52 DOI: 10.1002/jpln.200700043 J. Plant Nutr. Soil Sci.

2007, 170, 5260

Review Article
Contribution of dissolved organic matter to carbon storage in forest mineral
soils
Karsten Kalbitz1* and Klaus Kaiser2
1 Department of Soil Ecology, University of Bayreuth, 95440 Bayreuth, Germany
2 Soil Biology and Soil Ecology, Martin Luther University Halle-Wittenberg, Weidenplan 14,
06108 Halle (Saale), Germany

Abstract
Dissolved organic matter (DOM) is often considered the most labile portion of organic matter in
soil and to be negligible with respect to the accumulation of soil C. In this short review, we pre-
sent recent evidence that this view is invalid. The stability of DOM from forest floor horizons,
peats, and topsoils against microbial degradation increases with advanced decomposition of the
parent organic matter (OM). Aromatic compounds, deriving from lignin, likely are the most stable
components of DOM while plant-derived carbohydrates seem easily degradable. Carbohydrates
and N-rich compounds of microbial origin produced during the degradation of DOM can be rela-
tively stable. Such components contribute much to DOM in the mineral subsoil. Sorption of DOM
to soil minerals and (co-)precipitation with Al (and probably also with Fe), especially of the inher-
ently stable aromatic moieties, result in distinct stabilization. In laboratory incubation experi-
ments, the mean residence time of DOM from the Oa horizon of a Haplic Podzol increased from
<30 y in solution to >90 y after sorption to a subsoil. We combined DOM fluxes and mineraliza-
tion rate constants for DOM sorbed to minerals and a subsoil horizon, and (co-)precipitated with
Al to estimate the potential contribution of DOM to total C in the mineral soil of a Haplic Podzol in
Germany. The contribution of roots to DOM was not considered because of lack of data. The
DOM-derived soil C ranges from 20 to 55 Mg ha1 in the mineral soil, which represents
19%50% of the total soil C. The variation of the estimate reflects the variation in mineralization
rate constants obtained for sorbed and (co-)precipitated DOM. Nevertheless, the estimates indi-
cate that DOM contributes significantly to the accumulation of stable OM in soil. A more precise
estimation of DOM-derived C in soils requires mineralization rate constants for DOM sorbed to
all relevant minerals or (co-)precipitated with Fe. Additionally, we need information on the contri-
bution of sorption to distinct minerals as well as of (co-)precipitation with Al and Fe to DOM
retention.

Key words: biodegradation / chemical composition / (co-)precipitation / dissolved organic matter / DOM /
forest soils / sorption

Accepted May 31, 2007

1 Introduction
Growing concern about climate change evoked interest in the
role of dissolved organic matter (DOM) in the global C bal-
ance (Hedges, 2002). Fluxes of dissolved organic C (DOC)
into the marine environment via runoff from terrestrial settings
are estimated to 0.2 (Harrison et al., 2005) to 0.4 1015 g
(IPCC, 2001). These fluxes are small compared to the C
fluxes due to primary productivity and heterotrophic respira-
tion in terrestrial ecosystems. Total DOC in the oceans is
approx. 685 1015 g (Hansell and Carlson, 1998) and contri-
butes only to 1.8% of marine C storage. Therefore, DOC
fluxes are generally not considered to be of relevance for the
global C cycle (Neff and Asner, 2001). However, DOC fluxes
in terrestrial ecosystems are several folds larger than those
with rivers. In certain environmentsparticularly in peatlands
and at northern latitudessoluble C can substantially contri-
bute to the ecosystem C cycle (Neff and Asner, 2001). Large
internal DOC fluxes in soils relative to small outputs into
aquatic ecosystems may result from mineralization or stabili-
zation, thus, accumulation in the soil, as suggested by Kaiser
and Guggenberger (2000).

* Correspondence: Dr. K. Kalbitz;

e-mail: karsten.kalbitz@uni-bayreuth.de
Topical Issue Soils as a source and sink for CO Mechanisms
2
and regulation of organic matter stabilisation in soils (editors: I. Kgel-
Knabner and E. Matzner). Synthesis of the DFG Priority Program
SPP 1090 (German Research FoundationDeutsche Forschungs-
gemeinschaft).

