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Wiesmeier et al. (2016). Projected loss of soil organic carbon in temperate agricultural soils in the 21st century: effects of climate change and carbon input trends

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Projected loss of soil organic carbon 

in temperate agricultural soils in 

the 21

st

 century: effects of climate 

change and carbon input trends

Martin Wiesmeier

1

, Christopher Poeplau

2

, Carlos A. Sierra

3

, Harald Maier

4

, Cathleen Frühauf

5

, 

Rico Hübner

6

, Anna Kühnel

1

, Peter Spörlein

7

, Uwe Geuß

7

, Edzard Hangen

7

, Bernd Schilling

7

, 

Margit von Lützow

1

 & Ingrid Kögel-Knabner

1

,

8

Climate change and stagnating crop yields may cause a decline of SOC stocks in agricultural soils 
leading to considerable CO

2

 emissions and reduced agricultural productivity. Regional model-based 

SOC projections are needed to evaluate these potential risks. In this study, we simulated the future 

SOC development in cropland and grassland soils of Bavaria in the 21

st

 century. Soils from 51 study sites 

representing the most important soil classes of Central Europe were fractionated and derived SOC pools 
were used to initialize the RothC soil carbon model. For each site, long-term C inputs were determined 

using the C allocation method. Model runs were performed for three different C input scenarios as 

a realistic range of projected yield development. Our modelling approach revealed substantial SOC 

decreases of 11–16% under an expected mean temperature increase of 3.3 °C assuming unchanged C 
inputs. For the scenario of 20% reduced C inputs, agricultural SOC stocks are projected to decline by 
19–24%. Remarkably, even the optimistic scenario of 20% increased C inputs led to SOC decreases of 
3–8%. Projected SOC changes largely differed among investigated soil classes. Our results indicated 
that C inputs have to increase by 29% to maintain present SOC stocks in agricultural soils.

Soil organic carbon (SOC) represents the largest carbon pool in terrestrial ecosystems and is a key factor that 

controls important soil functions, e.g. the productivity of agricultural soils

1

. The maintenance of SOC stocks in 

croplands and grasslands of the world is thus of upmost importance for ensuring global food security and the 

prevention of substantial CO

2

 emissions. Under long-term constant management and environmental conditions, 

agricultural SOC stocks are in a dynamic equilibrium between C inputs, mainly in form of crop residues and 

organic fertilizers, and a loss of C due to decomposition of soil organic matter (SOM). However, as the decompo-

sition of SOM is strongly controlled by temperature and soil moisture, climate change imposes the risk of SOC 

losses

2–6

. Both a the global and regional scale decreasing SOC stocks were observed along temperature gradients 

from colder to warmer regions

7,8

. This suggests that SOM decomposition rates change faster as a function of 

temperature than does net primary production (NPP). Accordingly, rising temperatures in the course of climate 

change are discussed to cause significant declines of SOC. In particular, agricultural soils could be affected, as 

observed stagnation of crop yields and associated stagnation of C inputs in the last decades may aggravate climate 

change-induced SOC losses

9,10

. First indications for declining SOC stocks in agricultural soils were already found, 

but due to changes of land use and the agricultural management, a relationship with climate change could not be 

confirmed to date

11–17

.

1

Chair of Soil Science, TUM School of Life Sciences Weihenstephan, Technical University of Munich, 85354 Freising, 

Germany. 

2

Thuenen Institute for Agricultural Climate Research, Bundesallee 50, 38106 Braunschweig, Germany. 

3

Max Planck Institute for Biogeochemistry, Hans-Knöll-Str. 10, 07745 Jena, Germany. 

4

Deutscher Wetterdienst, 

Abteilung Agrarmeteorologie, Niederlassung Weihenstephan, Alte Akademie 16, 85350 Freising-Weihenstephan, 

Germany. 

5

Deutscher Wetterdienst, Abteilung Agrarmeteorologie, ZAMF, Bundesallee 50, 38116 Braunschweig, 

Germany. 

6

Chair for Strategic Landscape Planning and Management, TUM School of Life Sciences Weihenstephan, 

Technical University of Munich, 85354 Freising, Germany. 

7

Bavarian Environment Agency, 95030 Hof, Germany. 

8

Institute for Advanced Study, Technical University of Munich, 85748 Garching, Germany. Correspondence and 

requests for materials should be addressed to M.W. (email: wiesmeier@wzw.tum.de)

Received: 28 April 2016

a

ccepted: 08 August 2016

P

ublished: 02 September 2016

OPEN

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The potential risk of SOC losses induced by climate change calls for a projection of agricultural SOC stocks 

under future climate change scenarios on the basis of SOC models. A frequently applied model is the Rothamsted 

Carbon Model (RothC) which was specifically designed to simulate SOC dynamics in temperate cropland and 

grassland soils

18

. Several modelling approaches were conducted to simulate regional or global development of 

agricultural SOC stocks

19–25

. However, these SOC projections were often based on simple estimations of impor-

tant input variables or legacy data with a relatively low spatial resolution. In particular, rough estimations for the 

C input were used, which is one of the most decisive parameters in SOC projections. As the determination of 

the total C input, particularly the belowground components such as roots and rhizodeposition, is difficult and 

elaborate, values are generally estimated using an inverse application of the RothC model or simply gross values 

from the literature

18,20,26,27

. However, large discrepancies were observed between measured and estimated C input 

values and thus an independent, more reliable approach is needed

28

. Future projections of the development of C 

inputs in agricultural soils are even more challenging due to overlapping implications of crop and land use man-

agement, future trend of crop breeding and technology and climate change. In several SOC modelling studies it 

was assumed that the development of agricultural C inputs are simply related to NPP development, which can 

be simulated by vegetation models

19–22,25,29–31

. However, such an approach neglects the fact that the C input in 

agricultural soils is controlled by various factors including management practices. Moreover, reliable SOC pro-

jections using the RothC model require a reasonable initialization of the model. In terms of model initialization, 

SOM pools derived from soil fractionation is a laborious, but probably the most suitable approach as this method 

reflects the actual state of SOC pools

27,32–34

.

In this study, we simulated the future development of SOC stocks of cropland and grassland soils in Bavaria 

from 2000 to 2095 under a moderate climate scenario of the IPCC (A1B) using a large number of investigated 

sites (Fig. 1). In total, 21 cropland and 30 grassland sites representing most important soil classes of Central 

Europe were sampled and SOC pools were fractionated following the approach of Zimmermann 

et al.

34

, who were 

able to empirically link SOC fractions to RothC pools. In a preliminary study, C inputs of major crops and grass-

land in Bavaria were determined with a high spatial and temporal resolution

35

, which enabled a calculation of 

site-specific C input values. The RothC model was then used to simulate SOC development under three different 

C input scenarios, which covered the range of expected crop yield development in Bavaria.

Results

Climate conditions at study sites and projected changes. 

The ensemble approach on the basis of 19 

climate models revealed reliable spatially differentiated projections of air temperature, precipitation and evapo-

transpiration for all studied sites (Table 1, Fig. 2). In general, cropland sites exhibited average long-term values 

(1971 to 1999) for mean annual temperature of 8.3 to 8.9 °C, for mean annual precipitation of 831 to 873 mm and 

for mean annual evapotranspiration of 707 to 760 mm. Between 2000 and 2095 climate projections under the 

A1B climate scenario revealed increases in air temperature by 3.1 to 3.4 °C, in precipitation by 35 to 82 mm and 

in evapotranspiration by 80 to 123 mm. Due to their main distribution in pre-alpine regions, grassland sites had 

slightly lower mean annual temperatures of 7.6 to 8.7 °C, considerably higher values of mean annual precipitation 

of 846 to 1331 mm and slightly lower mean annual evapotranspiration of 663 to 739 mm. The climate projections 

Figure 1.

 

Locations of study sites under cropland and grassland in Bavaria (the map was generated using 

ESRI ArcMap 9.2,

 

www.esri.com

).

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indicated for grassland sites a temperature increase of 3.2 to 3.4 °C, a variable change of precipitation of 

−

 36  to 

95 mm and an increase of evapotranspiration of 77 to 143 mm.

Basic soil properties, SOC fractions and C input. 

Basic soil properties and SOC fractions for each soil 

unit under cropland and grassland are given in Table 2. In general, cropland soils revealed higher thicknesses 

of the A horizon (29 

±

  6 cm) compared to grassland soils (23 

±

  6 cm). Clay contents ranged between 21 and 

52% in soil units under cropland and from 19 to 45% under grassland. The contribution of OC in different soil 

fractions to bulk SOC stocks differed strongly among the investigated soil/land use units. Generally, DOC con-

tributed with only 2 to 7% to total SOC stocks of both cropland and grassland topsoils. Considerable differences 

between soil units were found for the POM fraction that contained 5 to 21% of total SOC stocks in soils under 

cropland and 3 to 16% under grassland. The highest POM-OC amounts were found for Cambisols from Tertiary 

material (C1) as well as from sandstone (C6, C7). For the S

+

 A fraction distinct differences were found between 

cropland and grassland soil units. Under cropland, S

+

 A contained only 2 to 10% of bulk SOC with the excep-

tion of groundwater-affected soils (G) that stored 16%. In contrast, grassland soil units revealed much higher 

OC amounts in the S

+

 A fraction (16 to 34% of bulk SOC) with highest amounts in groundwater-affected soils  

(G, 57%). The major part of SOC was found in the s

+

 c fraction that contained 50 to 83% of total SOC stocks inde-

pendent from land use (only groundwater-affected soils under grassland revealed a considerably lower contribu-

tion of 30%). The inert rSOC fraction contained OC amounts of 3% to 10% of total SOC stocks without distinct 

differences between cropland and grassland soil units. In total, soil units under cropland revealed significantly 

(P 

<

  0.05) lower SOC stocks in A horizons (59 

±

  21 t ha

−

1

) than grassland soils (72 

±

  24 t ha

−

1

). The calculated 

mean C input values derived from the period from 1995 to 1999 showed no significant differences among soil 

units under cropland and grassland. The C inputs ranged between 3.4 and 4.1 t ha

−

1

 in cropland soils and from 4.5 

to 5.3 t ha

−

1

 in grassland soils. Over a 16 year period from 1995 to 2010, C inputs in cropland showed no signifi-

cant trend but remained on a constant level (Fig. 3). In grasslands soils, C inputs revealed a slight increase of 8% 

(based on a linear trend with an R

2

 value of 0.23).

Projected development of SOC stocks under different C input scenarios. 

