Carbon sequestration potential of soils in southeastGermany derived from stable soil organic carbonsaturationMART IN WIESME IER * , R ICO HUBNER † , P ETER SP €ORLE IN ‡ , UWE GEUß ‡ ,
EDZARD HANGEN ‡ , ARTHUR RE I SCHL ‡ , B ERND SCH ILL ING ‡ , MARG IT VON L UTZOW*
and INGRID K €OGEL-KNABNER*§
*Lehrstuhl fur Bodenkunde, Department fur Okologie und Okosystemmanagement, Wissenschaftszentrum Weihenstephan fur
Ern€ahrung, Landnutzung und Umwelt, Technische Universit€at Munchen, Freising-Weihenstephan 85350, Germany, †Lehrstuhl
fur Wirtschaftslehre des Landbaues, Wissenschaftszentrum Weihenstephan fur Ern€ahrung, Landnutzung und Umwelt, Technische
Universit€at Munchen, Freising-Weihenstephan 85350, Germany, ‡Bavarian Environment Agency, Hof 95030, Germany,
§Institute for Advanced Study, Technische Universit€at Munchen, Garching 85748, Germany
Abstract
Sequestration of atmospheric carbon (C) in soils through improved management of forest and agricultural land is
considered to have high potential for global CO2 mitigation. However, the potential of soils to sequester soil organic
carbon (SOC) in a stable form, which is limited by the stabilization of SOC against microbial mineralization, is largely
unknown. In this study, we estimated the C sequestration potential of soils in southeast Germany by calculating the
potential SOC saturation of silt and clay particles according to Hassink [Plant and Soil 191 (1997) 77] on the basis of
516 soil profiles. The determination of the current SOC content of silt and clay fractions for major soil units and land
uses allowed an estimation of the C saturation deficit corresponding to the long-term C sequestration potential. The
results showed that cropland soils have a low level of C saturation of around 50% and could store considerable
amounts of additional SOC. A relatively high C sequestration potential was also determined for grassland soils. In
contrast, forest soils had a low C sequestration potential as they were almost C saturated. A high proportion of sites
with a high degree of apparent oversaturation revealed that in acidic, coarse-textured soils the relation to silt and clay
is not suitable to estimate the stable C saturation. A strong correlation of the C saturation deficit with temperature
and precipitation allowed a spatial estimation of the C sequestration potential for Bavaria. In total, about 395 Mt
CO2-equivalents could theoretically be stored in A horizons of cultivated soils – four times the annual emission of
greenhouse gases in Bavaria. Although achieving the entire estimated C storage capacity is unrealistic, improved
management of cultivated land could contribute significantly to CO2 mitigation. Moreover, increasing SOC stocks
have additional benefits with respect to enhanced soil fertility and agricultural productivity.
Keywords: agricultural management, climate change, CO2 mitigation, soil organic carbon stocks, soil fractionation, stabilization
of soil organic matter
Received 18 April 2013 and accepted 30 August 2013
Introduction
Sequestration of atmospheric carbon (C) in soils is
considered to contribute significantly to CO2 mitiga-
tion, and several management options for increasing
SOC stocks have been discussed. For forest ecosystems,
practices such as a change in tree species composition,
afforestation, thinning, drainage, fertilization, liming,
site preparation and harvest management are associ-
ated with an increase in SOC stocks and are conse-
quently viewed as having a high potential for soil C
sequestration (Goodale et al., 2002; Liski et al., 2002;
Karjalainen et al., 2003; Lal, 2005; Jandl et al., 2007; Ciais
et al., 2008; Lorenz & Lal, 2010; Luyssaert et al., 2010;
Carroll et al., 2012; Vesterdal et al., 2012; Wiesmeier
et al., 2013b). An even higher C sequestration potential
is assumed for agricultural soils because a distinct
depletion of SOC stocks has been observed in most
cultivated soils (Paustian et al., 1997; Lal, 2004; Smith,
2004). Among several agricultural practices that may
increase C sequestration in cultivated soils, promising
management options are promotion of organic inputs,
conservation/zero tillage, converting cropland to grass-
land, introduction of perennials, improved manage-
ment of farmed peatland and organic farming (Cole
et al., 1997; Paustian et al., 2000; Sauerbeck, 2001; Vlees-
houwers & Verhagen, 2002; Freibauer et al., 2004; Hol-
land, 2004; Lal, 2004; Johnson et al., 2007; Smith, 2012).Correspondence: Martin Wiesmeier, tel. +49 (0)8161 71 3679,
fax +49 (0)8161 71 4466, e-mail: [email protected]
© 2013 John Wiley & Sons Ltd 653
Global Change Biology (2014) 20, 653–665, doi: 10.1111/gcb.12384
However, C sequestration by improved management
of forest and agricultural soils reaches a new equilib-
rium at a higher SOC level after a certain period of
time. Several studies have shown that there is an upper
limit of SOC storage, confirming the hypothesis of soil
C saturation (Six et al., 2002; Goh, 2004; Stewart et al.,
2007, 2008; Chung et al., 2008). This is related to the lim-
ited potential of soils to stabilize soil organic matter
(SOM) against microbial mineralization (Baldock &
Skjemstad, 2000). There are three major SOM stabiliza-
tion mechanisms: selective preservation due to recalci-
trance of SOM, spatial inaccessibility of SOM due to
hydrophobicity or occlusion in soil aggregates, and
interaction with mineral surfaces (Sollins et al., 1996;
von Lutzow et al., 2006). The last is regarded as quanti-
tatively the most important in a wide range of soils, as
indicated by a strong correlation of SOC stocks with
clay contents (e.g. Oades, 1988; Arrouays et al., 2006).
Hassink (1997) assumed that the capacity of soils to
preserve SOC is limited by the proportion of silt and
clay particles (fine fraction <20 lm). He found a strong
correlation of SOC stored in the fraction containing silt
and clay particles in a wide range of uncultivated and
grassland topsoils of temperate and tropical regions
and proposed that this correlation could be used to esti-
mate the stable SOC saturation. The difference between
the potential C saturation of the fine fraction and the
actual measured C content of this fraction corresponds
to the C saturation deficit or, in other words, the C
sequestration potential. Hassink’s approach was used
in several experimental studies to estimate the degree
of C saturation in cultivated soils (Chan, 2001; Six et al.,
2002; Carter et al., 2003; Conant et al., 2003; Sparrow
et al., 2006; Zhao et al., 2006). However, these studies
included only a limited number of investigated loca-
tions, thus the results are not suitable to derive compre-
hensive conclusions for larger (landscape) scales.
Recently, Angers et al. (2011) estimated the C saturation
deficit of French agricultural topsoils using Hassink’s
approach on the basis of about 1.5 million SOC and 0.3
million particle-size determinations. However, the
results must be regarded as a very rough estimation of
the C saturation deficit as the actual C content of the
fine fraction was not measured; instead, an estimated
constant proportion of 85% of total SOC was assumed.
Moreover, no absolute amount of the C sequestration
potential could be derived due to missing data for bulk
density (BD). The need for quantitative studies estimat-
ing the C sequestration potential for various soils and
land uses on larger scales was highlighted in a recent
review of SOC sequestration (Stockmann et al., 2013).
In this study, we used a comprehensive soil data set
within the state of Bavaria in southeast Germany to
estimate the C sequestration potential of soils under the
main land uses of cropland, grassland and forest and to
gain insight into the controlling factors. For 516 soil
profiles, data on soil texture, SOC content, BD and
stone content (SC) enabled a quantification of the C
saturation potential according to Hassink (1997). The
determination of the current C concentration of all
major soil units and land uses in Bavaria allowed a
calculation of the C sequestration potential. The
objectives of the study were to do the following
1. Determine the C saturation deficit in agricultural
and forest soils.
