Soil Biology & Biochemistry 38 (2006) 20112024

Decomposition in peatlands: Reconciling seemingly contrasting results

on the impacts of lowered water levels
Raija Laiho

Peatland Ecology Group, Department of Forest Ecology, University of Helsinki, P.O. Box 27, FIN-00014 Helsinki, Finland
Received 9 September 2005; received in revised form 3 February 2006; accepted 22 February 2006
Available online 4 April 2006
Abstract
Northern peatlands represent about 30% of the global soil C pools. The C pool in peat is a result of a relatively small imbalance
between production and decay. High water levels and the consequent anoxia are considered the major causes for the imbalance. As such,
the C sink of a peatland is labile, and sensitive to disturbances in environmental conditions.
Changes in peatland ecosystem functions may be mediated through land-use change, and/or climatic warming. In both cases, lowering
of the water level may be the key factor. Logically, lowered water levels with the consequent increase in oxygen availability in the surface
soil may be assumed to result in accelerated rates of organic matter decomposition. Yet, earlier research has given highly contrasting
results concerning the effects of lowered water levels on the rates of decomposition and the C sink/source behaviour of peatlands. The
mechanisms controlling this variation remain unresolved.
This paper summarizes the changes observed in the biotic and abiotic controls of decomposition following natural or articial lowering
of peatland water levels and show that they are complex and their interactions have not been previously explored. Long-term changes in
the C cycle may differ from short-term changes. Short-term changes represent a disturbance in the ecosystem adapted to the pre-water-
level-lowering conditions, while long-term changes result from several adaptive mechanisms of the ecosystem to the new hydrological
regime. While in a short term, the disturbed system will always lose C, the long-term changes inherently vary among peatland types,
climates, and extents of change in the water level. The paper closes by identifying the gaps in our knowledge that need to be addressed
when proceeding towards a causal and unifying explanation for the C sink/source behaviour of peatlands following persistent lowering of
the water level.
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Keywords: C cycling; Decomposition dynamics; Decomposer communities; Hydrological change; Litter quality; Long-term change; Secondary succession;
Short-term change; Wetland ecology
1. Introduction
Peatlands represent a wide variety of wetlands that are
characterized by an organic soil, but differ in hydrology,
chemistry, and, consequently, vegetation composition.
Northern peatlands have been a signicant sink of carbon
(C) from the atmosphere, representing about 30% of the
global soil C pools with their estimated reservoir of 455 Pg
(10
15
g) (Gorham, 1991). This has been achieved by a
relatively small imbalance between production and decay:
only 216% of the net primary production of a peatland
ecosystem gets deposited as peat over centuries or millennia
(Pa iva nen and Vasander, 1994). High water levels and the
consequent anoxia, accompanied with low soil tempera-
tures are considered the major causes for the imbalance. As
such, the C sink of a peatland is labile, and sensitive to
variations in environmental conditions (Alm et al., 1999a;
Grifs et al., 2000; Bubier et al., 2003a).
Peatlands slowly change along their successional develop-
ment, which is regulated by both allogenic and autogenic
factors (e.g., Klinger and Short, 1996; Hughes and Du-
mayne-Peaty, 2002). Relatively rapid changes in peatland
ecosystem functions may be mediated through land-use
change, and/or climatic warming. In both cases, lowering of
the water level is a key factor (Gitay et al., 2001). Quite
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E-mail address: raija.laiho@helsinki..
logically, lowered water levels with the consequent increase in
oxygen availability in the surface soil may be assumed to
result in accelerated rates of organic matter decomposition.
During warm, dry summers that have resulted in temporarily
lowered water levels, substantial C losses from boreal or
subarctic bogs and fens have been observed (e.g., Schreader
et al., 1998; Alm et al., 1999a; Moore et al., 2002). These
short-term observations have strengthened the prevailing
paradigm, that persistent lowering of the water level will
reduce the C sink of a peatland, and eventually turn it into a
source of C into the atmosphere (Armentano and Menges,
1986; Silvola, 1986).
In accordance with the paradigm, increased CO
2
efuxes
have been measured from peatlands where a persistent
lowering of the water level has been induced by ditching
(i.e., those drained for forestry) (Glenn et al., 1993;
Martikainen et al., 1995; Alm et al., 1999b). In great
contrast, an extensive Finnish inventory on long-term (56
decades) changes in drained peatlands indicated that on
average, peat C stores had increased following drainage
(Fig. 1; Minkkinen and Laine, 1998b; Minkkinen et al.,
2002). In individual peatlands, both decreases and in-
creases in peat C stores following long-term drainage have
been observed (Sakovets and Germanova, 1992; Vompers-
ky et al., 1992; Minkkinen and Laine, 1998b; Minkkinen et
al., 1999). In accordance with these contradicting results, in
the few eld experiments on drainage-induced changes in
decomposition rates, increase, decrease, and no change
have all been observed (Lieffers, 1988; Minkkinen et al.,
1999; Domisch et al., 2000; Laiho et al., 2004).
Thus, earlier research may be summarized as follows:
after persistent lowering of the water level, a peatland site
may become a source of C into the atmosphere, remain a
sink, or become a stronger sink. To some extent, these
differences may be linked to peat soil nutrient level
(vegetation type) and climatic conditions (Minkkinen and
Laine, 1998b; Minkkinen et al., 1999); however, the
mechanisms controlling this variation remain unresolved.
The aim of this paper is to synthesize current informa-
tion on the changes, induced by natural or articial
lowering of the water level, in the biotic and abiotic factors
affecting decomposition processes in peatlands. As the
factors affecting aerobic decomposition in the surface
layers may be considered to experience more changes than
those affecting anaerobic decomposition deeper down,
more emphasis will be given on those. The ultimate goal is
to proceed towards a causal and unifying explanation for
the observed variation in the C sink/source behaviour of
peatlands following lowering of the water level. The review
will limit on changes that might be discernible during the
rst 12 centuries, and will not examine peat accumulation
and C sequestration over millennia, which is the time-scale
of peatland succession generally.
Decomposition means mass loss of organic matter as gas
or in solution, caused by either leaching or consumption by
saprotrophic organisms. Terms decomposition, decay,
breakdown and humication have been used in the
literature, some times with varying emphasis (see Clymo,
1984). Here, only the term decomposition is used irrespec-
tive of the original terminology used by the authors in the
studies referred to. Water level refers to the distance from
mire surface to the saturated layer, which shows as a free
water surface in a well, or tube inserted in the peat.
2. Constraints for decomposition in pristine peatlands
Generally, decomposition of organic matter depends on
four factors: substrate quality, environmental conditions,
decomposers present, and nutrient availability. Nutrient
availability is determined by both substrate and environ-
mental characteristics: If decomposer fungi do not get
enough of nutrients from the substrate that they utilize as
an energy source, they may in some cases translocate it
from the surroundings (e.g., Lindahl et al., 2001). These
four factors all interact: environmental conditions and
nutrient availability regulate the vegetation composition
(Wheeler and Proctor, 2000; kland et al., 2001), which in
turn largely determines substrate quality (Hobbie, 1996),
which then, together with environmental conditions and
possibly nutrient availability, regulates the composition of
the decomposer community (Borga et al., 1994; Panikov,
1999).
Substrate quality has usually been described as concen-
trations of C fractions (solubles, holocellulose, lignin), and/
or nutrients. The effects of substrate quality on decom-
position can be studied by incubating different litter types
in constant environmental conditions in the laboratory, or
in similar conditions within a site in the eld. Generally,
litters with high concentrations of soluble C compounds,
ARTICLE IN PRESS
Region
1 2 3 4 5
C
h
a
n
g
e

