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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>7</volume_number>
		<issue_number>1</issue_number>
		<publication_year>2010</publication_year>
	</journal>
	<doi>10.5194/bg-7-27-2010</doi>
	<article_url>http://www.biogeosciences.net/7/27/2010/</article_url>
	<abstract_html>http://www.biogeosciences.net/7/27/2010/bg-7-27-2010.html</abstract_html>
	<fulltext_pdf>http://www.biogeosciences.net/7/27/2010/bg-7-27-2010.pdf</fulltext_pdf>
	<start_page>27</start_page>
	<end_page>41</end_page>
	<publication_date>2010-01-05</publication_date>
	<article_title content_type="html">Measuring and modeling continuous quality distributions of soil organic matter</article_title>
	<authors>
		<author numeration="1" affiliations="1">
			<name>S. Bruun</name>
			<email>sab@life.ku.dk</email>
		</author>
		<author numeration="2" affiliations="2">
			<name>G. I. Ågren</name>
		</author>
		<author numeration="3" affiliations="3">
			<name>B. T. Christensen</name>
		</author>
		<author numeration="4" affiliations="1">
			<name>L. S. Jensen</name>
		</author>
	</authors>
	<affiliations>
		<affiliation numeration="1" content_type="html">Plant and Soil Science Laboratory, Department of Agriculture and Ecology, Faculty of Life Sciences, University of Copenhagen, Thorvaldsensvej 40, 1871 Frederiksberg C, Denmark</affiliation>
		<affiliation numeration="2" content_type="html">Department of Ecology, Swedish University of Agricultural Sciences, P.O. Box 7044, 750 07 Uppsala, Sweden</affiliation>
		<affiliation numeration="3" content_type="html">Department of Agroecology and Environment, Faculty of Agricultural Sciences, Aarhus University, 8830 Tjele, Denmark</affiliation>
	</affiliations>
	<abstract content_type="html">An understanding of the dynamics of soil organic matter (SOM) is important
for our ability to develop management practices that preserve soil quality
and sequester carbon. Most SOM decomposition models represent the
heterogeneity of organic matter by a few discrete compartments with
different turnover rates, while other models employ a continuous quality
distribution. To make the multi-compartment models more mechanistic in
nature, it has been argued that the compartments should be related to soil
fractions actually occurring and having a functional role in the soil. In
this paper, we make the case that fractionation methods that can measure
continuous quality distributions should be developed, and that the temporal
development of these distributions should be incorporated into SOM models.
The measured continuous SOM quality distributions should hold valuable
information not only for model development, but also for direct
interpretation. Measuring continuous distributions requires that the
measurements along the quality variable are so frequent that the
distribution approaches the underlying continuum. Continuous distributions
lead to possible simplifications of the model formulations, which
considerably reduce the number of parameters needed to describe SOM
turnover. A general framework for SOM models representing SOM across
measurable quality distributions is presented and simplifications for
specific situations are discussed. Finally, methods that have been used or
have the potential to be used to measure continuous quality SOM
distributions are reviewed. Generally, existing fractionation methods will
have to be modified to allow measurement of distributions or new
fractionation techniques will have to be developed. Developing the
distributional models in concert with the fractionation methods to measure
the distributions will be a major task. We hope the current paper will help
generate the interest needed to accommodate this.</abstract>
	<references>
		<reference numeration="1" content_type="text"> % vor jede Referenz Ågren, G. and Bosatta, E.: Theoretical ecosystem ecology: understanding element cycles, Cambridge University Press, Cambridge, 1996a. </reference>
		<reference numeration="2" content_type="text"> Ågren, G. I. and Bosatta, E.: Quality: a bridge between theory and experiment in soil organic matter studies, Oikos, 76, 522–528, 1996b. </reference>
		<reference numeration="3" content_type="text"> Ågren, G. I., Bosatta, E., and Balesdent, J.: Isotope discrimination during decomposition of organic matter: a theoretical analysis, Soil Sci. Soc. Am. J., 60, 1121–1126, 1996a. </reference>