2. Elucidate relationships between land use-specific C
saturation deficits and environmental factors.
3. Quantify the total C sequestration potential of soils
within Bavaria.
Materials and methods
Study area
The state of Bavaria, with an area of 70 550 km2, is located in
southeast Germany and comprises various landscapes. The
north-western part of Bavaria is dominated by the southern
German escarpment landscape that adjoins low mountain
ranges of the Bohemian Massif in the east. Southwards the
Molasse basin ascends to the mountain range of the Alps at
the southern border of Bavaria. Elevation ranges between 107
and 2962 m above sea level. Due to its location in central
Europe, Bavaria exhibits a suboceanic climate that is charac-
terized by a transitional situation between a maritime climate
in the north-west and subcontinental influences in the east.
Mean annual temperature and precipitation from the escarp-
ment landscape in the north-west to the Alps in the south
range between 9 and 4 °C and 550 and 2500 mm respectively.
Dominant soil classes are soils with well-developed B hori-
zons (Cambisols) at 45% of the total area, soils with initial soil
formation (Leptosols, Regosols) at 14% and soils with water
stagnation (Stagnosols, Albeluvisols, Planosols) at 13% accord-
ing to the German soil systematic (AD-HOC AG Boden, 2005)
and the equivalent Reference Soil Groups of the WRB system
(IUSS Working Group WRB, 2006).
Compilation of soil data
Available data from different soil surveys and permanent soil
observation sites in Bavaria overseen by the Bavarian Environ-
ment Agency and the Bavarian State Institute for Forestry
were screened to compile a representative data set. Sampling
locations were incorporated only where topsoil material had
been analysed for SOC, BD, SC and soil texture. The minimum
requirement for SOC analysis was a determination by dry
combustion using a CN elemental analyser. Generally, only
soil data collected after 1990 were considered to reduce the
impact of temporal SOC changes. The selected sampling loca-
tions that fulfilled all requirements comprised 516 sites. The
© 2013 John Wiley & Sons Ltd, Global Change Biology, 20, 653–665
654 M. WIESMEIER et al.
main land uses were adequately represented, with 115 loca-
tions (22% of the data) as cropland (34% of the total area), 110
locations (21%) as grassland (16%), 249 locations (48%) as
forest (35%) and 42 locations (8%) under other land uses
(15%). The main part of the data constituted a grid sampling
within Bavaria (Joneck et al., 2006). Between 2000 and 2004,
soil profiles were sampled using grids of 8 9 8 km within
Bavaria. For each soil profile, a representative location was
selected within a radius of 500 m around the grid node to
achieve a homogeneous sampling area in terms of vegetation,
relief, soil type and parent material as well as a central posi-
tion in the particular land use type. Anthropogenic distur-
bances in the subsoil were excluded in a pre-exploratory
survey using a soil auger. Topsoil material was collected as a
composite sample from eight sub-locations around one main
soil profile to cover the small-scale heterogeneity of the soils.
At the main soil profile, steel core samples with a diameter of
10 cm were extracted for topsoil horizons. A small number of
soil profiles originated from permanent soil monitoring sites
(Schubert, 2002) and other regional soil surveys.
Determination of soil properties
The proportion of SOC stored in the fraction <20 lm was
obtained by physical fractionation of available topsoil material
from 95 locations of the compiled soil data set which are rep-
resentative of the main land uses within Bavaria. For each
land use type, three sampling locations were selected for all
major soil units within Bavaria to cover the range of environ-
mental conditions in terms of soil type, parent material and
climate. For the fractionation, 30 g of soil material <2 mm was
suspended in 150 ml of deionized water and dispersed using
a calibrated ultrasonic probe-type (Bandelin, Berlin, Germany)
with an output energy of 22 J ml�1. This relatively low energy
was applied to disrupt only weakly stabilized soil macroag-
gregates and to prevent the disruption of mineral-associated
SOM (Amelung & Zech, 1999). The stable SOC fraction was
separated by sieving the suspension at 20 lm using pressure
filtration to overcome capillary forces at a 20 lm level
(Amelung & Zech, 1999; Spielvogel et al., 2006; Steffens et al.,
2009) followed by a subsequent extraction of labile dissolved
organic carbon (OC) using a 0.45 lm membrane filter. The
fraction <20 lm was dried at 40 °C, weighed, ground and
analysed in duplicate for OC concentration together with bulk
topsoil samples by dry combustion using a Vario EL elemental
analyser (Elementar, Hanau, Germany).
For the analyses of soil texture, soil samples (<2 mm) were
oxidized with H2O2 to remove organic matter (OM). The
remaining material was dispersed with Na4P2O7 and shaken
for 16:00 hours to 24:00 hours, followed by wet sieving to iso-
late sand fractions >63 lm. To determine silt and clay frac-
tions, approximately 3 g of the <63 lm fraction was
suspended in deionized water using Na4P2O7 and an ultraso-
nication for 3 min with 75 J ml�1 was conducted. Afterwards,
the distribution of silt and clay fractions was obtained by
measuring the X-ray absorption of the soil–water suspension
during sedimentation of the soil particles using a Micromeri-
tics Sedigraph 5100 (Micromeritics, Norcross, GA, USA)
(Sp€orlein et al., 2004). The proportion of the fine frac-
tion <20 lm consisted of medium silt (20–6.3 lm), fine silt
(6.3–2.0 lm) and clay (<2.0 lm). BD was quantified with the
mass of the oven-dry soil (105 °C) divided by the core volume
(Hartge & Horn, 1989). Soil pH values were measured in
0.01 M CaCl2 at a soil to solution ratio of 1 : 2.5 at room tem-
perature.
Calculation of C saturation and C sequestration potential
The potential C saturation of particles <20 lm was calculated
using the equation of Hassink (1997):
Csat�pot ¼ 4:09þ 0:37� particles � 20 lm(%) ð1Þwhere Csat-pot is the potential C saturation (mg g�1), which is
calculated using a linear regression with an intercept of 4.09
and a slope of 0.37 multiplied by the proportion of fine soil
particles <20 lm (%). For the calculation of the C saturation
deficit of a location, the current C concentration of the fine
fraction <20 lm has to be determined. Due to the laborious
soil fractionation, the current C concentration of the fine frac-
tion <20 lm was determined for representative soils under
main land uses of the study area at 95 locations (21 cropland,
32 grassland and 42 forest sites). The relative proportion of the
current C concentration of the fine fraction <20 lm was calcu-
lated for soils under the main land uses cropland, grassland,
forest and other uses and multiplied by the total SOC concen-
tration of topsoils of all 516 sampling locations, which were
allocated to these land uses. To account for uncertainty that
may result from natural variation in the measured current C
concentration of each land use type, a Monte Carlo simulation
was applied (Larocque et al., 2008). To calculate the C satura-
tion deficit of the sampling locations, the estimated current C
concentrations of the fine fraction were subtracted from the
potential C saturation:
Csat�def ¼ Csat�pot � Ccur ð2Þwhere Csat-def is the C saturation deficit (mg g�1) and Ccur is
the current mean C concentration of the fine fraction <20 lm(mg g�1). The total amount of the C sequestration potential
was calculated using the following equation:
Cseq ¼ Csat�def � BD� ð1� RFÞ � T� 10�2 ð3Þwhere Cseq is the C sequestration potential (kg m�2), Csat-def is
the C saturation deficit (mg g�1), BD is the bulk density
(g cm�3), RF is the volumetric fraction of rock fragments
>2 mm (%) and T is the topsoil thickness (10 cm).