i
n

s
o
i
l

C

p
o
o
l
,

k
g

C

m
-
2

-30
-20
-10
0
10
20
30
40
Intermediate
Moderately poor
Extremely poor
*
*
*
*
*
*
*
(South) (North)
Fig. 1. Change in total peat soil C pool over 5060 years following
drainage for forestry in pine fens representing different nutrient levels, as
reported by Minkkinen and Laine (1998). Stars show where the change
was signicantly different from zero. Region codes: 1 South Finland,
2 Central Finland, 3 Eastern Finland, 4 Northern Finland
excluding Lapland, 5 Lapland. Redrawn from Minkkinen and Laine
(1998). Nutrient levels: intermediate fens correspond to mesotrophic,
moderately poor to oligotrophic, and extremely poor to oligo-ombro-
trophic.
R. Laiho / Soil Biology & Biochemistry 38 (2006) 20112024 2012
low concentrations of lignin or lignin-like phenolic poly-
mers, and high nutrient concentrations decompose rela-
tively fast (Bartsch and Moore, 1985; Szumigalski and
Bayley, 1996; Preston and Trofymow, 2000; Scheffer et al.,
2001; Thormann et al., 2001; Limpens and Berendse, 2003).
The temporal patterns of these constraints may differ
(Berg, 1984; Sariyildiz and Anderson, 2003), and they may
also vary among sites (Berg et al., 1993). In the peatland
context, leaf and root litters from herbs and sedges
decompose relatively fast (5175% mass loss during 2
years), moss litters decompose slowly (932% mass loss
during 2 years) and arboreal litters stand in between (mass
loss examples from Szumigalski and Bayley, 1996; Scheffer
et al., 2001; Thormann et al., 2001). The material used in
the experiments: live, recently dead, or long dead, may
have a considerable effect on the results (Limpens and
Berendse, 2003).
The effects of environmental factors can be studied by
incubating cellulose and/or standard litter materials in
varying conditions. Laboratory studies may yield single
effects of different environmental factors, while eld
studies yield the sum of all factors in effect. The most
important environmental conditions that constrain decom-
position in peatlands are generally considered to be oxygen
availability, temperature, and acidity. Accordingly, labora-
tory experiments have shown that within the range of
environmental conditions found in peatlands, CO
2
evolu-
tion, an indicator of decomposition, from peat samples
generally increases with lowering water level (Blodau et al.,
2004), increasing pH and increasing temperature (Bergman
et al., 1999).
Extracellular enzymes excreted by certain fungi, actino-
myces and bacteria to decompose high molecular weight
complex compounds, are direct indicators of decomposer
activity. Freeman et al. (2001a, 2004) have shown that the
activity of phenol oxidase may be a key regulator of
peatland C cycling. Under low phenol oxidase activity,
phenolic compounds accumulate in the soil, which inhibits
hydrolase enzymes crucial for organic matter decomposi-
tion (Freeman et al., 2001a, 2004). The activity of phenol
oxidase is strongly constrained by the presence of
bimolecular oxygen (Freeman et al., 2001a, 2004). Further,
phenol oxidase activity increases with increasing tempera-
ture (Q10 1.36 in a thermal gradient of 2201C, Freeman
et al., 2001b), and is limited by low pH (Pind et al., 1994;
Williams et al., 2000). Yet, Williams et al. (2000) reported
that phenol oxidase activity in Sphagnum peat, especially,
was regulated less by aeration and more by pH, and when
pH was favourable, phenol oxidase activity depended more
on wetland vegetation type and botanical composition of
the peat than water level. Thus, the role of phenol oxidase
may vary among peatland types.
In general, comparisons among the plentiful eld studies
on decomposition rates in different pristine peatland sites
yield few generally valid conclusions on the impacts of the
environmental factors on decomposition. Differences in the
decomposition rates of standard litters have been observed
for sites with differing surface water chemistry: mass losses
have correlated negatively with water levels and, in
contrast to laboratory results, also pH, while positive
correlations with base cation status, soluble N and/or
soluble P have been observed (Verhoeven et al., 1990, 1996;
Verhoeven and Toth, 1995; Szumigalski and Bayley, 1996;
Thormann and Bayley, 1997; Scheffer and Aerts, 2000;
Thormann et al., 2001). The patterns have varied among
studies and litter types within studies. This may in part be
due to the decomposer communities being subject to
multiple limitations (Panikov et al. 1997). It would
generally seem that litter type actually exerts a greater
effect on aerobic decomposition than the variation in
environmental conditions among or within peatland sites
(Bartsch and Moore, 1985; Szumigalski and Bayley, 1996;
Scheffer et al., 2001). On the other hand, different litter
types are often produced under different environmental
conditions, and the combined effects of litter quality and
environment on decomposition may be complex (Belyea,
1996). Generally, more readily decomposable litters are
produced under relatively wet conditions (fens vs. bogs,
and hollows vs. hummocks within bogs), and C accumula-
tion may take place because the time for aerobic
decomposition remains short. In contrast, under relatively
dry conditions, less readily decomposable litters are
produced, and the rate of C accumulation is often faster
in spite of the longer time available for aerobic decom-
position (e.g., Turunen et al., 2002).
There are only few studies that link the decomposers
present in peatland sites to the decomposition of different
substrates. Thormann et al. (2002) observed variation in
the ability of nine species of lamentous microfungi
isolated from Sphagnum fuscum
1
to decompose different
C compounds or S. fuscum and wood chips. Further,
Thormann et al. (2003) found different fungal assemblages
decomposing different litter types in two peatland sites.
They concluded that litter quality variables were more
important to the variation in the fungal assemblages than
physical or surface water chemistry variables. Williams and
Crawford (1983) have demonstrated high physiological
diversity among peatland microbes: some decomposers
may metabolise well even in cold and very acid conditions.
Coulson and Buttereld (1978) suggested that a major
factor in the slow decomposition of Sphagnum mosses and
Eriophorum vaginatum is their unattractiveness to soil
animals. These results again emphasize the role of litter
quality in decomposition processes.
Decomposition rates usually decrease from the surface
downwards, the biggest change taking place when condi-
tions change to anoxic. This is due to both the substrate
