HYDROLOGICAL ASPECTS OF THE CARBON BALANCE IN A BOREAL

CATCHMENT: A MODEL STUDY

.
Document name
DOCTORAL DISSERTATION
Date of issue
2007-10-01
Organization
LUND UNIVERSITY
Geobiosphere Science Centre
Dep. of Physical Geography and Ecosystems Analysis,
Slvegatan 12, 223 62 Lund Sweden
Author(s)
Yurova Alla Yu.
Sponsoring organization
NECC
Title and subtitle
Hydrological aspects of the carbon balance in a boreal catchment: A model study. Meddelanden frn Lunds
Universitets Geogrfiska Institution. Avhandlingar 173
Abstract
The cycling of carbon in its gaseous, solid and dissolved forms is of central importance to ecosystem
science, particularly, when the natural carbon cycle has been perturbed by human activity. Mathematical
models are commonly used to asses and predict global and regional patterns of terrestrial carbon exchange.
At the same time, processes operating on smaller (sub-grid) scales are usually outside the scope of the large-
scale models exploring climatebiosphere interactions. However, if a sub-grid feature is found repeatedly in
a large number of particular realizations, we may expect that its characteristic dynamics are likely to be
important even at larger scales. This work examines some of these phenomena. First, a boreal landscape
differentiated into forests and peatlands is investigated. Second, the export of dissolved organic carbon
(DOC) in peatland streams is assessed.
The work described herein mainly addresses methodology, i.e. how approaches to hydrology and ecosystem
modelling can be combined. Specifically, the general ecosystem model GUESS was modified and supplied
with new modules to be used for peatland simulations. The model was also designed to reproduce soil
moisture and related ecosystem properties along a topographical catena. In addition, a completely new
model of DOC production, transport and transformation in peat was developed.
The data for model development and validation came from boreal catchments in the Vindelns Forest
Research Parks, The Swedish University of Agricultural Sciences, Northern Sweden. A relatively good fit
between the modelled and measured NEE at the Deger Stormyr peatland was demonstrated for 2001-2003.
Further, the model was used to demonstrate that the NEE was quite insensitive to changes in water table
level within the ranges typical for the dominant mire plant community. The newly constructed model was
able to reproduce DOC concentrations measured at the outlet of the Kallklsmyren peatland for 1993-2001.
The model suggested that the main drivers for interannual variability in DOC concentrations were the rates
of microbially mediated DOC production and mineralization and the flow intensity in the active surface
layer of the peatland. The amount of DOC in the peat water at a particular time was mainly determined by
(much larger) amounts of soluble carbon sorbed on the peat matrix. A dynamic, rather than static,
description of the adsorption and desorption processes was advocated.
The work makes some contribution to filling the gap that currently exists between field observations at the
plot scale and large-scale model analysis of the climatebiosphere interaction.
Keywords : model, ecohydrology, boreal peatland, dissolved organic carbon
Classification system and/or index terms (if any)
Supplementary bibliographical information Language
English
ISSN and key title
0346-6787
ISBN
978-91-85793-03-7
Recipients notes Number of pages
50
Price
Security classification
Distribution by (name and address) Alla Yurova, alla.yurova@nateko.lu.se
I, the undersigned, being the copyright owner of the abstract of the above-mentioned dissertation, hereby grant
to all reference sources permission to publish and disseminate the abstract of the above-mentioned
dissertation.
Signature: Date: 2007-10-01
Hydrological aspects of the carbon balance
in a boreal catchment: A model study
Alla Yurova
An academic thesis in fulfillment of the degree of Doctor of Philosophy in
Geobiosphere Science, discipline of Physical Geography and Ecosystem Analysis,
at the Faculty of Science of Lund University, which will be publically defended on
Friday 26
th
October 2007 at 10:15 in Vrlden, Geocentrum I, Slvegatan 12, Lund
Faculty opponent: Dr. Per-Erik Jansson, The Royal Institute of Technology,
Stockholm, Sweden
A doctoral thesis at a university in Sweden is produced either as a monograph or as a
collection of papers. In the latter case, the introductory part constitutes the formal thesis,
which summarizes the accompanying papers. These have either already been published
or are manuscripts at various stages (in press, submitted or in preparation).
Yurova, A. (2007). Hydrological aspects of the carbon balance in a boreal catchment: A
model study. Scripta Academica Lundensia. Meddelanden frn Lunds Universitets
Geogrfiska Institution. Avhandlingar 173. 50 pp. Lund.
MEDDELANDEN FRN LUNDS UNIVERSITETS
GEOGRAFISKA INSTITUTION. AVHANDLINGAR
Avhandlingsnr. 173
ISBN: 978-91-85793-03-7
ISSN: 0346-6787
2007 Alla Yurova
Print: KFS AB, Lund, 2007
To my teachers
CONTENTS
List of papers
Acknowledgements
INTRODUCTION...9
THE SUBJECT AND AIMS OF THE STUDY......14
METHODS.15
Model approach15
Field and laboratory materials..15
Modelling framework...19
Model formulation19
Parameterization and validation..24
Model experiments24
Sensitivity and uncertainty issues.24
RESULTS...26
Spatial variation in carbon storage along the catena. Dynamic aspects
of the soil moisturesoil C relationship (Paper I)26
Mechanism of NEE regulation by WT level on a mire (Paper II)...29
Hydrological and physico-chemical control of DOC production and
export from the mire (Papers III, IV)...29
CONCLUSIONS....31
POSSIBLE APPLICATIONS AND FURTHER DEVELOPMENT32
APPENDICES....35
REFERENCES..42
POPULRVETENSKAPLIG SAMMANFATTNING..48
Papers I-IV
List of doctoral thesis published in the present series
LIST OF PAPERS
This thesis is based on studies reported in the following papers, referred to by their
Roman numerals. Published papers are reprinted by kind permission from the copyright
holder
I. Yurova A., Lankreijer H. 2007. Carbon storage in the organic layers of boreal
forest soils under various moisture conditions: a model study for Northern
Sweden sites. Ecological Modelling, 204, 475-484
II. Yurova A., Wolf A., Sagerfors J., Nilsson M. 2007. Variations in Net
Ecosystem Exchange of Carbon Dioxide in a Boreal Mire: Modelling
Mechanisms Linked to Water Table Position. Journal of Geophysical
Research: Biogeosciences, 112, G02025, doi: 10.1029/2006JG000342
III. Yurova A., Buffam I., Bishop, K. A model of dissolved organic carbon
concentrations in peat and stream waters. I: Development and parameterization
(Submitted)
IV. Yurova A., Sirin A., Bishop K., Laudon, H. A model of dissolved organic
carbon concentrations in peat and stream waters. II: Application and the
sensitivity analysis (Submitted)
ACKNOWLEDGEMENTS
I greatly acknowledge generous financial support that came from NECC project and
Lund University. Special thanks to Anders Lindroth and Harry Lankreijer for providing
me with an excellent opportunity to be fully devoted to research during the last four
years.
Coming to Sweden I was amazed how well organized, esthetic and cozy a scientific
institution can be, and how easy it can be to solve all practical problems in a friendly
atmosphere. Thanks for everyone who keep it running, not least to administration,
librarians and technical staff at Lund University.
During these years I was much enriched by scientific communication with many people;
to list few who influenced me most: Annett Wolf, Mats Nilsson, Kevin Bishop, Ben
Smith, Oleg Chertov and Andrej Sirin. The great variety of things you told me was
overwhelming, but it was always a pleasure to note how understandable and well-
structured things get after I have discussed them with you. Thanks to all and each of
you!
Special thanks to many people at the Department of Forest Ecology at Ume University
(and others involved into Krycklan activity) who have inspired me to work on various
aspects of C modeling on a catchment scale and to look closer at peatland ecosystems.
Your data and expertise formed the basis for this PhD thesis. Thanks to Jrgen, Ishi and
Hjalmar for careful and almost immediate response to all my data queries and for
sharing your knowledge of an area. Ishi is also greatly acknowledged for setting and
running long-term laboratory peat incubations that were so needed for parameterization
of our DOC model.
I thank Harry and Ben for providing me an opportunity to participate in teaching
Ecosystem modeling course. Ive learned a lot from being beside such experienced
mentors as you are; and motivated and curious students have forced me to think more
carefully about many issues.
Sees-editing Ltd is acknowledged for quick, thoughtful and efficient English correction
of the manuscripts.
Thanks to my friends- Annett and Christian, Anna-Maria, Ejiang and Jin Chuan-with
you I felt like at home. Thanks also to my friends in Moscow, especially to Natasha U.
and Semionovs family-great that you know how to deal with me. Distant but so
familiar voices, reminding ironically that most of the ecosystems are in fact managed by
humans and the book from 1930s, which was finally found after some adventures in
Moscow libraries, may still be not exactly the one I need- thank you for all help and
support.
This thesis is dedicated to my teachers, whose deep knowledge, patience, enthusiasm,
fundamental view on science and abilities to ruin all unjustified common sense
thinking and yet to demonstrate elegant and integral theories instead have always been
an encouraging example for me. It becomes really so much easier to think things
through and to face difficult problems when one was trained by you, and finishing a
PhD is just another occasion to say thanks.
ABBREVIATIONS
GUESS general ecosystem simulator [Smith et al., 2001]
LPJ Lund-Potsdam-Jena dynamic global vegetation model [Sitch et al., 2003]
ROMUL model of raw humus, moder and mull [Chertov et al., 2001]
MMWH mixed mire water and heat model [Granberg et al., 1999]
GLUE generalized likelihood uncertainty estimation [Beven and Binley, 1992]
REA representative elementary area
DOC dissolved organic carbon
SSOC sorbed soluble organic carbon
O layers organic (soil) layers
NEE net ecosystem exchange
PET potential evapotranspiration
NECC Nordic Center for Studies of Ecosystem Carbon exchange and its
interaction with the Climate system
IPCC Intergovernmental Panel of Climate Change
INTRODUCTION
The rapid increase in atmospheric CO
2
concentration resulting from human activities
has raised concerns in the scientific community and society in general about possible
climate changes induced by altering the global carbon cycle. It has been shown that
even though anthropogenic CO
2
emission is insignificant on the geological time-scale, it
will affect the Earths climate for hundreds of years [e.g., Manabe, 1967; IPCC reports
1996, 2001]. At the same time, our current understanding of the climate system is
limited and it is not possible to predict future climate with any certainty. One remaining
scientific challenge has its origin in the nature of greenhouse gases that affect the
Earths energy balance. Water vapour, carbon dioxide and methane molecules not only
obey physical and chemical laws, but also cycle continuously through living organisms
and the products of their activity and decay. The complexity of the sphere of life the
biosphere [Vernadsky, 1926] provides numerous possibilities for feedbacks between
its components, biogeochemical cycles and climate. An understanding of the numerous
links between different ecosystem components, as well as reliable quantitative estimates
of carbon cycling in the biosphere, are needed to improve prediction of climate system
behaviour with increasing CO
2
.
Climate can affect an ecosystem by altering (i) radiative balance, (ii) heat balance and
therefore temperature regime and (iii) water balance. The focus of this work is on the
hydrological aspects of ecosystem functioning. Water provides environment for all
organisms that inhabit the oceans, freshwater ecosystems and soil and sediment
solutions. CO
2
, CH
4
and various organic molecules, including humic substances,
dissolve in it. Water also plays a critical role, from the molecular to the biome level, in
terrestrial ecosystems. Soil moisture has a significant effect on vegetation composition,
growth and disturbance regimes. Direct coupling between photosynthesis and
transpiration in plants suggests a close link between the carbon and water balances. The
process of decomposition and further transformation of soil organic matter is largely
dependent on the soil moisture [e.g., Davidson et al., 2006; Carrasco et al., 2006].