Environmental variables
Several environmental parameters that potentially impact the
C saturation deficit were compiled to gain insight into the
processes controlling the C sequestration potential of soils. On
the basis of a digital elevation model with a resolution of 25 m
from the Bavarian Surveying and Mapping Authority, differ-
ent topographical parameters were calculated (Wilson &
Gallant, 2000). As primary terrain attributes, elevation, slope
© 2013 John Wiley & Sons Ltd, Global Change Biology, 20, 653–665
SOC SATURATION AND SEQUESTRATION POTENTIAL 655
and curvature were determined. As secondary parameters,
the contributing area (CA) and the topographic wetness index
(TWI) were calculated using the following equation:
TWI ¼ lnCA
tan a
� �ð4Þ
where CA is the specific upslope contributing area derived by
a geographical information system and a is the slope. The TWI
is a topographical variable that indicates soil moisture condi-
tions (Beven & Kirkby, 1978; Sorensen et al., 2006). To include
geology as a potential parameter influencing the C saturation,
parent material data were assigned from a map with 35 parent
material classes (BAG500) with a resolution of 2 km from the
Bavarian Environment Agency. Information about the soil type
was included using a generalized soil map (B€UK1000N) with
28 superior soil classes (Leitbodenassoziationen) with a resolu-
tion of 2 km from the Federal Institute for Geosciences and
Natural Resources. The factor land use was incorporated by
using 2006 satellite data from the CORINE Land Cover project
(CLC2006) from the German Remote Sensing Data Center. For
climatic variables, annual precipitation and mean annual tem-
perature determined between 1981 and 2010 by the German
Weather Service with a resolution of 1 km were allocated. All
environmental parameters were assigned to 25 9 25 m cells.
Statistical analysis
Descriptive statistics were applied to describe the soil data sets
including mean, minimum and maximum values, median,
interquartile range, extremes and outliers, skewness and kur-
tosis. In order to scale current C concentrations of the fine frac-
tion <20 lm from relatively few 95 representative locations to
all 516 sampling locations within the study area, natural vari-
ability had to be accounted for (Landres et al., 1999). Larocque
et al. (2008) highlight the importance of input data amplitudes
for modelling, due to their effect on the model outcomes sensi-
tive to variations in the inputs. For our simple steady-state
model (output independent of time) thus some degree of sto-
chasticity was included by using a Monte Carlo approach. In
the first step, input data distributions were determined for
each land use type subset. In a second step, random generated
values were applied to the model and results of 1.000 itera-
tions were recorded. Model results were used further for the
prediction of the C saturation deficit and the respective stor-
age potential on the landscape level of the study region.
For the analysis of the importance of a variable to the C
saturation deficit, Pearson’s correlation coefficients among all
parameters were calculated. To include the nominally scaled
parameters soil class and parent material, the different classes
of soil and parent material were ranked according to their
potential to store SOM. For the factor soil class, the ranking
was done by considering the properties soil profile thickness,
soil texture and wetness of the respective soil class which
largely impact total SOC storage. Median values of the soil
profile thickness were calculated for all soil classes and
grouped into four classes (1 = >100 cm; 2 = 95–100 cm;
3 = 90–95 cm; 4 = <90 cm). For soil texture and wetness,
which control the stabilization and degradation of SOM,
empirical values for all soil classes based upon expert knowl-
edge and the literature (AD-HOC AG Boden, 2005) were used
to derive a texture/wetness gradient composed of four classes
ranging from very moist/high clay contents (class 1) to dry/
high sand contents (class 4). The soil classes were grouped into
five classes according to the combined assignment to profile
thickness and texture/wetness classes (the lower the cumula-
tive value of classes, the higher the potential for SOC storage).
To estimate the SOC storage potential of soils based on differ-
ent parent materials, the degree of weatherability and clay con-
tents in the parent material as well as in the weathering
product derived from expert knowledge were considered. For
the ranking, the different parent materials were grouped into
10 units ranging from highly weathered, clay-rich material
with a high potential for soils to store SOC to material with a
low degree of weatherability and low proportions of clay.
Incorporation of nominally scaled parameters using dummy
variables was dismissed due to the high number of soil classes
and parent materials. Due to strong intercorrelations between
environmental parameters, a principal component analyses
(PCA) was carried out to extract the main factors controlling
SOM storage. A stepwise multiple linear regression model was
calculated using the extracted factors. All statistical calcula-
tions were performed using the software IBM SPSS Statistics 19.
Results
Current C concentration and potential C saturation of thefine fraction
The determination of soil texture revealed similar
contributions of particles <20 lm among different land
uses. Topsoils under the main land uses cropland,
grassland and forest had almost identical proportions
of the fine fraction (44–45%; Table 1). For other land
uses, a slightly lower percentage (42%) was deter-
mined. Therefore, the potential C saturation of the fine
Table 1 Proportion of particles <20 lm, SOC concentration, pH value and potential C saturation (Csat-pot) for topsoils (0–10 cm)
under different land uses in Bavaria (median values with 25th and 75th percentile in parentheses)
n Particle<20 lm (%) SOC (mg g�1) pH (CaCl2) Csat-pot (mg g�1)
Cropland (C) 115 45 (34/56) 14.0 (11.2/19.9) 6.3 (5.6/6.8) 20.8 (16.8/24.7)
Grassland (G) 110 45 (32/63) 26.4 (17.7/34.6) 5.5 (4.8/6.1) 20.6 (15.9/27.3)
Forest (F) 249 44 (30/58) 49.0 (31.6/73.3) 3.5 (3.2/3.9) 20.2 (15.3/25.4)
Other use (O) 42 42 (30/63) 14.5 (12.1/20.7) 5.8 (5.3/7.0) 19.7 (15.1/27.3)
© 2013 John Wiley & Sons Ltd, Global Change Biology, 20, 653–665
656 M. WIESMEIER et al.
fraction, which was calculated according to the equa-
tion of Hassink (1997), was also similar between land
uses and ranged between 19.7 and 20.8 mg g�1.
The current C concentration of the fine fraction was
measured for soils under major land uses in Bavaria
(Fig. 1). For cropland soils, the proportion of OC
stored in the fraction <20 lm related to the total OC
content of the bulk soil was in a relatively narrow
range, with a median value of 77%. Soils under grass-
land and forest showed generally lower proportions as
well as a higher variability in fine fraction OC. For
grassland soils, OC contributions of the fine fraction
ranged between 49 and 68%, with a median of 60%.
Under forest, soils showed even lower proportions of
OC stored in the fine fraction, with values of 26–46%with a median of 38%. The estimation of the current C
concentrations of all 516 sampling locations using a
Monte Carlo simulation from the results of 95 fraction-
ated locations revealed considerable differences
between the land uses (Table 2). Cropland soils
showed a relatively low median current C concentra-
tion of 9.6 mg g�1. For grassland and forest soils,
significantly (P < 0.05) higher values of 14.9 and
18.5 mg g�1 were determined. Soils under other uses
showed a current C concentration of 10.6 mg g�1.
Compared with the potential C saturation of the fine
fraction, cropland soils had a mean C saturation of
47%, grassland soils of 73%, forest soils of 93% and
soils under other land uses of 62%.
The variability in the current C concentration was
determined as a function of the proportion of the fine
fraction (Fig. 2). No significant relationship between
the current C concentration of the fine fraction and the
proportion of the fine fraction was determined for soils
under agricultural use soils and soils under other land
uses. In contrast, forest sites showed increasing
current C contents of the fine fraction with increasing
proportions of the fine fraction. However, a generally
high variability in under- and oversaturated sites was
found. For sites which were apparently oversaturated
(current C concentration exceeded the potential C satu-
ration), no clear relationship with the contribution of
the fine fraction was found except for cropland soils,
where oversaturated sites were restricted to fine frac-
tion contents below 40%.