becoming more recalcitrant and the conditions becoming
less favourable (Belyea, 1996). There is often a secondary,
or even primary, peak in decomposition rates at the range
of the water level variation (Tuominen, 1981; Santelmann,
ARTICLE IN PRESS
1
Nomenclature follows Moore (1982) for vascular plants and Koponen
et al. (1977) for mosses.
R. Laiho / Soil Biology & Biochemistry 38 (2006) 20112024 2013
1992; Belyea, 1996), especially if the water level remains
below 20 cm for most of the year, as is often the case in
hummocks (Fig. 2; Tuominen, 1981).
The average water level in northern peatlands generally
varies from a few to around 40 cm below the surface (e.g.,
Gorham and Janssens, 1992; Laine et al., 2004). At a height
growth rate of 2.5 mmyear
1
of the mire surface, it would
take 100 years for litter deposited on the surface to reach a
depth of 25 cm. This calculation is grossly simplied, of
course, as the water level varies seasonally and moss and
root litters are deposited below the mire surface. In any
case, it suggests that in dry sites, many litter types (see
Latter et al., 1998; unfortunately, there are no such data for
mosses) have well reached their maximal mass losses (see
Berg and Meentemeyer, 2002) before entering the anoxic
zone, unlike in wet sites. If lowering in the water level, and
thickening of the oxic zone, were the only changes taking
place, the most important question from the decomposition
point of view would be how much decomposition
potential the substrate exposed for the oxic conditions
would still have.
3. Methods for studying the effects of persistent lowering of
the water level
The effects of persistent lowering of the water level can
be studied using three approaches. Short-term results may
be obtained from laboratory incubations. Because some of
the factors in effect in the nature or their interactions are
always excluded, laboratory incubations fail in producing
reliable long-term results. Long-term results may be
obtained in eld studies where the lowering of the water
level has been achieved by ditching. In such studies it is
important to be certain that the ecosystem has not been
manipulated in other ways, like by fertilization, or
vegetation removal or control. Forestry drainage methods
applied in northern Europe and Canada often, but not in
all cases, full this requirement. Mesocosm studies where
relatively large and intact peat monoliths with their natural
vegetation are brought under controlled conditions provide
an intermediate approach. This method allows monitoring
the existing system for a fairly long time; however, it does
not enable plant, possibly also microbial, community
dynamics in the way they would occur in natural
conditions. In the eld, invasion of species present in other
habitats of the peatland or nearby uplands is possible
unlike in the mesocosms. All approaches have their
limitations, and when drawing conclusions one has to
pay special attention to the time scale applied and to the set
of factors that can be considered to show their true
impacts.
Water levels in a peatland usually uctuate within a
certain range, depending on season and weather condi-
tions. Such uctuations have their specic effects on
decomposition processes, and these may differ from those
of a persistent change (Moore and Dalva, 1993; Nizovtseva
et al., 1995; Aerts and Ludwig, 1997; Corstanje and Reddy,
2004). The focus in the following will be on a persistent
change in average water levels.
4. Effects on decomposition rates
4.1. Peat soil
There are no studies where in situ dynamics of peat
decomposition following lowering of the water level would
have been reported; however, inferences on soil C balance
have been drawn from soil respiration studies. These are
complicated by the difculties in separating the specic
sources of the CO
2
uxes: autotrophic versus heterotrophic
respiration. Further, what can we consider peat soil?
Possibly a major part of soil respiration originates from the
recently deposited litter instead of peat proper (Malmer
and Holm, 1984). For instance, Hogg (1993) observed that
even under favourable conditions in the laboratory, CO
2
release from recently deposited Sphagnum litter was nearly
twice as high as that from older material just 10 cm below.
Generally, up to two-fold CO
2
emissions have been
observed in sites where water levels have been articially
lowered, as compared with the controls (Silvola et al.,
1996a; Chimner and Cooper, 2003). Much higher increases
have been observed in laboratory studies (Moore and
Knowles, 1989; Moore and Dalva, 1993; Blodau and
Moore, 2003). A linear relation between CO
2
emission and
water level has been observed in laboratory conditions
(Moore and Dalva, 1993). In situ, the increase in CO
2
emission has mostly been seen only with a lowering to a
certain depth, between 10 and 3040 cm depending on the
study, with no further increase with a further lowering
(Silvola et al., 1996a; Chimner and Cooper, 2003). Chimner
and Cooper (2003) suggested the lack of easily oxidized
labile C in the deeper soil layers as a reason for this pattern.
This is supported by the results of Hogg et al. (1992) that in
drained samples in vitro the release of CO
2
was about 10
times greater from 010 cm peat layer than from 3040 cm
ARTICLE IN PRESS
Water-level
Decomposition potential
D
e
p
t
h
Lawn
Hummock
Surface
Increases
Fig. 2. Schematic presentation of the decomposition potential relative to
water level in a hummock versus a lawn. Based on cellulose decomposition
values presented by Tuominen (1981) for a moderately poor, sedge pine
fen. Note that the actual position of the water level is not the same for the
two cases.
R. Laiho / Soil Biology & Biochemistry 38 (2006) 20112024 2014
layer, which they attributed to the relatively large pool of
non-structural carbohydrates in surface samples, deriving
from recently dead plant biomass.
Hogg et al. (1992) suggested that generally, C losses from
peat following water level drawdown essentially depend on
peat quality. Peats that have already been exposed to long
periods of aerobic decomposition may be highly resistant
to further decomposition (Bridgham and Richardson,
1992). Consequently, lowered water levels might not cause
a clear increase in C release from peatlands that already
have water tables 20 cm or more below the peat surface for
most of the summer. Many treed bogs and fens would have
such a water regime. In contrast, in peatlands where
organic strata near the surface have been continuously
inundated, the peat with easily oxidizable labile C could
decompose at increased rates. Wet open fens would most