The projected develop-

ment of SOC stocks of cropland and grassland soils under the reference scenario and different C input scenarios 

land 

use

soil 

class MAT (°C) MAP (mm)

Evap 

(mm)

∆

SOC C

0

  

(t ha

−

1

)

∆

SOC C

−

20%

 

(t ha

−

1

)

∆

SOC C

+

20%

  

(t ha

−

1

)

∆

MAT 

(°C)

∆

MAP 

(mm)

∆

Evap 

(mm)

C

G

8.9 

±

  0.2

862 

±

  63

747 

±

  9

−

 11.8 

±

  1.2

−

 17.1 

±

  1.3

−

 6.9 

±

  1.0

3.2 

±

  0.1

57 

±

  47

118 

±

  21

L2

8.9 

±

  0.2

831 

±

  20

747 

±

  28

−

 9.4 

±

  2.2

−

 14.0 

±

  2.9

−

 5.2 

±

  1.6

3.1 

±

  0.1

59 

±

  52

105 

±

  19

C1

8.9 

±

  0.1

834 

±

  55

760 

±

  11

−

 9.3 

±

  2.7

−

 13.9 

±

  3.6

−

 5.2 

±

  1.8

3.1 

±

  0.1

59 

±

  44

123 

±

  14

C3

8.3 

±

  0.1

873 

±

  79

713 

±

  12

−

 8.5 

±

  3.6

−

 13.4 

±

  4.4

−

 4.1 

±

  3.0

3.4 

±

  0

82 

±

  10

87 

±

  10

C4

8.8 

±

  0.1

845 

±

  70

754 

±

  17

−

 9.7 

±

  1.5

−

 15.0

±

 1.4

−

 4.8 

±

  1.7

3.2 

±

  0

50 

±

  8

113 

±

  1

C6

8.8 

±

  0.2

840 

±

  40

744 

±

  8

−

 8.6 

±

  1.7

−

 13.0 

±

  2.6

−

 4.4 

±

  1.0

3.3 

±

  0.1

51 

±

  11

114 

±

  31

C7

8.3 

±

  0.5

864 

±

  105

707 

±

  47

−

 4.8 

±

  4.6

−

 9.3 

±

  4.1

−

 0.7 

±

  5.0

3.3 

±

  0.1

62 

±

  5

87 

±

  28

V

8.4 

±

  0.2

831 

±

  31

710 

±

  25

−

 9.3 

±

  2.7

−

 14.3 

±

  3.4

−

 4.6 

±

  2.3

3.3 

±

  0

35 

±

  33

80 

±

  20

G

G

8.0 

±

  0.4

1112 

±

  260

683 

±

  50

−

 4.4 

±

  7.7

−

 10.6 

±

  7.3

1.3 

±

  8.1

3.3 

±

  0

−

 8 

±

  92

124 

±

  29

L1

7.6 

±

  0.2

1327 

±

  60

663 

±

  31

−

 1.0 

±

  5.3

−

 7.3 

±

  4.5

4.7 

±

  6.0

3.4 

±

  0

−

 27 

±

  12 143 

±

  10

L2

7.9 

±

  0.5

1129 

±

  273

686 

±

  50

−

 5.1 

±

  7.7

−

 11.6 

±

  7.6

0.8 

±

  7.8

3.4 

±

  0

2 

±

  83

128 

±

  33

C1

8.5 

±

  0.9

1019 

±

  231

731 

±

  48

−

 7.0 

±

  9.1

−

 13.4 

±

  9.1

−

 1.1 

±

  9.2

3.3 

±

  0.1

4 

±

  41

139 

±

  10

C2

7.9 

±

  0.7

1331 

±

  267

669 

±

  30

−

 4.0 

±

  6.4

−

 9.8 

±

  6.4

1.2 

±

  6.4

3.3 

±

  0.1

−

 36 

±

  22

142 

±

  3

C3

8.3 

±

  0.6

846 

±

  92

719 

±

  49

−

 11.4 

±

  5.6

−

 17.5 

±

  6.2

−

 5.7 

±

  5.1

3.3 

±

  0.1

60 

±

  26

94 

±

  21

C5

8.6 

±

  0.2

856 

±

  36

739 

±

  2

−

 11.7 

±

  1.6

−

 17.5 

±

  2.3

−

 6.3 

±

  0.9

3.2 

±

  0.1

95 

±

  16

105 

±

  14

C6

8.7 

±

  0.3

871 

±

  54

727 

±

  39

−

 9.3 

±

  1.0

−

 15.2 

±

  1.6

−

 3.8 

±

  0.6

3.3 

±

  0

28 

±

  33

112 

±

  35

C7

8.0 

±

  0.2

875 

±

  63

698 

±

  16

−

 10.8 

±

  2.3

−

 17.2 

±

  2.7

−

 4.8 

±

  2.0

3.4 

±

  0

71 

±

  12

77 

±

  6

V

8.2 

±

  0.3

906 

±

  41

708 

±

  24

−

 10.2 

±

  0.6

−

 16.5 

±

  0.8

−

 4.5 

±

  0.4

3.4 

±

  0

69 

±

  10

80 

±

  5

Table 1.

 

 Long-term (1971–1999) mean annual temperature (MAT), mean annual precipitation (MAP), 

mean annual evapotranspiration (Evap) projected SOC changes (2000–2095) under constant C input 

(

∆

SOC C

0

), decreased C input by 20% (

∆

SOC C

−

20%

) and increased C input by 20% (

∆

SOC C

+

20%

) as well as 

projected changes of MAT, MAP and Evap (2000–2095) of major soil units in Bavaria under cropland (C) 

and grassland (G) (mean values 

±

 standard deviation).

 G 

=

  groundwater-affected soils (Gleysols, Fluvisols); 

L1 

=

  shallow to intermediate soils with clay accumulation in the subsoil (Luvisols); L2 

=

  intermediate to deep 

soils with clay accumulation in the subsoil (Luvisols); C1 

=

  soils with well developed B horizons from Tertiary 

material (Cambisols); C2 

=

  soils with well developed B horizons from morainal material in places with clay 

accumulation in the subsoil (Cambisols, Luvisols); C3 

=

  shallow soils from limestone weathering with or 

without loess coverings (Cambisols, Luvisols, Leptosols); C4 

=

  intermediate to deep soils from limestone 

weathering with or without loess coverings (Cambisols, Luvisols); C5 

=

  soils with well developed B horizons 

from acidic material with low base saturation (Cambisols); C6 

=

  soils with well developed B horizons from 

sandstone with low base saturation (Cambisols); C7 

=

  soils with well developed B horizons from sandstone with 

initial podzolisation (Cambisols, Podzols); V 

=

  clay-rich soils (Cambisols, Vertisols, Stagnosols).

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are shown in Figs 4 and 5. In cropland soils, the reference climate scenario (constant average climate conditions 

1970–1999) showed an increase of mean SOC stocks from 58.5 

±

  20.8 t ha

−

1

 in 2000 to 66.3 

±

  13.9 t ha

−

1

 in 2095. 

The projection of SOC stocks under climate change and the constant C input scenario C0 indicated a slight 

increase of SOC stocks until 2020 up to 61.7 

±

  18.1 t ha

−

1

, followed by a slight decline down to 57.2 

±

  11.8 t ha

−

1

 

in 2095. Under climate change and C input scenario C20

−

 , a noticeable decline of SOC stocks to 52.4 

±

  11.2 t 

ha

−

1

 in 2095 was determined after a peak of 60.9 

±

  18.0 t ha

−

1

 in 2020. Climate change and the C input scenario 

C20

+

  revealed a slight SOC stock increase up to 61.6 

±

  12.4 t ha

−

1

 in 2095.

The calculated net SOC change as the difference between the reference scenario and climate/C input scenarios 

showed mean losses of SOC stocks of 

−

 9.1 t  ha

−

1

 (

−

 16% of initial SOC stocks) under climate change and C0, of 

−

 13.9 t  ha

−

1

 (

−

 24%) under climate change and C20

−

  and of 

−

 4.7 t  ha

−

1

 (

−

 8%) under climate change and C20

+

  

(Fig. 6). Distinct differences of SOC losses were found among the investigated soil units under cropland (Table 2). 

The highest relative SOC losses (compared to total SOC amount) were found for intermediate to deep soils with 

clay accumulation in the subsoil (L2) and Cambisols from Tertiary material (C1). For the scenarios C0, C20

−

   and 

C20

+

 , SOC stocks in these soil units decreased by 

−

 20  to 

−

 21%, 

−

 30  to 

−

 31%  and 

−

 11  to 

−

 12%,  respectively. 

For all other soil units under cropland the respective SOC decreases ranged between 

−

 12  to 

−

 17%, 

−

 19  to 

−

 26% 

and 

−

 2  to 

−

 9%.

For grassland sites, only a slight increase of mean SOC stocks from 71.6 

±

  24.0 t ha

−

1

 in 2000 to 74.9 

±

  13.7 t 

ha

−

1

 in 2095 was projected for the reference scenario (Fig. 5). The climate change and C0 scenario revealed a SOC 

stock decrease down to 67.4 

±

  10.6 t ha

−

1

 in 2095 after a peak of 73.4 

±

  17.5 t ha

−

1

 in 2020. SOC projections under 

climate change and C20

−

  showed a considerable decline down to 61.2 

±

  10.1 t ha

−

1

 in 2095. A slight increase 

of SOC stocks up to 73.1 

±

  10.9 t ha

−

1

 was found for climate change and the C20

+

  scenario. The resulting net 

SOC changes accounted for 

−

 7.5  t  ha

−

1

 for C0 (

−

 11% of initial SOC stocks), 

−

 13.7  t  ha

−

1

 for C20

−

   (

−

 19%) 

and 

−

 1.8  t  ha

−

1

 for C20

+

   (

−

 3%) (Fig. 6). Among different soil units under grassland, distinct differences of 

SOC changes were detected (Table 2). In general, soil units mainly distributed in the southern part of Bavaria 

(groundwater-affected soils, G; soils with clay accumulation in the subsoil L1, L2; Cambisols and Luvisols from 

moraine material, C2) showed considerably lower SOC decreases than other investigated soil units. For C input 

scenarios C0 and C20

−

 , relative SOC decreases ranged between 

−

 1  to 

−

 7%  and 

−

 9  to 

−

 16%, respectively. Under 

scenario C20

+

 , even SOC increases of 1 to 6% were projected for these soil units. In contrast, all other soil units 

under cropland revealed much higher declines of SOC of 

−

 11  to 

−

 23% for C0, 

−

 19  to 

−

 34% for C20

−

   and 

−

 2 

to 

−

 12% for C20

+

 .

Estimation of total SOC changes in agricultural soils of Bavaria. 

In order to estimate the total 

changes of SOC stocks for the investigated soil units on an area basis, projected mean SOC stocks from all studied 

Figure 2.

 

Projected mean annual temperature (MAT) and mean annual precipitation (MAP) for the study 

sites for the period 2000–2095 under the A1B scenario (median with interquartile range (IQR) derived 

from 19 climate models) 

.