2008 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim www.plant-soil.com

J. Plant Nutr. Soil Sci. 2008, 171, 5260
Litter layers and the upper, organic-rich mineral horizons are
the main sources of DOM in soils whereas the deeper mineral
horizons are the major sinks. In forest ecosystems, fluxes of
DOC from the forest floor into the mineral soil have been esti-
mated to 115500 kg C ha1 y1, representing up to 35% of
the annual litterfall C (Guggenberger and Zech, 1993; Currie
et al., 1996; Michalzik and Matzner, 1999; Solinger et al.,
2001; Kaiser et al., 2001; Michalzik et al., 2001). The fluxes
from forest floor and topsoil horizons down into deeper soil
horizons decrease because of degradation in soil solution,
sorption to Fe and Al oxides/hydroxides and clay minerals,
and (co-)precipitation by polyvalent cations (summarized by
Kalbitz et al., 2000). Therefore, concentrations of DOC in
deeper soil horizons as well as the export from mineral sub-
soil are usually small (Guggenberger and Zech, 1993; Michal-
zik and Matzner, 1999; Solinger et al., 2001). Retention in
mineral subsoils ranges from 40 to 370 kg DOC ha1 y1 (Cur-
rie et al., 1996; Guggenberger and Kaiser, 2003). Given that
most of the reduction in DOC is due to abiotic reactions and
not mineralization, DOC could be a major contributor to the
total C accumulation in soil, including O layers, which may
reach up to 1440 kg C ha1 y1 (Schulze et al., 2000). Based
on estimates of Michalzik et al. (2001) and Guggenberger
and Kaiser (2003), the retention of DOC in the mineral soil of
17 temperate forest sites studied in Europe and N America
averages to 191 kg DOC ha1 y1. The proportion to which
the decrease of DOC fluxes with soil depth is either due to
microbial consumption or abiotic storage via sorption or preci-
pitation, however, is mostly unknown. Besides knowledge
about DOM fluxes, we need robust estimates of DOM degra-
dation in soil solution and after sorption and (co-)precipitation
to rate the contribution of DOM to the formation of stable soil
organic matter (SOM).
In this short review, we summarize recent findings on the sta-
bilization of DOM in soils, defined as reduced respiratory C
release. We evaluate the stability of DOM against microbial
degradation in soil solution, after sorption to mineral subsoil
and minerals and after precipitation by polyvalent cations.
Furthermore, we estimate the portion of DOM-derived OM in
the mineral soil of a Podzol using DOM fluxes measured in
the field and degradation rates of DOM from that site.
2 Degradation of DOM in soil solution
Laboratory incubations showed 5%93% of the DOM in soil
solutions to be potentially microbially degradable (Jandl and
Sletten, 1999; Yano et al., 2000; Kalbitz et al., 2000; Sachse
et al., 2001; Kalbitz et al., 2003a; Don and Kalbitz, 2005;
Kiikkil et al., 2006). Marschner and Kalbitz (2003) consid-
ered intrinsic properties of DOM to control its biodegradation.
Dissolved organic matter with a large portion of C in the
XAD-8-adsorbable fraction (i.e., hydrophobic by definition),
rich in aromatic structures, and poor in carbohydrates is little
biodegradable (Qualls and Haines, 1992; Jandl and Sollins,
1997; Jandl and Sletten, 1999; Volk et al., 1997; Amon et al.,
2001; Kalbitz et al., 2003a; Kiikkil et al., 2006). Kalbitz et al.
(2003a) concluded that microbial degradation of DOM de-
rived from forest floors, peats, and A horizons decreased with
increasing degree of decomposition of the parent material.
That seems reasonable because of the preferential utilization
2008 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim
Stabilization and dissolved organic matter 53
of labile material at earlier states of decomposition
(Haider, 1992; Dai et al., 2001). In consequence, DOM
percolating into the mineral soil should be largely stable
because it derives predominately from the more decomposed
parts of the forest floor (Frberg et al., 2006). That also
suggests that microbial degradation cannot explain the
decrease in DOC concentrations in the mineral soil with
increasing depth.
Aromatic compounds, possibly deriving from lignin, which
survive biodegradation preferentially, seem to be the most
stable constituents of DOM from forest floors and topsoils
(Kalbitz et al., 2003b). Thus, the partial biodegradation leaves
behind compounds with favorable properties for sorption to
minerals (McKnight et al., 1992; Kaiser et al., 1997) and con-
sequently for further stabilization (Kalbitz et al., 2005; see
below).
In acid forest soils, Al and Fe can form stable complexes with
DOM. Schwesig et al. (2003a) found a decreased mineraliza-
tion after addition of Al to four different DOM solutions. How-
ever, DOM complexation by metals will not necessarily
reduce its biodegradability but instead may enhance micro-
bial activity by reducing the free metal-ion concentrations and
thus their toxicity (Marschner and Kalbitz, 2003).
The properties of DOM change with soil depth due to prefer-
ential sorption and (co-)precipitation of aromatic, lignin-de-
rived portions, resulting in increasing portions of carbohy-
drate- and N-rich compounds in solution (McKnight et al.,
1992; Kaiser et al., 1997; Kalbitz, 2001; Schwesig et al.,
2003b; Kaiser et al., 2004). Despite its large content of carbo-
hydrates, subsoil DOM is less degradable than the largely
stable DOM rich in aromatic compounds from Oa horizons
(Boyer and Groffman, 1996; Schwesig et al., 2003b). Where-
as plant-derived carbohydrates seem to be the preferred sub-
strate for microorganisms (Kalbitz et al., 2003b), microbial
carbohydrates and N-rich compounds released during OM
degradation can be hardly degradable (Ogawa et al., 2001;
Kalbitz et al., 2003b). Consequently, subsoil DOM, enriched
in carbohydrates, amino sugars, and other compounds of
mainly microbial origin (Guggenberger and Zech, 1994;
Kaiser et al., 2004), is not labile per se as assumed previous-
ly (e.g., Qualls and Haines, 1992).
3 Degradation of DOM after sorption to
minerals
Sorption to mineral subsoils, clay minerals, and Fe oxide/