The C sequestration potential as affected byenvironmental factors
The C sequestration potential, calculated as the differ-
ence between the current C concentration and the
potential C saturation, was considerably different
between the land uses (Table 2; Fig. 3). In cropland
soils, a low current C amount in the fine fraction of
1.3 kg m�2 compared with a potential C saturation of
3.0 kg m�2 resulted in a relatively high C sequestration
potential of 1.5 kg m�2. In grassland and forest soils,
current C amounts of 1.8 and 1.6 kg m�2 were close to
potential C saturation values of 2.4 and 1.7 kg m�2
respectively. Thus, grassland and forest soils had lower
C sequestration potentials of 0.6 and 0.1 kg m�2 respec-
tively. Soils under other uses showed a current C
amount of the fine fraction of 1.5 kg m�2 and a poten-
tial C saturation of 2.6 kg m�2, resulting in a C seques-
tration potential of 0.9 kg m�2. The C saturation deficit
of soils under agricultural use was positively correlated
Table 2 Monte Carlo simulation of the current C concentra-
tion of the fine fraction <20 lm (Ccur) and the C saturation
deficit (Csat-def) in topsoils (0–10 cm) under cropland (C),
grassland (G), forest (F) and other uses (O) derived from 95
locations with measured values for the current C concentra-
tion of the fraction >20 lm
C
n = 115
G
n = 110
F
n = 249
O
n = 42
Ccur
(mg g�1)
Minimum 4.4 4.5 2.0 3.1
Maximum 24.4 57.4 156.6 101.7
Mean 10.8 16.3 22.5 23.2
Median 9.6 14.9 18.5 10.6
Standard
deviation
4.0 8.7 17.9 25.9
Skewness 0.9 1.7 3.5 1.9
Kurtosis 0.4 4.4 19.8 2.7
Csat-def
(mg g�1)
Minimum �7.6 �35.6 �124.4 �75.4
Maximum 22.5 27.3 25.4 24.8
Mean 10.3 5.0 �2.1 �1.7
Median 11.1 5.0 1.1 4.5
Standard
deviation
6.4 9.6 16.6 23.3
Skewness 0.6 0.7 �3.3 �1.7
Kurtosis 0.1 2.3 19.6 2.9
Fig. 1 Relative proportion of the OC content of the fraction
<20 lm to the total OC content of the bulk soil for cropland (C),
grassland (G) and forest (F) soils of Bavaria.
© 2013 John Wiley & Sons Ltd, Global Change Biology, 20, 653–665
SOC SATURATION AND SEQUESTRATION POTENTIAL 657
with the proportion of the fine fraction, particularly
in cropland soils (Fig. 4). In contrast, forest soils and
soils under other land uses showed no significant
relationship with the fine fraction content. To gain
insight into the factors that control the C sequestration
potential, correlations between several environmental
Fig. 2 Correlation between the proportion of particles <20 lm and the current C concentration of this soil fraction (Ccur) for cropland (C),
grassland (G), forest (F) and other land uses (O). The black line indicates the potential C saturation (Csat-pot) according to Hassink (1997).
Fig. 3 Current C concentration of the fine fraction <20 lm (Ccur) derived by a Monte Carlo simulation of 95 locations as well as poten-
tial C saturation (Csat-pot) and the C sequestration potential (Cseq) of topsoils (0–10 cm) under cropland (C), grassland (G), forest (F) and
other uses (O). Lines within the boxes give the median, boxes represent the 25th and 75th percentile, whiskers show the lowest and
highest values excluding outliers, circles represent outliers (1.5–3.0 interquartile range), triangles represent extremes (more than 3.0
interquartile ranges).
© 2013 John Wiley & Sons Ltd, Global Change Biology, 20, 653–665
658 M. WIESMEIER et al.
parameters and the C saturation deficit were examined
(Table 3). Strong positive correlations (P < 0.01) were
found with mean annual temperature and pH and
strong negative correlations (P < 0.01) with annual pre-
cipitation, elevation, slope and soil class. However, a
multiple linear regression analysis that included two
factors derived from a PCA (Table 4) revealed that the
C saturation deficit was strongly controlled by one fac-
tor that showed high loadings of temperature, precipi-
tation and elevation.
Fig. 4 Correlation between proportion of particles <20 lm and the C saturation deficit (Csat-def) for cropland (C), grassland (G), forest
(F) and other land uses (O).
Table 3 Correlation matrix of the C saturation deficit (Csat-def) and different pedogenetic, topographical and environmental
parameters
Csat-def Temp. Prec. Elev. Slope Curv. CA TWI PM Soil C. pH
Temp. 0.418**
Prec. �0.354** �0.840**
Elev. �0.420** �0.940** 0.907**
Slope �0.130** �0.484** 0.534** 0.472**
Curv. 0.065 0.126** �0.144** �0.151** �0.106*
CA �0.056 0.032 �0.029 �0.032 �0.044 �0.011
TWI 0.066 0.372** �0.290** �0.316** �0.702** 0.000 0.312**
PM �0.117** �0.333** 0.531** 0.474** 0.151** �0.117** 0.034 0.074
Soil C. �0.254** �0.208** 0.144** 0.176** 0.116** �0.009 �0.023 �0.095* �0.023
pH 0.367** 0.295** �0.149** �0.241** �0.119** �0.045 0.063 0.213** 0.051 �0.310**
SC �0.014 �0.189** 0.052 0.088* 0.193** 0.025 �0.025 �0.189** �0.059 0.111* �0.049
Level of significance: *P ≤ 0.05; **P ≤ 0.01.
Csat-def, C saturation deficit (mg g�1); Temp., mean annual temperature (°C); Prec., annual precipitation (mm); Elev., elevation (m
a.s.l.); Curv., curvature; CA, contributing area; TWI, topographic wetness index; PM, parent material; Soil C., soil class according to
AD-HOC AG Boden (2005); SC, stone content.
© 2013 John Wiley & Sons Ltd, Global Change Biology, 20, 653–665
SOC SATURATION AND SEQUESTRATION POTENTIAL 659
Discussion
The C saturation deficit of agricultural soils
The estimation of the C saturation deficit in soils of
Bavaria revealed that it was dependent on the current
C content of the fine fraction. The potential C saturation
as a function of the proportion of particles <20 lmaccording to Hassink (1997) was almost identical
among the main land uses due to their having similar
proportions of this fraction (Table 1). The current C
concentration was distinctly different between agricul-
tural and forest soils, pointing towards a clear impact
of cultivation (Table 2; Fig. 3). Remarkably, in cropland
soils the current C content of the fine fraction was on a
constant median level of around 10 mg g�1 indepen-
dent from the fine fraction content (Fig. 2). Fine-
textured cropland soils with a high potential to stabilize
SOM showed almost the same C contents as sand-
dominated soils that were close to the potential C satu-
ration or even exceeded it. This probably indicates that
the investigated cropland soils, most of which were
under long-term cultivation, had reached a constant
equilibrium level of mineral-associated SOM that is not
(or no longer) related to the proportion of the fine frac-
tion. The fact that the relatively low amount of SOC
associated with the fine fraction in coarse-textured
cropland soils was only marginally depleted probably
reflects a more careful management of these soils in
terms of SOM supply. A study that estimated the C sat-
uration deficit of agricultural topsoils in France also
determined a high degree of C saturation in very sandy
soils (Angers et al., 2011). In contrast, fine-textured
cropland soils showed a clear depletion of silt- and
clay-associated SOC. This loss can be ascribed to the
effect of tillage that causes destruction of soil macroag-
gregates and leads to a subsequent mineralization of the
released OM (Mann, 1986; Balesdent et al., 2000; Post &
Kwon, 2000; Six et al., 2000). On the other hand, the
physical stabilization of OM is reduced due to a deterio-
ration of microaggregate formation (Six et al., 1999). Fur-
thermore, the input of OM is reduced due to removal of
crop residues with the harvest, resulting in relatively
low SOC concentrations in the topsoils of cultivated
soils (Table 1). However, analyses of a comprehensive
data set of cropland soils in Bavaria revealed that the
depletion of SOC in the uppermost 10 cm, as it was
determined in this study, was compensated by a translo-
cation of SOC with depth due to deepening of the
plough layer to 30 cm (Wiesmeier et al., 2012, 2013a).