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sites for a soil unit were multiplied with the respective area of the soil unit (Table 3, Fig. 7). In general, cropland 

and grassland topsoils of Bavaria stored 122.9 Mt and 192.9 Mt SOC in 2000. In croplands, the soil units interme-

diate to deep Luvisols (L2) and Cambisols from Tertiary material (C1) with the largest extend in Bavaria stored 

40% of total SOC stocks. Moreover, the soil units groundwater-affected soils (G), shallow soils from limestone 

weathering (C3) and clay-rich soils (V) contained 41% of total SOC stocks due to relatively high soil-unit-specific 

SOC stocks of 71.8 to 80.8 t ha

−

1

. Estimations of total projected SOC changes under the scenarios C0, C20

−

   and 

C20

+

  revealed SOC losses of 20.3 Mt, 30.9 Mt and 10.5 Mt for cropland soils, respectively. The highest absolute 

SOC losses were found for intermediate to deep Luvisols (L2) and Cambisols from Tertiary material (C1) due to 

their spatial extent and projected high decreases of SOC.

land 

use

soil 

class

sites 

(n)

A horizon 

(cm)

clay (%)

C input  

(t ha

−

1

)

DOC  

(t ha

−

1

)

SA (t ha

−

1

)

POM  

(t ha

−

1

)

SC (t ha

−

1

)

rSOC  

(t ha

−

1

) SOC t ha

−

1

)

C

G

3

32 

±

  7

30 

±

  15

4.1 

±

  0.3

2.5 

±

  0.3

12.6 

±

  18.7

4.6 

±

  2.0

55.5 

±

  14.8

5.7 

±

  4.5

80.8 

±

  2.5

L2

3

29 

±

  5

27 

±

  8

3.5 

±

  0.5

1.8 

±

  0.3

1.0 

±

  0.5

2.4 

±

  2.0

36.8 

±

  9.4

2.6 

±

  0.5

44.5 

±

  10.9

C1

3

29 

±

  5

21 

±

  6

3.5 

±

  0.5

1.3 

±

  0.3

1.9 

±

  0.8

9.9 

±

  8.4

29.3 

±

  11.5

4.3 

±

  1.7

46.8 

±

  2.3

C3

2

26 

±

  8

44 

±

  13

3.5 

±

  0.5

2.2 

±

  1.6

2.3 

±

  1.3

4.5 

±

  3.4

59.1 

±

  31.5

3.6 

±

  2.0

71.7 

±

  39.9

C4

2

28 

±

  4

47 

±

  2

3.8 

±

  0.2

1.5 

±

  0.9

1.9 

±

  0.3

3.8 

±

  0.6

45.3 

±

  4.9

4.9 

±

  1.5

57.4 

±

  8.1

C6

3

32 

±

  8

22 

±

  10

3.5 

±

  0.7

1.7 

±

  0.9

2.4 

±

  0.3

5.6 

±

  1.7

42.5 

±

  17.7

4.0 

±

  1.3

56.3 

±

  21.1

C7

3

27 

±

  6

30 

±

  16

3.4 

±

  0.4

1.3 

±

  0

3.6 

±

  2.0

4.2 

±

  2.0

24.1 

±

  4.1

2.8 

±

  0.9

36.0 

±

  5.0

V

2

26 

±

  4

52 

±

  13

3.6 

±

  0.5

2.3 

±

  1.5

3.6 

±

  1.0

5.7 

±

  1.9

54.4 

±

  20.3

5.1 

±

  3.6

71.0 

±

  27.3

G

G

3

16 

±

  6

33 

±

  13

5.0 

±

  0.7

1.6 

±

  0.3

45.5 

±

  33.3

6.4 

±

  2.6

23.8 

±

  6.9

2.6 

±

  0.9

80.0 

±

  27.3

L1

3

25 

±

  5

32 

±

  10

5.3 

±

  0.3

2.1 

±

  0.8

24.8 

±

  14.0

2.7 

±

  1.1

49.1 

±

  12.9

3.7 

±

  0.3

82.3 

±

  6.6

L2

3

25 

±

  6

32 

±

  12

5.2 

±

  0.3

2.8

±

 1.2

24.3 

±

  14.9

5.2 

±

  4.1

37.0 

±

  7.4

2.3 

±

  0.4

71.6 

±

  6.5

C1

3

19 

±

  2

23 

±

  9

5.1 

±

  0.2

2.4 

±

  0.5

16.6 

±

  4.0

4.1 

±

  1.5

37.6 

±

  8.0

4.2 

±

  1.5

65.0 

±

  10.6

C2

3

32 

±

  6

28 

±

  13

5.1 

±

  0.3

2.7 

±

  0.8

17.3 

±

  11.9

4.7 

±

  1.2

50.6 

±

  14.6

6.7 

±

  0.7

82.0 

±

  25.3

C3

3

24 

±

  6

37 

±

  16

4.5 

±

  0.5

1.5 

±

  0.8

10.6 

±

  16.2

4.7 

±

  2.6

47.1 

±

  22.1

2.3 

±

  0.7

66.3 

±

  32.0

C5

3

20 

±

  4

19 

±

  2

4.5 

±

  0.5

1.7 

±

  1.1

11.1 

±

  2.8

4.2 

±

  4.0

30.8 

±

  12.9

4.0 

±

  1.8

51.8 

±

  20.9

C6

3

23 

±

  7

26 

±

  6

4.5 

±

  0.5

3.4 

±

  3.4

9.5 

±

  4.6

7.9 

±

  5.2

25.2 

±

  8.9

4.7 

±

  5.2

50.7

±

 19.4

C7

3

24 

±

  9

36 

±

  23

4.9 

±

  0.1

2.6 

±

  1.9

18.8 

±

  18.4

6.9 

±

  6.1

44.1 

±

  11.9

8.3 

±

  9.8

80.6 

±

  35.8

V

3

23 

±

  8

45 

±

  9

4.6 

±

  0.3

2.5 

±

  1.4

17.1 

±

  4.9

4.1 

±

  2.9

58.0 

±

  30.7

3.6 

±

  1.2

85.2 

±

  38.5

Table 2.

 

 Depth of the A horizon, clay content, C input, OC amount of soil fractions (DOC 

=

 dissolved 

organic matter, SA 

=

 sand- and aggregate-associated SOM, POM 

=

 particulate organic matter, SC 

=

 silt- 

and clay-associated SOM less an inert fraction, rSOC 

=

 inert SOM) of major soil units in Bavaria under 

cropland (C) and grassland (G) (mean values 

±

 standard deviation).

 G 

=

  groundwater-affected  soils 

(Gleysols, Fluvisols); L1 

=

  shallow to intermediate soils with clay accumulation in the subsoil (Luvisols); 

L2 

=

  intermediate to deep soils with clay accumulation in the subsoil (Luvisols); C1 

=

  soils with well developed 

B horizons from Tertiary material (Cambisols); C2 

=

  soils with well developed B horizons from morainal 

material in places with clay accumulation in the subsoil (Cambisols, Luvisols); C3 

=

  shallow soils from 

limestone weathering with or without loess coverings (Cambisols, Luvisols, Leptosols); C4 

=

  intermediate  to 

deep soils from limestone weathering with or without loess coverings (Cambisols, Luvisols); C5 

=

  soils with well 

developed B horizons from acidic material with low base saturation (Cambisols); C6 

=

  soils with well developed 

B horizons from sandstone with low base saturation (Cambisols); C7 

=

  soils with well developed B horizons 

from sandstone with initial podzolisation (Cambisols, Podzols); V 

=

  clay-rich soils (Cambisols, Vertisols, 

Stagnosols).

Figure 3.

 

C input in cropland and grassland sites between 1995 and 2010 (mean values with standard 

deviation). 

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Soil units under grassland uniformly contributed to total SOC stocks. For the scenarios C0, C20

−

  and C20

+

 ,  

total SOC losses of 11% of initial SOC stocks (7.6 Mt), 20% (13.7 Mt) and 3% (2.0 Mt) were estimated for grass-

land soils, respectively. However, for the scenario C20

+

  marginal increases of total SOC stocks of 0.1 to 0.3 Mt 

were estimated for grassland soil units in the vicinity of the Alps (groundwater-affected soils, G; soils with clay 

accumulation in the subsoil L1, L2; Cambisols and Luvisols from moraine material, C2). In total, the projections 

revealed absolute SOC losses of 27.9 Mt for the scenario C0, 44.6 Mt for C20

−

  and 12.5 Mt for C20

+

  in agricul-

tural soils of Bavaria. These SOC losses correspond to total emissions of CO

2

-equivalents of 102.3 Mt, 163.6 Mt 

and 45.9 Mt, respectively.

Discussion

Reliable projections of SOC stock evolution using the RothC model require a reasonable estimation of actual 

SOC pool size by soil fractionation

34

. The application of proposed pedotransfer functions or equilibrium model 

runs as alternative methods for SOC pool size estimation may be adequate in long-term undisturbed land use 

systems or for long projection periods

26,36

, but are probably only partly suitable for intensively managed croplands 

and accurate projections for coming decades. Moreover, the determination of the C input on the basis of the C 

allocation method using crop-specific C allocation coefficients and county-specific crop yields is a promising 

method to estimate C inputs precisely

35,37

. This was confirmed by the reference scenarios (SOC projections under 

constant climatic conditions) that indicated only a slight overestimation of C inputs (Figs 4 and 5), assuming that 

SOC stocks at all investigated sites were in equilibrium. Therefore, the overall accuracy of our modelling approach 

can be assessed as relatively high.

Our projections of SOC stocks under future climate change and constant C inputs revealed substantial SOC 

losses for cropland (

−

 16%) and grassland (

−

 11%) in Bavaria until 2095 (Table 1, Fig. 6). The projected mean 

temperature increase of 3.3 °C at the study sites until 2095 and slightly increased precipitation under the A1B 

climate scenario will obviously lead to an increased mineralization of SOC stocks

3,5

. An anticipated decline of C 

inputs by 20% (C20

−

 ) would result in even higher decreases of SOC by 24 and 19% for cropland and grassland, 

respectively. Under cropland, projected SOC losses affected all investigated soil units, particularly Cambisols 

from Tertiary material (C1). This is mainly attributed to the fact, that these soils are characterized by relatively 

low clay contents (21 

±

  6%). As a result, these soils contain a much lower proportion of SOC stabilized in soil 

aggregates and the s

+

 c fraction and an equivalent higher contribution of labile SOC (DOC 

+

  POM) than other 

cropland soils (Table 2). Under grassland, slightly lower projected SOC losses compared to cropland are due to 

generally higher C inputs (Fig. 3) and considerably higher proportions of aggregate-protected SOC (Table 2). 

Among grassland soil units, noticeably lower SOC declines were projected for soils in relatively cool, pre-alpine 

Figure 4.

 

Projected development of SOC stocks of cropland soils in Bavaria between 2000 and 2095 under 

current climate and land use conditions (reference scenario, RS), climate change and constant C input (C0), 

climate change and decreased C input by 20% (C20

−

) and climate change and increased C input by 20% 

(C20

+

) (mean values with standard deviation from 21 sites). 

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regions with precipitation amounts 

>

 1000 mm and relatively high C input values 

>

 5 t  ha

−

1

. Remarkably, even 

under the optimistic assumption of an increase of C inputs by 20%, a decline of SOC stocks by 8 and 3% was 

projected for cropland and grassland, respectively (Table 1, Fig. 6). Only grassland soil units in pre-alpine regions 

revealed a slight SOC increase.

The finding of generally declining SOC stocks under future climate change is in line with some other SOC 

projection approaches for agricultural soils. Xu 

et al.