hydroxides results in strong stabilization of DOM, as indi-
cated by smaller mineralization rate constants for sorbed OM
than for DOM (Tab. 1; Kalbitz et al., 2005; Mikutta et al.,
2007). The mean residence time of OM derived from the Oa
horizon of a Podzol increased from <30 y in solution to 91 y
after sorption (Kalbitz et al., 2005). These results are in
agreement with stabilization, sometimes even complete sup-
pression of mineralization, of simple organic compounds like
glucose, citrate, oxalate, and malate after sorption to miner-
als and mineral soils (Jones and Edwards, 1998; Strm et al.,
2001; van Hees et al., 2003). In turn, Keil et al. (1994) and
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54 Kalbitz, Kaiser
Nelson et al. (1994) found mineralization rates for OM to
increase up to five orders of magnitude after desorption from
marine sediments and soils, respectively. Fluorescence spec-
tra, UV absorbance, and 13C signature suggest that aromatic
compounds were preferentially stabilized by sorption (Kalbitz
et al., 2005).
The two main reasons responsible for the stabilization of
DOM by sorption are: (1) increased stability of organic com-
pounds due to interactions with the mineral phase (Kgel-
Knabner et al., 2008a, this issue, pp. 6182) and (2) selective
sorption of inherently more stable components. Formation of
strong chemical bonds between the mineral phase and
organic molecules likely reduces the mineralization of C after
sorption (Saggar et al., 1994; Kaiser and Guggenberger,
2000, 2007). Furthermore, physical separation of OM from
microorganisms and hydrolytic enzymes by sorption into
small pores (Mayer, 1994) and sterical changes due to multi-
ple ligand attachment to the sorbing surface (Kaiser and
Guggenberger, 2003) can contribute to the reduced C miner-
alization.
Sorption is strongly selective for the inherently recalcitrant
aromatic and complex compounds (Kaiser and Guggenber-
ger, 2000; Kalbitz et al., 2005). Sorptive stabilization of stable
compounds is consequently much more efficient than of the
less strongly sorbing labile ones (Ransom et al., 1998; Kaiser
and Guggenberger, 2000; Kalbitz et al., 2005). Therefore, the
stronger the sorption of a given DOM, the stronger is the
sorptive accumulation and stabilization of OM (Kaiser and
Guggenberger, 2000; Kalbitz et al., 2005).
4 Degradation of DOM after (co-)precipitation
Precipitation with Al and Fe can be efficient in immobilizing
DOM in acidic soils (Zysset and Berggren, 2001; Nierop et
al., 2002; Jansen et al., 2005). At molar Al : C ratios typical
Table 1: Results of the three incubation experiments used in this study. Parameters were calculated on a basis of 50 d incubation at 20C. For
all incubation experiments, the same DOM (from the Oa horizon of a Podzol; Kalbitz et al., 2005) was used. A and k were calculated by a single
exponential model: Cmineralized (%) = A (1 exp[kt]); k: mineralization rate constant (d1); t: time (d); A: mineralizable DOC (%).
Study / treatment Mineralized C
(%)
Results of Kalbitz et al. (2005) b
OM in solution 18.6
OM sorbed to mineral soil 1.4
Results of Mikutta et al. (2007)
OM in solution 8.7
OM sorbed to goethite c 2.1
OM sorbed to pyrophyllite c 3.5
Results of Scheel et al. (2007)
OM in solution 4.8
OM (co-)precipitated with Al d 0.5
a ratio between the mineralization rate constant of OM sorbed to the mineral soil and the constant of OM sorbed to minerals or (co-)precipitated
with Al
b results of the long-term incubation of Kalbitz et al. (2005); data after 50 d incubation
c sorption with NaCl as background electrolyte (Mikutta et al., 2007)
d precipitation at pH 3.8; molar Al : C ratio: 0.1 (Scheel et al., 2007)
2008 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim
J. Plant Nutr. Soil Sci. 2008, 171, 5260
for acidic forest soils (0.1), up to 80% of DOC can (co-)preci-
pitate (Nierop et al., 2002; Scheel et al., 2007). Co-precipita-
tion (also termed co-flocculation) means precipitation of the
mineral phase (e.g., Al hydroxide) with simultaneous adsorp-
tion of organic molecules (e.g., Jekel, 1986). It is difficult to
distinguish between true precipitation, i.e., formation of inso-
luble organicmetal complexes, and co-precipitation. Both
processes may occur simultaneously. Therefore, we use the
term (co-)precipitation. As for the adsorption of DOM to
minerals, spectroscopic analyses indicated preferential (co-)
precipitation of aromatic compounds, suggesting that mainly
inherently stable compounds tend to precipitate (Rmkens
and Dolfing, 1998; Blaser et al., 1999; Dolfing et al., 1999;
Sharpless and McGown, 1999; Scheel et al., 2007). Schwe-
sig et al. (2003a) noticed formation of flocculated OM and
simultaneously decreasing mineralization of DOM after addi-
tion of Al to soil solutions.
Mineralization of DOM (co-)precipitated with Al is small as
compared to DOM in solution (Tab. 1; Boudot et al., 1989;
Scheel et al., 2007). It depends on the Al species involved in
(co-)precipitate formation (Scheel et al., 2007). The propor-
tion of hydrous Al species seems to be crucial, and it in-
creases with increasing pH and Al : C ratios. Stabilization
decreased with increasing proportions of hydrous Al species
involved in (co-)precipitation as indicated by larger minerali-
zation of precipitates formed at pH 4.5 in comparison to 3.8
and with increasing Al : C ratios in the precipitates (Scheel
et al., 2007). Probably, OM interacts stronger with Aln+ than
with Al hydroxides.
The observed stability of (co-)precipitates could contribute
significantly to stabilization of DOM in soils. The Al concentra-
tions in the solution of acidic soils increase with depth
(Matzner et al., 2004). Besides adsorption on Al and Fe oxi-
des, (co-)precipitation with Al (and other polyvalent cations)
could explain the decreasing DOC concentrations with soil
A k Conversion factor a
(%) (d1)
17.4 0.1033
1.5 0.0482 1.000
8.5 0.1675
2.4 0.0397 1.214
4.0 0.0571 0.844
4.1 0.2466
0.7 0.0212 2.274
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J. Plant Nutr. Soil Sci. 2008, 171, 5260 Stabilization and dissolved organic matter 55

