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Microbial Biomass

Chapter · June 2009

DOI: 10.1007/b82392_7

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Chapter 6
Microbial Biomass

R. Brumme, M. Raubuch, J. Priess, C. P. Wang, and T.–H. Anderson

6.1 Introduction

Micro-organisms (bacteria, fungi) contribute to more than 90% of the carbon

dioxide evolved during decomposition of forest litter (Schaefer 1991), indicating
their role in the element cycling in forest ecosystems. Nitrogen is mostly cycled
between primary producers and decomposer biota. The activity and growth of
micro-organisms depend on carbon and nutrient supply and the physico-chemical
environment because of their high surface area/volume ratio of microbial bodies
(Hattori and Hattori 1976; Paul and Clark 1996). Soil acidity has often been shown
to reduce microbial biomass (Cmic) and to increase metabolic respiration (qCO2,
ratio between microbial-respiration-C and microbial-biomass-C) in the surface
mineral soil which was interpreted as an increased stress on micro-organisms living
in acid soils (Anderson and Domsch 1993; Anderson, Chap. 20, this volume). In
addition to the chemical stress, soil acidity may change the litter quality, litter
amount and their distribution in the soil profile by acid sensitive earthworms.
Quality and amount of litter determine the amount of micro-organisms that would
live on the organic matter, and is indicated by the fraction of Cmic in organic carbon
(Corg) (Anderson and Domsch 1986). Another effect of soil acidity relates to an
increase in bacterial respiration with increasing pH. Less is known of microbial
biomass and the mentioned relationships in the surface organic layer and deeper soil
horizons. Here, we provide depthwise distribution of microbial carbon (Cmic),
microbial nitrogen (Nmic), metabolic quotient (qCO2), the fraction of Cmic in Corg
(Cmic-to-Corg ratio) and the fraction of fungal respiration of soil respiration in three
beech forest soils, the Göttinger Wald, Zierenberg, and the Solling sites (for site
description see Part A). The following questions will be addressed. (1) Does soil
acidity reduce the microbial biomass and the qCO2 value in the acid soil profile at
Solling site compared to the less acid soils at Göttinger Wald and Zierenberg sites?
(2) Does better substrate quality at Göttinger Wald and Zierenberg sites increase the
Cmic-to-Corg ratio? (3) What is the contribution of bacteria and fungi to soil
respiration and how does soil acidity affect these contributions?

R. Brumme and P.K. Khanna (eds.), Functioning and Management of European Beech 87
Ecosystems, Ecological Studies 208,
DOI: 10.1007/978‐3‐642‐00340‐0_6, # Springer-Verlag Berlin Heidelberg 2009
88 R. Brumme et al.

6.2 Microbial Carbon and Nitrogen

Microbial carbon (Cmic), measured by substrate-induced respiration method (SIR)

(Anderson and Domsch 1978; Heilmann and Beese 1992) in May 1995, was the
highest in the L layers (17–18.9 mg Cmic g 1) and showed no difference among the
three beech forest sites (Table 6.1). In the moder profile microbial biomass-C
decreased from 18.9 mg g 1 in the L layer to 0.97 mg g 1 in the H layer at the
Solling site. In the mineral soils, the lowest Cmic value was found at the Solling site
(about 0.1 mg Cmic g 1), and it did not change down to 20 cm depth. The microbial
biomass values in the uppermost base-rich Göttinger Wald and Zierenberg soils
were up to 10 times higher than those at the Solling site and decreased to about
0.2 mg Cmic g 1 in the 15–20 cm depth. These values are in the range observed for
soils under beech and oak forests in northern Germany. Anderson (Chap. 20, this
volume) reported values for microbial biomass in the range of 0.18–3.12 mg Cmic
g 1 in 0–5 cm depth.
Similar microbial biomass values were found in the base-rich soils at the
Göttinger Wald (988 kg Cmic ha 1) and Zierenberg (919 kg Cmic ha 1) sites for L
to 20 cm soil depth out of which 11% and 36% were attributed to the surface
organic layers of the two sites, respectively. The microbial biomass at the Solling
site was 42% and 48% lower (532 kg Cmic ha 1) than in the base-rich Zierenberg
and Göttinger Wald soils. A high fraction of microbial biomass was present in the
surface organic layer (67%) of the Solling site but it did not cover for the low values
in the mineral soil. Similar values for the microbial biomass were reported for other
European forest soils (148–619 kg Cmic ha 1 in L to 22 cm depth, micro-calorime-
ter method) by Raubuch and Beese (1995). They reported ratios of Cmic in the
surface organic layer and the surface mineral soils (0–22 cm) between 0.3 in less
acid to 1.9 in the highly acid soils, and emphasised the contribution of microbial
decomposition in the surface organic layer.