Water provides specific conditions for certain groups of microorganisms, both aerobic
and anaerobic. Water content in the soil therefore determines the dominant ecosystem
type, forest or wetland, in an area. Wetlands are of particular concern in the context of
greenhouse gas balance because of specific C exchange. During the Holocene they have
constantly sequestered C in peat, but, at the same time, they emit CH
4
and may, under
certain conditions, emit large quantities of CO
2
[e.g., Moore and Knowles, 1987;
Gorham, 1991]. Water and carbon cycling in wetlands are closely interconnected [e.g.,
Ivanov, 1981; Hilbert et al., 2000]. While the net peat accumulation is controlled by the
balance between the rates of net C uptake and release, small changes in either of these
processes caused by changes in the water balance may substantially alter the rate of net
peat accumulation.
9
Besides shaping the landscape, water in the soil acts as a conduit between terrestrial and
aquatic ecosystems. Dissolved organic carbon (DOC) is formed as a product of the
decomposition and transformation of organic matter in soils and peat. DOC is
transported in moving water from the land to aquatic ecosystems, where it has many
important functions; including providing an energy source for heterotrophic
microorganisms [e.g., McKnight et al., 1998; Steinberg et al., 2006]. Thus, C originally
taken up by plants may be either deposited in ocean and freshwater sediments [e.g.,
Dean and Gorham, 1998] or mineralised and emitted back to the atmosphere in
significant amounts [e.g., Cole et al., 1994; Cole et al., 2007].
If the climate changes, it will affect water balance, and therefore certain ecosystem
processes. The responses of ecosystems may in turn affect climate, and could involve
both physical and biochemical aspects. Surface reflectance, latent and sensible heat
fluxes, local atmospheric instability, cloudiness, and precipitation can be altered due to
changes in plant physiology, stand level canopy structure, phenology and vegetation
composition [e.g., Charney, 1975; Shukla and Mintz, 1982; Bonan et al., 1992; Bonan
2002]. Biochemical effects may caused by changes in photosynthetic CO
2
uptake
affected by soil moisture and changes in the rate of soil respiration. The latter is likely if
relatively liable C, which is currently protected from decomposition by high soil
moisture, is exposed to drier conditions [e.g., Carrasco et al., 2006]; alternatively,
drought can inhibit decomposition in water-limited regions [e.g., Schaphoff et al.,
2005]. Potential peatland feedback mechanisms deserve special consideration, because
of the possibility of autogenic non-reversible lateral peatland expansion [e.g., Ivanov,
1981; Inisheva et al., 2006] and abrupt changes in C balance that these ecosystems may
exhibit if the water balance is altered [e.g., Hilbert et al., 2000]. Changes in the amount
of DOC transported from soils and peat can influence DOC exports to the ocean
[Freeman et al., 2001b] and CO
2
emissions from rivers and lakes [e.g., Sobek, 2005].
Our knowledge of the structure and functioning of ecosystems can be formalized in
mathematical equations, thus providing opportunities to study interconnected and non-
linear internal dynamics of the biosphere and its responses to external drivers. Today,
terrestrial vegetation models are widely used in ecosystem studies at various scales from
individual plots to the entire globe. The current generation of dynamic vegetation
models has the ability to capture the effects of environmental drivers on plant
physiology and biochemistry, vegetation composition and soil organic matter dynamics
[e.g., Cramer et al., 2001; Sykes et al., 2001]. Global versions of dynamic vegetation
models are now being coupled to general atmospheric circulation models and used for
improving future climate predictions [e.g., Foley et al., 1998; Cox et al., 2000]. Results
produced using such coupled models have stimulated a growing interest in the impact of
the biosphere on the Earths climate. However, those results are not without controversy
[e.g., Moorcroft, 2006]. Future changes predicted by biosphereatmosphere modelling,
such as the disappearance of the Amazonian forests [Cox et al., 2000], may seem too
10
drastic. The intuitive feeling is that ecosystems are more adaptive than the models
sometimes suggest. Whether the soil C pool will be influenced by future climate change
to such an extent and in such a manner that models predict, is also open to debate [e.g.,
Agren and Hyvonen, 2003; Davidson and Janssens, 2006; Carrasco et al., 2006;
Moorcroft, 2006]. One way to test the validity of any model is to investigate how well it
describes the current situation. However global models are very generalized, while the
landscape we observe in reality is much more diverse. It is not an easy or
straightforward task to choose an average or standard ecosystem from the wide
variety of ecosystem types. In addition, valuable information about the ecosystems
flexibility can be derived by studying the adaptations of vegetation and soils to the local
climatic, hydrological or geochemical factors acting at scales much smaller than the
model resolution. Downscaling of models is, therefore, one way to check a models
value and is necessary for better understanding and prediction of the biosphere
atmosphere interaction.
Models at sub-grid scales are valuable on their own, and can be used as a tool for
studying particular ecosystem types, e.g., forests and wetlands. However, it is also
important to examine how information derived from such models can be assimilated
into those representing a larger-scale. Many variants are possible, and the simplest
approach is to divide the large-scale model cell into portions dominated by a particular
ecosystem type. Another approach is to try to use a representative elementary area
[REA, Wood et al., 1988]. It has been suggested that at the scale of an REA, the
variability between different catchments is minimised and it is statistical distribution,
not actual spatial patterns, of soil characteristics within the REA that determines runoff
and, potentially, other associated parameters (soil C storage, DOC fluxes). At smaller
scales, variability is much greater and model results are strongly dependent on the
choice of scale. At the REA scale, the catchment area is large enough so that all
important characteristics are sufficiently sampled to represent the mixture of landscape
units. At larger scales, variability may become important due to differences in geology,
geomorphology and other large-scale factors.
The focus of this work was on various effects that hydrological processes may have on
C cycling, and landscape differentiation as a result of soil moisture was of interest.
Some approaches to the coupled modelling of C and water balances were considered as
potentially useful for this study (Table 1).
11
Table 1. Models of carbon dynamics at different scales, noting how hydrological
processes are incorporated
Can models simulate? Model type How hydrology is
modelled Veget.
C
Soil
C
Wetland
C
DOC
No.
of
pub
a
.
Global
Phenomenological eq.
for infiltration and
runoff
Yes Yes Yes
b
No 740
Regional
1D solution of Richards
or another eq. for
vertical flow, lateral
flow parameterized or
not included
Yes Yes Yes
c
? 81
Catchment
various, from complete
3D solution of flow eq.
to lumped
phenomenological
runoff eq., many
combinations of
intermediate
complexity possible
Yes ? ? Yes 16
Catena
Various, mainly
analytical or numerical
solutions of kinematic
wave or other 1D
hillslope flow eq.
Yes ? No Yes 3
Site
1D solution of Richards
or another eq. for
vertical flow, lateral
flow parameterized or
not included
Yes Yes Yes ? 858
? in the table indicates that the author of this thesis is not aware of models that include the specified
component. The possibility that someone has undertaken such a study is, however, not excluded
a: 1969-2007, ISI Web of KnowledgeSM search from 16/08/07
b: on-going work Wania et al. (LPJ model), de Noblet et al. (ORCHIDEE model)
c: on-going work Roulet et al. (Canadian Land Surface Scheme)
Due to their development history and the scale at which they operate, models of
different types found in the literature have different ways of describing hydrological
processes (Table 1). Many models based on 1D differential equations of water flow are
designed to be used at the site scale, and direct measurements of hydrological variables
can be easily obtained for validation. In contrast, many models operating at a large scale
utilize rather generalized phenomenological expressions to simulate an area-average
runoff and soil moisture. However, it can be seen (Table 1) that there is a gap between
12
the field scale (m) and the regional-to-global grid cells (typically more than 5050 km).
The scale least represented in the modern literature devoted to ecosystem modelling is
small or medium catchments (from 1km
2
). Yet, this scale is one of the most interesting
from many perspectives. In the boreal region in particular, a number of factors support
this contention:
- Much of the spatial variability in soil carbon storage can be found within small
catchments. The gradient from dry sites, usually with poor O layers (typical mor),
to wet sites on fully developed humic podzol (rich O layers, hydromor) or peat
soil, is a characteristic, repeated feature of such catchments
- Peatlands often occupy a significant proportion of a catchment, saturated due to
the high water table position. Peatlands contain the greatest amounts of carbon per
unit area of all the terrestrial ecosystems, and as much as 20-30% of the global soil
C is currently stored in the boreal peatlands [Schlesinger, 1997]. Although the
overall likelihood of peatland formation and growth in the region is a factor of
climate, the presence of peatland at a certain location is largely determined by the
configuration of a particular catchment [Ivanov, 1981]
- DOC fluxes are strongly controlled by water routing at this scale. Export of DOC
is likely to occur mainly from the areas where flowing water can bypass mineral
soils, i.e. surface and subsurface flow through riparian zones and peatlands [e.g.,
McKnight et al., 1998; Qualls, 2000; Bishop et al., 2004].
- The size of the representative elementary area (REA) in till soils in Northern
Europe is 1 to 10 km
2
. [Beldring et al., 1999; Temnerud et al., 2007].
It is therefore important to (i) look at specific landscape units, including the peatlands,
and (ii) try to find methods for quantifying moisture-related gradients for ecosystem
properties, starting with small experimental catchments (1-2km
2
). Such studies will
foster understanding and will help with the parameterization of processes that are of
great importance at much larger scales.
13
THE SUBJECT AND AIMS OF THE STUDY
This PhD thesis was supported by NECC (Nordic Center for Studies of Ecosystem
Carbon exchange and its interaction with the Climate system), with the goal of
investigating the features characteristic of Nordic ecosystems and integrating the data
collected into regional C balance assessments. The PhD research involved a numbers of
steps aimed at constructing a process-based model, which would:
- describe the C balance of a small boreal catchment
- do this explicitly, so that each main terrestrial ecosystem type, including peatland,
is simulated separately
- use a dynamic hydrological approach allowing moisture accounting and solute
(DOC) tracing
The following research questions were addressed as model development progressed:
- How does soil C storage in organic layers differ with respect to local soil moisture
conditions within a landscape? Are there new explanations for this spatial
variation? (Paper I)
- How is net ecosystem CO
2
exchange in a Sphagnum-dominated peatland related
to water table position? (Paper II)
- How large and how variable is the export of DOC from a peatland? Which factors
determine temporal variability in DOC fluxes at different time scales? (Paper IV)
Three of the four papers included in this thesis (Papers II, III, IV) were devoted to a
particular ecosystem type, namely peatland. In addition, as presented in Paper I and
partly here in the thesis summary, an attempt was made to consider the full range of
ecosystem types along the soil moisture gradient.
14
METHODS
Model approach
Models were the main tools of this study; they were modified and developed, as
follows:
- The dynamic vegetation model LPJ-GUESS was coupled to a dynamic soil
organic matter model, ROMUL (Paper I)
- The combined GUESS-ROMUL model was linked to a simple hydrological
scheme allowing simulation of vegetation and soil dynamics along a forested
catena (Paper I, thesis)
