil, 238, 97–110, 2002. </reference>
		<reference numeration="24" content_type="text"> % vor jede Referenz Mattsson, M., Häusler, R. E., Leegood, R. C., Lea, P., and Schjoerring, J. K.: Leaf-atmosphere ammonia exchange in barley mutants with reduced activities of glutamine synthetase, Plant Physiol., 114, 1307–1312, 1997. </reference>
		<reference numeration="25" content_type="text"> % vor jede Referenz Mattsson, M., Husted, S., and Schjoerring, J. K.: Influence of nitrogen nutrition and metabolism on ammonia volatilization in plants, Nutr. Cycl. Agroecosys., 51, 35–40, 1998. </reference>
		<reference numeration="26" content_type="text"> % vor jede Referenz Mattsson, M. and Schjoerring, J. K.: Dynamic and steady state responses of inorganic nitrogen pools and NH&lt;sub&gt;3&lt;/sub&gt; exchange in leaves of \textitLolium perenne and \textitBromus erectus to changes in root N supply, Plant Physiol., 128, 742–750, 2002. </reference>
		<reference numeration="27" content_type="text"> % vor jede Referenz Mattsson, M., Herrmann, B., David, M., Loubet, B., Riedo, M., Theobald, M. R., Sutton, M. A., Bruhn, D., Neftel, A., and Schjoerring, J. K.: Temporal variability in bioassays of ammonia exchange potential in relation to plant and soil nitrogen parameters in intensively managed grassland, Biogeosciences Discuss., 5, 2749–2772, 2008. </reference>
		<reference numeration="28" content_type="text"> % vor jede Referenz Milford, C., Theobald, M. R., Nemitz, E., Hargreaves, K. J., Horvath, L., Raso, J., Dämmgen, U., Neftel, A., Jones, S. K., Hensen, A., Loubet, B., Cellier, P., and Sutton, M. A.: Ammonia fluxes in relation to cutting and fertilization of an intensively managed grassland derived from an inter-comparison of gradient measurements, Biogeosciences Discuss., 5, 4699–4744, 2008. </reference>
		<reference numeration="29" content_type="text"> % vor jede Referenz Mountford, J. O., Lakhani, K. H., and Kirkham, F. W.: Experimental assessment of the effects of nitrogen addition under hay-cutting and aftermath grazing on the vegetation of medows on a Somerset peat moor, J. Appl. Ecol., 30, 321–332, 1993. </reference>
		<reference numeration="30" content_type="text"> % vor jede Referenz Nielsen, K. H. and Schjoerring, J. K.: Regulation of apoplastic NH$_4^+$ concentration in leaves of oilseed rape, Plant Physiol., 118, 1361–1368, 1998. </reference>
		<reference numeration="31" content_type="text"> % vor jede Referenz Oelmann, Y., Wilcke, W., Temperton, V. M., Buchmann, N., Roscher, C., Schumacher, J., Schulze, E. D., and Weisser, W. W.: Soil and plant nitrogen pools as related to plant diversity in an experimental grassland, Soil Sci. Soc. Am. J., 71, 720–729, 2007. </reference>
		<reference numeration="32" content_type="text"> % vor jede Referenz Pontes, L. S., Carrere, P., Andueza, D., Louault, F., and Soussana, J. F.: Seasonal productivity and nutritive value of temperate grasses found in semi-natural pastures in Europe: responses to cutting frequency and N supply, Grass Forage Sci., 62, 485–496, 2007. </reference>
		<reference numeration="33" content_type="text"> % vor jede Referenz Pywell, R. F., Bullock, J. M., Tallowin, J. B., Walker, K. J., Warman, E. A., and Masters, G.: Enhancing diversity of species-poor grasslands: an experimental assessment of multiple constraints, J. Appl. Ecol., 44, 81–94, 2007. </reference>
		<reference numeration="34" content_type="text"> % vor jede Referenz Schjoerring, J. K. and Mattsson, M.: Quantification of ammonia exchange between agricultural cropland and the atmosphere: Measurements over two complete growth cycles of oilseed rape, wheat, barley and pea, Plant Soil, 228, 105–115, 2001. </reference>
		<reference numeration="35" content_type="text"> % vor jede Referenz Silvertown, J: The dynamics of a grassland ecosystem: botanical equilibrium in the Park grass experiment, J. Appl. Ecol., 17, 491–504, 1980. </reference>
		<reference numeration="36" content_type="text"> % vor jede Referenz Silvertown, J., Poulton, P., Johnston, E., Edwards, G., Heard, M., and Biss, P. M.: The Park Grass Experiment 1856–2006: Its contribution to ecology, J. Ecol., 94, 801–814, 2006. </reference>
		<reference numeration="37" content_type="text"> % vor jede Referenz Stevens, C. J., Dise, N. B., Gowing, D. J. G., and Mountford, J. O.: Loss of forb diversity in relation to nitrogen deposition in the UK: regional trends and potential controls, Global Change Biol., 12, 1823–1833, 2006. </reference>
		<reference numeration="38" content_type="text"> % vor jede Referenz Stevens, C. J., Dise, N. B., Mountford, J. O., and Gowing, D. J.: Impact of nitrogen deposition on the species richness of grasslands, Science, 303, 1876–1879, 2004. </reference>
		<reference numeration="39" content_type="text"> % vor jede Referenz Sutton, M. A., Nemitz, E., Theobald, M. R., Milford, C., Dorsey, J. R., Gallagher, M. W., Hensen, A., Jongejan, P. A. C., Erisman, J. W., Mattsson, M. E., Schjoerring, J. K., Cellier, P., Loubet, B., Roche, R., Neftel, A., Hermann, B., Jones, S., Lehman, B. E., Horvath, L., Weidinger, T., Rajkai, K., Burkhardt, J., Löpmeier, F. J., and Daemmgen, U.: Dynamics of ammonia exchange with cut grassland: strategy and implementation of the GRAMINAE Integrated Experiment, Biogeosciences Discuss., 5, 3347–3407, 2008. </reference>
		<reference numeration="40" content_type="text"> % vor jede Referenz van Hove, L. W. A., Heeres, P., and Bossen, M. E.: The annual variation in stomatal ammonia compensation point of rye grass (\textitLolium perenne L.) leaves in an intensively managed grassland, Atmos. Environ., 36, 2965–2977, 2002. </reference>
		<reference numeration="41" content_type="text"> % vor jede Referenz Whitehead, D. C.: Grasses: Uptake of nitrogen and effects on morphology and physiology, in: Grassland nitrogen, CAB International, edited by: Whitehead, D. C., UK, 16–34, 1995. </reference>
		<reference numeration="42" content_type="text"> % vor jede Referenz Wichink Kruit, R. J., van Pul, W. A. J., Otjes, R. P., Hofschreuder, P., Jacobs, A. F. G., and Holtslag, A. A. M.: Ammonia fluxes and derived canopy compensation points over non-fertilized agricultural grassland in The Netherlands using the new gradient ammonia-high accuracy-monitor (GRAHAM), Atmos. Environ., 41, 1275–1287, 2007. </reference>
		<reference numeration="43" content_type="text"> % vor jede Referenz Wilman, D. and Wright, P. T.: The effect of interval between harvests and nitrogen application on the concentration of nitrate-nitrogen in the total herbage, green leaf and &quot;stem&quot; of grasses, J. Agric. Sci., Cambridge, 106, 467–475, 1986. </reference>
	</references>
</article>