often fall into this category. These postulates are supported
by the results of Silvola et al. (1996a), Minkkinen et al.
(1999) and Laeur et al. (2005).
It seems evident based on the results reviewed that
lowering of the water level is followed by at least short-
term C losses from the soil, when recently deposited
relatively easily decomposable organic matter is decom-
posed. Further, CO
2
stored in the soil prole may be
released as the water-level drops (Moore and Dalva, 1993).
In the laboratory studies of Moore and Dalva (1993) and
Blodau and Moore (2003) the increased CO
2
emissions
lasted for a few weeks following water level drawdown,
when obviously the readily utilizable substrate was
consumed. The Scottish peatlands studied by Hargreaves
et al. (2003) acted as sources of C for 24 years after
draining, ploughing and afforestation; 48 years after tree
planting they acted again as sinks. Nonetheless, there are
little bases yet for estimating the extent and duration of the
increased C losses from different types of peat in situ, that
are caused by different kinds of changes in the hydrological
regime (extent of lowering in the water-level, extent of
uctuations). Further, the properties of the substrate
available for decomposition will change dynamically with
the inputs of new litter.
Attempts have also been made to directly estimate the
changes that have taken place in soil C pools over time
following drainage (Sakovets and Germanova, 1992;
Vompersky et al., 1992; Minkkinen and Laine, 1998b;
Minkkinen et al., 1999). These comparisons, that involve
many assumptions, cover both the decomposition of the
peat soil that existed at the time of drainage, and the inputs
and decomposition of the litterfall that took place after
drainage. The old peat can only decompose, but
estimating the rate of peat decomposition specically
would require means for estimating the inputs and
decomposition of the new litters (Vompersky et al.,
1992). Rather surprisingly, the results of these studies
suggest that in most cases, the C balance would have
remained positive. This would mean that the new litter
inputs exceeded the decomposition losses from both the
old peat and the new litters (Sakovets and Germano-
va, 1992; Vompersky et al., 1992). The C pool estimates
include living ne roots, which may contribute up to
1 kg m
2
of C (Sjo rs, 1991; Laiho and Fine r, 1996).
Silvola et al. (1996b) estimated that the contribution of
plant roots to the CO
2
uxes was 3545% of total soil
respiration in the middle and late summer at sites with
abundant vegetation. The root contribution correlated
with tree stand volume. The root proportions were likely
underestimates; the respiration from the plots where roots
had been excluded would still include extra C from the
killed roots during the summer after root isolation. The
lack of a long-term response of CO
2
emissions to water
level in the treeless mesocosms of Updegraff et al. (2001)
further suggests that tree roots are a major source of CO
2
emissions in forested sites (see also Ho gberg et al., 2001). If
the root contribution were about 50% in the sites of Silvola
et al. (1996a) with the highest soil respiration, the
remainder would be approximately in balance with the
above-ground litter inputs estimated by Laiho et al. (2003)
for a most closely corresponding site (pine-dominated, with
ample moss and shrub vegetation). This comparison lends
some support to the observations mentioned above that in
spite of increased CO
2
emissions, the C balance of peat
may not always be negative in drained sites.
4.2. Litter and cellulose
In all studies comparing rates of decomposition in
drained versus pristine peatlands so far, either cellulose
(Lieffers, 1988; Minkkinen et al., 1999) or standard litter
materials (Lieffers, 1988; Domisch et al., 2000; Laiho et al.,
2004) have been used. Thus, the results largely describe the
changes in decomposition potential, i.e., they summarize
the changes in environmental conditions.
Most of these studies have been conducted in treed fens
that have been common objects for forestry drainage. Yet,
the patterns of change that arise from these studies are not
unambiguous. In an Alberta site drained 2 years pre-
viously, both Sphagnum warnstori sections and cellulose
decomposed faster than in its pristine counterpart, at the
depth of 30 cm (Lieffers, 1988). At the depth of 10 cm, there
was no difference between sites. In Finland, Minkkinen
et al. (1999) found that cellulose decomposition was in
general faster in sites drained 30 years previously than
in their undrained counterparts, but the depth patterns
of these differences varied among sites (Fig. 3). In the study
of Domisch et al. (2000), drainage increased the mass
loss from Scots pine (Pinus sylvestris) needle litter in Carex-
peat but not in Sphagnum-peat, while mass loss from
Scots pine ne root litter was not clearly affected by
drainage in either peat type. Laiho et al. (2004) found
that mass loss from Scots pine needle litter, as well as
from Scots pine ne root litter incubated at 1020 cm
depth, was faster in a pristine site than in its drained
counterparts, while mass loss from ne roots incubated
at 1020 cm and small roots at either depth did not differ
between sites.
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R. Laiho / Soil Biology & Biochemistry 38 (2006) 20112024 2015
It is difcult to draw any consistent conclusions from
these studies. Both Lieffers (1988) and Laiho et al. (2004)
attributed the relatively slow decomposition at the surface
of the drained sites to moisture stress. It is noteworthy that
all of these studies dealt with the early phases of
decomposition (max. 1224 months) only. Long-term
patterns may differ from short-term observations, and will
be more crucial from the C sequestration point of view
(Latter et al., 1998; Moore et al., 2005; Prescott, 2005).
Further, with the exception of the work by Laiho et al.
(2004), the experimental designs did not include replica-
tions over time, i.e., the impacts of annual variation in
climatic factors could not be accounted for.
5. Effects on the abiotic and biotic factors regulating
decomposition processes
5.1. Physical and chemical environment for decomposition
Following a lowering of the water level, the water
content in the surface peat will decrease. Simultaneously,
the air-lled porosity and oxygen content increase (Boggie,
1977), and the oxic limit extends deeper (La hde, 1969;
Silins and Rothwell, 1999). These changes will generally
improve the conditions for the aerobic decomposition of
organic matter, which is faster than anaerobic decomposi-
tion (e.g., Bergman et al., 1999; S