27

 modelled SOC changes in eight Irish grassland soils from 

2021 to 2060 assuming constant C inputs on the basis of RothC and two different initialization methods. They 

estimated a decrease of SOC stocks by 2 to 6% for different climate change scenarios, but speculated that future 

summer droughts could reduce C inputs in grasslands. In a study in Australia, a SOC decrease of 10 to 11% was 

predicted under the A2 climate scenario for the period 2008 to 2100 for 12 grassland sites with a constant C input 

using different initialization methods of RothC

26

. For cropland and grassland soils in Louisiana, a RothC sim-

ulation under different climate scenarios and unchanged C input revealed SOC declines of 11 to 18% and 12 to 

Figure 5.

 

Projected development of SOC stocks of grassland soils in Bavaria between 2000 and 2095 under 

current climate and land use conditions (reference scenario, RS), climate change and constant C input (C0), 

climate change and decreased C input by 20% (C20

−

) and climate change and increased C input by 20% 

(C20

+

) (mean values with standard deviation from 30 sites). 

Figure 6.

 

Projected average SOC changes of cropland (C) and grassland (G) sites between 2000 and 2095 

under climate change (A1B) and different C input scenarios. 

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17%, respectively

24

. An estimation of SOC changes of agricultural soils of Italy within the 21

st

 century using legacy 

soil data indicated SOC decreases of 3.6 to 11.5% for different climate scenarios and unchanged C inputs

23

. A 

European-wide study on the basis of legacy data and different climate scenarios estimated SOC declines between 

1980 and 2080 of 10 to 14% for croplands and 6 to 10% for grasslands without C input changes

19

. However, these 

authors also incorporated a simulation of NPP changes in order to estimate C input changes and found a lower 

decrease of croplands SOC stocks of 3 to 4% and even a slight increase for grassland due to a strong projected 

increase of NPP.

Similar studies that assumed increased C inputs due to increased NPP generally found SOC gains. For agri-

cultural soils of northeastern Spain, a mean increase of SOC stocks of 6.3% was estimated for different climate 

projections and agricultural systems (with a wide range of SOC changes of 

−

 12.3 to 32.8%) for the period 2007 

to 2087 under the assumption of large C input increases of up to 44%

22

. Lugato 

et al.

21

 also estimated a slight SOC 

gain of 2% for European agricultural soils in the 21

st

 century. In this study, higher soil respiration under climate 

change was counterbalanced by an assumed 22% higher crop productivity. In SOC projection studies including 

all land uses on the global or the European scale, both increases and decreases of SOC stocks were projected for 

coming decades, but it was generally assumed that NPP increases and associated C input gains would offset or 

even outperform SOC losses induced by global warming, also in agricultural soils

20,25,38,39

.

From our point of view, assumed C input increases in agricultural soils under climate change is a rather opti-

mistic scenario given rising evidence for negative effects of climate change on crop productivity. In several stud-

ies, an increased occurrence of droughts and high temperatures above the optimum of crops and a shortening of 

the growing season were associated with a reduction of NPP

40–46

. The exceptional dry and hot year 2003 provided 

a first hint for such a development, as a dramatic decline of C inputs was determined for 2003 (Fig. 4)

47

. As such 

years are projected to increase under climate change, the possibility of stagnating or even reduced C inputs should 

be considered more strongly in SOC projections. Moreover, socioeconomic reasons, e.g. a change of agricultural 

management as a consequence to changes in the common agricultural policy (CAP) in the EU in the 1990s, were 

quoted to contribute to observed yield stagnation in Europe in the last 25 years

9,42,48

. Some authors assumed that 

land use soil class SOC (Mt)

∆

SOC C0

∆

SOC C20

−

∆

SOC C20

+

(Mt)

(CO

2

-eq.)

(Mt)

(CO

2

-eq.)

(Mt)

(CO

2

-eq.)

C

G

11.0

−

 1.6

−

 5.9

−

 2.3

−

 8.5

−

 0.9

−

 3.4

L2

31.0

−

 6.6

−

 13.7

−

 9.8

−

 20.4

−

 3.6

−

 7.6

C1

18.7

−

 3.7

−

 24.0

−

 5.6

−

 35.8

−

 2.1

−

 13.2

C3

19.4

−

 2.3

−

 8.5

−

 3.6

−

 13.3

−

 1.1

−

 4.0

C4

8.4

−

 1.4

−

 5.2

−

 2.2

−

 8.0

−

 0.7

−

 2.6

C6

9.1

−

 1.4

−

 9.6

−

 2.1

−

 14.7

−

 0.7

−

 4.7

C7

5.3

−

 0.7

−

 5.1

−

 1.4

−

 7.7

−

 0.1

−

 2.6

V

20.0

−

 2.6

−

 2.6

−

 4.0

−

 5.0

−

 1.3

−

 0.4

total

122.9

−

 20.3

−

 74.5

−

 30.9

−

 113.4

−

 10.5

−

 38.5

G

G

5.8

−

 

−

 0.3

−

 1.2

−

 0.8

−

 2.9

0.1

0.3

L1

5.5

−

 0.1

−

 0.3

−

 0.5

−

 1.8

0.3

1.1

L2

4.6

−

 0.3

−

 3.7

−

 0.7

−

 7.1

0.1

−

 0.6

C1

9.4

−

 1.0

−

 2.6

−

 1.9

−

 6.3

−

 0.2

0.8

C2

14.6

−

 0.7

−

 1.2

−

 1.7

−

 2.7

0.2

0.2

C3

3.8

−

 0.7

−

 2.4

−

 1.0

−

 3.7

−

 0.3

−

 1.2

C5

9.6

−

 2.2

−

 3.0

−

 3.3

−

 4.8

−

 1.2

−

 1.3

C6

3.7

−

 0.7

−

 8.0

−

 1.1

−

 11.9

−

 0.3

−

 4.3

C7

6.2

−

 0.8

−

 2.5

−

 1.3

−

 4.1

−

 0.4

−

 1.0

V

6.7

−

 0.8

−

 3.0

−

 1.3

−

 4.8

−

 0.4

−

 1.4

total

70.0

−

 7.6

−

 27.8

−

 13.7

−

 50.2

−

 2.0

−

 7.3

total

192.9

−

 27.9

−

 102.3

−

 44.6

−

 163.6

−

 12.5

−

 45.9

Table 3.

 

 Total storage of SOC (2000) and total projected SOC changes in Bavaria (2000–2095) under 

constant C input (

∆

SOC C0), decreased C input by 20% (

∆

SOC C20

−

) and increased C input by 20% 

(

∆

SOC C20

+

) according to major soil units under cropland (C) and grassland (G).

 G 

=

  groundwater-

affected soils (Gleysols, Fluvisols); L1 

=

  shallow to intermediate soils with clay accumulation in the subsoil 

(Luvisols); L2 

=

  intermediate to deep soils with clay accumulation in the subsoil (Luvisols); C1 

=

  soils  with 

well developed B horizons from Tertiary material (Cambisols); C2 

=

  soils with well developed B horizons from 

morainal material in places with clay accumulation in the subsoil (Cambisols, Luvisols); C3 

=

  shallow soils from 

limestone weathering with or without loess coverings (Cambisols, Luvisols, Leptosols); C4 

=

  intermediate  to 

deep soils from limestone weathering with or without loess coverings (Cambisols, Luvisols); C5 

=

  soils with well 

developed B horizons from acidic material with low base saturation (Cambisols); C6 

=

  soils with well developed 

B horizons from sandstone with low base saturation (Cambisols); C7 

=

  soils with well developed B horizons 

from sandstone with initial podzolisation (Cambisols, Podzols); V 

=

  clay-rich soils (Cambisols, Vertisols, 

Stagnosols).

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Figure 7.

 

Regional distribution of projected SOC changes in Bavaria between 2000 and 2095 under climate 

change and constant C input (C0), climate change and decreased C input by 20% (C20

−

) and climate change 

and increased C input by 20% (C20

+

) (the maps were generated using ESRI ArcMap 9.2,

 

www.esri.com/

).

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future improvement of technology, which was mainly responsible for the obtained yield increases until the 1990s, 

could lead to a strong increase of crop productivity, but as its drivers are not clear, future technology development 

is afflicted with high uncertainty

26,49

. Despite ongoing technological improvement in the last three decades, par-

ticularly related to plant breeding

50

, yields of many crops showed no equivalent increase.

The projected decrease of agricultural SOC stocks is associated with a substantial emission of CO

2

 into the 

atmosphere. A rough estimation for the entire agriculturally used land in Bavaria revealed emissions of 46 to 

164 Mt CO

2

-equivalents for the period 2000 to 2095 (depending on the C input scenario) (Table 3, Fig. 7). On a 

yearly basis, this would increase the CO

2

 emissions of the agricultural sector of Bavaria by 4 to 12% (based on an 

estimated CO

2

 emission of approximately 14 Mt in 2012). Given the high projected amount of CO

2

 emitted by 

agricultural soils, climate-change-induced SOC changes should be included in national C accounting programs. 

Besides the emission of CO

2

, projected losses of SOM in agricultural soils may have negative effects on several 

important soil functions, e.g. retention of pollutants, buffering capacity, erosion control, water holding capacity 

and nutrient supply. Overall, the estimated SOC losses could have detrimental effects on soil fertility and eco-

system resilience, which would in turn aggravate crop productivity and thus residue-derived C input – a positive 

feedback. This calls for an extension of soil monitoring programs and the implementation of early warning indi-

cators in order to verify projected SOC losses.

In view of these potential risks, there is the need to increase C inputs in order to maintain present SOC stocks. 

A comparison of relative C inputs (related to the SOC stock) for the reference scenario (current climate condi-

tions) and the climate change scenario indicated a future increase of C inputs of 29% needed to counterbalance 

increased decomposition of SOM (Fig. 8).

Several options were proposed to enhance C inputs in agricultural soils. Promising agricultural practices 

comprise increased return of crop residues and incorporation of other organic inputs, improved crop rotation 

including legumes and catch crops, organic farming with clover prominent in the rotation, increased cultivation 

of bioenergy and perennial crops, improved pasture and livestock management, agroforestry and conversion of 

cropland to grassland

51–60

. Recently, such C sequestration options were proposed to offset global anthropogenic 

CO

2

 emissions in form of the “4‰ concept” at the climate conference in Paris (COP21). This concept is based 

on the assumption that a yearly increase of global SOC stocks (first 40 cm of soils except permafrost) of 4‰ by 

improved management would largely contribute to counterbalance human-induced CO

2

 emissions. However, 

our results indicate that a distinct increase of C inputs by improved agricultural management would be necessary 

in future decades only to counterbalance enhanced SOM decomposition. Therefore, the potential contribution 

of improved management of agricultural soils to decrease the atmospheric rise in CO

2

 should be evaluated more 

carefully.

Although our results provide a good estimate on the potential trajectories of SOC storage in temperate agri-

cultural soils under the given assumptions of climate change and C inputs, model predictions need independent 

verification and a calculation of uncertainty. There are different sources of uncertainty that may contribute to the 

final uncertainty in predictions, for instance, uncertainty in model parameters, and in model structure. Although 

the RothC model was developed for temperate agroecosystems and has been applied successfully in regions under 

similar climatic and pedogenic conditions in the past, it is still unknown whether RothC’s default parameters may 

need to be adapted for the study region. Also, the existence of a completely inert pool has been challenged

61,62

, and 

the potential slow dynamics of this pool may become relevant for long-term predictions.