		<reference numeration="4" content_type="text"> Ågren, G. I., Kirschbaum, M. U. F., Johnson, D. W., and Bosatta, E.: Ecosystem physiology – Soil organic matter, in: Global change: Effects on coniferous forests and grasslands, edited by: Breymeyer, A. I., Hall, D. O., Melillo, J. M., and Ågren, G. I., John Wiley &amp; Sons Ltd, Chichester, 1996b. </reference>
		<reference numeration="5" content_type="text"> Arah, A. R. M. and Gaunt, J. L.: Questionable assumptions in the current soil organic matter transformation models, in: Sustainable management of soil organic matter, edited by: Rees, R. M., Ball, B. C., Campbell, C. D., and Watson, C. A., CABI publishing, Oxon, 83–89, 2001. </reference>
		<reference numeration="6" content_type="text"> Arshad, M. A., Starnaud, R. J., and Huang, P. M.: Characterization of electrophoretic separates of soil clays, Soil Sci., 112, 46–52, 1971. </reference>
		<reference numeration="7" content_type="text"> Baisden, W. T. and Amundson, R.: An analytical approach to ecosystem biogeochemistry modeling, Ecol. Appl., 13, 649–663, 2003. </reference>
		<reference numeration="8" content_type="text"> Baisden, W. T., Amundson, R., Cook, A. C., and Brenner, D. L.: Turnover and storage of C and N in five density fractions from California annual grassland surface soils, Global Biogeochem. Cy., 16, 1117, doi:10.1029/2001GB001822, 2002. </reference>
		<reference numeration="9" content_type="text"> Balesdent, J.: The significance of organic separates to carbon dynamics and its modelling in some cultivated soils, Eur. J. Soil Sci., 47, 485–493, 1996. </reference>
		<reference numeration="10" content_type="text"> Balesdent, J. and Mariotti, A.: Measurement of soil organic matter turnover using $^13$C natural abundance, in: Mass spectrometry of soils, edited by: Boutton, T. W. and Yamasaki, S. I., Marcel Dekker, New York, 83–111, 1996. </reference>
		<reference numeration="11" content_type="text"> Bernaux, M., Cerri, C. C., Neill, C., and de Moraes, J. F. L.: The use of stable carbon isotopes for estimation soil organic matter turnover rates, Geoderma, 82, 43–58, 1998. </reference>
		<reference numeration="12" content_type="text"> Bird, M., Santr\`ucková, H., Lloyd, J., and Lawson, E.: The isotopic composition of soil organic carbon on a north-south transect in western Canada, Eur. J. Soil Sci., 53, 393–403, 2002. </reference>
		<reference numeration="13" content_type="text"> Bosatta, E. and Ågren, G. I.: Theoretical analysis of decomposition of heterogeneous substrates, Soil Biol. Biochem., 17, 601–610, 1985. </reference>
		<reference numeration="14" content_type="text"> Bosatta, E. and Ågren, G. I.: Dynamics of carbon and nitrogen in the organic matter of the soil: a generic theory, Am. Nat., 138, 227–245, 1991. </reference>
		<reference numeration="15" content_type="text"> Bosatta, E. and Ågren, G. I.: Exact solutions to the continuous-quality equation for soil organic matter turnover, J. Theor. Biol., 224, 97–105, 2003. </reference>
		<reference numeration="16" content_type="text"> Boudreau, B. P.: A kinetic model for microbic organic matter decomposition in marine sediments, FEMS Microbiol. Ecol., 102, 1–14, 1992. </reference>
		<reference numeration="17" content_type="text"> Bruun, S., Jensen, E. S., and Jensen, L. S.: Microbial mineralization and assimilation of black carbon: Dependency on degree of thermal alteration, Org. Geochem., 39, 839–845, 2008a. </reference>
		<reference numeration="18" content_type="text"> Bruun, S. and Luxhøi, J.: Letter to the editor on &quot;Can incubations be used to measure meaningful pools of soil organic matter?&quot;, Soil Sci. Soc. Am. J., 70, 2164, 2006. </reference>
		<reference numeration="19" content_type="text"> Bruun, S., Six, J., and Jensen, L. S.: Estimating vital statistics and age distributions of measurable soil organic carbon fractions based on their pathway of formation and radiocarbon content, J. Theor. Biol., 230, 241–250, 2004. </reference>
		<reference numeration="20" content_type="text"> Bruun, S., Six, J., Paustian, K., and Jensen, L. S.: Estimating turnover of measurable soil organic carbon fractions based on radiocarbon measurements, Radiocarbon, 47, 99–113, 2005a. </reference>