antru c kova et al., 2004).

Simultaneously, the density of the surface peat may
increase due to several factors (Minkkinen and Laine,
1998a; Silins and Rothwell, 1998; Laiho et al., 1999). Peat
compaction, in turn, is associated with changes in pore size
distribution: the proportion of small pores with high water
retention capacity increases (Pa iva nen, 1973; Silins and
Rothwell, 1998). This may partly counteract the changes in
water and oxygen contents initially brought about by a
lowering of the water level (Rothwell et al., 1996).
If the water level drops deep enough, moisture stress may
counter the positive effects of drainage on decomposition
in the topmost soil layers (Lieffers, 1988). Peat soils
generally have a capillary fringe that reaches the surface
when the water level is within 3040 cm, only in highly
decomposed soils may it reach the surface when the water
level is at 60 cm (Verry, 1997). Laiho et al. (2004) showed
low soil matric potentials in the top 5 cm of soil in a
drained fen site when the water level was 40 cm below the
surface. Even if the topmost soil becomes sub-optimally
dry, it seems likely that there is a layer between the dry
surface and the new water level where increased aeration
and decreased moisture result in improved conditions for
aerobic decomposition.
There are also other noteworthy changes in the abiotic
conditions that may affect decomposition. First of all, soil
thermal properties and temperature regime change. Initi-
ally, during 13 years, the growing-season temperatures in
surface peat may increase (Lieffers and Rothwell, 1987;
Lieffers, 1988); however, along with an increase in tree
cover, temperatures in drained boreal peatlands remain
colder than in pristine sites (Ho kka et al., 1997; Vena la inen
et al., 1999). Minkkinen et al. (1999) observed that this
difference extended at least to a depth of 50 cm, at which
depth the temperature remained below +5 1C approxi-
mately a month longer in the drained site than its pristine
counterpart. The temperature sum (0 1C threshold) of the
surface soil (025 cm) may be 400 d.d. higher in pristine,
wet conditions than after long-term water-level drawdown
(Laine et al., 2004) (Fig. 4). Such a difference in air
temperature sums would correspond to a 500700 km
geographical distance in northsouth direction. Also, soil
frost tends to melt later in drained than in undrained sites
(Eurola, 1975). These indirect impacts of lowered water
levels are noteworthy as they may counteract the effects of
increasing air temperatures.
Further, the acidity of the surface soil increases
following water-level drawdown (Lukkala, 1929; Laiho
and Laine, 1990; Laine et al., 1995a; Minkkinen et al.,
1999). This has been attributed to several factors such as
reduction in the inowing groundwater that has a
neutralizing effect in pristine minerotrophic sites, enhanced
oxidation of both organic and inorganic compounds, and
increased uptake of base cations by tree stands (Laine et
al., 1995b). In the treed fens studied by Laiho and Laine
(1990) the decrease in soil pH extended down at least to the
2535 cm layer, being in the range of 0.51 units in Carex
peat during 5 decades, and less in Sphagnum peat (dry peat
to 0.01 M CaCl
2
1:2.5).
The changes in temperature and soil acidity may be
considered to have a retarding effect on decomposition
(e.g., Ivarson, 1977; Bergman et al., 1999), even though
ARTICLE IN PRESS
Decomposition potential
D
e
p
t
h
Surface
Increases
Before
After, option 1
After, option 2
Old water-level
New water-level
Fig. 3. Schematic presentation of the change in the decomposition
potential caused by a lowering of the water level. Based on cellulose
decomposition values presented by Minkkinen et al. (1999), and needle
decomposition values presented by Laiho et al. (2004), for moderately
poor, sedge pine fens. The Before case relates to old water-level, and
the After-cases, which represent situations where the surface layer either
does (option 1) or does not (option 2) experience moisture stress or other
changes retarding decomposition (hummock conditions), relate to new
water-level.
R. Laiho / Soil Biology & Biochemistry 38 (2006) 20112024 2016
many peatland microbes may tolerate quite acid conditions
(Williams and Crawford, 1983). However, environmental
factors have interactions that complicate the interpreta-
tions of these changes. For instance, the effect of changes in
temperature depends on moisture conditions. Hogg et al.
(1992) observed no change in soil respiration with higher
incubation temperature under inundated conditions. On
the other hand, Laiho et al. (2004) observed that higher
temperature sums during xed-length incubation periods
led to lower mass losses from needle and ne root litter in
drained treed fens. They concluded that this was because
higher temperatures enhanced moisture stress in the surface
layers of drained sites. This conclusion was supported by
positive correlations between mass losses and summertime
precipitation sum (Laiho et al., 2004).
The interactions between temperature and moisture
should also be considered when estimating the effects of
increasing temperatures, caused by climate change, on
decomposition rates. An increase in temperature will have
an unambiguously positive effect on decomposition only
when the moisture conditions are favourable. In sub-
optimally dry conditions, decomposition may be retarded
because of moisture stress, and in waterlogged conditions
because of lack of oxygen (La hde, 1969; La hde, 1971;
Hogg et al., 1992).
Interestingly, and surprisingly, Coulson and Buttereld
(1978) and Moore et al. (2005) have reported that there
were no consistent differences in mass loss rates of several
standard litter materials between upland and nearby
peatland sites. This emphasizes again the signicance of
litter quality in addition to environmental conditions for
the progress of decomposition.
5.2. Vegetation and organic matter inputs
There are two major biotic agents in the C cycle of a
peatland ecosystem: vegetation, which produces the or-
ganic matter (litter), and decomposers, which consume the
organic matter to a varying extent. Vegetation composition
is largely determined by water level and the solution pH
(Wheeler and Proctor, 2000; kland et al., 2001). A
persistent change in the water level induces acclimatization
and adaptation, rst within the existing community
(Weltzin et al., 2000, 2003), followed by slower but more
drastic changes when species better adapted to the new
conditions gain dominance (Laine et al., 1995a). Changes
in both species abundances and species composition may
be critical turning points for the C balance of a site
(Malmer et al., 2003). Litter quality is the key: different
ARTICLE IN PRESS
Fig. 4. Peat temperature (bottom) and temperature sum (accumulated daily average temperatures40 1C) (top) at different depths in a pristine, moderately
poor, sedge fen (left) and its counterpart drained for forestry 30 years earlier (right). The undrained site was treeless, while the drained one had developed a
tree stand of about 110 m
3
ha
1
following drainage. Source: Laine et al. (2004; see also Minkkinen et al., 1999).
R. Laiho / Soil Biology & Biochemistry 38 (2006) 20112024 2017
litter materials may have vastly differing rates of decom-
position (Hobbie, 1996; Thormann et al., 2001).
Generally, following a lowering of the water level species
adapted to wet conditions rapidly decline, while hummock
species benet. Sedges such as Carex lasiocarpa and
C. rostrata, which are important for peat formation in