Table 6.1 Microbial biomass carbon (substrate-induced respiration method, SIR) and pH(H2O)
values in the soil profile of three beech forests (standard error in parentheses) (n = 5) (adopted from
Chodak et al. 2003)
Göttinger Wald Zierenberg Solling
pH C pH C pH C
mic mic mic

mg g 1
kg ha 1 mg g 1 kg ha 1 mg g 1 kg ha 1

L – 17.0 (0.57) 104 – 17.9 (0.54) 329 – 18.9 (1.31) 209

F – – – – – – – 2.75 (0.37) 94
H – – – – – – 3.4 0.97 (0.05) 53
0–5 5.0 1.18 (0.17) 370 5.6 0.83 (0.05) 250 3.4 0.11 (0.05) 37
5–10 4.7 0.51 (0.07) 233 5.4 0.51 (0.06) 202 3.7 0.13 (0.01) 52
10–15 4.8 0.30 (0.04) 146 5.5 0.28 (0.03) 85 4.0 0.12 (0.04) 49
15–20 5.1 0.26 (0.03) 135 5.7 0.18 (0.02) 53 4.2 0.09 (0.1) 38
P
L 20 988 919 532
6 Microbial Biomass 89

Table 6.2 Microbial biomass nitrogen and Cmic/Nmic ratios (chloroform fumigation extraction
method, CFE) in the soil profile of Göttinger Wald and Solling forests (standard deviation in
parentheses) (n = 3)
Göttinger Wald Solling
Nmic Nmic
1 1 1 1
mg g kg ha Cmic/Nmic mg g kg ha Cmic/Nmic
L 2.06 (0.65) 1.26 8.6 1.92 (1.71) 2.12 10.8
F – – – 0.98 (0.41) 3.35 7.4
H – – – 0.40 (0.10) 2.20 8.6
0–5 0.30 (0.04) 9.47 5.3 0.06 (0.01) 1.87 10.5
5–10 0.18 (0.05) 8.12 6.1 0.03 (0.02) 1.07 10.2
10–20 0.10 (0.04) 10.4 7.0 0.02 (0.02) 1.91 8.8
P
L 20 29.3 12.5

Microbial nitrogen, measured with the chloroform fumigation and extraction

method (Brookes et al. 1985a, b), showed a similar distribution in the soil profiles at
the Göttinger Wald and Solling sites as microbial carbon. However, microbial
nitrogen was very low in the acid Solling soil as indicated by higher Cmic/Nmic
ratios than those at the Göttinger Wald site in all soil layers (Table 6.2). Mostly,
Cmic/Nmic ratios of 3–6 in agricultural and forest soils have been reported which are
independent of land use pattern (Jenkinson 1988; Paul and Clark 1996). However,
acid forest soils have much higher Cmic/Nmic ratios of up to 17, and a much larger
variation, than the agricultural soils (Joergensen et al. 1995). The fungi, which are
the primary decomposers of forest litter, could have a much higher C/N ratio (4–15,
Paul and Clark 1996) which may explain the higher Cmic/Nmic ratio observed in the
acid forest soil of the Solling site. In a study on ten acid and ten neutral surface
mineral soils under beech, a high fungal-to-bacterial respiratory fraction ranging
from 74 to 26 at high pH of 6 and from 94 to 6 at low pH of 3 were reported by
selective inhibition of either fungal or bacterial metabolism (Blagodatskaya and
Anderson 1998). The Zierenberg site showed a similar pH effect on the fungal-to-
bacterial respiration from the basalt-dominated upper hill (main research area) to
the lime dominated lower hill area. With increasing pH from the basalt to the lime-
dominated soil, the fungal-to-bacterial respiration fraction decreased from 87 to 13
(pH(KCl) 4.1), 76 to 24, (pH(KCl) 5.5), and to 55 to 45 (pH(KCl) 6.4) (Fig. 6.1).

6.3 Metabolic Quotient and Cmic-to-Corg Relationship

Microbial carbon contributes between 0.3 and 7% of soil organic carbon (SOC)
(Cmic-to-Corg) (Anderson and Domsch 1989) with mean values of 1.6% at Göttinger
Wald, 1.9% at Zierenberg and 0.68% at Solling for L–20 cm depth in the soils. The
L layers contained the highest content between 3.6% and 4.7% and low values of
0.6%–1.1% in the 15–20 cm depth (Fig. 6.2). The Cmic-to-Corg ratio is an indicator
90 R. Brumme et al.