		<reference numeration="21" content_type="text"> Bruun, S., Stenberg, B., Breland, T. A., Gudmundsson, J., Henriksen, T., Jensen, L. S., Korsaeth, A., Luxhøi, J., Palmason, F., Pedersen, A., and Salo, T.: Empirical predictions of plant material C and N mineralization patterns from near infrared spectroscopy, stepwise chemical digestion and C/N ratios, Soil Biol. Biochem., 37, 2283–2296, 2005b. </reference>
		<reference numeration="22" content_type="text"> Bruun, S., Thomsen, I. K., Christensen, B. T., and Jensen, L. S.: In search of stable soil organic carbon fractions: a comparison of methods applied to soils labelled with $^14$C for 40 days or 40 years, Eur. J. Soil Sci., 59, 247–256, 2008b. </reference>
		<reference numeration="23" content_type="text"> Buffle, J. and Leppard, G. G.: Characterization of aquatic colloids and macromolecules .2. Key role of physical structures on analytical results, Environ. Sci. Technol., 29, 2176–2184, 1995. </reference>
		<reference numeration="24" content_type="text"> Buyanovsky, G. A., Aslam, M., and Wagner, G. H.: Carbon turnover in soil physical fractions, Soil Sci. Soc. Am. J., 58, 1167–1173, 1994. </reference>
		<reference numeration="25" content_type="text"> Carpenter, S. R.: Decay of heterogenous detritus: a general model, J. Theor. Biol., 89, 539–547, 1981. </reference>
		<reference numeration="26" content_type="text"> Christensen, B. T.: Physical fractionation of soil and organic matter in primary particle size and density separates, Adv. Soil Sci., 20, 1–90, 1992. </reference>
		<reference numeration="27" content_type="text"> Christensen, B. T.: Matching measurable soil organic matter fractions with conceptual pools in simulation models: Revision of model structure, in: Evaluation of soil organic matter models, using existing long term datasets, edited by: Powlson, D. S., Smith, P., and Smith, J. U., Springer, Berlin, 143–159, 1996. </reference>
		<reference numeration="28" content_type="text">Christensen, B. T.: Physical fractionation of soil and structural and functional complexity in organic matter turnover, Eur. J. Soil Sci., 52, 345–353, 2001. </reference>
		<reference numeration="29" content_type="text"> Coleman, K. and Jenkinson, D. S.: Roth C-26.3 – A model for the turnover of carbon in soil, in: Evaluation of soil organic matter models, using existing long-term datasets, edited by: Powlson, D. S., Smith, P., and Smith, J. U., Springer, Berlin, 237–246, 1996. </reference>
		<reference numeration="30" content_type="text"> Dalal, R. C. and Mayer, R. J.: Long-term trends in fertility of soils under continuous cultivation and cereal cropping in Southern Queensland .IV. Loss of organic carbon from different density fractions, Aust. J. Soil Res., 24, 301–309, 1986. </reference>
		<reference numeration="31" content_type="text"> Dauwe, B. and Middelburg, J. J.: Amino acids and hexosamines as indicators of organic matter degradation state in North Sea sediments, Limnol. Oceanogr., 43, 782–798, 1998. </reference>
		<reference numeration="32" content_type="text"> Dauwe, B., Middelburg, J. J., Herman, P. M. J., and Heip, C. H. R.: Linking diagenetic alteration of amino acids and bulk organic matter reactivity, Limnol. Oceanogr., 44, 1809–1814, 1999. </reference>
		<reference numeration="33" content_type="text"> David, C.: Thermal degradation of polymers, in: Degradation of polymers, edited by: Bamford, C. H. and Tipper, C. F. H., Elsevier, Amsterdam, 1–173, 1975. </reference>
		<reference numeration="34" content_type="text"> DosRamos, J. G. and Silebi, C. A.: The determination of particle size distribution of submicrometer particles by capillary hydrodynamic fractionation (CHDF), J. Colloid. Interf. Sci., 135, 165–177, 1990. </reference>
		<reference numeration="35" content_type="text"> Elliott, E. T., Paustian, K., and Frey, S. D.: Modeling the measurable or measuring the modelable: A hierarchical approach to isolating meaningful soil organic matter fractionations, in: Evaluation of soil organic matter models, using existing long term datasets, edited by: Powlson, D. S., Smith, P., and Smith, J. U., Springer, Berlin, 161–179, 1996. </reference>
		<reference numeration="36" content_type="text"> England, L. S., Lee, H., and Trevors, J. T.: Bacterial survival in soil - Effect of clays and protozoa, Soil Biol. Biochem., 25, 525–531, 1993. </reference>