- Modifications were made to the model in order to quantify the CO
2
flux in a
Sphagnum-dominated peatland. The MMWH model was then used to simulate
hydrology and the peat thermal regime (Paper II)
- A completely new model of DOC concentration and export from a peatland was
developed (Papers III and IV).
Field and laboratory materials
Although there was no geographical reference in the model formulation (the scheme
was kept rather general), this study was closely linked to a few geographical locations
situated in the Vindeln Forest Research Park, Northern Sweden. Field data used in the
thesis for model development and validation (summary in Table 2, p.18) were kindly
provided by the Department of Forest Ecology at SLU (Swedish University of
Agricultural Sciences) in Ume.
Study area
The Vindelns Forest Research Park is situated 60 km northwest of Ume, Northern
Sweden (Figure 1). There is a long history of geomorphological, hydrological, botanical
and ecological studies in an area, and mires have always been of a particular interest.
The area is located in the boreal zone with a cold temperate humid climate. The
reference mean annual temperature, measured at the nearby Kulbcksliden
meteorological station, is +1.2
o
C [Alexandersson et al., 1991] and the mean annual
precipitation is 523 mm, 34% of which falls as snow [Alexandersson et al., 1991]. The
spring flood is the dominant hydrological event of the year, accounting for up to 60% of
the annual stream discharge [Bishop et al., 1995]. Precipitation and runoff have been
higher than normal in a last decade [Lindstrm and Alexandersson, 2004]. The
underlying geology is blocky glacial till from the quaternary; the landscape includes
moraine features with hills and former melt water beds. Soil types are iron-podzol in dry
upslope areas, iron humic podzol in wetter locations and humic podzol with a distinct
peat horizon in areas with high ground water levels. The most common tree species are
15
Scots pine (Pinus sylvestris) in dry areas and Norway spruce (Picea abies) in wetter
areas. The most extensive mire type is aapa (a mire complex differentiated into
oligotrophic and minerotrophic sections). Mires are rather common and can be found on
gentle slopes, in drained and undrained topographical depressions and in riparian zones
of streams and rivers. The peat largely consists of Sphagnum remains, but sedge
remains are also commonly found.
Two experimental catchments were studied: the drainage basin of the Deger Stormyr
peatland (Figure 1b) and the Nynget catchment that includes the Kallklsmyren
peatland (Figure 1c). A topographical transect selected to represent soil C storage in
organic layers along the catena is situated within the Nynget catchment. This was
drawn on a national topographical map (Lantmteriet, 2002), across the Kallklsmyren
peatland, perpendicular to the stream (Figure 1c). Transect crosses the main landscape
elements: a dry upland forest, a peatland, mesic-moist and mesic forests on a slope and
a stream riparian zone. Field data collected from two peatlands (Deger Stormyr and
Kallklsmyren) have been used for model development and validation.
Deger Stormyr (6411'N, 1933'E) is a large (6.5km
2
) oligotrophicminerotrophic
mire complex. Peat depth is mostly between 3 and 4 m, with maximum depths up to 8 m
[Malmstrm, 1923]. The hydrology of the system is rather complex, with ground water
being routed into pipe flow and springs [Malmstrm, 1923]. The mire has distinct
microlandscape differentiation and only the lawn plant community was studied here.
The water table position usually varies from 5 to 19 cm below the peat surface (90%
percentile-10% percentile), but flooding occurs at every snowmelt while the peat is still
frozen. Species dominating the lawn plant community are Sphagnum balticum Russ. C.
Jens. with sparse Sphagnum lindbergii. The vascular plant community is dominated by
Eriophorum vaginatum L., Vaccinium oxycoccos L., Andromeda polifolia L., with
Scheuchzeria palustris L occurring more sparsely.
Kallklsmyren (6414'N, 1946'E) is a small (9ha) complex mire occupying a
topographic depression. The mire is 2 to 5m deep. It is drained by a single stream,
which originates on its southern edge. Flow in the surface peat layer is accompanied by
a minor, but not insignificant, ground water component [Sirin et al., 1998]. The main
section of the mire is raised, sloping towards the edges, and has distinct ombrotrophic
features, with vegetation composed of a sparse Scots pine stand with dwarf-shrubs and
Sphagnum mosses, with Carex on the edges. Mineratrophic fen vegetation occupies the
southern area of the mire adjacent to the stream and is found in two belts situated on the
eastern and western sides of the raised section.
16
64
o
N, 19
o
E
Figure 1. (a) Map of Sweden showing the location of the study area. (b) Map of
the area near Vindeln, Vsterbotten in Northern Sweden, showing the locations
of the Deger mire, the eddy tower measurement site, and the area of a plot (c).
(c) Map showing the location of the Kallkllsmyren mire, the measurement site
at the stream origin, and the location of transect AB used to simulate soil C
storage along the catena. Peatland areas are shown in dark gray.
* MarkInfo [2006] http://www-markinfo.slu
Part of the dataset used in this publication was made available by the Swedish National Survey of
Forest Soils and Vegetation performed by the Department of For est Soils, SLU. The author are
solely responsible for the interpretation of data.
**Monthly averaged air temperature, precipitation and runoff measured at
Svartberget meteo station in 1981-2001 [Ottosson-Lfvenius et al., 2003].
17
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18
Measurements (Table 2)
Net ecosystem exchange was measured with an eddy-flux technique in a lawn section of
Deger Stormyr peatland since 2001 [Sagerfors et al., 2007] (site location on Figure
1b). DOC concentrations and stream discharge were observed at the outlet of the
Kallklsmyren peatland during the long-term monitoring program started in 1993
[Bishop and Laudon, unpublished data] (site location on Figure 1c). Unfortunately, an
integral assessment of both vertical (CO
2
) and horizontal (DOC) carbon fluxes for
Deger Stormyran [Sagerfors et al., 2007] has been published only at a late stage of this
PhD work, when recently established measurements of DOC concentrations at this
peatland became available [Laudon et al., unpublished data]. Those data were not
included in the thesis.
Laboratory materials
To obtain estimates of microbial DOC production and mineralization rates in peat, long-
term laboratory peat incubations were designed (Paper III). Experiments were done in
cooperation with the laboratory of the Department of Forest Ecology at SLU.
Methodology and information related to the laboratory experiments can be found in
Appendix of Paper III.
Modelling framework
Model formulation
The model supporting this work can be presented in two variants: forest and peatland
(Figure 2). Coupled to the peatland version, but not integral to it, is a model of DOC
concentration in peat water; in the course of this study, this was both formulated and the
computer programme developed (Figure 3).
The GUESS model [Smith et al., 2001; Sitch et al., 2003] was used here as a framework
for ecosystem simulations. GUESS (also known in its global version as LPJ) is a model
that describes population dynamics, with the focus on plant functional types competition
and replacement. GUESS is based on gap theory and simulates individual plant growth
in a number of replicate patches, the patch average representing the forest stand. For
each patch, stochastic generic disturbance is applied. The model follows generally
accepted schemes to simulate photosynthesis [Farquhar et al., 1980; Halexeltine and
Prentice, 1996a] and plant respiration [Sprugel et al., 1995; Halexeltine and Prentice,
1996a]. Soil biochemistry in the model is somewhat simplified: three pools of soil
organic matter are specified and the turnover rates are fixed as a simple function of
temperature. Intermediate complexity is assumed for other important ecosystem
processes, such as plant carbon allocation and light interception [Sitch et al., 2003]. The
hydrology subroutine utilises an empirical infiltration model [Haxeltine and Prentice,
1996b] and runoff is defined as any excess over the field capacity of a soil. Coupling
19
between evapotranspiration and photosynthesis is explicitly modelled [e.g., Sitch et al.,
2003]. The LPJ-GUESS model develops dynamically and is valuable in global and
regional C balance assessments [e.g., Morales et al., 2007; Prentice et al., 2007; Zaehle
et al., 2007], on a national level, particularly for Sweden [Koca et al., 2006], in research
projects related to plant population dynamics in changing environmental conditions
[e.g., Lucht et al., 2002], in paleostudies and for investigating site-specific ecosystem
phenomena. Recently, [Smith et al., on-going work] the model has also been coupled to
the regional version of the Swedish climate model RCA [latest version. Kjellstrm et
al., 2005].
In this study the ROMUL model replaced the standard formulation of soil biochemistry
in GUESS based on a simple compartment model. ROMUL [Chertov et al., 2001] is
also a compartment model, but the compartments are specified using the concept of
humus type. The humus of organic and mineral soil horizons is distinguished. The
rate of mineralization and the flow of matter between different compartments are
described by kinetic coefficients. Each coefficient is specifically dependent on both the
organic matter quality (C/N ration) and the environmental conditions (temperature and
moisture). The coefficients used for litter and humified forest floor were derived from
long-term laboratory studies [Chertov, 1985], and the rates of organic matter turnover in
the mineral horizon were deduced from the data on the activity of specific groups of soil
organisms [Chertov and Komarov, 1997]. The model has been evaluated against
numerous long-term experimental observations and provides good results for real forest
data sets [e.g., Chertov et al., 1997]
Forest modification for transect simulations (Figure 2a)
In this set up (Paper I, thesis) GUESS-ROMUL was run for a number of sites along the
topographical transect. A steady-state ground water level along the transect was
estimated and scaling method was used to correct the surface moisture to correspond
with the ground water depth (for a description of the methodology, see below). Each
simulated site, therefore, had a specific moisture regime, ranging from dry to
waterlogged. If the simulated steady-state ground water level was at or above the land
surface, indicating permanently waterlogged conditions, the peatland modification of
the model was used. Other environmental conditions, such as air temperature and
incoming solar radiation were the same for all the simulated sites, which is of course a
simplification.
Data collected from till soils within the Nordic region have demonstrated that it is
appropriate (at least in a statistical sense) to estimate surface soil moisture from the
local depth to the ground water table [Beldring et al., 1999]. Following Beldring et al.
[1999], a modified Brooks and Corey equation [Brooks and Corey, 1966] was used in
this study to calculate the surface soil moisture at a given site, based on (i) ground water
level and (ii) surface soil moisture at a reference ground water level (derivation and
20
Precipitation
Temperature
Light
availability
CO
2
Geology,
topography
Water availability
establ.:
tree and
grass PFTs
NPP
Leaves
and
roots
decomp. litter
mortality
wood
Water table
CO
2
soil
OM
Soil
texture
Precipitation
Temperature
Light
availability
CO
2
Water content
Sphagnum
NPP
decomp. litter
Water
table
CO
2
peat Topography
DOC
sorption
SSOC
export
a: forest model
b: peatland model
Figure 2. Schematic diagram of the model structure . Model core is an ecosystem
simulator GUESS . Shaded are extensions applied in this study: A-scheme for
simulation of ground water depth and soil moisture for selected topographical transect;
B-ROMUL soil organic matter dynamics model; C-Mixed Mire Water and Heat
model, MMWH; D-Model of DOC concentration in peatland. Model abbreviations are
explained in the text.
A
B
C
D
21
A gradient of the
the DOC flux
density
Zero-order
production,
first-order
mineralization
Linear
kinetics
How
modelled
D
0
, P,
1
k
s
Q
10
,, k
anP ,
k
an
K
D
, t
des
Parameters
Hydrological Microbial Sorption Processes
Any, dependent
on flow rate
Long/weeks-
years
Short/hours Time scale
MMWH model
Paper III. Implementation: numerical
or analytical solution of CDE
Paper IV. Implementation: numerical
solution of CDE
W
a
t
e
r