Particularly relevant sources of uncertainty for these types of predictions are the temperature- and 

moisture-dependent functions implemented in this model to modify decomposition rates. The rate modification 

by temperature is identical for each SOM pool in RothC despite indications for varying temperature sensitivity 

of labile and stable SOM pools. The temperature sensitivity of differently stabilized SOM pools is an element of 

uncertainty in current SOM turnover models and is thus a highly topical debate with regard to global change

2,63

. 

The feedback between climate change and C losses from soil would be much stronger than suggested by equal 

temperature sensitivities of all SOM pools – like in our calculations, if stable SOM pools react more sensitively 

to warming than more labile SOM pools. In our studied soils the stable SOM pools amounted for 66 to 90% of 

Figure 8.

 

Relative C input in cropland (C) and grassland (G) sites vs. relative projected SOC change under 

current climate conditions (baseline, 2095) and climate change (A1B, 2095). 

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total SOC. According to the Arrhenius equation stable SOM pools are thought to be more temperature sensitive 

than less stabilized SOM pools

3,64

. Results from laboratory incubations and long-term experiments indicate that 

the more stable SOM pools are indeed more temperature sensitive

65–67

. A higher temperature sensitivity of the 

stable SOM pools implies that the projected mean temperature increase of 3.3 °C may lead to even stronger and 

sustained C losses from soils than suggested by our scenarios. Therefore, there is the need to integrate information 

on temperature sensitivity in soil carbon models.

Further critical issues are related to C input projections. Although our scenarios cover the range of expected 

C input development based on yield projections, more spatially and temporarily precise C input scenarios are 

needed. A combination of sophisticated yield models together with an estimation of future changes of C alloca-

tion patterns of crops could be a promising approach to derive more precise C input estimates.

Materials and Methods

Study area. 

The state of Bavaria comprises a total area of 70550 km

2

 and is located in southeast Germany. 

The northwestern part of Bavaria is dominated by the southern German escarpment landscape that adjoins the 

low mountain ranges of the Bohemian Massif in the east. Southwards the Molasse basin ascends to the mountain 

range of the Alps. Elevation ranges between 107 and 2962 m above sea level. Due to its location in central Europe, 

Bavaria exhibits a sub-oceanic climate that is characterized by a transitional situation between a maritime climate 

in the northwest and sub-continental influences in the east. Mean annual temperature and precipitation from the 

escarpment landscape in the northwest to the Alps in the south range between 9° and 4 °C and 550 and 2500 mm, 

respectively. Around half of the area of Bavaria is under agricultural use, with cropland and grassland accounting 

for areas of 22843 km

2

 (32% of the total area) and 9897 km

2

 (14% of the total area), respectively (Fig. 1). Dominant 

soil classes within agricultural land are soils with well-developed B horizons (Cambisols), soils with clay accu-

mulation in the subsoil (Luvisols), soils from limestone weathering with or without loess coverings (Cambisols, 

Luvisols, Leptosols), clay-rich soils (Cambisols, Vertisols, Stagnosols) and groundwater-affected soils (Gleysols, 

Fluvisols) according to the German soil system and the equivalent Reference Soil Groups of the WRB system

68

.

Due to the heterogeneous nature in terms of geology, soils and climate, the conditions for agriculture differ 

considerably within Bavaria

69

. In the south of Bavaria, the region of the Alps and Pre-Alps is characterized by high 

mean annual precipitation (MAP) of 

>

 1000 mm and relatively low mean annual temperatures (MAT) between 

5.4 and 7.5 °C. Intensive grassland use for cattle husbandry prevails with up to six swaths per year. The scarce 

arable land is predominantly cultivated with corn for silage (

Zea mays

). To the north, between the Pre-Alps and 

the Danube River, the Tertiary Hills Region encompasses a small structured mixed landscape. A more temperate 

climate with MAT of 7.0–7.7 °C, MAP of 700 to 1000 mm and deep, loamy soils provide good farming conditions. 

Here, animal husbandry has decreased tremendously over the last 20 years and the conversion of grassland to ara-

ble land has been strongest within Bavaria. Corn for biogas production has become a major crop in the rotation 

together with a mix of other cereals and rapeseed (

Brassica napus

). Within the northern part of the Tertiary Hills 

Region and further northwest in Bavaria, small, isolated Loess Regions are prominent and form the most produc-

tive areas in Bavaria, accompanied by favourable climatic conditions, leading to intensive agricultural production 

of e.g. sugar beet (

Beta vulgaris

) and winter wheat (

Triticum aestivum

). The East-Bavarian Mid-Range Mountains 

are characterized again by rather unfavourable climatic and pedological conditions for most high-output cere-

als. While small areas are used as grassland, potatoes (

Solanum tuberosum

), triticale (

×

 

Triticosecale

), a hybrid 

of wheat and rye, spring barley (

Hordeum vulgare

) and oats (

Avena sativa

) find good conditions. In the central 

part of northern Bavaria, the region of Jurassic sediments is characterized by MAT of 7.1–7.2 °C, MAP of around 

800 mm and fertile soils. To the northwest, the large northern Bavarian Hill Area exhibits a slightly warmer 

and drier climate, but sandy, acidic soils and low water availability during the summer months inhibit intensive 

agricultural production, allowing the cultivation of winter rye (

Secale cereale

) and triticale. Further to the west, 

the Franconian Lowlands are characterized by lower precipitation (

<

 700 mm), higher MAT (7.5–9.2 °C) and 

clay-rich, fertile soils, which restrict agriculture to drought-resistant crops like spelt (

Triticum spelta

) and durum 

wheat (

Triticum durum

) together with winter rye. These grains are also cultivated in the very north-eastern part 

of Bavaria forming the Franconian Mid-Range Mountains that exhibit a cooler climate (5.2–8.0 °C) and acidic 

soils but the main use of agricultural land is grassland. Overall, there has been a general trend of intensification 

of arable land, mainly on more or less productive sites in Bavaria, enhanced through the lifting of the obligatory 

set-aside policy in 1992, the introduction of a series of renewable energy subsidies since 2003, leading to a huge 

demand for green maize for biogas production and, last but not least, technological progress in general. On the 

other hand, there is a visible trend of extensification of marginal sites, where there is a tendency of farmland 

abandonment, further decrease in livestock numbers and remaining unclear perspectives of farming in the future. 

Additionally, the steady demand for land for urbanisation in the booming regions of Bavaria is increasing the 

pressure on agricultural land. However, the share of arable land remained since 20 years and only grassland has 

decreased substantially, mainly due to compensatory effects for the developments.

Selection of study sites and soil sampling. 

Study sites were selected for all major soil units and land 

uses in Bavaria using a map that combines predominant soil units with main land uses

70

. The map comprises 38 

soil units within Bavaria resulting from an intersection of soil type and parent material with macroclimate. These 

soil units were subdivided to 61 units according to cropland and grassland use on the basis of CORINE Land 

Cover data. Due to the great effort of the fractionation approach to derive RothC pools, all soil/land use units 

were excluded which were smaller than 1% of the area of Bavaria. For each of the remaining 18 soil/land use units  

(8 soil units under cropland and 10 soil units under grassland) representative locations were selected on the basis 

of available soil material and data from different soil surveys and permanent soil observation sites in Bavaria 

compiled by the Bavarian Environment Agency (Fig. 1). Although grassland covers only about half the area of 

cropland in Bavaria, grassland use is distributed over a much wider climatic range and thus more grassland soil 

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units were incorporated in this study. For the majority of soil/land use units, three soil profiles provided repre-

sentative data. For three of the soil units under cropland (shallow soils from limestone weathering with or without 

loess coverings; intermediate to deep soils from limestone weathering with or without loess coverings; soils with 

well developed B horizons from sandstone with initial podzolisation) only two representative soil profiles were 

available. In total, 51 study sites (21 long-term cropland sites and 30 permanent grassland sites) were selected. The 

selected soil profiles comprise permanent soil monitoring sites and a grid sampling of 8 

×

  8 km within Bavaria 

between 2000 and 2004. 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 position in the particular land use type. Anthropogenic disturbances in the subsoil 

were excluded in a pre-exploratory survey using a soil auger. Within the topsoil (A horizons), soil material was 

collected as a composite sample from four sub-locations located with a radius of 6.6 m and four sub-locations with 

a radius of 3.8 m around the main soil profile in order to cover the small-scale heterogeneity of the soils. At the 

main profile, soil horizons were additionally sampled using steel cylinders for the determination of bulk density 

(BD). The content of rock fragments 

>

 2 mm was estimated visually in the soil profiles.

Fractionation of SOC pools and determination of basic soil properties. 

For each topsoil horizon 

(A horizons), soil material was fractionated according to the approach of Zimmermann 

et al.

34

 and Poeplau  

et al.

71

 (Fig. 9). Soil material was sieved to 2 mm and 30 g were suspended in 150 ml of deionised water and dis-

persed 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 macroaggregates and to prevent the 

disruption of mineral-associated SOM

72

. The suspension was wet sieved over a 63 

μ

 m sieve until the rinsing water 

was clear and subsequently filtered through a 0.45 

μ

 m membrane filter. The rinsing water was collected and dis-

solved organic carbon (DOC) was measured using a TOC analyzer 5050A (Shimadzu, Duisburg, Germany). The 

fractions 

>

 0.45 

μ

 m  and 

>

 63 

μ

 m were dried at 40 °C and weighted. Afterwards, 50 ml of a sodium polytungstate 

solution with a density of 2.0 g cm

−

3

 

71

 was added to the fraction 

>

 63 

μ

 m and the floating particulate organic 

matter (POM) was separated after centrifugation at 1000 g for 15 minutes. The remaining fraction 

>

 63 

μ

 m  (sand 

and stable aggregates, S

+

 A) and the POM fraction were washed with deionised water to remove the sodium 

polytungstate, dried at 40 °C and weighted. The fraction 

>

 0.45 

μ

 m (silt and clay, s

+

 c) was homogenized and 

a subsample of 1 g was oxidized with sodium hypochlorite (NaOCl) to separate a stable SOC fraction. 50 ml of 

6% NaOCl, adjusted to pH 8 with concentrated HCl, was added to the subsample and after 18 hours at 25 °C 

the sample was centrifuged at 1000 g for 15 minutes and washed with deionised water. The oxidation proce-

dure was repeated twice. The residual SOC fraction (rSOC) was dried at 40 °C and weighted. All solid fractions 

were measured for SOC and N concentration by dry combustion on an EA 3000 analyser (Hekatech, Wegberg, 

Germany). Soil fractions that contained CaCO

3

 were heated to 500 °C for 4 hours to remove organic carbon, and 

the concentration of inorganic C of the residual material was determined by dry combustion. The SOC con-

tent was calculated by subtracting inorganic C from the total C concentration of the untreated material. For the 

Figure 9.

 

Fractionation scheme of SOC pools according to the method of Zimmermann 

et al.