many fens, are the rst to suffer from drought (Bubier et
al., 2003a,b), and disappear relatively fast after a persistent
drop in the water level (Laine et al., 1995a). Eriophorum
vaginatum is generally the only graminoid that can thrive in
dry sites, until the shading from the tree stand becomes too
much at canopy closure. Sphagnum mosses, another species
group important for peat formation, generally decline
following a persistent lowering of the water level, while
forest mosses such as Pleurozium schreberi and Hyloco-
mium splendens increase in abundance (Laine et al., 1995a).
In some treed fen types, it has been observed that both the
coverage (Laine et al., 1995a) and biomass (Laiho et al.,
2003) of Sphagnum mosses may remain high, thanks to
hummock species such as S. angustifolium, S. magellanicum
and, especially, S. russowii (Laine et al., 1995a).
Shrubs are the rst to benet from lowered water levels
(Laine et al., 1995a; Bubier et al., 2003b; Laiho et al.,
2003), while trees may gain dominance over a longer period
of time. Following persistent lowering of the water level,
mire species are at least to some extent gradually replaced
by forest species (Laine et al., 1995a; Vasander et al., 1997).
The extent and rapidity of this secondary succession
depend on the nutrient regime and initial wetness of the
site. The changes are the greatest in initially wet, nutrient-
rich sites (Laine et al., 1995a). In extremely nutrient-poor
ombrotrophic conditions the changes are small: in initially
dry sites they may remain almost non-existent (Vasander,
1982), while in initially wet sites or microforms, lichens
may replace the specialized mosses (Jauhiainen et al.,
2002).
Total plant biomass in peatland sites has a strong
positive correlation with tree stand biomass (or volume),
while the biomass of lower vegetation layers is negatively
correlated with tree stand biomass (Reinikainen et al.,
1984). Thus, total plant biomass generally increases
following lowering of the water level (Reinikainen et al.,
1984; Laiho et al., 2003) unless the site is too poor in
nutrients to support tree growth (Vasander, 1982). Total
production is also higher in dry than wet sites, again with
the exception of the most nutrient-poor sites (Reinikainen
et al., 1984).
The effects of vegetation changes on the quantity and
quality of litter inputs has received little attention, even
though it was recognized already by Lieffers (1988) that the
type and mass of litter will change following drainage,
which should have an effect on decomposition processes.
Laiho et al. (2003) estimated that in treed fens, the total
annual litterfall (including moss and below-ground litter)
did not change radically during 60 years following
drainage, and ranged from ca. 600 g m
2
year
1
(2025
years after drainage) to 960 g m
2
year
1
(8 years after
drainage) (Fig. 5). In contrast, there were dramatic changes
in litterfall composition. In pristine state, half of the total
annual litterfall derived from sedges, one third from
mosses, and less than 20% from trees and shrubs. Within
20 years after water-level drawdown, shrubs and trees had
become the major litter source, producing more than 80%
of the total litterfall. At 55 years after drainage, 68% of
total litterfall derived from trees and shrubs, and 30% from
mosses. There was also a change in the composition of the
arboreal litterfall in the course of time, as the proportions
of woody debris and cones increased.
The changes in the tissue type composition of the
litterfall are likely to have a negative effect on decomposi-
tion rates. Generally, arboreal litters decompose slower
than graminoid and herbaceous litters (Taylor et al., 1991;
Hobbie, 1996; Moore et al., 1999; Thormann et al., 2001).
This holds for foliage also, but woody materials: branches,
cones and eventually tree trunks, decompose especially
slowly (Taylor et al., 1991; Hobbie, 1996; Laiho and
Prescott, 1999).
Of Sphagnum mosses, the species adapted to dry
conditions (e.g., S. fuscum, S. magellanicum) decompose
slower than the species of wet habitats (e.g., S. cuspidatum,
ARTICLE IN PRESS
Fig. 5. Evolution of litter inputs into a moderately poor, sedge pine fen
following persistent lowering of the water level. Based on a chronose-
quence study. Source: Laiho et al. (2003). FWD ne woody debris,
other nonw. other nonwoody debris.
R. Laiho / Soil Biology & Biochemistry 38 (2006) 20112024 2018
S. fallax) (Johnson and Damman, 1991; Limpens and
Berendse, 2003). Sphagnum mosses are generally consid-
ered as poor, slowly decomposing materials (Standen and
Latter, 1977), but also the forest mosses common in
drained sites (e.g., Pleurozium schreberi, Hylocomium
splendens) have proven to decompose very slowly (Hobbie,
1996; Karsisto et al., 1996). Thus, the abundance and
vigour of the moss layer may signicantly affect the carbon
balance of a given site.
While in pristine peatlands sedge roots may grow down
to depths of more than 2 m (Saarinen, 1996), the arboreal
communities dominating dry/drained sites are generally
shallow-rooted (Paavilainen, 1966a,b). This means that
following water level drawdown, there may be a reduction
in the priming effect of root exudates and fresh root litter
on the anaerobic decomposition in deeper peat layers.
5.3. Decomposer communities
The structure of the microbial and faunal communities
varies with the peatland plant community (Borga et al.,
1994; Drouk, 1995; Fisk et al., 2003). Thus, it is likely that
such changes in hydrology that affect the plant community
will also induce adaptation in the microbial and faunal
communities (Markkula, 1986a,b), and the patterns of
adaptation may change over time since the hydrological
change.
Unfortunately, only sporadic information exists on the
changes in the decomposer communities and their func-
tioning caused by changes in the water level. Paarlahti and
Vartiovaara (1958) reported an increase in cellulose-
decomposing fungi and bacteria down to a depth of about
30 cm in intensively drained sites. Karsisto (1979) observed
an increased number of aerobic bacteria in sites where the
water level had been lowered down to 70 cm, as compared
to similar sites with water levels in 10 and 30 cm depths.
The increase was limited to the 010 cm peat layer. In
contrast, she observed generally lower lengths of fungal
mycelia in the sites with the deepest water levels. As yet, the
responses of decomposer communities to changing water
levels have not been analysed using PLFA proles
(phospholipid fatty acids; Ratledge and Wilkinson, 1988)
or molecular methods.
The food webs in peatlands are, as a whole, still
relatively poorly known. It has been suggested that
enchytraeids drive the processes of early stage decomposi-
tion in peat soils (Standen, 1973, 1978; Cole et al., 2000).
The majority of soil invertebrates, including enchytraeids,
occur in the surface horizons where they are vulnerable to
changes in environmental conditions (Briones et al., 1997;
Cole et al., 2002a). They thrive in a relatively moist and
cool environment (Briones et al., 1997), and are vulnerable
to drought (e.g., Cole et al., 2002a). During dry, warm
periods, some species such as Cognettia sphagnetorum may
migrate downwards (Springett et al., 1970; Briones et al.,
1997); however, they may not tolerate extended periods of
anoxia (Healy, 1987). Further, the more decomposed
substrate of deeper soil layers may not be suitable for
enchytraeids. Briones and Ineson (2002) showed that
enchytraeids assimilated carbon that had been removed
from the atmosphere 510 years earlier. Correspondingly,
Cole et al. (2000) suggested that downward migration of