f
l
u
x
T, water content
Figure 3. A schematic representation of formulation and structure of the peatland DOC
model. Source: Paper III.
Modifier to account for the effect of oxygen supply on
1
k
an
Dispersivity
Molecular diffusion coefficient D
0
,,
Modifier to account for the effect of oxygen supply on P k
anP
Modifier to account for the effect of temperature on P and
1
Q
10
The constant reducing the microbial mineralization rate when the soluble OC is sorbed k
s
The rate of the microbial mineralization of DOC at 20
0
C
1basal
The rate of microbial DOC production at 20
0
C P
basal
Partitioning coefficient describing the sorption equilibrium K
D
Desorption kinetic constant t
des
22
discussion are presented in Appendix C). The reference ground water depth
corresponded to the standard GUESS soil moisture simulations. Corrections then
decreased or increased soil moisture at locations where ground water differed from the
reference, but the daily variability in soil moisture remained the same as in the standard
simulation.
Depth to the ground water table was estimated using an equation describing the steady-
state ground water level in a topographical transect between two aquatic bodies. The
method was originally proposed by Ivanov [Ivanov and Shumkova, 1967; Ivanov, 1981]
as a rough tool for peatland delineation and for predicting changes in peatland area in
response to altered water regime. Here a derivation of this equation (eq. 12 in Paper I)
starting from the Boussinesq equation for the subsurface one-dimensional flow
[Boussinesq, 1904] is presented for two cases: horizontal and tilted bedrock (Appendix
B). Surface topography, depth to the bedrock, effective precipitation, hydraulic
conductivity and porosity are required for the calculations. Sample calculations for the
profile crossing the studied peatland, Kallklsmyren, are included in this thesis. Bedrock
depth was taken from Sirin et al. [1998] and the hydraulic conductivity was chosen
from the normal range observed in Scandinavian tills [Lind and Lundin, 1990].
Peatland modification (Figure 2b)
To make GUESS-ROMUL applicable to peatland simulations, the MMWH model
[Granberg et al., 1999] was used to reconstruct soil thermal and water regimes.
Designed for boreal peatlands, the model provides a process-based description of heat
transfer and snow dynamics. The model also simulates daily fluctuations in water table
depth. An empirical, but justifiable, function [Weiss et al., 1998] was used here to
describe the steady-state moisture distribution in the unsaturated zone above the water
table.
Sphagnum moss was included as a new PFT within the model and equations that allow
moss photosynthesis rates to be controlled by variation in moss water content were
added [Williams and Flanagan, 1998]. Specific coefficients were used to describe
decomposition of the Sphagnum litter and peat and a simple parameterization of
decomposition under anoxic conditions was applied to a soil organic matter dynamics
module.
Model of DOC concentration in peatland water
The model was designed to reconstruct DOC concentration and fluxes in peatland
water. The starting point for model construction was a mass balance for dissolved
organic carbon in the form of convectiondispersion equation. In conjunction, a mass
balance equation was also written for the soluble organic carbon sorbed on a peat matrix
at a given time. Microbial DOC production and mineralization, sorptive exchange
between the solid and dissolved phases and hydrological transport with flowing water
are the main processes included in the model (Figure 3). Unlike previous DOC models
23
[e.g., Grieve, 1991; Boyer et al., 1996; Michalzik et al., 2003] the model proposed here
is based on a dynamic rather than a static description of sorption. The state variables
included in the simulation of DOC concentration, namely water fluxes (vertical and
lateral), water content in the unsaturated zone and peat temperature, were modelled
using the MMWH. The DOC model was applied to two situations: one to describe
stagnant water conditions in a laboratory peat incubation (Paper III), and another for the
field situation with flowing water in a peatland (Paper IV).
The derivations of two analytical solutions for the mass balance equations for dissolved
and sorbed soluble organic carbon in stagnant water conditions (eq. 5 and 6, Paper III)
are presented in Appendix A.
Parameterization and validation
Some of the DOC model parameters were derived from long-term laboratory peat
incubations that were conducted using peat collected from the Kallklsmyren peatland
(Paper III).
All model variables that were validated (Papers II, IV) are included in Table 2, p.18.
Mean square error, mean bias error and the Willmott index of agreement [Willmott,
1982] were chosen to characterize the model performance.
Model experiments
In this work, model tests were widely used. In such cases, the model was not run to
represent a real situation, but in order to study a specified factor. For example, it was
investigated how the simulated response of NEE to water table depth in a mire depends
on the parameterization of (i) photosynthesis and (ii) heterotrophic respiration in
relation to water content (Paper II). The model was also used to partition the integral
fluxes into contributory components (photosynthesis and heterotrophic respiration;
terms contributing to DOC balance).
Sensitivity and uncertainty issues
All the papers in this thesis address the issue of model sensitivity and uncertainty, but
more advanced methodology is used in the later papers. In Paper I it was found that the
disturbance regime regulates long-term litter input into the soil and, therefore, affects
soil C storage. A single sensitivity test was conducted to demonstrate the dependency of
simulated soil C storage in O horizons on a specified fire return interval (Figure 2, Paper
I). Simulating a large number of patches, with disturbance occurring randomly, also
gave some estimate of the uncertainty associated with the mean C storage (Figure 1,
Paper I). In addition, the paper included an illustration (Figure 5, Paper I) of the
24
potential size of the uncertainty associated with the moisture response function for the
soil organic matter transformation rates close to and above water saturation.
In Paper II global sensitivity analysis [e.g., Saltelli, 2000] was examined using Monte-
Carlo analysis for all model parameters introduced or modified for the peatland
simulations (Table 2, Paper II). It was both proposed and proved that the studied
variable, NEE, is characterized by a non-linear response to parameters (Table 5, Paper
II). The variance-based Sobol sensitivity indices were therefore chosen as a measure of
the parameters importance [Sobol, 1990; Saltelli, 2000]. One important result of the
sensitivity analysis was to show, through parameterizations, the key role that
autotrophic respiration plays in determining NEE.
In Paper III the kinetics of DOC sorption were studied by differential sensitivity
analysis [e.g., Saltelli, 2000]. The partial derivatives (local sensitivity coefficients) of
the DOC concentration with respect to the model parameters were expressed from the
original model equations. Normalized sensitivity coefficients were then plotted and
compared on the basis of hourly time intervals (Figure 4, Paper III). It was then possible
to separate two phases in the laboratory peat incubation: one ruled mostly by sorption
kinetics (the kinetic sorption coefficient has the highest effect on model results) and
another when microbial processes played the dominant role, though modulated by the
steady-state sorption partitioning.
In Paper IV the GLUE methodology [Beven and Binley, 1992] was used to estimate
uncertainty in DOC simulations associated with uncertainties in parameter values.
Outcomes of the Monte Carlo simulations were compared to the observed DOC
concentrations and a Willmott index calculated for each run as a measure of model
performance with a particular set of parameters. The prediction bounds (upper and
lower 10% quantiles) were then estimated for a daily time interval from the distribution
of all model outputs weighted by the Willmott index of agreement (Figure 3, Paper IV).
Global sensitivity analysis was also conducted, as described in Paper IV (Table 2, Paper
IV).
25
RESULTS
The main goal of this thesis was to explore different aspects of the interactions between
hydrology and carbon dynamics in boreal catchments. To achieve this goal a general
ecosystem model, GUESS, was linked to a dynamic soil organic matter model,
ROMUL, and modified so that new applications of the model to wet and water-logged
conditions were possible. A novel approach to modelling DOC concentrations was also
introduced and developed into an original model of DOC in peatland water. One
important result of this work was that relatively good model performance was
demonstrated both for NEE simulated at Deger Stormyr mire for 2001-2003 (Paper II,
Table 3, Figure 2) and for DOC concentration at the outlet of Kallklsmyren mire for
1993-2001 (Paper IV, Table 3, Figure 3). More specific results are described below.
Spatial variation in carbon storage along the catena. Dynamic aspects of the soil
moisturesoil C relationship (Paper I)
In Paper I the GUESS-ROMUL model was run for two contrasting soil moisture
conditions (mesic and mesic-to-wet). Some unpublished results from the model runs for
a number of sites (from dry to waterlogged) along the chosen topographical transect are
presented here in Figure 4.
1. Soil C storage in organic layers (litter, humified litter and peat) differs notably
depending on local soil moisture conditions. In areas where the water table
comes close to the surface (WT depth<1m), the GUESS-ROMUL model
predicts significantly greater O layer accumulation than in mesic and dry areas
(WT depth>1m) (Figure 4). Considering the high proportion of wet areas in the
boreal zone, simulations for well drained or average soil moisture conditions
may not provide the optimal estimate of soil C storage
2. Both organic matter decomposition rate and fire disturbance frequency
contribute to the difference in soil C storage in organic horizons between sites
with different moisture conditions (Figure 4). The model shows that, with a
natural fire regime, as much as 40% of the difference between mesic and
mesic-to-wet sites can be explained by the difference in litter input related to
fire frequency. Drier forests burn more often, and due to frequent disturbance
they are generally less productive because the age distribution shifts to
younger, less productive classes.
3. It is important to study, both under laboratory conditions and in the field, how
water content affects decomposers and their ability to utilize and transform
different fractions of litter and soil organic matter. The subject is
underrepresented in the current literature.
26
300 400 500 600 700 800 900 1000
10
20
30
40
50
Distance, m
E
l
e
v
a
t
i
o
n
,