		<reference numeration="37" content_type="text"> Eusterhues, K., Rumpel, C., and Kögel-Knabner, I.: Stabilization of soil organic matter isolated via oxidative degradation, Org. Geochem., 36, 1567–1575, 2005. </reference>
		<reference numeration="38" content_type="text"> Fontaine, S., Mariotti, A., and Abbadie, L.: The priming effect of organic matter: a question of microbial competition?, Soil Biol. Biochem., 35, 837–843, 2003. </reference>
		<reference numeration="39" content_type="text"> Fraunhofer, W. and Winter, G.: The use of asymmetrical flow field-flow fractionation in pharmaceutics and biopharmaceutics, Eur. J. Pharm. Biopharm., 58, 369–383, 2004. </reference>
		<reference numeration="40" content_type="text"> Gelinas, Y., Prentice, K. M., Baldock, J. A., and Hedges, J. I.: An improved thermal oxidation method for the quantification of soot/graphitic black carbon in sediments and soils, Environ. Sci. Technol., 35, 3519–3525, 2001. </reference>
		<reference numeration="41" content_type="text"> Giddings, J. C.: A system based on split-flow lateral transport thin (Splitt) separation cells for rapid and continuous particle fractionation, Sep. Sci. Technol., 20, 749–768, 1985. </reference>
		<reference numeration="42" content_type="text"> Gillon, D., Joffre, R., and Ibrahima, A.: Can litter decomposability be predicted by near infrared reflectance spectroscopy?, Ecology, 80, 175–186, 1999. </reference>
		<reference numeration="43" content_type="text"> Goldberg, E. D.: Black carbon in the environment, John Wiley and Sons, New York, 1985. </reference>
		<reference numeration="44" content_type="text"> Guggenberger, G. and Kaiser, K.: Dissolved organic matter in soil: challenging the paradigm of sorptive preservation, Geoderma, 113, 293–310, 2003. </reference>
		<reference numeration="45" content_type="text"> Gustafsson, O., Bucheli, T. D., Kukulska, Z., Andersson, M., Largeau, C., Rouzaud, J. N., Reddy, C. M., and Eglinton, T. I.: Evaluation of a protocol for the quantification of black carbon in sediments, Global Biogeochem. Cy., 15, 881–890, 2001. </reference>
		<reference numeration="46" content_type="text"> Gustafsson, O., Haghseta, F., Chan, C., MacFarlane, J., and Gschwend, P. M.: Quantification of the dilute sedimentary soot phase: Implications for PAH speciation and bioavailability, Environ. Sci. Technol., 31, 203–209, 1997. </reference>
		<reference numeration="47" content_type="text"> Hassink, J.: Decomposition rate constants of size and density fractions of soil organic matter, Soil Sci. Soc. Am. J., 59, 1631–1635, 1995. </reference>
		<reference numeration="48" content_type="text"> Hassink, J. and Dalenberg, J. W.: Decomposition and transfer of plant residue $^14$C between size and density fractions in soil, Plant Soil, 179, 159–169, 1996. </reference>
		<reference numeration="49" content_type="text"> Helfrich, M., Flessa, H., Mikutta, R., Dreves, A., and Ludwig, B.: Comparison of chemical fractionation methods for isolating stable soil organic carbon pools, Eur. J. Soil Sci., 58, 1316–1329, 2007. </reference>
		<reference numeration="50" content_type="text"> Hunter, R. J.: Zeta potential in colloid science, Academic Press, London, 1981. </reference>
		<reference numeration="51" content_type="text"> Hyvönen, R., Ågren, G. I., and Dalias, P.: Analysing temperature response of decomposition of organic matter, Glob. Change Biol., 11, 770–778, 2005. </reference>
		<reference numeration="52" content_type="text"> Joffre, R., Ågren, G. I., Gillon, D., and Bosatta, E.: Organic matter quality in ecological studies: theory meets experiment, Oikos, 93, 451–458, 2001. </reference>
		<reference numeration="53" content_type="text"> Joffre, R., Gillon, D., Dardenne, P., Agneessens, R., and Biston, R.: The use of near-infrared reflectance spectroscopy in litter decomposition studies, Ann. Sci. For., 49, 481–488, 1992. </reference>
		<reference numeration="54" content_type="text"> Kiem, R., Knicker, H., and Kögel-Knabner, I.: Refractory organic carbon in particle-size fractions of arable soils I: distribution of refractory carbon between the size fractions, Org. Geochem., 33, 1683–1697, 2002. </reference>
		<reference numeration="55" content_type="text"> Kiem, R. and Kögel-Knabner, I.: Contribution of lignin and polysaccharides to the refractory carbon pool in C-depleted arable soils, Soil Biol. Biochem., 35, 101–118, 2003. </reference>