Compared with the potential C saturation, cropland
soils of Bavaria lost more than 50% of silt- and clay-
associated SOC in the uppermost 10 cm, resulting in a
median C saturation deficit of 11.1 mg g�1. This is in
line with several studies that investigated C saturation
of cropland soils. A comprehensive study of agricul-
tural topsoils in France revealed a slightly lower med-
ian C saturation deficit of 8.1 mg g�1 (Angers et al.,
2011). However, only a very rough estimation of the
current C amount of the fine fraction (85% of bulk SOC)
was applied. Long-term cultivated temperate cropland
soils in China showed a silt- and clay-associated C
saturation of around 50% (Zhao et al., 2006). For agri-
cultural experimental sites in eastern Canada, coarse-
textured topsoils with silt and clay contents <40% were
nearly saturated, whereas fine-textured soils (>60% silt
and clay) had a C saturation of 65–70% (Carter et al.,
2003). In cropland soils of Australia, a lower C
saturation of around 35% was determined, which was
related to low precipitation and high temperatures
(Hassink, 1997; Chan, 2001). In contrast, agricultural
soils in Tasmania under a cool, temperate climate were
close to or even exceeded the potential C saturation
(Sparrow et al., 2006).
Grassland soils showed a level of potential C satura-
tion similar to that of cropland soils but a considerably
higher current C concentration of the fine fraction
(Tables 1 and 2; Fig. 3). Although the current C concen-
tration was also not significantly related to the propor-
tion of silt and clay particles, a higher median value of
14.9 mg g�1 and generally a higher variability was
detected. This is presumably attributed to a much
lower intensity of the land use and a higher above- and
belowground input of OM into grassland soils. More-
over, the higher variability in the current C concentra-
tion of grassland soils is related to a wider range of
environmental conditions compared with cropland
sites and also includes areas with a cool, humid climate
or water-affected soils, such as fens, where a high pro-
portion of SOC is not associated with the fine fraction
(Fig. 1). Due to the higher current C contents of the fine
fraction, the C saturation of 73% is distinctly higher
Table 4 Rotated component matrix derived from a principal
component analysis of variables controlling the C saturation
deficit in soils of Bavaria
Factor 1 Factor 2
Temperature �0.915 �0.088
Precipitation 0.905 0.281
Elevation 0.928 0.229
Slope 0.702 �0.310
Curvature �0.178 �0.252
Contributing area �0.108 0.417
Topographic wetness index �0.548 0.628
Parent material 0.458 0.620
Soil class 0.264 �0.285
pH value �0.322 0.367
© 2013 John Wiley & Sons Ltd, Global Change Biology, 20, 653–665
660 M. WIESMEIER et al.
compared with cropland soils. This is in the range
reported for pastures in the south-eastern United
States, where a silt- and clay-associated C saturation of
60% was determined (Conant et al., 2003). A higher C
saturation in pastures compared with cropland was
also detected in Australia, though on a lower C satura-
tion level (Chan, 2001).
In summary, both cropland and grassland soils in
Bavaria have a substantial potential to sequester addi-
tional amounts of C in a stable form. A positive rela-
tionship of the C saturation deficit with the silt and clay
content (Fig. 4) revealed that soils that are particularly
fine textured have a large potential for C sequestration.
C saturation in forest soils and limitations of Hassink’sequation
In forest soils, a distinctly different level of C saturation
in the fine fraction was observed as compared with
agricultural soils. The current C concentration
increased slightly with the proportion of particles
<20 lm. However, there was a high variability in the
current C concentration in forest soils, with a high
proportion of sites having current C contents above the
potential C saturation and apparently being oversatu-
rated (Fig. 2). As the median current C concentration of
the fine fraction was close to the potential C saturation,
forest soils showed a high C saturation of 93% and a
resulting small C saturation deficit of only 1.1 mg g�1.
However, more than 60% of the total SOC of
investigated forest soils was not associated with the silt
and clay fraction (Fig. 1) and the C saturation deficit
showed no significant relationship with the proportion
of silt and clay (Fig. 4). Moreover, a high number of
forest sites showed a marked degree of apparent over-
saturation. Presumably, C saturation in forest soils is
not strongly related to the proportion of silt and clay as
was found for agricultural soils and thus, Hassink’s
equation provides no reliable estimation of the C satu-
ration in most forest soils. In acidic forest topsoils with
a median pH value of 3.5 (Table 1), other C stabiliza-
tion mechanisms are probably more relevant than an
association with clay minerals.
In several studies, the importance of amorphous iron
(Fe) and aluminium (Al) oxides for the stabilization of
OM in acidic soils was emphasized (Kaiser & Zech,
2000; Kaiser et al., 2002; Kleber et al., 2005; Sch€oning
et al., 2005; Wiseman & Puttmann, 2005; K€ogel-Knabner
et al., 2008; Spielvogel et al., 2008; Dumig et al., 2011).
Reactive surfaces of Fe and Al oxides may have a stron-
ger impact on stabilization of SOM in these soils than
clay minerals. Therefore, Kleber et al. (2005) and
Wiseman & Puttmann (2005) suggested that the C
sequestration potential of acidic soils could be related
to the content of poorly crystalline mineral phases. In a
study that investigated the relationship between the C
content of silt and clay particles and the proportion of
silt and clay particles in forest topsoils, Six et al. (2002)
assumed that Fe and Al oxides could play an important
role in explaining differences between cropland and
forest soils. However, the association of SOM with oxi-
des was predominantly related to subsoils (Kaiser et al.,
2002; Kleber et al., 2005; Spielvogel et al., 2008; Rumpel
& K€ogel-Knabner, 2011). In topsoils, preservation due
to recalcitrance of aliphatic forest OM might further be
a more relevant C stabilization mechanism (Eusterhues
et al., 2005; von Lutzow et al., 2006). Moreover, Baldock
& Skjemstad (2000) pointed out that Hassink’s equation
excluded the potential OM that may be associated with
the <20 lm fraction due to physical protection within
soil aggregates. These C stabilization mechanisms could
be the basis for an alternative method to estimate the C
sequestration potential in acidic soils using soil proper-
ties which are associated with these processes (e.g. reac-
tive surface of oxides or content of recalcitrant
compounds). An alternative method to Hassink’s equa-
tion could not only be used for acidic forest soils but
also for coarse-textured agricultural soils, where satu-
rated or apparently oversaturated conditions also indi-
cated insufficient detection of the C stabilization
processes (Fig. 2). A promising approach to estimate
the capacity of soils to preserve C by its association with
silt and clay more precisely was proposed by Six et al.
(2002), who developed specific regression equations for
cultivated grassland and forest soils as well as for dif-
ferent types of clay minerals. Moreover, Feng et al.