m
2
4
6
8
C

i
n

O

l
a
y
e
r
s
,

k
g
C

m
-
2
100 300 500
3
4
5
6
7
C

i
n

O

l
a
y
e
r
s
,

k
g
C

m
-
2
Fire return interval, years
Dry Peatland Mesic Mesic- Mesic
moist
Site class Fire return *
interval
____________________
Mesic-moist 300 years
Mesic 100 years
Dry 50 years
140 a
c
water table depth,sim.
b
peatland border, map
Figure 4. (a) Simulated soil C storage in organic horizons for sites with
various moisture conditions along the catena illustrated in (b);
(b) steady-state estimates of average ground water elevations;
(c) relationship between fire-return intervals and C storage in O soil
layers at the mesic site. The lines of boxes in plots (a) and (c) indicate
the lower, median, and upper quartiles, and the whiskers show the
lowest and highest values. The location of topographical transect AB
is shown in Figure 1c. Mire borders taken from the map are shown in (b),
and calculated locations of points where the water table rises above the
land surface roughly match them. *Engelmark [1987]. Source: (a), (b)-
unpuplished model results, (c)-Paper I.
27
-30 -20 -10 0
-2
-1
0
1
C
O
2

f
l
u
x
,


g
C
m
-
2
d
a
y
-
1
-30 -20 -10 0
-2
-1
0
1
-30 -20 -10 0
-2
-1
0
1
-30 -20 -10 0
-2
-1
0
1
WT depth, cm
stand model
test1(psn)
test2(resp)
obs
model
a
b
d c
Figure 5. (a) Relationship between simulated daily photosynthesis rate
and water table depth for the lawn at Degero Stormyr. (b) Relationship
between simulated daily heterotrophic respiration and water table
depth (c)-(d) Relationship between NEE and water table depth
(c) Trend lines (best-fits by second-order polynomial regression) based
on measured data (solid) and standard model simulations (dashed);
(d) Trend lines based on the data from the standard model simulation
(bold solid), test run 1 (thin solid with crosses), and test run 2 (dash-
dotted). Test run 1: no effect of water table depth on photosynthesis.
Test run 2: decay constant, corresponding to anoxic conditions,
regardless of the water table depth. Source: (a), (b)-unpublished
model results, (c),(d)-Paper II.
psn model resp model
28
Mechanism of NEE regulation by WT level on a mire (Paper II)
In Paper II, net ecosystem exchange at Deger Stormyr was modelled using a modified
version of GUESS-ROMUL. The relationship between NEE and water table depth was
investigated
1. Model simulations showed that any changes in the water balance that do not
cause the water table to shift outside the normal range (10-20 cm) are likely to
have only minor effects on the annual rates of landatmosphere CO
2
exchange
in the studied peatland
2. When the water table level was either higher (> - 10 cm) or lower (< - 20 cm)
than normal the mire net CO
2
uptake decreased (Figure 5c)
3. According to model simulations the reductions in net CO
2
uptake rates that
occurred when the water table level was higher than -10 cm are attributable to
reductions in photosynthetic rate (Figure 5a, d)
4. The reductions in net CO
2
uptake when the lower water table level was lower
than -20 cm resulted from increases in heterotrophic respiration, rather than
reductions in photosynthesis (Figure 5b, d).
Hydrological and physico-chemical control of DOC production and export from
the mire (Papers III, IV)
Paper III introduces the concept of modelling DOC concentrations with a convection
dispersion equation. Some model parameters were derived on the basis of long-term
laboratory peat incubations and model behaviour for stagnant water conditions was
studied. In Paper IV DOC concentrations and fluxes at the outlet of Kallklsmyren were
simulated using a newly constructed model.
1. Soluble organic carbon is present in the mire system in two states: dissolved
(DOC) and sorbed, potentially soluble, but solid at a time (SSOC). SSOC
contributes a much higher proportion to the total store than does DOC (the
latter accounts for no more then 20%).
2. Sorption is one of the major processes determining DOC concentration and
therefore fluxes. Externally induced changes in the DOC concentration in the
peat pore water may be counteracted by sorption or desorption. Release of
DOC from the SSOC store occurs if DOC concentration is decreased by
dilution, for example, if precipitation water is added. DOC removal may occur
if the DOC concentration is increased, for example, if DOC-rich water from
the surface peat is advected into the layers below.
3. The balance between the two phases (DOC and SSOC) is time-dependent, and
adsorption and desorption have to be described as dynamic rather than static.
29
1993 1995 1997 1999 2001
10
20
30
40
50
60
70
80
S
S
O
C
&
D
O
C