34

 and 

Poeplau 

et al.

71

 (s

+

c 

=

 silt- and clay-associated SOM less an inert fraction; DOC 

=

 dissolved organic 

matter; S

+

A 

=

 sand- and aggregate-associated SOM; POM 

=

 particulate organic matter; rSOC 

=

 inert 

SOM) and assignment to RothC pools (BIO 

=

 microbial biomass; HUM 

=

 humified organic matter; 

DPM 

=

 decomposable plant material; RPM 

=

 resistant plant material) using splitting ratios. 

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determination of clay contents, soil samples (

<

 2 mm) were oxidized with H

2

O

2

 to remove organic matter (OM). 

The remaining material was dispersed with Na

4

P

2

O

7

 and shaken for 16 to 24 hours, followed by wet sieving to 

isolate sand fractions 

>

 63 

μ

 m. To determine silt and clay fractions, approximately 3 g of the 

<

 63 

μ

 m  fraction 

was suspended in deionised water using Na

4

P

2

O

7

 and an ultrasonication for 3 minutes with 75 J mL

−

1

 was con-

ducted. 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 Micromeritics Sedigraph 5100 

(Micromeritics, Norcross, GA, USA). For the determination of BD, the mass of the oven-dry soil (105 °C) was 

divided by the volume of the soil cores.

Estimation of the C input. 

In a preliminary study

35

, annual C input values were estimated for most impor-

tant crops as well as grassland in Bavaria on the county-scale for the period from 1951 to 2010. The C input 

was calculated on the basis of the C allocation approach proposed by Bolinder 

et al.

37

. This method is based on 

the allocation of NPP to four plant fractions: the agricultural product (R

P

), aboveground biomass excluding R

P

 

(R

S

), belowground biomass excluding RP (R

R

) and extra-root C (R

E

). Due to possible changes of the proportions 

of the C allocation coefficients R

P

, R

S

, R

R

 and R

E

 in the period between 1951 and 2010 induced by progress in 

plant breeding, improved crop management and fertilization, C allocation coefficients were estimated both for 

the beginning (1951–1955) and the end (1995–2010) of the considered period. For the period 1951 to 1955, C 

allocation coefficients were derived from numerous agricultural field studies and reports that were conducted in 

the post-war period. The proportions of R

P

 and R

S

 were derived from agricultural statistics for Germany from 

1951 to 1955 and from Köhnlein and Vetter

73

. The contribution of R

R

 to NPP was estimated on the basis of sev-

eral field studies which determined the root biomass of different crop plants gravimetrically by washing of soil 

monoliths

74–82

. The contribution of R

E

 was estimated to be two-thirds of root biomass according to Bolinder  

et al.

37

. Recent C allocation coefficients for the period 1995–2010 were derived from agricultural statistical data 

for Germany as well as literature data from 1995 onwards

37,83–88

. If no data was provided for R

E

, the contribution 

of rhizodeposition was estimated as described above. Between 1955 and 1995, allocation coefficients were linearly 

interpolated for each year according to the linear trend of crop yields within this period.

The total C input as the sum of the C input of all plant fractions except the agricultural product (C

S

+

 C

R

+

 C

E

) 

can be calculated using the relative C allocation coefficients R

P

, R

S

, R

R

 and R

E

 as well as the crop yield:

=

×

C

R
R

C

(1)

S

S

P

P

=

×

C

R
R

C

(2)

R

R

P

P

=

×

C

R
R

C

(3)

E

E

P

P

where C

S

 is the C input of aboveground biomass excluding the agricultural product (t ha

−

1

 yr

−

1

), C

R

 is the C 

input of belowground biomass (roots) (t ha

−

1

 yr

−

1

), C

E

 is the C input of rhizodeposition (t ha

−

1

 yr

−

1

) and C

P

 is 

the amount of C in the agricultural product (t ha

−

1

 yr

−

1

). The amount of C in the agricultural product (C

P

) can 

be calculated using dry-matter yields of respective crops obtained from agricultural statistics assuming that the 

C content of different plants and plant parts is 0.45 kg kg

−

1

 

37

. For the estimation of C inputs of the selected study 

sites, information on the crop rotation, return of harvest residues and the input of organic fertilizers were gained 

from farmers for each site for a 16-year period (1995–2010). Mean C input values for the initialization of RothC 

were calculated using C input data derived from 1995 to 1999 before the start of the soil sampling campaign.

Climate projections. 

Projections of climate parameters for each study site for the period 2000 to 2095 were 

derived by using a multi-model ensemble approach on the basis of climate change scenario A1B. The ensemble 

consists of 19 climate predictions for Germany. They are a combination of different regional climate models 

(RCM) that are driven by several global climate models (GCM) (Table 4). The multi-model ensemble had been 

developed in the European framework project ENSEMBLES

89

. As climate models represent reduced natural pro-

cesses, one model cannot give the full range of information about uncertainties of future climate. To handle 

these model-based uncertainties the use of ensembles for climate impact studies is widely accepted and recom-

mended

90

. The result of every member of the ensemble is equally probable and represents one possible future but 

not the only one. The climate data set for the 51 study sites in Bavaria, provided by the German Weather Service 

(DWD), covers the period from January 2000 to December 2095 and consists of monthly data on air temperature, 

precipitation and evapotranspiration of the FAO

91

. The latter refers to a short green grass completely shading the 

ground and with adequate water supply. The spatial resolution of climate data is 25 

×

  25 km. Every site is repre-

sented by a grid cell of 3 

×

  3 grid points with the study site in the middle. For each grid point median values were 

calculated for temperature, precipitation and evapotranspiration from the 19 climate projections. To account for 

spatial insecurity the arithmetic mean of the results of the nine grid points was used for this study.

Running the RothC model for different carbon input scenarios. 

To predict the SOC stock devel-

opment in the topsoil, the RothC 26.3 model was used, which was run in the R environment using the package 

SoilR

92

. The model simulates the turnover of SOC on the basis of five conceptual pools. Incoming plant material 

is separated into decomposable plant material (DPM) and resistant plant material (RPM), which are decomposed 

to humified organic matter (HUM), microbial biomass (BIO) and CO

2

. The HUM and BIO pools undergo further 

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14

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 | 6:32525 | DOI: 10.1038/srep32525

decomposition. Each of these compartments decomposes by first-order kinetics with decay rate constants of 10, 

0.3, 0.66 and 0.02 per year for DPM, RPM, BIO and HUM, respectively. Only the inert organic matter pool (IOM) 

is resistant to decomposition. The decay rate constants are modified by temperature and soil moisture deficit 

(difference between potential evapotranspiration and precipitation with a maximum deficit defined by the clay 

content). For model initialization, the measured fractions from the A horizons were summed up and converted to 

model pools according to the approach of Zimmermann 

et al.

34

 using average RPM/DPM and BIO/HUM split-

ting ratios for temperate croplands (0.0102 and 0.0272) and grasslands (0.1271 and 0.0259) (Fig. 9). The selected 

51 sites were modelled in monthly time steps using the default parameter set of RothC. In order to predict the 

evolution of SOC stocks from 2000 to 2095, the model was run under unchanged climate conditions (reference 

scenario, mean climate conditions from the period 1971 to 1999) and a moderate climate change scenario (A1B). 

The reference scenario served as validation of the estimated C input values. The difference of SOC projections 

between the reference and the climate change scenario represented the projected SOC change. As a simplification, 

the measured SOC stock obtained from soil samples taken between 2000 and 2004 was assumed to resemble the 

stock of the year 2000.

The SOC projections were performed for three different C input scenarios as a realistic range of possi-

ble yield development: i) constant C input over time (mean C input from the period 1995 to 1999) (C0), ii) 

decreasing C input over time (reaching a 20% lower C input in 2095) (C20

−

 ) and iii) increasing C input over 

time (reaching a 20% higher C input in 2095) (C20

+

 ). These scenarios were based on present and projected 

crop yields in this study, which showed a strong correlation with agricultural C inputs so far

35,93

. The scenario 

C0 (constant C input) is based on recent yield stagnation of several crops in Bavaria. Evidence was found that 

yields of most important crops of Bavaria are stagnating since the 1990s along with stagnating C inputs

9,35

. 

Besides the Status quo of stagnating C inputs, several crop yield modeling approaches projected yield changes 

within the 21

st

 century. For some study regions in southern Germany, a decline of yields of winter wheat and 

corn for silage by around 10 to 20% until the mid-century was projected

94,95

. On the other hand, an estimation 

of changes of crop productivity in Europe assumed an increase of yields of major European crops by 15 to 32% 

from 2020 until 2080 under different climate scenarios (without considering technology development)

49

. In 

order to cover the full range of expected crop yield development, the scenarios C20

−

  and C20

+

  were included 

in this study.

Statistics. 

Descriptive statistical methods were used to analyze the soil and climate data sets for maximum 

and minimum values, mean, median, standard deviation, interquartile range, skewness, kurtosis, variance and 

coefficient of variation. In order to test the significance of differences of soil properties between the investi-

gated soil units and land uses one-way analyses of variance (ANOVA) combined with post hoc tests (Tukey and 

Scheffé) and Student’s t-tests were applied. Linear regression analyses were conducted to characterize the trend of 

C inputs. All statistical calculations were carried out using the software IBM SPSS Statistics 19 (2010, IBM Corp., 

Armonk).

Country/Institution

GCM

RCM

1

Northern Ireland/Community Climate Change 

Consortium for Ireland

HADCM3Q16

RCA 3.0

2

France/Meteo France

ARPEGE

ALADIN RM5.1

3

Denmark/Danish Meteorological Institute 

ARPEGE

HIRMAM 5

4

BCM

5

ECHAM 5_r3

6

Switzerland/ETH Zurich (Eidgenössische Technische 

Hochschule)

HADCM3Q0

CLM 2.4.6

7

Germany/Helmholtz-Zentrum Geesthacht, 

Centre for Materials and Coastal Research

ECHAM 5_r1

CLM 2.4.11

8

ECHAM 5_r2

9

England/Hadley Centre for Climate Prediction 

and Research

HADCM3Q0

HADRM3Q0

10

HADCM3Q3

HADRM3Q3

11

HADCM3Q16

HADRM3Q16

12

Italy/Intern. Centre for Theoretical Physics

ECHAM 5_r3

RegCM 3

13

Netherlands/Royal Netherlands Meteorological Institute

ECHAM 5_r3

RACMO 2.1

14

Germany/Max Planck Institute for Meteorology

ECHAM 5_r1

REMO 2005

15

ECHAM 5_r2

REMO 2009

16

ECHAM 5_r3

REMO 5.7

17

Sweden/Swedish Meteorological and 

Hydrological Institute, Rossby Centre

BCM

RCA 3.0

18

ECHAM 5_r3

19

HADCM3Q3

Table 4.

 

 Overview over the ensemble of 19 regional climate projections as a combination of global 

circulation models (GCM) and regional circulation models (RCM).

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15

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 | 6:32525 | DOI: 10.1038/srep32525

References

1.  Scharlemann, J. P. W., Tanner, E. V. J., Hiederer, R. & Kapos, V. Global soil carbon: understanding and managing the largest terrestrial 

carbon pool. 