depth as well as accumulation and stabilization of OM in sub-
soils. Co-precipitates rich in OM can also form with Fe
(Schwertmann et al., 2005), especially if weathering of Fe(II)-
bearing minerals is rapid or if interchanging redox conditions
prevail. However, so far no studies addressed the stability of
OM precipitated with cations such as Fe3+ and Ca2+.
A(t) = F (1 exp[kt ])/k (Olson, 1963).
The maximum value of OM accumulation from the input flux
(Aend) can be calculated by assuming time to approach
infinity:
Aend = F / k.

5 Estimation of DOM contribution to C in the
subsoil of a forested site
The contribution of DOM to the accumulation of OM in the
mineral soil depends on the amount of C retained and its
mineralization. The retained C is the difference between the
input of DOC into and its export from the mineral soil. The
decreases in DOM in the mineral soil should be caused by
abiotic processes such as sorption and (co-)precipitation
because microbial degradation of OM in solution likely is of
minor importance (see above).
We estimated the contribution of DOM to the organic C (OC)
in the mineral horizons of a Podzol under Norway spruce in
the Fichtelgebirge, Germany (Kgel-Knabner et al., 2008b,
this issue, pp. 513; Tab. 2). To the best of our knowledge, it
is the only site where DOM retention and DOM mineralization
in soil solution, after sorption to the mineral soil and different
minerals as well as after precipitation by Al, has been studied.
The observed retention of 260 kg C ha1 y1 between 0 and
90 cm depth (Kalbitz et al., 2004a) is well within the range of
the 17 sites (40370 kg C ha1 y1) mentioned in the introduc-
tion. According to Frberg et al. (2006), much of the retention
of DOC occurs in the B horizons. All mineralization rate con-
stants used (Tab. 1, 3) were obtained by incubation of DOM
from the Oa horizon of the study site. Also, the mineral soil
horizon (Bw) used to test the effect of DOM sorption (Kalbitz
et al., 2005) is from that site. Therefore, we have a unique set
of data that allows for estimating the contribution of DOM to
OM in the mineral soil.
5.1 Methods
Generally, the pool size of OM (A) with constant input flux
(F; e.g., DOM) and constant exponential mineralization rate
constant (k) is determined as
As described earlier, mineralization of DOM decreases after
adsorption to minerals or after (co-)precipitation with poly-
valent cations. Therefore, mineralization of the retained DOM
can be described by different mineralization rate constants.
The pool size of OM derived from the constant DOM input (A)
can be calculated as follows:
A(t) = F1 / k1 (1 exp[k1t ] + F2 / k2 (1 exp[k2t ]) + ......
+ Fn / kn (1 exp[knt ]).
F1 to Fn represent the amounts of DOM retained by sorption
to different soil minerals and (co-)precipitation; k1 to kn are
the respective mineralization rate constants. Considering the
different mineralization rate constants, the limit value of OM
accumulation from DOM input (Aend) can be calculated:
Aend = F1 / k1 + F2 / k2 + ...... + Fn / kn.
To estimate the long-term contribution of DOM to the OM
pool in the mineral soil, we used the mineralization rate con-
stants of the stable C pool (kslow) for OM sorbed to the
mineral soil (Bw horizon), as obtained by long-term (375 d)
incubation (Tab. 3; Kalbitz et al., 2005). The mineralization
rate constants for OM after sorption to soil minerals (Mikutta
et al., 2007) and after (co-)precipitation by Al (Scheel et al.,
2007) could not be used directly because of shorter incuba-
tion periods (precipitated OM: 50 d; sorbed OM: 90 d).
Furthermore, mineralization rate constants computed with a
double-exponential model were partly not significant (p >
0.1), and the standard errors were often larger than the con-
stants themselves. Also Sleutel et al. (2005) observed that
double-exponential models did not fit data obtained in degra-
dation experiments with incubation periods <100 d. There-
fore, we normalized all mineralization rate constants to those
of obtained for the long-term incubation of OM sorbed to
mineral soil. To do so, we calculated a mineralization rate
constant for the first 50 d of the long-term incubation of OM