s
t
o
r
e
d
,

g
m
-
2

D
O
C
,

m
g
L
-
1
1993 1995 1997 1999 2001
0
10
20
30
40
50
60
year
S
S
O
C
&
D
O
C

s
t
o
r
e
d
,

g
m
-
2
SSOC&DOC stored previous year, model
vol.weighted stream DOC, model
vol.weighted stream DOC, obs.
SSOC&DOC stored current year, model
net production, model
stream export, model
a
b
Figure 6. (a) Interannual variations in the amounts of SSOC and DOC
stored before the start of the hydrological year (spring flood) simulated
by the model. Interannual variation in the volume-weighted annual
stream DOC concentration, simulated by the model and calculated from
the interpolated observations. (b) Simulated components of the annual
DOC-SSOC balance: microbially-mediated net production and DOC
stream export. Changes from year to year in the SSOC amounts stored
in the acrotelm are also shown (estimated for the dates before the first
autumn frost).
30
4. Interannual variation in the average volume-weighted DOC concentration
generally resembles that of the SSOC and DOC store within the acrotelm, as
reproduced by the model (Figure 6a).
5. According to the model, some memory is characteristic of the system: the
store in the preceding year affects the concentration and the fluxes in the
following year (Figure 6a). The same is true when the seasons are considered
separately. The correlation between the simulated volume-weighted stream
DOC concentration in a particular season and the store of DOC and SSOC in
the preceding season is high for the winter (r=0.75, P=5.8 10
-5
) and summer
(r=0.87, P=1.9 10
-5
), but lower for the spring (r=0.40, P=0.023)
6. Interannual variability in the SSOC and DOC stores within the acrotelm, in
turn, depends on the conditions for microbially mediated DOC production and
mineralization during the current year (temperature, aeration) (Figure 6b: net
production) and flow intensity (Figure 6b: stream export).
7. It is important to include vertical peatland inhomogeneity in the model to give
an adequate description of the processes that are either strongly dependent on
depth (production, horizontal transport) or driven by vertical inhomogeneity
(sorption, dispersion)
CONCLUSIONS
The following conclusions can be drawn from this work:
- Patterns of soil C storage in a boreal landscape resulting from differences in soil
moisture are advantageous objects to model. Repeated and spatially predictable,
such patterns are helpful in a broader understanding of ecosystem dynamics and
functioning; particularly so, if we expect water balance to change in response to
altered precipitation patterns.
- Peatlands that occupy extreme habitats and exhibit specific self-regulating
mechanisms can be described using the same simulation framework as other
ecosystem types. It is important, however, to incorporate an adequate model of
peatland hydrology. With general models such as GUESS, that were not originally
designed for peatland simulations, predictions for one vegetation type are feasible,
but more work is required to parameterize competition between plant types.
- Wherever peatlands are present and drained by surface flow, a substantial flux of
DOC always accompanies the water flow, but seasonal and annual variations in
DOC concentration can be large. According to our model, the amount of soluble
organic carbon sorbed on a peat matrix strongly influences the annual volume-
weighted DOC concentration at a peatland outlet. The rate of microbial organic
matter transformation determines net production of soluble C, and sorption
partitions it between the solid and liquid phases. Most of the soluble C is stored in
a sorbed state and a part of it is released into flowing water when the sorption
equilibrium is shifted.
31
POSSIBLE APPLICATIONS AND FURTHER DEVELOPMENT.
The model developed in this study could be applied as a diagnostic and predictive tool.
Potentially, it can also offer an approach to the parameterization of smaller scale
processes in C balance models to be applied at larger scales. A few possible model
applications are presented below.
Diagnostic applications
It has been proposed [H. Grip, personal communication] that the approach used here to
reconstruct the depth to the water table along the selected topographical profile [Ivanov,
1981] could be applied to a study of ecosystem development in systems of newly
emerging islands. Specifically, this approach provides a rough estimate of the steady
state distribution of peatlands, defined as locations where the calculated water table is
situated above ground level. It is, therefore, possible to examine whether newly formed
landscape features gradually develop towards this theoretical steady state and how fast
this happens. Even more interesting could be to run the whole GUESS-ROMUL model
to see if it can reconstruct vegetation and soil dynamics over longer time intervals.
Peatland modification of the model can be used to study the relationship between NEE
and the depth to the water table for other peatland microtopographical features, in
addition to the lawn type studied here, but only under the assumption of unchanging
plant species composition.
New tests could be conducted using the DOC model to see how changes in peat sorption
properties affect DOC concentration [K. Bishop, personal communication]. Additional
development is needed, however, to introduce the dependency of kinetic sorption
coefficients on chemical characteristics of the soil solution, such as ionic strength [e.g.,
Skyllberg and Magnusson, 1995] and SO4
2-
concentration [e.g., Clark et al., 2006].
Predictive applications
Some estimates of the effect of increased precipitation on the soil C storage in the
organic horizons were presented in Paper I. To evaluate the net change in the soil C
store more sites and climate scenarios need to be included in the analysis. The influence
of moisture conditions on ecosystem functions may be as important as that of
temperature. Although there is great uncertainty associated with predicting precipitation
using climate models [e.g., Palmer et al., 2005], it is very important to further evaluate
the role of changed soil moisture conditions on the soil C store. Such a study could also
consider the effect of precipitation change on DOC fluxes.
The peatland modification of the model in its current version does not describe all
aspects of plant competition, and long-term changes in NEE related to shifts in species
32
composition cannot be predicted by the model. It is, however, possible to run the model
for various climate scenarios and see whether the present vegetation types are still stable
under new conditions, i.e. if NEE stays within its normal range in the specific plant
community [M. Nilsson, personal communication].
The DOC model can be used to predict the effects of climate and human activity on
DOC fluxes from peatlands. The model could also be used to test some of the widely
discussed hypotheses such as the enzymatic latch mechanism, i.e. the increase in
anaerobic peat decomposition resulting from a temperature increase [Freeman et al.,
2001a, 2001b; questioned by Tranvik and Jansson, 2002], to eliminate the role of
resistance time, intensity and pathways of water flow.
Scaling
The question is sometimes posed: Are big basins just the sum of small catchments?
[e.g., Shaman et al., 2004]. The question is not only relevant to runoff modelling, but
also to all aspects of C balance modelling where local water flow and surface moisture
patterns become important (identification of wetlands and wet areas emitting methane,
modelling carbon accumulation in wet soils, studying cycling and loss of DOC etc). It is
important to know whether small basins already contain most of the variability in the
properties studied, and thus whether statistical information about the distribution of
these properties on smaller scales is enough to extrapolate to larger scale assessments.
The existence of a representative elementary area (REA) has yet to be theoretically
proven and may not be possible [e.g., Fan and Bras, 1995]. Intuitively, however, it is
clear that patterns of soil C storage in a boreal catchment, regulated by soil moisture, are
spatially predictable and repetitious. The spatial variability in DOC concentration in
aquatic bodies, which is a reflection of water routing through the patterns of organic
soils [e.g., Bishop et al., 2004], could be expected to have a characteristic pattern too.
Some information about these features at the catchment scale could, therefore, improve
regional and global C predictions. Descriptions of a large-scale picture can be built on
generalizations, rather than neglecting small-scale complexity. Potentially, there are two
alternatives, presented below, in which the model developed in this study could be
placed in the broader context and applied to scaling issues
Alternative I. Explicit catchment hydrology model
In this option the GUESS-ROMUL model could be coupled to the catchment hydrology
model. Important criteria for model choice are: explicit moisture accounting in the
unsaturated zone; water table fluctuations; the interaction between the saturated and
unsaturated zones; and explicit solutions of the flow equation in the modelling domain
(the latter is important for DOC predictions). The TOPOG model [e.g., Vertessy et al.,
1993; Zhang et al., 1999] may be a good option here. It uses an analytical solution of
the Richards equation to describe vertical flow in the unsaturated zone and an explicit
33
kinematic wave solution for the lateral fluxes in the saturated zone. The TOPOG-
Dynamic model [Silberstein et al., 1999] can also simulate plant growth. In addition, a
package that solves the convectiondispersion equation is available, and can be used to
simulate DOC concentrations. One advantage of this approach is that it is possible to
describe explicitly both flow and surface soil moisture within a catchment at a relatively
good spatial resolution. Therefore, straightforward validation of the model is possible
where data is available. One disadvantage, however, may be that a lot of computer time
is needed to run such a model. It is doubtful whether on-line nesting of the catchment-
scale model into the larger-scale C balance models is practically possible.
Alternative II. Statistical generalization of the catena approach
In a statistical sense it is valid to relate the surface soil moisture to the depth to the
ground water [Beldring et al., 1999]. If the area is drained by a series of parallel streams
then the approach used here to estimate, roughly, the steady-state ground water depth is
potentially applicable. It is then possible to collect sufficient information about patches
with various water table depths by sampling a number of transects perpendicular to the
streams in a catchment. Ensemble averaging or other statistical procedures could be
used to generate a catena that is not a real transect, but rather a statistical generalization
for the whole area or a whole cell within a large-scale model. Instead of a number of
random patches currently simulated by GUESS, patches with characteristic soil
moisture and fire disturbance regimes could be modelled. Changes in the distribution of
patches due to changes in the water budget can be predicted using the equation for the
average water table depth. Analytical solutions for the kinematic wave equation [Troch
et al., 2002] can be used to simulate daily water fluxes at specific locations along the
catena if daily accounting of DOC fluxes is required. The accompanying convection
dispersion equation can also be applied to the DOC, and an analytical solution is
possible under some simplifying assumptions. It has yet to be proven whether the
proposed generalization is mathematically correct. However, such an intuitive approach
is found in classical ecological studies, where patterns and relationships found in sample
catena studies are extrapolated to the whole geographical region of interest.
34
APPENDICES
Appendix A
A1. Derivation of Equation (5) in Paper III
Starting from equations (3)-(4) in Paper III (symbol definitions in the paper)
, a M P a
V
M a
K
t
a
c 1 s des
c
D des
c
+ +

=
c
c
t t
(1)
, a a
V
M a
K
t
a
s 2 s des
c
D des
s

=
c
c
t t
(2)
(1)+(2)
, a a M P
t
) a a (
s 2 c 1
s c
=
c
+ c

(3)
by definition , k
1 s 2
= (4)
using (4) in (3)
), a k a ( M P
t
) a a (
s s c 1
s c
+ =
c
+ c

(5)
at the sorption steady-state
, a
V
M a
K
s des
c
D des
=

t t
,
V
M
K a a
D c s
=
(6)
using (6) in (5)
),
V
M
K k 1 ( a M P
t
)
V
M
K 1 ( a
D s c 1
D c
+ =
c
+ c

introducing
V
M
K 1 1 K
D
+ = and )
V
M
K k 1 ( 2 K
D s 1
+ = , (7)
c
c
c
c
a
1 K
2 K
M
1 K
P
t
a
, a 2 K PM
t
a
1 K
=
c
c
=
c
c
Introducing
1 K
PM
3 K = and
1 K
2 K
4 K = , (8)
35
,
4 K
3 K )) t t ( 4 K exp( ) a 4 K 3 K (
a
)), t t ( 4 K exp(
a 4 K 3 K
a 4 K 3 K
, t 4 K
a 4 K 3 K
) a 4 K 3 K (
, a 4 K 3 K
t 4 K
) a 4 K 3 K (
, a 4 K 3 K
t
a
1 1 c
c
1
1
c
c
t
1 t
t
1 t
c
c
c
c
c
c

=
=


c =

c
=
c
c
=
c
c
) )
,
4 K
3 K
)) 1 t t ( 4 K exp( )
4 K
3 K
a ( a
1 c c
+ = (9)
from (8) and (7)
V
M
K 1
)
V
M
K k 1 (
1 K
2 K
4 K
D
D s 1
+
+
= =

,
)
V
M
K k 1 (
PM
2 K
PM
4 K
3 K
D s 1
+
= =

,
by definition V ) t ( c a
c
= and V c a
1
1
c
= ,
,
V
M
K k 1
M P
)) t t (
V
M
K 1
V
M
K k 1
exp( )
V
M
K k 1
M P
V c ( V ) t ( c
D s 1
1
D
D s
1
D s 1
1
|
.
|

\
|
+

+
|
|
|
|
.
|

\
|
+
+

|
.
|

\
|
+

A2. Derivation of Equation (6) in Paper III

If volume of water, V, and mass of peat, M, are constant during the experiment, then
from the mass conservation:
V c M s V c M s
0 0 1 1
+ = + ,
where c
0
and c
1
are DOC concentration
and s
0
and s
1
are SSOC concentration at the start of experiment and at t
1
, respectively
0 0 1 1
c
V
M
s c
V
M
s + = + ,
At steady-state
1 D 1
c K s = ,
where K
D
is an equilibrium sorption partitioning constant
0 0 D 1
c
V
M
s ) 1
V
M
K ( c + = + ,
36
,
1
V
M
K
c
V
M
s
c
D
0 0
1
+
+
=
Appendix B. Derivation of the equation for the annual average steady-state depth
to the water table on a 1D hillslope
Subsurface flow from a one-dimensional hillslope with a sloping aquifer can be
described by the Boussinesq equation [Boussinesq, 1904]
f
P
x
h
i sin
x
h
h
x
i cos
f
k
t
h
+
(

c
c
+ |
.
|

\
|
c
c
c
c
=
c
c
, (1)
where h is elevation of the water table measured perpendicular to the underlying
impermeable bedrock
i is a slope angle of the bedrock
k is hydraulic conductivity
f is drainable porosity
P is recharge to the saturated area.
When averaged over a year, the changes in unsaturated zone storage can be ignored. In
till soils surface discharge by overland flow is seldom observed. P then equals effective
precipitation, the difference between precipitation and actual evapotranspiration.
Equation (1) is written using Cartesian coordinates, xh (Figure A), so that the x-axis
runs parallel to the bedrock
For slightly sloping bedrock , i cos i sin <<
f
P
x
h
h
x
i cos
f
k
t
h
+
(

|
.
|

\
|
c
c
c
c
=
c
c
,
A steady-state water table depth can be calculated from:
i cos k
P
x
h
h
x
= |
.
|