enchytraeids during drought might have little effect on
decomposition.
The responses of soil mesofauna to potential climate
change have been studied in the blanket peats in UK
(Briones et al., 1997, 2004; Cole et al., 2002a). Only
temperature changes have been considered; no water level
treatments have been applied. Cole et al. (2002b) suggested
that soil warming might reduce the functional role of
enchytraeids with respect to their ability to enhance
decomposition.
Silvan et al. (2000) observed that the total numbers of
enchytraeids, collembolans and mites correlated positively
with water level depths in a drainage-succession continuum
of sedge pine fens. In sites where the water level was at a
depth of 2530 cm, the populations were generally about
ten times higher than in sites where the water level was at
5 cm. Simultaneously, the proportion of oribatid mites
decreased and that of collembolans increased. Laiho et al.
(2001) further observed that when the average water level
of a site was below 20 cm, it did not have a signicant effect
on the within-site distribution of mites, but still affected
that of enchytraeids and, especially, collembolans. Species
composition, activity, or role in the food web, were not
accounted for in these papers.
The implications of the observed changes in decomposer
communities following lowering of the water level for
organic matter decomposition are currently still somewhat
difcult to estimate. It seems likely that they would be
enhancing decomposition.
6. Synthesis from a C sink/source point of view
Based on the results reviewed, it is clear that although
high water levels in peatlands (relative to upland sites)
result in C sequestration, within a peatland, the relations
between water level, decomposition and C sequestration
are not that simple (see also Waddington and Roulet, 2000;
Bubier et al., 2003b).
Mechanistically, whether a peatland will remain a C
sink, become a stronger sink, or become a source of C to
the atmosphere will depend on 1) the rate of decomposition
of the old, pre-water-level-lowering, peat, and 2) the
rates of inputs and decomposition of the new organic
matter entering the system as litter produced under the new
environmental conditions (Fig. 6). Quite simply, if the
accumulation (inputs decomposition losses) of the new
organic matter exceeds the decomposition losses from the
old peat, the peatland will remain a sink of C. If not, then
the peatland will become a source of C to the atmosphere.
The old peat can only decompose. The C balance of the
new litter inputs will largely depend on the vegetation
composition. Easily decomposable tissue types may be
ARTICLE IN PRESS
R. Laiho / Soil Biology & Biochemistry 38 (2006) 20112024 2019
preferred by decomposers over the older substrate (Wil-
liams and Crawford, 1983), while in case of lowered litter
quality (e.g., tree litter vs. graminoid litter), the situation
may be the opposite. The role of the changing vegetation in
the C cycling following a persistent lowering of the water
level has in previous research been largely neglected.
The rst postulate regarding the effects of lowered water
levels on decomposition processes and C sink/source
behaviour is that long-term changes differ from short-term
changes that merely reect a disturbance in the old system
adapted to the pre-water-level-lowering conditions. In a
short term, the disturbed system will probably always lose
C. An essential question here is, for how long? Obviously,
the C loss from peat is at its greatest during and
immediately after the lowering in the water level, when
drought is not yet limiting decomposition, and slows down
with time (e.g., Blodau and Moore, 2003), due to
decreasing substrate quality. The photosynthetic ow of
C into the soil, in turn, may also decrease (Bubier et al.,
2003a,b; Blodau et al., 2004), but may recover with the
response of the plant community, such as transition from
sedges to shrubs that may occur rather rapidly (Bubier et
al., 2003b). The rapidity of the response of the plant
community may depend on the site nutrient regime
(Tuittila, E.-S., Alm, J. and Laine, J. unpublished data).
In Scotland, newly drained peatlands acted as sources of C
during the rst 4 years, after which they became sinks again
(Hargreaves et al., 2003).
The second postulate is that long-term changes in the C
cycle result from several adaptive mechanisms of the
ecosystem to the new hydrological regime. Hence, the
long-term changes may inherently vary among peatland
types, climates, and extents of change in the water level.
The determinants must in one way or the other derive from
the interactive effects of the changes in not only water level
and aeration (Blodau and Moore, 2003), but also vegeta-
tion composition and organic matter inputs (Bubier et al.,
2003b; Laiho et al., 2003), and possibly soil temperature
and acidity (Minkkinen et al., 1999).
The reviewed studies emphasize the paramount impor-
tance of organic matter quality in determining the rate of
decomposition in peatlands (see also Bauer, 2004). Actually,
such observations led already Coulson and Buttereld (1978)
to suggest that the position of the water level or the pH
differences in soils were of only indirect consequence for the
rates of decomposition, through determining the composi-
tion of the plant community. However, when evaluating the
effects of persistent lowering of the water level on C cycling,
accounting for changes in species composition only would be
an oversimplication just like assuming that the change in
water level is the only determinant. On deep-peat sites the
lower parts of the peat deposit remain oxygen-decient
despite of a drop in the water level, and maintain a zone
where new material may be accumulating or at least old
material may be preserved. Thin-peated sites, if they lose
altogether an anoxic layer, may be at the greatest risk for
losing considerable amounts of C.
The few studies that have so far systematically looked at
the effect of water level on decomposition within one
vegetation type, were all done on rather nutrient poor treed
fen or bog sites in southern or middle boreal conditions
(Lieffers, 1988; Minkkinen et al., 1999; Domisch et al.,
2000; Laiho et al., 2004). From these studies it may be
concluded that in such peatland types, the site may turn
into a large hummock-system where several factors,
including litter quality, relative moisture deciency, lower
soil temperature, higher acidity, and in deeper layers also
oxygen deciency, may interact to constrain organic matter
decomposition. This may explain the continuing sink
behaviour that has been observed in these kinds of sites.
Currently, we do not know what are the critical limits in
the properties of vegetation (litter inputs) and peat that
switch the site either to a source or a sink. A critical factor
may be the quality and thickness of the intermediate
peat layer where the rate of decomposition is increased in
any case (Fig. 3).
7. Conclusions
Based on the literature reviewed, the following critical
gaps have been identied in our knowledge of the C cycle in
peatlands under change. There is a lack of information on:
1. how the amounts and quality parameters of litter inputs
change in different peatland sites after short- and long-
term change in the water level,
2. how the litters produced by the successional vegetation
communities decompose under the changed environ-
mental conditions following persistent lowering of the
water level in long-term,
ARTICLE IN PRESS
Before After 1 After 2 After 3 After 4 After 5
S
o
i
l