Table 2: Soil properties, C storage (Kalbitz et al., 2004b), 14C contents, and radiocarbon age (Rumpel et al., 2002) at the Waldstein site (Fich-
telgebirge, Germany; Kgel-Knabner et al., 2008b, this issue, pp. xxxx).
Horizon Thickness Sand Silt Clay pH C stock 14C Radiocarbon age
(cm) (%) (CaCl2) (t C ha1) (pmC) (y BP)
Oi 0.5 3.60 3.6
Oe 5 2.90 25.4
Oa 3 2.60 31.0
EA 10 51.6 38.0 10.4 2.90 27.4 93.6 525 30
Bh 2 34.0 49.6 16.4 3.30 8.5 98.5 120 25
Bs 18 44.7 44.8 10.4 3.90 51.1 91.1 745 40
Bw 25 45.8 43.4 10.8 4.30 20.7 82.2 1570 25
C1 15 56.4 34.0 9.6 4.20 1.5 62.0 3840 70
C2 15+ 50.8 38.0 11.2 4.10 0.9
R 170.1

2008 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim www.plant-soil.com

56 Kalbitz, Kaiser
sorbed to the mineral soil (Tab. 1). That mineralization rate
constant was divided by those of sorbed and precipitated OM
obtained in short-term incubations (50 d). The calculated
ratios (conversion factors, Tab. 1) were then divided by the
mineralization rate constant (kslow) determined for OM sorbed
to the mineral soil by long-term incubation (Tab. 3). We
assume the difference in rate constants between the soil
minerals and precipitates in the short-term incubation to hold
through also for long-term incubation.
Table 3: Input parameters (kslow) to estimate the contribution of DOM
to OM in the mineral soil at the Waldstein site (Fichtelgebirge,
Germany) after long-term incubation (375 d; Kalbitz et al., 2005). The
decomposition rate constants kslow were converted to 5C according
to Tjoelker et al. (2001) and Kalbitz et al. (2005); kslow values derived
from a double-exponential model: mineralized C (% of total C) = a (1
exp[kfast t]) + (100 a) (1 exp[kslow t]); a: labile C (%), 100 a:
stable C (%), kfast: mineralization rate constant of labile C (d1), kslow:
mineralization rate constant of stable C (d1), t: time (d). The amount
of retained DOC is 260 kg ha1 y1 (Kalbitz et al., 2004a).
kslow of OM sorbed to mineral soil 0.000030
kslow of OM matter sorbed to goethite 0.000025a
kslow of OM sorbed to pyrophyllite 0.000035a
kslow of OM (co-)precipitated with
Al at pH 3.8 and Al : C = 0.1 0.000013a
a calculated by dividing the kslow of OM sorbed to mineral soil
(determined by long-term incubation) by the conversion factor
(Tab. 1)
From the study of Mikutta et al. (2007), we selected two soil
minerals, goethite and pyrophyllite (an illite-like material but
with smaller permanent charge), with NaCl as the back-
ground electrolyte for sorption. From Scheel et al. (2007), we
selected the precipitates obtained at pH of 3.8 and a molar
Al : C ratio of 0.1; because these values reflect average con-
ditions in the different mineral soil horizons (Tab. 2).
Our data set did not allow for distinguishing between different
mineral soil horizons. Instead, we assumed a uniform reten-
tion throughout the mineral soil and neglected preferential
flow (Kalbitz et al., 2005). Furthermore, all mineralization rate
constants were adjusted to the mean annual temperature of
our study site (5C) using the approach of Tjoelker et al.
(2001) and Kalbitz et al. (2005):
kx = kx / Q10 (Tx Tx) / 10C,
Tx = 5C (specific temperature: aim of conversion),
Tx = 20C (reference temperature: incubation),
kx = mineralization rate constant at 5C,
kx = mineralization rate constant at 20C.
As we do not know to which extent sorption by different soil
minerals and (co-)precipitation with Al contributed to the
DOM retention, we tested each variant independently by allo-
cating the total retention of 260 kg C ha1 y1 (Kalbitz et al.,
2004a) to the various processes. This procedure gives the
range of the contribution of DOM to OC in the mineral soil.
The four scenarios tested were:
2008 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim
J. Plant Nutr. Soil Sci. 2008, 171, 5260
(1) DOM retention only by sorption to a mineral soil horizon
(Bw; Kalbitz et al., 2005);
(2) DOM retention only by sorption to goethite (Mikutta et al.,
2007);
(3) DOM retention only by sorption to pyrophyllite (Mikutta
et al., 2007);
(4) DOM retention only by (co-)precipitation with Al at pH of
3.8 and a molar Al : C ratio of 0.1 (Scheel et al., 2007).
The study soil is characterized by large stocks of hydrous Fe
and Al phases and large concentrations of dissolved Al, thus
sorption as well as (co-)precipitation of DOM are both likely.
5.2 Results
Using the mineralization rate constants for DOM sorbed to
the Bw horizon, the soil OC deriving from DOM was esti-