(2013) suggested an organic C loading method based on
the measurement of soil mineral-specific surface area as
well as a boundary line analysis using only data from
soils that have reached the maximal organic C stabiliza-
tion (upper tenth percentile) as alternative methods to
Hassink’s linear regression. The suitability of these
methods to estimate the C sequestration potential of
different soils should be tested in further studies.
Estimation of the soil C sequestration potential in Bavaria
A PCA and a multiple linear regression of the extracted
factors revealed that the C saturation deficit was mainly
controlled by climate (Tables 3 and 4). Therefore, the C
saturation deficit of soils under different land uses was
correlated with temperature and precipitation. Positive,
significant (P < 0.01) correlations with temperature and
negative correlations with precipitation were found for
all land uses (Figs 5 and 6). These relationships were
used for a rough estimation of the total C sequestration
potential of Bavarian soils. A similar approach was
used to estimate the C sequestration potential of
© 2013 John Wiley & Sons Ltd, Global Change Biology, 20, 653–665
SOC SATURATION AND SEQUESTRATION POTENTIAL 661
degraded grasslands worldwide (Conant & Paustian,
2002). For each land use, thresholds of temperature and
precipitation were derived (point of intersection of the
regression line at a C saturation deficit of 0) that divide
the areas of the respective land uses into regions with
saturated and unsaturated conditions (Table 5). The C
sequestration potential for each land use was estimated
by multiplying the area of unsaturated conditions by
the median value of the C saturation deficit of this area.
A regionalization of the C sequestration potential using
geostatistical methods was not conducted as the C satu-
ration deficit of the investigated locations was calcu-
lated using only an estimation of the current C
saturation of the fine fraction, which was based on a
smaller data set.
The results revealed that cropland and grassland
soils of Bavaria could potentially sequester 32 and 6 Mt
C in the uppermost 10 cm respectively. The high poten-
tial of cropland soils is related to the high C saturation
deficit of intensively cultivated soils and a large area
with unsaturated conditions. Less than 1% of the total
cropland area of Bavaria was assigned to saturated
conditions as cultivation is not feasible in cool, humid
areas. For forest soils, a C sequestration potential of
only 4 Mt was estimated with a high uncertainty as
previously explained. Other land uses have a potential
to sequester 9 Mt C. The low potential of forest soils to
sequester C can be ascribed to almost saturated condi-
tions in forest soils and the fact that only half of the
total forest area was associated with unsaturated condi-
tions. About 50% of Bavarian forests are located in
regions where cool, humid conditions result in a
complete C saturation of silt and clay particles.
The C sequestration potential estimated for the first
10 cm of the soil was extrapolated to the median depth
of A horizons of each land use, assuming that soil
texture and SOC contents are comparable within the A
horizon. The C sequestration potential of A horizons
under cropland, grassland, forest and other uses was
estimated to be 96, 12, 4 and 22 Mt respectively. In total,
soils of Bavaria could additionally sequester 134 Mt C,
18% of total SOC stocks of 764 Mt. This amount corre-
sponds to 490 Mt CO2-equivalents (CO2-eq.), which is
more than five times higher than the annual greenhouse
gas emission (in 2009) in Bavaria of 94 Mt CO2-eq.
(UGRdL, 2012). The majority, 395 Mt CO2-eq.
Fig. 5 Correlation between mean annual temperature and the C saturation deficit (Csat-def) for cropland (C), grassland (G), forest (F)
and other land uses (O). Values above the dashed line refer to a deficit of C saturation, values below the dashed line refer to an oversat-
uration of C.
© 2013 John Wiley & Sons Ltd, Global Change Biology, 20, 653–665
662 M. WIESMEIER et al.
(approximately 80%), could be sequestered in agricul-
tural soils. An increase in C saturation in forest soils
and soils under other land uses is associated with high
uncertainty. Assuming that due to an improved man-
agement of cultivated soils the theoretical stable C stor-
age potential is reached after a mean period of 30 years
(West & Six, 2007), a mean annual amount of 13 Mt
CO2-eq. could be sequestered in Bavarian soils over this
period of time, which is 14% of the annual emission of
greenhouse gases in Bavaria (in 2009). On an area basis,
4.1 t CO2-eq. ha�1 yr�1 could potentially be seques-
tered in agricultural soils, which is considerably higher
than observed and modelled C accumulation rates
through various management options aimed at increas-
ing SOC stocks in cultivated soils (Vleeshouwers &
Verhagen, 2002; West & Post, 2002; Freibauer et al.,
2004; Smith et al., 2008).
Outlook
A comparison of the current C amount with the poten-
tial C saturation of silt and clay particles according to
Hassink (1997) revealed high C sequestration potential
of agricultural topsoils in Bavaria. Although there are
some large uncertainties regarding the efficiency and
practicability of proposed management options to
Fig. 6 Correlation between annual precipitation and the C saturation deficit (Csat-def) for cropland (C), grassland (G), forest (F) and
other land uses (O). Values above the dashed line refer to a deficit of C saturation, values below the dashed line refer to an oversatura-
tion of C.
Table 5 Threshold of mean annual temperature (MATunsat) and annual precipitation (MAPunsat) for unsaturated soils, area of satu-
rated (Areasat) and unsaturated (Areaunsat) soils, C sequestration potential to a depth of 10 cm (Cseq-0-10) and extrapolated for the A
horizon (Cseq-A) for different land uses within Bavaria
MATunsat (°C) MAPunsat (mm) Areasat (km2) Areaunsat (km
2) Cseq (t ha�1) Cseq-0-10 (Mt) Cseq-A (Mt)
Cropland (C) >6.4 <1450 105 22817 13.9 32 96
Grassland (G) >7.0 <1150 458 9267 6.7 6 12
Forest (F) >8.1 <850 11561 11699 3.1 4 4
Other use (O) >8.0 <1000 3104 11312 7.9 9 22
Total 15228 55095 50 134
© 2013 John Wiley & Sons Ltd, Global Change Biology, 20, 653–665
SOC SATURATION AND SEQUESTRATION POTENTIAL 663
increase SOC stocks, the estimated high potential of
agricultural soils for C sequestration justifies optimized
SOM management of cultivated soils. One has to bear
in mind that besides the stable C sequestration in the
fine fraction, a significant additional amount of labile
SOC will also be sequestered as a result of improved
agricultural management. Furthermore, it is important
to note that there are benefits associated with C seques-
tration beyond CO2 mitigation because increased SOM
is associated with improved soil fertility, soil structure,
water holding capacity and thus a higher productivity.
Further important aspects are reduced risk of soil
erosion, decreased eutrophication and water contami-
nation as well as reduced costs for fossil fuel and fertil-
izer inputs (Paustian et al., 1998; Lal, 2007). Further
studies are needed to connect the estimated C seques-
tration potential of Bavarian soils with the economical
and political feasibility of agricultural practices aimed
at increasing SOC stocks. Such considerations should
include not only the possible range of CO2 mitigation,
but also additional benefits of SOM increases such as
improved soil fertility and productivity.
Acknowledgements
We thank Alfred Schubert from the Bavarian State Institute forForestry for providing forest soil data. Ulrike Maul, NadineEheim, Wiebke Wehrmann and Sigrid Hiesch are acknowledgedfor laboratory work. We are grateful to the Bavarian State Minis-try of the Environment and Public Health for funding theproject ‘Der Humusk€orper bayerischer B€oden im Klimawandel– Auswirkungen und Potentiale’.
References
AD-HOC AG Boden (2005) Bodenkundliche Kartieranleitung. Bundesanstalt fur Geowis-
senschaften und Rohstoffe (Ed.). E. Schweizerbart’sche Verlagsbuchhandlung, Stutt-
gart.