		<reference numeration="56" content_type="text"> Kleber, M., Mikutta, R., Torn, M. S., and Jahn, R.: Poorly crystalline mineral phases protect organic matter in acid subsoil horizons, Eur. J. Soil Sci., 56, 717–725, 2005. </reference>
		<reference numeration="57" content_type="text"> Kleber, M., Sollins, P., and Sutton, R.: A conceptual model of organo-mineral interactions in soils: self-assembly of organic molecular fragments into zonal structures on mineral surfaces, Biogeochemistry, 85, 9–24, 2007. </reference>
		<reference numeration="58" content_type="text"> Kögel-Knabner, I.: The macromolecular organic composition of plant and microbial residues as inputs to soil organic matter, Soil Biol. Biochem., 34, 139–162, 2002. </reference>
		<reference numeration="59" content_type="text"> Kretzschmar, R., Borkovec, M., Grolimund, D., and Elimelech, M.: Mobile subsurface colloids and their role in contaminant transport, Adv. Agron., 66, 121–193, 1999. </reference>
		<reference numeration="60" content_type="text"> Kristensen, E.: Characterization of biogenic organic-matter by stepwise thermogravimetry (Stg), Biogeochem., 9, 135–159, 1990. </reference>
		<reference numeration="61" content_type="text"> Kuzyakov, Y., Friedel, J. K., and Stahr, K.: Review of mechanisms and quantification of priming effects, Soil Biol. Biochem., 32, 1485–1498, 2000. </reference>
		<reference numeration="62" content_type="text"> Laevitt, S. W., Follet, R. F., and Paul, E. A.: Estimation of slow- and fast cycling soil organic carbon pools from 6N HCL hydrolysis, Radiocarbon, 38, 231–239, 1996. </reference>
		<reference numeration="63" content_type="text"> Lindahl, V. and Bakken, L. R.: Evaluation of methods for extraction of bacteria from soil, FEMS Microbiol. Ecol., 16, 135–142, 1995. </reference>
		<reference numeration="64" content_type="text">Lopez-Capel, E., Bol, R., and Manning, D. A. C.: Application of simultaneous thermal analysis mass spectrometry and stable carbon isotope analysis in a carbon sequestration study, Rapid Commun. Mass Spectrom., 19, 3192–3198, 2005a. </reference>
		<reference numeration="65" content_type="text"> Lopez-Capel, E., Sohi, S. P., Gaunt, J. L., and Manning, D. A. C.: Use of thermogravimetry-differential scanning calorimetry to characterize modelable soil organic matter fractions, Soil Sci. Soc. Am. J., 69, 136–140, 2005b. </reference>
		<reference numeration="66" content_type="text"> Magid, J., Gorissen, A., and Giller, K.: In search of the elusive &quot;active&quot; fraction of soil organic matter: three size-density fractionation methods for tracing the fate of homogeneously $^14$C-labeled plant materials, Soil Biol. Biochem., 28, 89–99, 1996. </reference>
		<reference numeration="67" content_type="text"> Manzoni, S. and Porporato, A.: Theoretical analysis of nonlinearities and feedbacks in soil carbon and nitrogen cycles, Soil Biol. Biochem., 39, 1542–1556, 2007. </reference>
		<reference numeration="68" content_type="text"> Martel, Y. A. and Paul, E. A.: The use of radiocarbon dating of organic matter in the study of soil genesis, Soil Sci. Soc. Am. Proc., 38, 501–506, 1974. </reference>
		<reference numeration="69" content_type="text"> Martens, H. and Naes, T.: Multivariate calibration, John Wiley and Sons, Chichester, 1989. </reference>
		<reference numeration="70" content_type="text">Masiello, C. A.: New directions in black carbon organic geochemistry, Mar. Chem., 92, 201–213, 2004. </reference>
		<reference numeration="71" content_type="text"> McGill, W. B.: Review and classification of ten soil organic matter (SOM) models, in: Evaluation of soil organic matter models, using existing long term datasets, edited by: Powlson, D. S., Smith, P., and Smith, J. U., Springer, Berlin, 111–132, 1996. </reference>
		<reference numeration="72" content_type="text"> Mikutta, R., Kleber, M., Kaiser, K., and Jahn, R.: Review: Organic matter removal from soils using hydrogen peroxide, sodium hypochlorite, and disodium peroxodisulfate, Soil Sci. Soc. Am. J., 69, 120–135, 2005. </reference>
		<reference numeration="73" content_type="text"> Neill, C. and Gignoux, J.: Soil organic matter decomposition driven by microbial growth: A simple model for a complex network of interactions, Soil Biol. Biochem., 38, 803–811, 2006. </reference>