\
|
c
c
c
c
,
) )
=
c
c
c
x
0
x
0
dx
i cos k
P
x
h
h ,
37
, c ) x x c (
i cos k
P
h
, c x c
i cos k 2
Px
2
h
, x ) c
i cos k
P
( h h
, c x
i cos k
P
x
h
h
4
2
3
2
2 1
2 2
x
0
x
0
1
1
+ =
+ + =
c + = c
+ =
c
c
) )
, c ) x x c (
i cos k
P
h
4
2
3
+ = (2)
where c
1
, c
2
, c
3
and c
4
are integration constants
In the case when elevation of the water level above the bedrock is the same in left and
right draining streams (Figure A1), equation (2) is symmetrical. The point with the
highest elevation of the water table along the hillslope is an equal distance, a, from the
left and right streams. Constants in (2) can then be determined:
At the left stream location , h c , c h , 0 x
2
0 4 4
= = =
Where h
o
is the average elevation of the stream water level above the bedrock
At the right stream location , h h , a 2 x
0
= =
, a 2 c
0 ) ) a 2 ( a 2 c (
i cos k
P
3
2
3
=
=
Replacing c
3
and c
4
in (2) , h ) x ax 2 (
i cos k
P
h
2
0
2
+ = (3)
Equation (3) for the case of horizontal bedrock (cos i=1) was used in Paper I with
reference to Ivanov [1981], who published it without derivation, but referring to
Kamensky [1935].
If another coordinate system h
~
x
~
is chosen so that elevation h
~
is calculated from a fixed
reference depth, which is absolute rather then related to the bedrock (Figure A1), then
equation (3) can be transformed:
, i tan ) x a 2 (
i cos
h
h
~
+ =
Similar to the symmetrical case is a derivation for the non-symmetrical case (Figure
A2), when beds associated with the left and right streams have different elevations from
the bedrock. Separate equations are then written for the portions of the hillslope
draining into the left and right streams respectively:
38
x
x
h
h h
0
h
0
h
max
a
h
0
i
h
0l
L
h
0r
h
l
h
0l
h
l
x
l
x
l
h
0r
h
r
h
r
h
max
a
l
a
r
x
r
x
r
i
Fig.A. Catchment characteristics used for the water table depth calculations
(Appendix B). Symbol definitions are in the text.
A1: symmetric case a=L/2;
left and right channels have the
same water level relative to the
impermeable bed
A2: asymmetric case a
l
+a
r
=L;
left and right channels have
different water level relative
to the impermeable bed
free ground water surface positions
impermeable horizon
39
Left portion:
, h ) x x a 2 (
i cos k
P
h
2
l
0
2
l l l l
+ = (4)
Right portion:
, h ) x x a 2 (
i cos k
P
h
2
r 0
2
r r r r
+ = (5)
The divide location
r r l l
a x , a x = = (Figure A2) can be found from
r l
h h = , and L a a
r l
= +
2
L
i cos k P L 2
h h
a
2
l 0
2
r 0
l
+

= , (6)
If another coordinate system h
~
x
~
is chosen so that elevation h
~
is calculated from the
fixed reference depth, which is absolute rather then related to the bedrock (Figure A2),
then equations (5)-(6) can be transformed:
, i tan x
i cos
h
h
~
, i tan ) x L (
i cos
h
h
~
r r
l l
+ =
+ =
Equations (5)-(6) were used in the thesis to reconstruct the water table position along
the sample catena (Figure 4 thesis).
Appendix C. Derivation of the equation for the correction factor for soil moisture
related to the local water table depth
The Brooks and Corey equation [1966] gives the following expression to connect
capillary potential ( ) and soil water content (u ):
,
n
a
r o
r

u
u u
|
|
.
|

\
|
=

(1)
where n
0
-porosity
a
-air entry tension head
- a parameter that depends on the pore size distribution of the soil
u -residual water content
Application of the Brooks-Corey characteristic function for simulating surface moisture
(
0
u ) from the depth to the ground water (z
WT
) assumes the steady-state distribution of
soil moisture in the unsaturated zone above the water table*. In steady-state conditions
40
(no vertical flow) the total potential is zero and the capillary potential equals the
gravitational potential. At the soil surface:
WT
z =
Equation (1) can then be modified for the surface water content, assuming zero residual
water content:
a WT
WT
a
0 0
z if ,
z
n

>
|
|
.
|

\
|
= (2)
Surface water content,
0
u , simulated by phenomenological equations in ecosystem
model GUESS can be related to a certain average water table depth. This can be
achieved by fitting simulated
0
u in (1) and estimating z
WT
for a longer time interval (a
year), with subsequent averaging. The reference water table depth, z
R
, obtained this way
can then be used to introduce a correction factor to the surface water content for sites
with different local water table depths.
a WT
WT
R
R
R WT
R a
0
z if ,
z
z
z z
z
n u

u

>
|
|
.
|

\
|
=
|
|
.
|

\
|
=
where u is the water content at a specified location and
R
u is the surface water content
simulated by GUESS, assuming free drainage. This correction does not change the daily
variability in simulated water content, but rather the absolute value becomes smaller for
dry locations with a deep water table and larger for wet and semi-wet areas with
impeded drainage and a shallow water table. Estimates are rough, but soil moisture
observations in till soils at sites with various water table depths [Beldring et.al, 1999]
show that they are not without value.
For
a WT
z s surface water content is assumed to be equal to porosity
* The following factors have to be considered to justify the validity of the steady-state
approximation [e.g., Grip and Rodhe, 1994]:
- Hydraulic conductivity should be high enough so that any shift from the steady-
state moisture distribution caused by precipitation or evapotranspiration would
lead to a new steady-state being established within a relatively short time period
- Groundwater should not be too deep. If the groundwater is deep, a zone with low
water content and low water conductivity may form between the surface layer and
the capillary fridge above the water table. In this case, no equilibration between
the two would occur.
Till soils generally satisfy the above conditions. Because of relatively low effective
porosity, the water table in till soils is usually shallow [Grip and Rodhe, 1994]. Water
table depth also varies greatly over the seasons [Grip and Rodhe, 1994]. No drastic
reduction in hydraulic conductivity occurs as a result of drying, and, in general,
equilibration within the unsaturated zone is relatively fast [hours according to numerical
experiments using Richards equation]. The capillary fridge usually reaches the soil
surface and variations in the surface soil moisture follow variations in water table depth
[e.g., Beldring et al., 1999].
41
42
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48
POPULRVETENSKAPLIG SAMMANFATTNING
Koncentrationen av vxthusgaser i atmosfren har kat kontinuerligt under de senaste
200 ren, och kningen tillskrivs huvudsakligen antropogena utslpp. FNs senaste
klimatrapport slr vidare fast att den kade halten av vxthusgaser i atmosfren leder till
klimatfrndringar. Bde frhjda halter av vxthusgaser i atmosfren och ett ndrat
klimat pverkar dock omsttningen av vxthusgaser i olika ekosystem, s.k.
terkopplingsmekanismer. Frndringarna i ekosystemens funktion kan leda till
antingen kad eller minskad produktion eller upptag av vxthusgaser. D.v.s att
frndringarna i ekosystemens funktion kan antingen ytterligare accelerera eller bromsa
vxthuseffekten. Ett av de viktigaste redskapen fr att frst hur olika ekosystem svarar
p klimatfrndringar r modeller.
Syftet med den hr avhandlingen har varit att utveckla och att anpassa
ekosystemmodeller till boreala ekosystem. Arbetet har utgtt ifrn en generell
ekosystemmodell, GUESS, till vilken en srskild modell fr markens biogeokemiska
processer, ROMUL, har kopplats. Arbetet har huvudsakligen inriktats p att utveckla
modeller fr att studera interaktioner mellan vattnets omsttning, hydrologin, och
omsttningen av kol. Fltdata fr svl parametrisering som testning av modellerna
kommer frn Vindelns Skogliga Frsksparker, Sveriges Lantbruksuniversitet, ca 6 mil
frn Ume, norra Sverige, och har tillhandahllits av olika forskargrupper. Mer precist
har fljande frgestllningar behandlats:
- Hur varierar mngden organiskt material i marken som en funktion av
varierande markfuktighet i ett skogslandskap?
- Hur pverkas utbytet av koldioxid mellan en myr och atmosfren av
vattenytans position, d.v.s. vad hnder om vattenbalansen ndras?
- Vad kontrollerar mngd och variabilitet av lst organiskt kol i avrinningen frn
torvmarker? ven vilka faktorer som styr variationen av lst organiskt material
p olika tidsskalor har studerats.
Simuleringarna av mngden organiskt material som en funktion grundvattenytans lge
utmed olika transekter i ett skogslandskap genererar samma variation i organiskt
material som terfinns i landskapet. Dock visar modellresultaten att olikheter i mngd
organiskt material bara till viss del kan frklaras med enbart variation i markfuktighet.
En vl s viktig faktor r frekomsten av skogsbrand. Om man antar naturlig frekomst
av skogsbrand s hrrr ca 40% av variationen i humuslagrets tjocklek frn brand. I och
med att mnniskan i dag, i Sverige, mycket framgngsrikt bekmpar skogsbrnder tyder
detta p att humustckets tjocklek i Svenska skogar sannolikt kommer att ka.
Ytterligare ett viktigt bidrag i avhandlingen har varit inkorporeringen av torvbildande
ekosystem, myrar, i den generella ekosystemmodellen GUESS. Den nya versionen av
modellen anvndes b.la. fr att studera hur varierande vattenyteniv samt temperatur
pverkar utbytet av koldioxid mellan myren och atmosfren. Modellen visar tydligt att
s lnge vattenytan varierar inom det intervall som r typiskt fr det dominerande
49
vxtsamhllet s r fldet av koldioxid relativt stabilt. Om vattenytan blir betydligt
hgre eller lgre n normalt s pverkas utbytet av koldioxid i frsta hand genom att
fotosyntesen pverkas och i mindre utstrckning genom ndrad nedbrytning.
Ytterligare en utveckling av modellen gjordes fr att kunna modellera frekomsten av
lst organiskt material i markvatten i myrar samt i avrinningen frn myrar.
Modelleringen av koncentrationen av lst organiskt material baseras p en ekvation fr
advektion och dispersion. Modellen inkluderar vidare bde produktion, nedbrytning
samt adsorption av det lsta organiska materialet. Eftersom mngden humus i sjar och
vattendrag har stor betydelse fr mnga av vattnens biogeokemiska funktioner och dess
kvalit r det av stor vikt kunna modellera mngden lst organiskt material och hur
denna pverkas av olika mnskliga aktiviteter svl som av klimatfrndringar.
Modellkrningar visar bl.a. att de huvudsakliga faktorerna som bestmmer nivn p hur
mycket lst organiskt material som kan finnas i vattnet r hastigheten p bildningen
samt nedbrytningen av det lsta organiska materialet. Hur mycket lst organiskt
material som finns vid en given tidpunkt bestms dremot huvudsakligen av
adsorptionsprocesser till det fasta organiska materialet.
De olika utvecklingarna och anpassningarna av ekosystemmodellen GUESS-ROMUL
r ett viktigt bidrag bde fr att ytterligare kunna frst kolomsttningen i olika
ekosystem men ocks fr att gra uppskattningar av storleken p olika processer eller
flden ver strre omrden.
50
I
I
ecologi cal modelli ng 2 0 4 ( 2 0 0 7 ) 475484
avai l abl e at www. sci encedi r ect . com
j our nal homepage: www. el sevi er . com/ l ocat e/ ecol model
Carbon storage in the organic layers of boreal forest soils
under various moisture conditions: A model study for
Northern Sweden sites
Alla Yu. Yurova