mated to 24 Mg C ha1, which represents 22% of the total soil
OC (Fig. 1; Kalbitz et al., 2005). The maximum contribution of
DOM-derived C was estimated to 50% (55 Mg C ha1),
assuming that all retained DOM is (co-)precipitated with Al.
This is because of the 2.3-fold larger mean residence time of
OM (co-)precipitated with Al than sorbed to the Bw horizon
(Tab. 1). Sorption of all retained DOM to goethite would result
in an accumulation of 28 Mg C ha1 in the mineral soil, which
corresponds to 26% of the stored OC. The larger mineraliza-
tion rate constant for OM sorbed to pyrophyllite results in a
smaller contribution to soil C (20 Mg ha1, corresponding to
19% of soil OC). Variations of the amounts of retained DOM
and of the mineralization rate constants will result in propor-
tional changes in the final contribution of DOM to SOM
because of the linearity of the used equation when approach-
ing the point of infinity.
120000
measured C stock in the mineral soil
C stock in the mineral soil (kg C ha-1)
110000
60000 (co)precipitation by Al
50000
40000
sorption to Goethite
30000
sorption to the mineral soil
20000
sorption to pyrophilite
10000
0
0 200 400 600 800 1000
time (years)
Figure 1: Contribution of DOM to the build-up of OM in the mineral
soil at the site Waldstein (Fichtelgebirge, Germany) assuming a
constant DOM retention of 260 kg C ha1 y1 and constant
mineralization rate constants. Dissolved organic matter retention
was allocated to four different scenarios. Mineralization rate
constants for sorption and (co-)precipitation are given in Tab. 3.
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J. Plant Nutr. Soil Sci. 2008, 171, 5260
5.3 Discussion
Our results highlight the contribution of DOM to the accumu-
lation of OM in the mineral soil. The similar chemical compo-
sition of DOM in forest floor percolates and of OM in acidic
subsoil is another indication that OM in the mineral soil can
derive from retained DOM (e.g., Kaiser and Guggenberger,
2000). In many acidic subsoils, most of the OM dissolves
upon dissolution of the mineral soil matrix with hydrofluoric
acid (Eusterhues et al., 2003). That means the OM is poten-
tially soluble, likely because representing sorbed or precipi-
tated DOM. Decomposition of roots is likely also linked to
production of DOM, which can be subsequently sorbed or
precipitated, thus contributing to OM in the subsoil.
Although our data do not allow to allocate the estimated stabi-
lization of DOM to the soil horizons of the study site, we can
assume that most of DOM-derived OM is located in B and C
horizons. The estimated DOM-derived C stock (2055 Mg C
ha1) comprises 25%66% of the OC in B and C horizons of
the study soil. Organic matter in the EA horizon is mainly par-
ticular, of high age and with large contribution of insoluble,
long-chain alkyl compounds (Rumpel et al., 2004). Mineral-
associated OM derived from sorption or (co-)precipitation is
only of minor importance in this horizon. Distribution, compo-
sition, and age of C in particle-size fractions and the large
proportion of mineral-associated OC indicate a predominant
influence of DOM on SOM in B and C horizons of the study
soil and also of a Dystric Cambisol under European beech
(Rumpel et al., 2004). The large abundance of carboxyl C in
the Bs horizon is a further proof that the OM derives from
water-soluble OM from the forest floor (Rumpel et al., 2004).
The apparent mean radiocarbon age in the Bh horizon (120 y;
Tab. 2) is between the mean residence time of sorbed and
(co-)precipitated OM as determined by incubation studies
(Fig. 1; Kalbitz et al., 2005). This is another indication for OM
in the Bh horizon to derive largely from DOM.
The different scenarios tested suggest that DOM-derived C in
the mineral soil can vary over a wide range (2055 Mg C
ha1), depending on the mineralization rate constants. The
ultimate contribution of DOM to OC in the mineral soil
depends on the portions of OM sorbed to different minerals
and (co-)precipitated. These portions, however, we do not
know. Adsorption to hydrous oxides (and acidic subsoils) as
well as (co-)precipitation with Al and Fe can remove DOM
from Oa horizons almost entirely from solution (Kaiser et al.,
1996; Kaiser and Zech, 1997; Nierop et al., 2002; Scheel
et al., 2007). In the review of Kalbitz et al. (2000), sorption in
the mineral soil is considered to be the main process of DOM
retention. On the basis of measured DOC fluxes, Guggenber-