		<reference numeration="74" content_type="text"> Oades, J. M. and Waters, A. G.: Aggregate hierarchy in soils, Aust. J. Soil Res., 29, 815–828, 1991. </reference>
		<reference numeration="75" content_type="text"> Parton, W. J., Schimel, D. S., Cole, C. V., and Ojima, D. S.: Analysis of factors controlling soil organic matter levels in Great Plains grasslands, Soil Sci. Soc. Am. J., 51, 1173–1179, 1987. </reference>
		<reference numeration="76" content_type="text"> Paul, E. A., Follett, R. F., Leavitt, S. W., Halvorson, A., Peterson, G. A., and Lyon, D. J.: Radiocarbon dating for determination of soil organic matter pool sizes and dynamics, Soil Sci. Soc. Am. J., 61, 1058–1067, 1997. </reference>
		<reference numeration="77" content_type="text"> Paul, E. A., Morris, S. J., Conant, R. T., and Plante, A. F.: Does the acid hydrolysis-incubation method measure meaningful soil organic carbon pools?, Soil Sci. Soc. Am. J., 70, 1023–1035, 2006. </reference>
		<reference numeration="78" content_type="text"> Plante, A. F., Chenu, C., Balabane, M., Mariotti, A., and Righi, D.: Peroxide oxidation of clay-associated organic matter in a cultivation chronosequence, Eur. J. Soil Sci., 55, 471–478, 2004. </reference>
		<reference numeration="79" content_type="text"> Plante, A. F., Conant, R. T., Paul, E. A., Paustian, K., and Six, J.: Acid hydrolysis of easily dispersed and microaggregate-derived silt- and clay-sized fractions to isolate resistant soil organic matter, Eur. J. Soil Sci., 57, 456–467, 2006. </reference>
		<reference numeration="80" content_type="text"> Plante, A. F., Fernández, J. M., and Leifeld, J.: Application of thermal analysis techniques in soil science, Geoderma, 153, 1–10, 2009. </reference>
		<reference numeration="81" content_type="text"> Poage, M. A. and Feng, X. H.: A theoretical analysis of steady state $\delta ^13$C profiles of soil organic matter, Global Biogeochem. Cy., 18, GB2016, doi:10.1029/2003GB002195,, 2004. </reference>
		<reference numeration="82" content_type="text"> Ranville, J. F., Chittleborough, D. J., and Beckett, R.: Particle-size and element distributions of soil colloids: Implications for colloid transport, Soil Sci. Soc. Am. J., 69, 1173–1184, 2005. </reference>
		<reference numeration="83" content_type="text"> Reichstein, M., Ågren, G., and Fontaine, S.: Is there a theoretical limit to soil carbon storage in old-growth forests?, in: Old-Growth forests, edited by: Wirth, G., Gleixner, G., and Heimann, M., Springer, Berlin, 267–281, 2009. </reference>
		<reference numeration="84" content_type="text"> Saggar, S., Parshotam, A., Sparling, G. P., Feltham, C. W., and Hart, P. B. S.: $^14$C-labelled ryegrass turnover and residence times in soils varying in clay content and mineralogy, Soil Biol. Biochem., 28, 1677–1686, 1996. </reference>
		<reference numeration="85" content_type="text"> Saito, T., Koopal, L. K., van Riemsdijk, W. H., Nagasaki, S., and Tanaka, S.: Adsorption of humic acid on goethite: Isotherms, charge adjustments, and potential profiles, Langmuir, 20, 689–700, 2004. </reference>
		<reference numeration="86" content_type="text"> Schimel, J. P. and Weintraub, M. N.: The implications of exoenzyme activity on microbial carbon and nitrogen limitation in soil: a theoretical model, Soil Biol. Biochem., 35, 549–563, 2003. </reference>
		<reference numeration="87" content_type="text"> Schmidt, M. W. I., Skjemstad, J. O., Czimczik, C. I., Glaser, B., Prentice, K. M., Gelinas, Y., and Kuhlbusch, T. A. J.: Comparative analysis of black carbon in soils, Global Biogeochem. Cy., 15, 163–167, 2001. </reference>
		<reference numeration="88" content_type="text"> Schulten, H. R.: Analytical Pyrolysis of Humic Substances and Soils – Geochemical, Agricultural and Ecological Consequences, J. Anal. Appl. Pyrol., 25, 97–122, 1993. </reference>
		<reference numeration="89" content_type="text"> Schulten, H. R. and Leinweber, P.: Thermal stability and composition of mineral-bound organic matter in density fractions of soil, Eur. J. Soil Sci., 50, 237–248, 1999. </reference>
		<reference numeration="90" content_type="text"> Schulten, H. R. and Leinweber, P.: New insights into organic-mineral particles: composition, properties and models of molecular structure, Biol. Fert. Soils, 30, 399–432, 2000. </reference>