Carbon Manag.

 

5,

 81–91 (2014).

2.  von Lützow, M. & Kögel-Knabner, I. Temperature sensitivity of soil organic matter decomposition-what do we know? 

Biology and 

Fertility of Soils

 

46,

 1–15 (2009).

3.  Davidson, E. A. & Janssens, I. A. Temperature sensitivity of soil carbon decomposition and feedbacks to climate change. 

Nature

 

440,

 

165–173 (2006).

4.  Conant, R. T. 

et al.

 Temperature and soil organic matter decomposition rates - synthesis of current knowledge and a way forward. 

Glob. Change Biol.

 

17,

 3392–3404 (2011).

5.  Kirschbaum, M. U. F. The temperature-dependence of soil organic-matter decomposition, and the effect of global warming on soil 

organic-C storage. 

Soil Biol. Biochem.

 

27,

 753–760 (1995).

6.  Knorr, W., Prentice, I. C., House, J. I. & Holland, E. A. Long-term sensitivity of soil carbon turnover to warming. 

Nature

 

433,

 

298–301, doi: 10.1038/nature03226 (2005).

7.  Wiesmeier, M. 

et al.

 Storage and drivers of organic carbon in forest soils of southeast Germany (Bavaria) - Implications for carbon 

sequestration. 

For. Ecol. Manage.

 

295,

 162–172 (2013).

8.  Jobbagy, E. G. & Jackson, R. B. The vertical distribution of soil organic carbon and its relation to climate and vegetation. 

Ecol. Appl.

 

10,

 423–436 (2000).

9.  Wiesmeier, M., Hübner, R. & Kögel-Knabner, I. Stagnating crop yields: An overlooked risk for the carbon balance of agricultural 

soils? 

Sci. Total Environ.

 

536,

 1045–1051 (2015).

10.  Grassini, P., Eskridge, K. M. & Cassman, K. G. Distinguishing between yield advances and yield plateaus in historical crop 

production trends. 

Nat. Commun.

 

4,

 11 (2013).

11.  Chen, L. Y., Smith, P. & Yang, Y. H. How has soil carbon stock changed over recent decades? 

Glob. Change Biol.

 

21,

 3197–3199 

(2015).

12.  Heikkinen, J., Ketoja, E., Nuutinen, V. & Regina, K. Declining trend of carbon in Finnish cropland soils in 1974–2009. 

Glob. Change 

Biol.

 

19,

 1456–1469 (2013).

13.  Lettens, S., van Orshoven, J., van Wesemael, B., Muys, B. & Perrin, D. Soil organic carbon changes in landscape units of Belgium 

between 1960 and 2000 with reference to 1990. 

Glob. Change Biol.

 

11,

 2128–2140 (2005).

14.  Meersmans, J. 

et al.

 Changes in organic carbon distribution with depth in agricultural soils in northern Belgium, 1960–2006. 

Glob. 

Change Biol.

 

15,

 2739–2750 (2009).

15.  Reijneveld, A., van Wensem, J. & Oenema, O. Soil organic carbon contents of agricultural land in the Netherlands between 1984 and 

2004. 

Geoderma

 

152,

 231–238 (2009).

16.  Riley, H. & Bakkegard, M. Declines of soil organic matter content under arable cropping in southeast Norway. 

Acta Agriculturae 

Scandinavica Section B-Soil and Plant Science

 

56,

 217–223 (2006).

17.  Taghizadeh-Toosi, A. 

et al.

 Changes in carbon stocks of Danish agricultural mineral soils between 1986 and 2009. 

European Journal 

of Soil Science

 

65,

 730–740 (2014).

18.  Coleman, K. 

et al.

 Simulating trends in soil organic carbon in long-term experiments using RothC-26.3. 

Geoderma

 

81,

 29–44 

(1997).

19.  Smith, J. 

et al.

 Projected changes in mineral soil carbon of European croplands and grasslands, 1990–2080. 

Glob. Change Biol.

 

11,

 

2141–2152 (2005).

20.  Gottschalk, P. 

et al.

 How will organic carbon stocks in mineral soils evolve under future climate? Global projections using RothC for 

a range of climate change scenarios. 

Biogeosciences

 

9,

 3151–3171 (2012).

21.  Lugato, E., Panagos, P., Bampa, F., Jones, A. & Montanarella, L. A new baseline of organic carbon stock in European agricultural soils 

using a modelling approach. 

Glob. Change Biol.

 

20,

 313–326 (2014).

22.  Alvaro-Fuentes, J., Easter, M. & Paustian, K. Climate change effects on organic carbon storage in agricultural soils of northeastern 

Spain. 

Agric. Ecosyst. Environ.

 

155,

 87–94 (2012).

23.  Mondini, C., Coleman, K. & Whitmore, A. P. Spatially explicit modelling of changes in soil organic C in agricultural soils in Italy, 

2001-2100: Potential for compost amendment. 

Agric. Ecosyst. Environ.

 

153,

 24–32 (2012).

24.  Zhong, B. & Xu, Y. J. Predicting climate change effects on surface soil organic carbon of Louisiana, USA. 

Environ. Monit. Assess.

 

186,

 

6169–6192 (2014).

25.  Zaehle, S. 

et al.

 Projected changes in terrestrial carbon storage in Europe under climate and land-use change, 1990–2100. 

Ecosystems

 

10,

 380–401 (2007).

26.  Senapati, N. 

et al.

 Projections of changes in grassland soil organic carbon under climate change are relatively insensitive to methods 

of model initialization. 

European Journal of Soil Science

 

64,

 229–238 (2013).

27.  Xu, X. L., Liu, W. & Kiely, G. Modeling the change in soil organic carbon of grassland in response to climate change: Effects of 

measured versus modelled carbon pools for initializing the Rothamsted Carbon model. 

Agric. Ecosyst. Environ.

 

140,

 372–381 

(2011).

28.  Heitkamp, F., Wendland, M., Offenberger, K. & Gerold, G. Implications of input estimation, residue quality and carbon saturation 

on the predictive power of the Rothamsted Carbon Model. 

Geoderma

 

170,

 168–175 (2012).

29.  Cramer, W. 

et al.

 Global response of terrestrial ecosystem structure and function to CO2 and climate change: results from six 

dynamic global vegetation models. 

Glob. Change Biol.

 

7,

 357–373 (2001).

30.  Sitch, S. 

et al.

 Evaluation of ecosystem dynamics, plant geography and terrestrial carbon cycling in the LPJ dynamic global vegetation 

model. 

Glob. Change Biol.

 

9,

 161–185 (2003).

31.  Jones, R. J. A., Hiederer, R., Rusco, E. & Montanarella, L. Estimating organic carbon in the soils of Europe for policy support. 

European Journal of Soil Science

 

56,

 655–671 (2005).

32.  Leifeld, J., Zimmermann, M., Fuhrer, J. & Conen, F. Storage and turnover of carbon in grassland soils along an elevation gradient in 

the Swiss Alps. 

Glob. Change Biol.

 

15,

 668–679 (2009).

33.  Skjemstad, J. O., Spouncer, L. R., Cowie, B. & Swift, R. S. Calibration of the Rothamsted organic carbon turnover model (RothC ver. 

26.3), using measurable soil organic carbon pools. 

Australian Journal of Soil Research

 

42,

 79–88 (2004).

34.  Zimmermann, M., Leifeld, J., Schmidt, M. W. I., Smith, P. & Fuhrer, J. Measured soil organic matter fractions can be related to pools 

in the RothC model. 

European Journal of Soil Science

 

58,

 658–667 (2007).

35.  Wiesmeier, M. 

et al.

 Estimation of past and recent carbon input by crops into agricultural soils of southeast Germany. 

Eur. J. Agron.

 

61,

 10–23 (2014).

36.  Weihermüller, L., Graf, A., Herbst, M. & Vereecken, H. Simple pedotransfer functions to initialize reactive carbon pools of the 

RothC model. 

European Journal of Soil Science

 

64,

 567–575 (2013).

37.  Bolinder, M. A., Janzen, H. H., Gregorich, E. G., Angers, D. A. & VandenBygaart, A. J. An approach for estimating net primary 

productivity and annual carbon inputs to soil for common agricultural crops in Canada. 

Agric. Ecosyst. Environ.

 

118,

 29–42 (2007).

38.  Jones, C. 

et al.

 Global climate change and soil carbon stocks; predictions from two contrasting models for the turnover of organic 

carbon in soil. 

Glob. Change Biol.

 

11,

 154–166 (2005).

39.  Nishina, K. 

et al.

 Quantifying uncertainties in soil carbon responses to changes in global mean temperature and precipitation. 

Earth 

Syst. Dynam.

 

5,

 197–209 (2014).

40.  Asseng, S. 

et al.

 Rising temperatures reduce global wheat production. 

Nature Clim. Change

 

5,

 143–147 (2015).

Page 16

Figures and page graphics, page 16

www.nature.com/scientificreports/

16

Scientific

 

RepoRts

 | 6:32525 | DOI: 10.1038/srep32525

41.  Battisti, D. S. & Naylor, R. L. Historical Warnings of Future Food Insecurity with Unprecedented Seasonal Heat. 

Science

 

323,

 

240–244 (2009).

42.  Brisson, N. 

et al.

 Why are wheat yields stagnating in Europe? A comprehensive data analysis for France. 

Field Crop. Res.

 

119,

 

201–212 (2010).

43.  Chmielewski, F. M., Muller, A. & Bruns, E. Climate changes and trends in phenology of fruit trees and field crops in Germany, 

1961–2000. 

Agric. For. Meteorol.

 

121,

 69–78 (2004).

44.  Estrella, N., Sparks, T. H. & Menzel, A. Trends and temperature response in the phenology of crops in Germany. 

Glob. Change Biol.

 

13,

 1737–1747 (2007).

45.  Lobell, D. B. & Field, C. B. Global scale climate - crop yield relationships and the impacts of recent warming. 

Environ. Res. Lett.

 

2,

 7 

(2007).

46.  Supit, I. 

et al.

 Recent changes in the climatic yield potential of various crops in Europe. 

Agric. Syst.

 

103,

 683–694 (2010).

47.  Ciais, P. 

et al.

 Europe-wide reduction in primary productivity caused by the heat and drought in 2003. 

Nature

 

437,

 529–533 (2005).

48.  Finger, R. Evidence of slowing yield growth - The example of Swiss cereal yields. 

Food Policy

 

35,

 175–182 (2010).

49.  Ewert, F., Rounsevell, M. D. A., Reginster, I., Metzger, M. J. & Leemans, R. Future scenarios of European agricultural land use I. 

Estimating changes in crop productivity. 

Agric. Ecosyst. Environ.

 

107,

 101–116 (2005).

50.  Olesen, J. E. 

et al.

 Impacts and adaptation of European crop production systems to climate change. 

Eur. J. Agron.

 

34,

 96–112 (2011).

51.  Smith, P. Carbon sequestration in croplands: the potential in Europe and the global context. 

Eur. J. Agron.

 

20,

 229–236 (2004).