Amelung W, Zech W (1999) Minimisation of organic matter disruption during parti-
cle-size fractionation of grassland epipedons. Geoderma, 92, 73–85.
Angers DA, Arrouays D, Saby NPA, Walter C (2011) Estimating and mapping the
carbon saturation deficit of French agricultural topsoils. Soil Use and Management,
27, 448–452.
Arrouays D, Saby N, Walter C, Lemercier B, Schvartz C (2006) Relationships between
particle-size distribution and organic carbon in French arable topsoils. Soil Use and
Management, 22, 48–51.
Baldock JA, Skjemstad JO (2000) Role of the soil matrix and minerals in protecting nat-
ural organic materials against biological attack. Organic Geochemistry, 31, 697–710.
Balesdent J, Chenu C, Balabane M (2000) Relationship of soil organic matter dynamics
to physical protection and tillage. Soil & Tillage Research, 53, 215–230.
Beven KJ, Kirkby MJ (1978) A physically based, variable contributing area model of
basin hydrology. Hydrological Sciences Bulletin, 24, 43–69.
Carroll M, Milakovsky B, Finkral A, Evans A, Ashton MS (2012) Managing carbon
sequestration and storage in temperate and boreal forests. In: Managing Forest
Carbon in a Changing Climate (eds Ashton MS, Tyrrell ML, Spalding D, Gentry B),
pp. 205–226. Springer, New York.
Carter MR, Angers DA, Gregorich EG, Bolinder MA (2003) Characterizing organic
matter retention for surface soils in eastern Canada using density and particle size
fractions. Canadian Journal of Soil Science, 83, 11–23.
Chan KY (2001) Soil particulate organic carbon under different land use and manage-
ment. Soil Use and Management, 17, 217–221.
Chung HG, Grove JH, Six J (2008) Indications for soil carbon saturation in a temperate
agroecosystem. Soil Science Society of America Journal, 72, 1132–1139.
Ciais P, Schelhaas MJ, Zaehle S et al. (2008) Carbon accumulation in European forests.
Nature Geoscience, 1, 425–429.
Cole CV, Duxbury J, Freney J et al. (1997) Global estimates of potential mitigation of
greenhouse gas emissions by agriculture. Nutrient Cycling in Agroecosystems, 49,
221–228.
Conant RT, Paustian K (2002) Potential soil carbon sequestration in overgrazed grass-
land ecosystems. Global Biogeochemical Cycles, 16, 9.
Conant RT, Six J, Paustian K (2003) Land use effects on soil carbon fractions in the
southeastern United States. I. Management-intensive versus extensive grazing.
Biology and Fertility of Soils, 38, 386–392.
Dumig A, Smittenberg R, K€ogel-Knabner I (2011) Concurrent evolution of organic
and mineral components during initial soil development after retreat of the
Damma glacier, Switzerland. Geoderma, 163, 83–94.
Eusterhues K, Rumpel C, K€ogel-Knabner I (2005) Stabilization of soil organic matter
isolated via oxidative degradation. Organic Geochemistry, 36, 1567–1575.
Feng WT, Plante AF, Six J (2013) Improving estimates of maximal organic carbon
stabilization by fine soil particles. Biogeochemistry, 112, 81–93.
Freibauer A, Rounsevell MDA, Smith P, Verhagen J (2004) Carbon sequestration in
the agricultural soils of Europe. Geoderma, 122, 1–23.
Goh KM (2004) Carbon sequestration and stabilization in soils: implications for soil
productivity and climate change. Soil Science and Plant Nutrition, 50, 467–476.
Goodale CL, Apps MJ, Birdsey RA et al. (2002) Forest carbon sinks in the Northern
Hemisphere. Ecological Applications, 12, 891–899.
Hartge KH, Horn R (1989) Die physikalische Untersuchung von B€oden. Enke Verlag,
Stuttgart.
Hassink J (1997) The capacity of soils to preserve organic C and N by their association
with clay and silt particles. Plant and Soil, 191, 77–87.
Holland JM (2004) The environmental consequences of adopting conservation tillage
in Europe: reviewing the evidence. Agriculture Ecosystems & Environment, 103,
1–25.
IUSS Working Group WRB (2006) World Reference Base for Soil Resources 2006. World
Soil Resources Reports No. 103, FAO, Rome.
Jandl R, Lindner M, Vesterdal L et al. (2007) How strongly can forest management
influence soil carbon sequestration? Geoderma, 137, 253–268.
Johnson JMF, Franzluebbers AJ, Weyers SL, Reicosky DC (2007) Agricultural oppor-
tunities to mitigate greenhouse gas emissions. Environmental Pollution, 150,
107–124.
Joneck M, Hangen E, Martin W et al. (2006) Wissenschaftliche Grundlagen fur den
Vollzug der Bodenschutzgesetze in Bayern (GRABEN) - ein Projekt stellt sich vor.
Bodenschutz, 2, 32–38.
Kaiser K, Zech W (2000) Dissolved organic matter sorption by mineral constituents of
subsoil clay fractions. Journal of Plant Nutrition and Soil Science, 163, 531–535.
Kaiser K, Eusterhues K, Rumpel C, Guggenberger G, K€ogel-Knabner I (2002) Stabil-
ization of organic matter by soil minerals - investigations of density and particle-
size fractions from two acid forest soils. Journal of Plant Nutrition and Soil Science,
165, 451–459.
Karjalainen T, Pussinen A, Liski J, Nabuurs GJ, Eggers T, Lapvetelainen T, Kaipainen
T (2003) Scenario analysis of the impacts of forest management and climate change
on the European forest sector carbon budget. Forest Policy and Economics, 5,
141–155.
Kleber M, Mikutta R, Torn MS, Jahn R (2005) Poorly crystalline mineral phases
protect organic matter in acid subsoil horizons. European Journal of Soil Science, 56,
717–725.
K€ogel-Knabner I, Guggenberger G, Kleber M et al. (2008) Organo-mineral associa-
tions in temperate soils: integrating biology, mineralogy, and organic matter chem-
istry. Journal of Plant Nutrition and Soil Science, 171, 61–82.
Lal R (2004) Agricultural activities and the global carbon cycle. Nutrient Cycling in
Agroecosystems, 70, 103–116.
Lal R (2005) Forest soils and carbon sequestration. Forest Ecology and Management, 220,
242–258.
Lal R (2007) Carbon management in agricultural soils. Mitigation and Adaption Strate-
gies for Global Change, 12, 303–322.
Landres PB, Morgan P, Swanson FJ (1999) Overview of the use of natural variability
concepts in managing ecological systems. Ecological Applications, 9, 1179–1188.
Larocque GR, Bhatti JS, Boutin R, Chertov O (2008) Uncertainty analysis in carbon
cycle models of forest ecosystems: research needs and development of a theoreti-
cal framework to estimate error propagation. Ecological Modelling, 219, 400–412.
Liski J, Perruchoud D, Karjalainen T (2002) Increasing carbon stocks in the forest soils
of western Europe. Forest Ecology and Management, 169, 159–175.
© 2013 John Wiley & Sons Ltd, Global Change Biology, 20, 653–665
664 M. WIESMEIER et al.
Lorenz K, Lal R (2010) Carbon Sequestration in Forest Ecosystems. Springer, New York.
von Lutzow M, K€ogel-Knabner I, Ekschmitt K, Matzner E, Guggenberger G, Marsch-
ner B, Flessa H (2006) Stabilization of organic matter in temperate soils: mecha-
nisms and their relevance under different soil conditions - a review. European
Journal of Soil Science, 57, 426–445.