		<reference numeration="91" content_type="text"> Sexstone, A. J., Revsbech, N. P., Parkin, T. B., and Tiedje, J. M.: Direct measurement of oxygen profiles and denitrification rates in soil aggregates, Soil Sci. Soc. Am. J., 49, 645–651, 1985. </reference>
		<reference numeration="92" content_type="text"> Shang, C. and Tiessen, H.: Organic matter lability in a tropical oxisol: Evidence from shifting cultivation, chemical oxidation, particle size, density, and magnetic fractionations, Soil Sci., 162, 795–807, 1997. </reference>
		<reference numeration="93" content_type="text"> Shang, C. and Tiessen, H.: Carbon turnover and carbon-13 natural abundance in organo-mineral fractions of a tropical dry forest soil under cultivation, Soil Sci. Soc. Am. J., 64, 2149–2155, 2000. </reference>
		<reference numeration="94" content_type="text"> Shaymukhametov, M. S., Titova, N. A., Travnikova, L. S., and Labenets, Y. M.: Use of physical fractionation methods to characterize soil organic matter, Soviet Soil Science, 16, 117–128, 1984. </reference>
		<reference numeration="95" content_type="text"> Shenk, J. S., Workman, J. J., and Westerhaus, M. O.: Application of NIR spectroscopy to agricultural products, in: Handbook of Near-Infrared Analysis, edited by: Burns, D. A. and Ciurczak, E. W., Marcel Dekker, Inc., 419–473, 2001. </reference>
		<reference numeration="96" content_type="text"> Siewert, C.: Rapid screening of soil properties using thermogravimetry, Soil Sci. Soc. Am. J., 68, 1656–1661, 2004. </reference>
		<reference numeration="97" content_type="text"> Simpson, M. J. and Hatcher, P. G.: Determination of black carbon in natural organic matter by chemical oxidation and solid-state 13C nuclear magnetic resonance spectroscopy, Org. Geochem., 35, 923–935, 2004. </reference>
		<reference numeration="98" content_type="text"> Six, J., Conant, R. T., Paul, E. A., and Paustian, K.: Stabilization mechanisms of soil organic matter: Implications for C-saturation of soils, Plant Soil, 241, 155–176, 2002. </reference>
		<reference numeration="99" content_type="text"> Six, J., Elliott, E. T., Paustian, K., and Doran, J. W.: Aggregation and soil organic matter accumulation in cultivated and native grassland soils, Soil Sci. Soc. Am. J., 62, 1367–1377, 1998. </reference>
		<reference numeration="100" content_type="text"> Six, J., Merckx, R., Kimpe, K., Paustian, K., and Elliott, E. T.: A re-evaluation of the enriched labile soil organic matter fraction, Eur. J. Soil Sci., 51, 283–293, 2000. </reference>
		<reference numeration="101" content_type="text"> Skjemstad, J. O., Clarke, P., Colchin, A., and Oades, J. M.: Characterization of soil organic matter by solid-state $^13$C NMR spectroscopy, in: Driven by nature. Plant litter quality and decomposition, edited by: Cadisch, G. and Giller, K. E., CAB International, Wallingford, 3–30, 1997. </reference>
		<reference numeration="102" content_type="text"> Skjemstad, J. O., Janik, L. J., Head, M. J., and McClure, S. G.: High energy ultraviolet photooxidation: a novel technique for studying physically protected organic matter in clay-sized and silt-sized aggregates, J. Soil Sci., 44, 485–499, 1993. </reference>
		<reference numeration="103" content_type="text"> Skjemstad, J. O., Spouncer, L. A., Cowie, B., and Swift, R. S.: Calibration of the Rothhamsted organic turnover model (Roth ver 26.3), using measureable soil organic carbon pools, Aust. J. Soil Res., 42, 79–88, 2004. </reference>
		<reference numeration="104" content_type="text"> Skjemstad, J. O., Taylor, J. A., and Smernik, R. J.: Estimation of charcoal (char) in soils, Comm. Soil Sci. Plant Anal., 30, 2283–2298, 1999. </reference>
		<reference numeration="105" content_type="text"> Smith, J. U., Smith, P., Monaghan, R., and MacDonald, A. J.: When is a measured soil organic matter fraction equivalent to a model pool?, Eur. J. Soil Sci., 53, 405–416, 2002. </reference>
		<reference numeration="106" content_type="text"> Sohi, S. P., Mahieu, N., Arah, J. R. M., Powlson, D. S., Madari, B., and Gaunt, J. L.: A procedure for isolating soil organic matter fractions suitable for modeling, Soil Sci. Soc. Am. J., 65, 1121–1128, 2001. </reference>