52.  Dendoncker, N., Van Wesemael, B., Rounsevell, M. D. A., Roelandt, C. & Lettens, S. Belgium’s CO

2

 mitigation potential under 

improved cropland management. 

Agric. Ecosyst. Environ.

 

103,

 101–116 (2004).

53.  Smith, P. Agricultural greenhouse gas mitigation potential globally, in Europe and in the UK: what have we learnt in the last 20 

years? 

Glob. Change Biol.

 

18,

 35–43 (2012).

54.  Paustian, K. 

et al.

 Agricultural soils as a sink to mitigate CO

2

 emissions. 

Soil Use Manage.

 

13,

 230–244 (1997).

55.  Freibauer, A., Rounsevell, M. D. A., Smith, P. & Verhagen, J. Carbon sequestration in the agricultural soils of Europe. 

Geoderma

 

122,

 

1–23 (2004).

56.  Holland, J. M. The environmental consequences of adopting conservation tillage in Europe: reviewing the evidence. 

Agric. Ecosyst. 

Environ.

 

103,

 1–25 (2004).

57.  Vleeshouwers, L. M. & Verhagen, A. Carbon emission and sequestration by agricultural land use: a model study for Europe. 

Glob. 

Change Biol.

 

8,

 519–530 (2002).

58.  Gattinger, A. 

et al.

 Enhanced top soil carbon stocks under organic farming. 

Proc. Natl. Acad. Sci. USA

 

109,

 18226–18231 (2012).

59.  Whitmore, A. P., Kirk, G. J. D. & Rawlins, B. G. Technologies for increasing carbon storage in soil to mitigate climate change. 

Soil Use 

Manage

. 

31,

 62–71 (2014).

60.  Poeplau, C. & Don, A. Carbon sequestration in agricultural soils via cultivation of cover crops - A meta-analysis. 

Agric. Ecosyst. 

Environ.

 

200,

 33–41 (2015).

61.  Sierra, C. A. & Müller, M. A general mathematical framework for representing soil organic matter dynamics. 

Ecol. Monogr.

 

85,

 

505–524 (2015).

62.  Sanderman, J., Baisden, W. T. & Fallon, S. Redefining the inert organic carbon pool. 

Soil Biol. Biochem.

 

92,

 149–152 (2016).

63.  Agren, G. I. & Wetterstedt, J. A. M. What determines the temperature response of soil organic matter decomposition? 

Soil Biol. 

Biochem.

 

39,

 1794–1798 (2007).

64.  Bosatta, E. & Agren, G. I. Soil organic matter quality interpreted thermodynamically. 

Soil Biol. Biochem.

 

31,

 1889–1891 (1999).

65.  Leifeld, J. & Fuhrer, J. The temperature response of CO2 production from bulk soils and soil fractions is related to soil organic matter 

quality. 

Biogeochemistry

 

75,

 433–453 (2005).

66.  Conant, R. T. 

et al.

 Experimental warming shows that decomposition temperature sensitivity increases with soil organic matter 

recalcitrance. 

Ecology

 

89,

 2384–2391 (2008).

67.  Rey, A., Pegoraro, E. & Jarvis, P. G. Carbon mineralization rates at different soil depths across a network of European forest sites 

(FORCAST). 

European Journal of Soil Science

 

59,

 1049–1062 (2008).

68.  IUSS Working Group WRB (2006).

69.  Wiesmeier, M. 

et al.

 Amount, distribution and driving factors of soil organic carbon and nitrogen in cropland and grassland soils of 

southeast Germany (Bavaria). 

Agric. Ecosyst. Environ.

 

176,

 39–52 (2013).

70.  Richter, A., Adler, G. H., Fahrak, M. & Eckelmann, W. 

Erläuterungen zur nutzungsdifferenzierten Bodenübersichtskarte der 

Bundesrepublik Deutschland BÜK 1000 N

 (Bundesanstalt für Geowissenschaften und Rohstoffe (BGR) (2007).

71.  Poeplau, C. 

et al.

 Reproducibility of a soil organic carbon fractionation method to derive RothC carbon pools. 

European Journal of 

Soil Science

 

64,

 735–746 (2013).

72.  Amelung, W. & Zech, W. Minimisation of organic matter disruption during particle-size fractionation of grassland epipedons. 

Geoderma

 

92,

 73–85 (1999).

73.  Köhnlein, J. & Vetter, H. Ernterückstände und Wurzelbild - Menge und Nährstoffgehalt der auf dem Acker verbleibenden Reste der 

wichtigsten Kulturpflanzen. 

Verlag Paul Parey

 (1953).

74.  Bohne, H. Untersuchungen über die Ernterückstände des Getreides. 

Zeitschrift für Pflanzenernährung, Düngung und Bodenkunde

 

51(2),

 147–150 (1950).

75.  Bohne, H. Untersuchungen über die Bedeutung der Ernterückstände des Getreides für die Humusversorgung 

Zeitschrift für 

Pflanzenernährung, Düngung und Bodenkunde

 

55(2),

 170–178 (1951).

76.  Bommer, D. Untersuchungen über die Ernterückstände von Feldfutterpflanzen in verschiedenen Höhenlagen. 

Zeitschrift für Acker- 

und Pflanzenbau

 239–258 (1955).

77.  Gericke, S. Die Bedeutung der Ernterückstände für den Humusgehalt des Bodens. 2. Teil: Die Ergebnisse eigener Untersuchungen. 

Zeitschrift für Pflanzenernährung, Düngung und Bodenkunde

 

37,

 46–61 (1945).

78.  Könekamp, A. H. Die Rolle von Klee und Gras bei der Humusversorgung der Böden. 

Zeitschrift für Acker- und Pflanzenbau

 

104,

 

89–102 (1957).

79.  Könekamp, A. H.Teilergebnisse von Wurzeluntersuchungen. 

Zeitschrift für Pflanzenernährung, Düngung und Bodenkunde

 

60,

 

113–124 (1953).

80.  Könnecke, G. Forschungsaufgaben und Feldversuche 1953–1955 des Instituts für Acker- und Pflanzenbau der MLU Halle-

Wittenberg. (Martin-Luther-Universität Halle-Wittenberg, 1957).

81.  Simon, W., Eich, D. & Zajonz, A. Vorläufiger Bericht über Beziehungen zwischen Wurzelmenge und Vorfruchtwert bei 

verschiedenen Klee- und Grasarten als Hauptfrucht auf leichten Böden. 

Zeitschrift für Acker- und Pflanzenbau

 

104,

 71–88 (1957).

82.  Simon, W. Luzerne, Klee und Kleegras. 

VEB Deutscher Landwirtschaftsverlag

 (1960).

83.  Bolinder, M. A., Angers, D. A. & Dubuc, J. P. Estimating shoot to root ratios and annual carbon inputs in soils for cereal crops. 

Agric. 

Ecosyst. Environ

. 

63,

 61–66 (1997).

84.  Bolinder, M. A., Angers, D. A., Giroux, M. & Laverdiere, M. R. Estimating C inputs retained as soil organic matter from corn (Zea 

Mays L.). 

Plant Soil

 

215,

 85–91 (1999).

85.  Gan, Y. T. 

et al.

 Carbon input to soil from oilseed and pulse crops on the Canadian prairies. 

Agric. Ecosyst. Environ.

 

132,

 290–297 

(2009).

86.  Johnson, J. M. F., Allmaras, R. R. & Reicosky, D. C. Estimating source carbon from crop residues, roots and rhizodeposits using the 

national grain-yield database. 

Agron. J

. 

98,

 622–636 (2006).

Page 17

Figures and page graphics, page 17

www.nature.com/scientificreports/

17

Scientific

 

RepoRts

 | 6:32525 | DOI: 10.1038/srep32525

87.  Li, C. S. 

et al.

 Simulating trends in soil organic carbon in long-term experiments using the DNDC model. 

Geoderma

 

81,

 45–60 

(1997).

88.  Williams, J. D. 

et al.

 Root:shoot ratios and belowground biomass distribution for Pacific Northwest dryland crops. 

J. Soil Water 

Conserv.

 68, 349–360 (2013).

89.  Van der Linden, P. & Mitchell, J. F. B. ENSEMBLES: Climate Change and its Impacts: Summary of research and results from the 

ENSEMBLES project. 

Met Office Hadley Centre

 (2009).

90.  Christensen, J. H., Kjellstrom, E., Giorgi, F., Lenderink, G. & Rummukainen, M. Weight assignment in regional climate models. 

Clim. Res.

 

44,

 179–1794 (2010).

91.  Allen, R. G., Smith, M., Perrier, A. & Pereira, L. S. An update for the definition of reference evapotranspiration. 

ICID Bulletin

 

43,

 

1–34 (1994).

92.  Sierra, C. A., Müller, M. & Trumbore, S. E. Models of soil organic matter decomposition: the SoilR package, version 1.0. 

Geosci. 

Model Dev.

 

5,

 1045–1060 (2012).

93.  Lokupitiya, E. 

et al.

 Carbon balances in US croplands during the last two decades of the twentieth century. 

Biogeochemistry

 

107,

 

207–225 (2012).

94.  Kersebaum, K. C. & Nendel, C. Site-specific impacts of climate change on wheat production across regions of Germany using 

different CO

2

 response functions. 

Eur. J. Agron.

 

52,

 22–32 (2013).

95.  Aurbacher, J. 

et al.

 Influence of climate change on short term management of field crops - A modelling approach. 

Agric. Syst.

 

119,

 

44–57 (2013).

Acknowledgements

We would like to thank Ulrike Maul, Nadine Eheim, Wiebke Wehrmann und Sigrid Hiesch for laboratory 

analysis. We are grateful to the Federal Ministry for Education and Research for funding the BonaRes-Centre for 

Soil Research (BonaRes-Zentrum für Bodenforschung, Teilprojekt C, FKZ 031A608C) and to the Bavarian State 

Ministry of the Environment and Consumer Protection for funding the project “Der Humuskörper bayerischer 

Böden im Klimawandel – Auswirkungen und Potentiale”. This work was supported by the German Research 

Foundation (DFG) and the Technical University of Munich within the funding programme Open Access 

Publishing.

Author Contributions

M.W., M.v.L. and I.K.-K. designed the study. P.S., U.G., E.H. and B.S. provided soil samples and soil data. C.P. 

and C.A.S. performed the modeling and H.M. and C.F. provided climate projections. M.W., C.P., R.H. and A.K. 

compiled and analyzed data. M.W. wrote the main manuscript with contributions of C.P., H.M., C.F., R.H. and 

M.v.L. All authors reviewed the manuscript.

Additional Information

Competing financial interests: 

The authors declare no competing financial interests.

How to cite this article

: Wiesmeier, M. 

et al.

 Projected loss of soil organic carbon in temperate agricultural soils 

in the 21

st

 century: effects of climate change and carbon input trends. 

Sci. Rep.

 

6

, 32525; doi: 10.1038/srep32525 

(2016).

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users will need to obtain permission from the license holder to reproduce the material. To view a copy of this 

license, visit http://creativecommons.org/licenses/by/4.0/

 

© The Author(s) 2016


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