C

p
o
o
l
,

%
0
20
40
60
80
100
120
Peat
Litter
Fig. 6. Schematic presentation of the sink/source behaviour of a peatland
site following a persistent lowering of the water level. Peat C pool in the
peat deposit; Litter C balance (input-decomposition) of annual litter
inputs. Before before the lowering of the water level. The different
After-options depict situations where peat decomposition clearly increases
(1, 2 and 5) or is little affected (3 and 4), and the C balance of litter inputs
remains the same (1 and 3), increases (2 and 4) or decreases (5). The soil C
pool (y-axis scale) is in per cent of the pool in peat before water-level
drawdown. Thus, an After-column below the dotted 100% line implies a
switch to a C source, while a column above this line implies continued sink
function.
R. Laiho / Soil Biology & Biochemistry 38 (2006) 20112024 2020
3. how the changes in litter inputs affect the community
structure and functioning of microbes responsible for
the aerobic decomposition of organic matter, and
4. how the quality parameters of peat affect its decom-
position rates under changed environmental conditions.
Further, the interactions of different environmental
factors have not been systematically analysed: how are
different water level regimes related to oxygen supply,
temperature, acidity, and nutrient supply within a given
site? The variations in physical factors could be simulated
with existing models (Granberg et al., 1999; Zhang et al.,
2002) but we need test data for situations with lowered
water levels.
Consequently, we need more successional studies where
we consider the interactions of all changing factors
(vegetation, microbes, several abiotic) on sites with
different old peat substrates. Especially, we need
experimental research that has been designed to contribute
to an overall synthesis.
Each case study will bring in valuable new information
on changes taking place in single peatland types and
climates. Yet, because of the multiple interactions between
the factors in effect, generalizations should not be based on
limited data sets. Moreover, we should generally be more
specic when discussing decomposition of organic matter,
and attribute our statement to both the subject of
decomposition (peat, cellulose strips, litter from different
plant species) and the time scale involved.
Acknowledgements
My sincere thanks are due to Hannu Fritze, Krista
Jaatinen, Marjut Karsisto, Veikko Kitunen, Jukka Laine,
Kari Minkkinen, Taina Pennanen and Petra Va vr ova for
inspirational discussions, Tim Moore, Timo Penttila , Eeva-
Stiina Tuittila and Harri Vasander for critical and
constructive comments, and the Academy of Finland for
funding (projects 203585, 205090).
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