		<reference numeration="107" content_type="text"> Sollins, P., Homann, P., and Caldwell, B. A.: Stabilization and destabilization of soil organic matter: Mechanisms and controls, Geoderma, 74, 65–105, 1996. </reference>
		<reference numeration="108" content_type="text"> Sollins, P., Kramer, M. G., Swanston, C., Lajtha, K., Filley, T., Aufdenkampe, A. K., Wagai, R., and Bowden, R. D.: Sequential density fractionation across soils of contrasting mineralogy: evidence for both microbial- and mineral-controlled soil organic matter stabilization, Biogeochem., 96, 209–231, 2009. </reference>
		<reference numeration="109" content_type="text"> Sollins, P., Swanston, C., Kleber, M., Filley, T., Kramer, M., Crow, S., Caldwell, B. A., Lajtha, K., and Bowden, R.: Organic C and N stabilization in a forest soil: Evidence from sequential density fractionation, Soil Biol. Biochem., 38, 3313–3324, 2006. </reference>
		<reference numeration="110" content_type="text"> Stevens, K. A. and Jaykus, L. A.: Bacterial separation and concentration from complex sample matrices: A review, Crit. Rev. Microbiol., 30, 7–24, 2004. </reference>
		<reference numeration="111" content_type="text"> Thomsen, I. K., Bruun, S., Jensen, L. S., and Christensen, B. T.: Assessing soil carbon lability by near infrared spectroscopy and NaOCl oxidation, Soil Biol. Biochem., 41, 2170–2177, 2009. </reference>
		<reference numeration="112" content_type="text"> Tipping, E. and Cooke, D.: The effects of adsorbed humic substances on the surface charge of goethite (alpha-FeOOH) in fresh waters, Geochim. Cosmochim. Ac., 46, 75–80, 1982. </reference>
		<reference numeration="113" content_type="text"> Tisdall, J. M. and Oades, J. M.: Organic matter and water-stable aggregates in soils, J. Soil Sci., 33, 141–163, 1982. </reference>
		<reference numeration="114" content_type="text"> Torn, M. S., Trumbore, S. E., Chadwick, O. A., Vitousek, P. M., and Hendricks, D. M.: Mineral control of soil organic carbon storage and turnover, Nature, 389, 170–173, 1997. </reference>
		<reference numeration="115" content_type="text"> Trumbore, S. E., Chadwick, O. A., and Amundson, R.: Rapid exchange between soil carbon and atmospheric carbon dioxide driven by temperature change, Science, 272, 393–396, 1996. </reference>
		<reference numeration="116" content_type="text"> Trumbore, S. E. and Zheng, S. H.: Comparison of fractionation methods for soil organic matter $^14$C analysis, Radiocarbon, 38, 219–229, 1996. </reference>
		<reference numeration="117" content_type="text"> van Veen, J. A. and Kuikman, P. J.: Soil structural aspects of decomposition of organic matter by micro-organisms, Biogeochem., 11, 213–233, 1990. </reference>
		<reference numeration="118" content_type="text"> Vandewiele, S., Cowie, G., Soetaert, K., and Middelburg, J. J.: Amino acid biogeochemistry and organic matter degradation state across the Pakistan margin oxygen minimum zone, Deep-Sea Res. II, 56, 376–392, 2009. </reference>
		<reference numeration="119" content_type="text"> Veeman, W. S.: Nuclear magnetic resonance, a simple introduction to the principles and applications, Geoderma, 80, 225–242, 1997. </reference>
		<reference numeration="120" content_type="text"> von Lützow, M., Kögel-Knabner, I., Ekschmitt, K., Flessa, H., Guggenberger, G., Matzner, E., and Marschner, B.: SOM fractionation methods: Relevance to functional pools and to stabilization mechanisms, Soil Biol. Biochem., 39, 2183–2207, 2007. </reference>
		<reference numeration="121" content_type="text"> von Lützow, M., Kögel-Knabner, I., Ekschmitt, K., Matzner, E., Guggenberger, G., Marschner, B., and Flessa, H.: Stabilization of organic matter in temperate soils: mechanisms and their relevance under different soil conditions – a review, Eur. J. Soil Sci., 57, 426–445, 2006. </reference>
		<reference numeration="122" content_type="text"> Vor, T., Dyckmans, J., Loftfield, N., Beese, F., and Flessa, H.: Aeration effects on CO&lt;sub&gt;2&lt;/sub&gt;, N&lt;sub&gt;2&lt;/sub&gt;O, and CH&lt;sub&gt;4&lt;/sub&gt; emission and leachate composition of a forest soil, J. Plant Nutr. Soil Sci., 166, 39–45, 2003. </reference>
		<reference numeration="123" content_type="text"> Zimmermann, M., Leifeld, J., Schmidt, M. W. I., Smith, P., and Fuhrer, J.: Measured soil organic matter fractions can be related to pools in the RothC model, Eur. J. Soil Sci., 58, 658–667, 2007. </reference>
	</references>
</article>