ger and Kaiser (2003) estimated the sorption capacity to be
exhausted within a few years, given no neoformation of reac-
tive mineral phases. However, field data did not indicate a
decrease in DOC retention. In contrast, DOC concentrations
decreased at 20 and 90 cm depth at the study site during the
last 10 y despite constant input from the forest floor (Kalbitz
et al., 2004a). On the other hand, it needs approx. 300 y to
reach the equilibrium between sorption and mineralization of
DOM (Fig. 1). Mineral weathering and neoformation of Fe
and Al oxides/hydroxides may provide sufficiently large sorp-
2008 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim
Stabilization and dissolved organic matter 57
tion capacity over such a period of time. Thus, the actual
sorption capacity cannot be used to predict the long-term
potential of soils to stabilize DOM via sorption. Also, weather-
ing releases metals such as Al and Fe which can induce (co-)
precipitation of DOM, thus, maintaining steady retention as
well as stabilization of DOM (Zysset and Berggren, 2001;
Kalbitz et al., 2005; Scheel et al., 2007).
The apparent mean 14C age of DOM in the B horizon of two
podzolic soils in Sweden was 50 and 300 y, respectively, and
exceeded the age of DOM in the forest floor by far (Frberg
et al., 2006). Also NaCl-extractable OC was much older in
the B horizon of the study soil (14C content: 99.9 pmC) than
in the Oe (114.4 pmC) and in the Oa horizon (102.6 pmC;
Michalzik et al., 2003). The age of DOM might reflect the time
to reach equilibrium between retention and mineralization of
DOM which is approx. 300400 y, depending on the minerali-
zation rate constant of the DOM retained. A younger age of
DOM than the equilibrium time may indicate a portion of
DOM having the 14C bomb signal to be directly leached from
the forest floor into deeper mineral horizons.
The mineralization rate constants of (co-)precipitated OM
were lower than those of sorbed OM (Scheel et al., 2007;
Mikutta et al., 2007; Tab. 1). Therefore, a stronger stabiliza-
tion of OM by (co-)precipitation than by sorption seems prob-
able. However, also sorption of DOM might result in much lar-
ger stabilization of OM than measured by Kalbitz et al. (2005)
and Mikutta et al. (2007). So, DOM sorption to a multi-domain
goethite and to amorphous Al hydroxide resulted in an almost
complete inhibition of C mineralization (Kaiser and Schneider,
unpublished data). The same observation was made by
Jones and Edwards (1998) after adsorption of simple organic
compounds to certain minerals such as ferrihydrite. Also, we
have no data on the mineralization of DOM co-precipitated
with Fe. Considering the strong stabilization of organic com-
pounds by ferrihydrite, the dominant mineral phase in such
co-precipitates (Schwertmann et al., 2005), the protection of
the contained OM is likely rather efficient. These examples
imply that our estimate of the potential contribution of DOM to
soil C is largely conservative.
5.4 Limitations and uncertainties of the estimation
One limitation of our study is due to the fact that we do not have
estimates of the contribution of each of the individual com-
pounds (different minerals, [co-]precipitates) to total DOM reten-
tion in the mineral soil. OM sorbed by different minerals or con-
tained in (co-)precipitates is mineralized at different rates (see
above). Our estimate of DOM-derived C stocks in the mineral
soil is only 1/3 to 1/2 the estimate by Michalzik et al. (2003) who
modeled the DOM dynamics in forested ecosystems. They
found that 73%89% of the OM in the mineral soil of two sites
derived from DOM, the rest from roots. In contrast, DOC con-
tributed to 25% of total soil C according to a modeling study
by Neff and Asner (2001). This range of estimates points to
other DOM sources like roots, not included in our estimation,
and to the conservative character of our calculations.
Roots may be an important source of DOM in soil, either by
releasing exudates or due to water-soluble compounds pro-
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58 Kalbitz, Kaiser
duced during their decomposition (Kalbitz et al., 2000). The
portion of root-derived DOM, however, is largely unknown. It
is not included or at least underestimated in soil solution
sampled by lysimeters in the field because of their small size
relative to the large spatial variability of root exudation and
decomposition. It is even impossible to estimate the contribu-
tion of roots to SOM although root litter is generally less