Luyssaert S, Ciais P, Piao SL et al. (2010) The European carbon balance. Part 3: forests.
Global Change Biology, 16, 1429–1450.
Mann LK (1986) Changes in soil carbon storage after cultivation. Soil Science, 142,
279–288.
Oades JM (1988) The retention of organic matter in soils. Biogeochemistry, 5, 35–70.
Paustian K, Andren O, Janzen HH et al. (1997) Agricultural soils as a sink to mitigate
CO2 emissions. Soil Use and Management, 13, 230–244.
Paustian K, Cole CV, Sauerbeck D, Sampson N (1998) CO2 mitigation by agriculture:
an overview. Climatic Change, 40, 135–162.
Paustian K, Six J, Elliott ET, Hunt HW (2000) Management options for reducing CO2
emissions from agricultural soils. Biogeochemistry, 48, 147–163.
Post WM, Kwon KC (2000) Soil carbon sequestration and land-use change: processes
and potential. Global Change Biology, 6, 317–327.
Rumpel C, K€ogel-Knabner I (2011) Deep soil organic matter-a key but poorly under-
stood component of terrestrial C cycle. Plant and Soil, 338, 143–158.
Sauerbeck DR (2001) CO2 emissions and C sequestration by agriculture - perspectives
and limitations. Nutrient Cycling in Agroecosystems, 60, 253–266.
Sch€oning I, Knicker H, Kogel-Knabner I (2005) Intimate association between O/N-
alkyl carbon and iron oxides in clay fractions of forest soils. Organic Geochemistry,
36, 1378–1390.
Schubert A (2002) Bayerische Waldboden-Dauerbeobachtungsfl€achen - Bodenuntersuchun-
gen. Wissenschaftszentrum Weihenstephan fur Ern€ahrung, Landnutzung und
Umwelt der Technischen Universit€at Munchen und Bayerische Landesanstalt fur
Wald und Forstwirtschaft, Freising.
Six J, Elliott ET, Paustian K (1999) Aggregate and soil organic matter dynamics under
conventional and no-tillage systems. Soil Science Society of America Journal, 63,
1350–1358.
Six J, Elliott ET, Paustian K (2000) Soil macroaggregate turnover and microaggregate
formation: a mechanism for C sequestration under no-tillage agriculture. Soil Biol-
ogy & Biochemistry, 32, 2099–2103.
Six J, Conant RT, Paul EA, Paustian K (2002) Stabilization mechanisms of soil organic
matter: implications for C-saturation of soils. Plant and Soil, 241, 155–176.
Smith P (2004) Carbon sequestration in croplands: the potential in Europe and the
global context. European Journal of Agronomy, 20, 229–236.
Smith P (2012) Agricultural greenhouse gas mitigation potential globally, in Europe
and in the UK: what have we learnt in the last 20 years? Global Change Biology, 18,
35–43.
Smith P, Martino D, Cai Z et al. (2008) Greenhouse gas mitigation in agriculture.
Philosophical Transactions of the Royal Society B-Biological Sciences, 363, 789–813.
Sollins P, Homann P, Caldwell BA (1996) Stabilization and destabilization of soil
organic matter: mechanisms and controls. Geoderma, 74, 65–105.
Sorensen R, Zinko U, Seibert J (2006) On the calculation of the topographic wetness
index: evaluation of different methods based on field observations. Hydrology and
Earth System Sciences, 10, 101–112.
Sparrow LA, Belbin KC, Doyle RB (2006) Organic carbon in the silt plus clay fraction
of Tasmanian soils. Soil Use and Management, 22, 219–220.
Spielvogel S, Prietzel J, K€ogel-Knabner I (2006) Soil organic matter changes in a
spruce ecosystem 25 years after disturbance. Soil Science Society of America Journal,
70, 2130–2145.
Spielvogel S, Prietzel J, K€ogel-Knabner I (2008) Soil organic matter stabilization in
acidic forest soils is preferential and soil type-specific. European Journal of Soil
Science, 59, 674–692.
Sp€orlein P, Dilling J, Joneck M (2004) Pilot study to test the equivalence or compara-
bility of soil-particle-size analysis according to E DIN ISO 11277: 06.94 (pipette
method) and by the use of the sedigraph. Journal of Plant Nutrition and Soil Science,
167, 649–656.
Steffens M, K€olbl A, K€ogel-Knabner I (2009) Alteration of soil organic matter pools
and aggregation in semi-arid steppe topsoils as driven by organic matter input.
European Journal of Soil Science, 60, 198–212.
Stewart CE, Paustian K, Conant RT, Plante AF, Six J (2007) Soil carbon saturation:
concept, evidence and evaluation. Biogeochemistry, 86, 19–31.
Stewart CE, Paustian K, Conant RT, Plante AF, Six J (2008) Soil carbon saturation:
evaluation and corroboration by long-term incubations. Soil Biology & Biochemistry,
40, 1741–1750.
Stockmann U, Adams MA, Crawford JW et al. (2013) The knowns, known unknowns
and unknowns of sequestration of soil organic carbon. Agriculture, Ecosystems &
Environment, 164, 80–99.
UGRdL (2012) Umwelt€okonomische Gesamtrechnung der L€ander, Energieverbrauch und
Treibhausgasemissionen - Analysen und Ergebnisse. Arbeitskreis Umwelt€okonomische
Gesamtrechnung der L€ander im Auftrag der Statistischen €Amter der L€ander.
Available at: www.ugrdl.de (accessed 22 August 2013).
Vesterdal L, Elberling B, Christiansen JR, Callesen I, Schmidt IK (2012) Soil respira-
tion and rates of soil carbon turnover differ among six common European tree
species. Forest Ecology and Management, 264, 185–196.
Vleeshouwers LM, Verhagen A (2002) Carbon emission and sequestration by
agricultural land use: a model study for Europe. Global Change Biology, 8, 519–
530.
West TO, Post WM (2002) Soil organic carbon sequestration rates by tillage and crop
rotation: a global data analysis. Soil Science Society of America Journal, 66, 1930–1946.
West TO, Six J (2007) Considering the influence of sequestration duration and carbon
saturation on estimates of soil carbon capacity. Climatic Change, 80, 25–41.
Wiesmeier M, Sp€orlein P, Geuss U et al. (2012) Soil organic carbon stocks in southeast
Germany (Bavaria) as affected by land use, soil type and sampling depth. Global
Change Biology, 18, 2233–2245.
Wiesmeier M, Hubner R, Barthold FK et al. (2013a) Amount, distribution and driving
factors of soil organic carbon and nitrogen in cropland and grassland soils of
southeast Germany (Bavaria). Agriculture Ecosystems & Environment, 176, 39–52.
Wiesmeier M, Prietzel J, Barthold FK et al. (2013b) Storage and drivers of organic car-
bon in forest soils of southeast Germany (Bavaria) - Implications for carbon
sequestration. Forest Ecology and Management, 295, 162–172.
Wilson J, Gallant J (2000) Terrain Analysis: Principles and Applications. John Wiley &
Sons, Inc., New York.
Wiseman CLS, Puttmann W (2005) Soil organic carbon and its sorptive preservation
in central Germany. European Journal of Soil Science, 56, 65–76.
Zhao LP, Sun YJ, Zhang XP, Yang XM, Drury CF (2006) Soil organic carbon in clay
and silt sized particles in Chinese mollisols: relationship to the predicted capacity.
Geoderma, 132, 315–323.
© 2013 John Wiley & Sons Ltd, Global Change Biology, 20, 653–665
SOC SATURATION AND SEQUESTRATION POTENTIAL 665