100 7

6
80
% fungal respiration % fungal
respiration 5
pH
60

pH [KCl]
4

3
40

2
20
1

0 0
basalt intermediate lime
dominated dominated
soil soil

Fig. 6.1 Fraction of fungal respiration to total respiration (bacterial plus fungal) determined by
selective inhibition with standard deviation and soil pH (KCl) at Zierenberg in three areas: from
basalt-dominated (main research area), lime-dominated area and intermediate area containing both
basalt and limestone (n = 5 6)

L
F
H
0–5
SO
5–10
ZB
10–15
15–20 GW

0 1 2 3 4 5
% Cmic in Corg

Fig. 6.2 %Cmic in Corg in the soil profiles at the Solling (SO), Zierenberg (ZB) and Göttinger Wald
(GW) sites

of how much of the organic carbon is used for the growth of microbial cell mass
(Anderson and Domsch 1986). The higher the Cmic-to-Corg fractions, the higher the
amount of carbon that is present in available form and the higher the amount of
micro-organisms that can live on the substrate. Thus, low Cmic-to-Corg ratio with
increasing soil depth indicated low substrate availability. Fast incorporation of leaf
litter by earthworms provided micro-organisms with easily available substrate in
the base-rich Göttinger Wald and Zierenberg mineral soils (Schaefer and Schauermann,
Chap. 7, this volume) especially from the luxuriant ground vegetation (Schmidt,
Chap. 5, this volume), and explained the slow decrease of the Cmic-to-Corg ratio with
soil depth in contrast to no change in the mineral layers of the acid Solling site
6 Microbial Biomass 91

L
SO
F
ZB
H
0–5 GW
5–10
10–15
15–20

0 1 2 3 4 5
qCO2 (µg CO2-C mg–1 Cmic h–1)

Fig. 6.3 Metabolic quotient (qCO2, mg mg 1 Cmic h 1) in the soil profiles at the Solling (SO),
Zierenberg (ZB), and Göttinger Wald (GW) sites (adopted from Chodak et al. 2003)

(Fig. 6.2). The strong decrease in Cmic-to-Corg ratio from L to F layers at the Solling
site was surprising because of the still discernible structure of the F layer suggesting
that the substrate was highly available for micro-organisms. However, similar
values for Cmic-to-Corg in the acid mineral soil and the F layer indicated that the
microbial availability of the F substrate was similar to that of SOM in the mineral
soil.
High metabolic quotient values (CO2–C/Cmic or qCO2) were found in the
mineral soil layers of the acid Solling site (3.7–4.4 g CO2–C mg 1 Cmic h 1)
(Fig. 6.3). The base-rich soils at the Göttinger Wald and Zierenberg sites had
about 2 times lower qCO2 values in the mineral soils indicating a much lower
soil chemical stress to micro-organisms by aluminum, protons and heavy metals
(Anderson and Domsch 1993; Duxbury 1995; Wood 1995).

6.4 Conclusions and Indications of Human Impacts

l The acid soil at the Solling site has the lowest microbial biomass, the lowest
Cmic-to-Corg ratio and the highest qCO2 value in all mineral soil layers of the
three beech forest soils. High soil acidity, low bioturbation and sparse ground
vegetation at the Solling site affected these parameters.
l Total microbial biomass was about 45% lower at the acid Solling site than at the
base-rich Göttinger Wald and Zierenberg sites. About two-thirds of the total
microbial biomass was present in the surface organic layer at Solling site
whereas more than two-thirds were present in the base-rich mineral soils of
the Göttinger Wald and Zierenberg sites.
l The luxuriant herb layer increased the substrate availability for micro-organisms
(high Cmic-to-Corg ratio) in all mineral soil layers at the Göttinger Wald and
Zierenberg sites by increasing earthworm activity. Low Cmic-to-Corg ratios
indicated a low substrate availability in the acid mineral soil layers and the
 92 R. Brumme et al.

strong decrease from L to the F/H layers suggested a rapid stabilisation of the
remaining litter in the F layer.
l The fraction of fungal respiration decreased with increasing pH from a basalt-
dominated soil (87% at pH(KCl) of 4.1) to a lime-dominated soil (55% at pH
(KCl) of 6.4) along an acidity gradient at Zierenberg site.

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