decomposable than leaf litter (reviewed by Rasse et al.,
2005). Rumpel et al. (2004) found large amounts of root-
derived compounds in SOM of the mineral soil. These find-
ings suggest a larger contribution of roots to SOM than pre-
viously assumed. Thus, root-derived compounds may also
contribute significantly to DOM in the subsoil.
Mineralization of DOM results in microbial production of
water-soluble compounds (Kalbitz et al., 2003b). This pro-
cess cannot be quantified but might be important particularly
in the deeper mineral soil where other C sources are sparse.
The 14C content in subsoil horizons decreases with depth
(Rumpel et al., 2002); this is paralleled by a large contribution
of smaller molecules of microbial origin to SOM in the deeper
mineral soil (Rumpel et al., 2004). These findings support the
assumption of microbial DOM production directly in the sub-
soil. Sorbed or (co-)precipitated OM is slowly mineralized
(k values: Tab. 1, 3) which likely is accompanied by the
release of water-soluble microbial carbohydrates and pep-
tides (Kalbitz et al., 2003b). These compounds will be lea-
ched to deeper soil horizons.
Besides of quantitatively unknown other DOM sources in the
mineral soil, our estimation of the contribution of DOM to OC
storage might be conservative because we used mineraliza-
tion rate constants obtained in the laboratory under optimum
conditions. We assume that mineralization in the laboratory is
faster than in the field although the rate constants were
adapted to the mean annual temperature of the study site.
Furthermore, mineralization rate constants depend on the
time of incubation. The rate constants for the mineralization
of C in DOM solutions from the Oi and Oa horizon of the
study site obtained in short-term incubations (50 d) were 2.6-
to 4.6-fold larger than those derived from long-term incuba-
tion (375 d) (data from Kalbitz et al., 2005). From a long-term
perspective, we have to consider that the mineralization rate
constants used in our calculations are probably too large.
The mean residence time of SOM in the Bh horizon according
to 14C analysis (120 y) is somewhat longer than that of
adsorbed OM obtained by long-term incubation (91 y; Kalbitz
et al., 2005). The true mean 14C age might be even higher
taking possible inputs of bomb 14C into account. If we
assume the true mineralization rates in the field to be half of
those obtained in the laboratory, the DOM-derived soil C
would double. Then, the proportion of DOM-derived C stock
in the mineral soil would be within the range of proportions
estimated by modeling (Michalzik et al., 2003).
6 Conclusions
Dissolved OM is an important contributor to the OC in mineral
soils of forest sites. Adsorption onto minerals and (co-)preci-
pitation with Al results in stabilization against microbial decay.
At least 20 to 55 Mg C ha1 in the mineral soil of the test site
2008 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim
J. Plant Nutr. Soil Sci. 2008, 171, 5260
derives from DOM, which represents 19%50% of total soil
OC. These numbers are probably underestimates because of
the unknown contribution of roots to the DOM pool and addi-
tional DOM production in the mineral soil. (Co-)precipitation
with Al seems to reduce the mineralization stronger than
sorption to phyllosilicates and goethite. However, reactive
minerals with large surface areas (multi-domain goethite, fer-
rihydrite, amorphous Al hydroxide) can inhibit C mineraliza-
tion of sorbed OM almost completely. Therefore, the type of
minerals and the formation of (co-)precipitates determine the
DOM-derived reservoir of OC in the mineral soil. An improved
evaluation of DOM-derived soil C therefore requires robust
estimates on contribution of individual minerals and (co-)pre-
cipitation with Al and Fe to the total DOM retention. Special
emphasis has to be given to obtain mineralization rate con-
stants under field conditions.
Acknowledgments
We gratefully acknowledge the financial support by the
Deutsche Forschungsgemeinschaft Priority Program 1090
Soils as sinks and source of CO2 mechanisms and regula-
tion of organic matter stabilisation in soils. C. Drfler,
L. Haumaier, R. Mikutta, T. Scheel, J. Schmerwitz, M. Schnei-
der, and D. Schwesig contributed much to the scientific back-
ground of the manuscript. For discussions and support we
thank G. Guggenberger, B. Marschner, and E. Matzner.
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