, Harry Lankreijer
Department of Physical Geography and Ecosystems Analysis, Lund University, S olvegatan 12, 223 62 Lund, Sweden
a r t i c l e i n f o
Article history:
Received 31 October 2005
Received in revised form
29 January 2007
Accepted 1 February 2007
Published on line 23 March 2007
Keywords:
Soil moisture
Carbon sequestration
Boreal forest
Vegetation dynamics
Model
ROMUL
GUESS
a b s t r a c t
A typical feature of the boreal forest landscape is a gradient from dry to wet sites, with
associated increases in the depth of the soil organic layers. In this study, the coupled
ecosystemsoil biogeochemistry model GUESSROMUL is used to explore how the spe-
cic features of soil organic matter decomposition and vegetation dynamics account for an
observed difference between the soils formed under contrasting moisture conditions. Two
sites, one mesic and one mesic-to-wet, representative of the natural forest in Northern Swe-
den, are simulated. In addition to the assumptions underlying the GUESSROMUL model,
it is assumed that the re frequency was higher at the mesic site. The model shows that
with a natural re regime, the soil organic layers at the mesic-to-wet site store 6.0kgCm
2
compared to 3.1kgCm
2
at the mesic site. Forty-seven percent of the difference between
the sites in this respect is explained by suppressed decomposition under higher moisture
conditions, 37%by the decreased litter input into the soil (more frequently disturbed ecosys-
tems have lower productivity) and 16% by direct consumption of the forest oor in res. It
is predicted that due to anthropogenic re suppression the organic soil layers of mesic sites
will, in the future, sequester carbon at an average rate of 0.0103kgCm
2
year
1
and have
an equilibrium storage capacity of 5.4kgCm
2
. For the mesic-to-wet site, the model pre-
dicts an extremely slow sequestration rate of 0.0022kgCm
2
year
1
. The effect of increased
precipitation on the carbon storage at the landscape level is also investigated.
2007 Elsevier B.V. All rights reserved.
1. Introduction
The soil, which represents a large and dynamic carbon pool,
is a key element that must be considered when discussing
how carbon storage in boreal ecosystems may be inuenced
by future climate change. Some fractions of soil carbon, such
as humic substances bound to mineral matrices, have a very
slow turnover rate and therefore changes in carbon storage
capacity should be considered over a long time scale (e.g.
Trumbore and Harden, 1997). However, the soil organic lay-

Corresponding author. Tel.: +46 46 2223749; fax: +46 46 2220321.

E-mail address: alla.yurova@nateko.lu.se (A.Yu. Yurova).
ers (O layers) are much more labile and the carbon content in
them can vary considerably, even within a decade (Bormann
et al., 1995; Yu et al., 2002). Among the factors affecting soil C
dynamics, changes in litter production and soil temperature
are most often considered. Following anticipated increases
in atmospheric CO
2
levels and temperature, primary produc-
tion is expected to increase, leading to increases in organic
material inputs to the soil. On the other hand, rises in soil
temperature may compensate for, or even overbalance this
effect by increasing respiratory losses from the soil. Currently,
0304-3800/$ see front matter 2007 Elsevier B.V. All rights reserved.
doi:10.1016/j.ecolmodel.2007.02.003
476 ecologi cal modelli ng 2 0 4 ( 2 0 0 7 ) 475484
there is no general agreement regarding how these two forces
are balanced in the boreal region, but soil organic matter mod-
els are widely used in attempts to explore the relationships,
and mechanisms, involved (Liski et al., 1999; Hyvonen et al.,
2002;

Agren and Hyvonen, 2003; Davidson and Janssens, 2006;
Carrasco et al., 2006).
Model simulations typically assume that soil moisture
conditions are average, or optimal, despite the fact that obser-
vations suggest that there is a strong correlation between the
amount of carbon in the O layers and soil moisture content.
A typical boreal landscape can be divided into the following
structural units according to the soil moisture regime: (i) the
driest watershed parts and convex slopes, (ii) mesic sites, sit-
uated mainly on the straight slopes, (iii) the hollows, lower
parts of the catchment adjacent to the streams and the con-
cave slopes withfavourable moisture conditions and (iv) mires
with water-logged conditions. The mires store the greatest
amounts of carbon per unit area, but they represent ecosys-
tems with distinct vegetation and hydrological regimes that
are beyond the range of the moisture gradient explored in this
study, so they are not considered here. Instead, we focus on
the gradient encompassing the other three units from dry
sites, with usually poor O layers (typical mor), to wet sites, on
fully developed humic podzol (rich Olayers, hydromor) or peat
soil that is a basic feature of boreal landscapes where mires
have not developed.
The well-drained and wet soil types are so quantitatively
and qualitatively different that they should be expected to
respond differently to a changing environment. Therefore, in
order to reliably predict the future carbon storage capacity of
the boreal zone, soil organic matter models should be tested
and further developed by incorporating the effects of varia-
tions in moisture conditions.
Acrucial issue is why the Olayers of moist forest soils accu-
mulate more C than those of well-drained soils. The accepted
explanation is that decomposition is suppressed under high
moisture conditions. However, there is an alternative hypoth-
esis linking the observed differences to the historical patterns
of re disturbance (M. Nilsson, pers. commun., 2004). Moist
sections of the boreal forest used to burn less frequently than
dry sections (e.g. Engelmark, 1987). Consequently they tended
to have higher proportions of mature trees (e.g. Wardle et al.,
2003) and, hypothetically, higher long-term average rates of
litter production. The fact that the amount of carbon stored in
the Olayers canbe as highas 27kgCm
2
(equivalent toadepth
of ca. 0.5m) in boreal ecosystems where the disturbance fre-
quency is apparently low provides support for this hypothesis
(Hesselman, 1926; Wardle et al., 2003). These unique soils were
found in old nature reserves, or unmanaged and rarely burned
forests on islands, and were not characterised by high soil
moisture content. For comparison, a meanestimate of the car-
bon storage capacity for the global boreal forest is 3.3kgCm
2
(Vogt et al., 1986).
It has been reported that as the disturbance frequency
in natural forest is reduced, soil begins to act as a C sink
(Kurz and Apps, 1996; Bhatti et al., 2002). Therefore, we could
speculate that anthropogenic re suppression could lead to
the increased sequestration of carbon by the O soil layers,
thus reducing the differences betweenmoist and well-drained
soils.
The aim of the study presented here was to use the cou-
pled ecosystemsoil organic matter model GUESSROMUL to
examine the effects of two moisture conditions, mesic and
mesic-to-wet, on carbon storage in organic layers of the natu-
ral Northern Sweden boreal forest soils. The model simulates
vegetation dynamics, including the disturbance regime, and
decomposition of various types of organic matter in relation
to soil/litter moisture levels. Analysis of the present situa-
tionandfuture predictions are made. Some roughcalculations
concerning the effect of current changes in precipitation on
carbon storage in the soil are also included in the study, by
changing the relative proportions of sites with different mois-
ture regimes in the simulated landscape.
2. Description of the modelling system
In the present study the general ecosystem simulator GUESS
(Smith et al., 2001) has been fully coupled with the soil organic
matter dynamics model ROMUL (Chertov et al., 2001); the
combined modelling system is called GUESSROMUL. It is
important to note that the GUESS model incorporates a formu-
lation of litter and soil biogeochemistry that was not applied
here.
GUESS is a terrestrial ecosystem model that simulates veg-
etation dynamics and carbon and water exchange. The key
processes in the model are vegetation establishment, growth,
mortality and the competition for light among plant individ-
uals, which determines vegetation composition and natural
succession. Each simulated woody individual belongs to a cer-
tain species or plant functional type (PFT) with a specic set
of parameters controlling its potential climatic space, ability
to survive under various stress conditions, scaling factors for
photosynthesis and respiration, allometry and carbon alloca-
tion, phenology, establishment and mortality. GUESS can be
applied to any given ecosystem by specifying these general
parameters rather than by using a direct validation proce-
dure. The driving force is soil texture and climate, provided
as model inputs, therefore there is always a generic connec-
tion between the variability of environmental parameters and
changes in ecosystem structure and functioning (concerning
the latter, changes in productivity are the most frequently
discussed). The model has been recommended for, and suc-
cessfully applied to, landscape-to-regional scale simulations
of various systems, including the Swedish boreal forest (Koca
et al., 2006). For a comprehensive description the reader is
referred to Smith et al. (2001). Here only the features of the
model that are directly linked to the simulation of soil carbon
dynamics (such as litter production, the disturbance regime,
soil temperature and moisture) are outlined.
2.1. Litter production
Net primary production (NPP) is modelled in GUESS by explic-
itly representing photosynthesis and maintenance respiration
(Haxeltine and Prentice, 1996; Haxeltine et al., 1996; Sitch et
al., 2003). At the end of the simulation year, after subtract-
ing the reproduction costs from NPP, the remaining fraction is
partitioned amongst leaf mass, ne root mass and sapwood
using allometric relationships. Leaf and ne root biomass are
ecologi cal modelli ng 2 0 4 ( 2 0 0 7 ) 475484 477
transferred into the litter pool with a given turnover rate. The
mortality of leaves (needles), root and woody biomass also
provides inputs into the litter pool. Mortality in GUESS is a
stochastic process with a background rate related to plant
longevity and growth efciency.
2.2. Disturbance
GUESS simulates individual plant growth in a number of repli-
cate patches, the patch average representing the forest stand.
For each patch, stochastic generic disturbance is applied with
a prescribed frequency, with disturbed biomass being trans-
ferred to the litter pool. This accounts for disturbances such
as wind or insect damage but does not include re, which is
modelled explicitly in GUESS as described by Thonicke et al.
(2001). Two basic assumptions are made in the model: (i) a
re can only occur if the amount of litter stored in a forest
oor exceeds a threshold value, and (ii) soil moisture is a key
environmental variable determining the probability of re. Fire
resistance prescribed for the PFT or plant species indicates the
percentage of individuals consumed in a re. A fraction of the
litter is also burned if the re occurs, and the re resistance of
the litter is assumed to be the same as for living individuals.
For comparison with historical data and reconstructions the
re probability estimated in the model can be recalculated as
an average re return interval.
2.3. Soil temperature and water balance
Soil temperature is assumed to follow surface temperature
according to a Fourier annual sinusoidal cycle with a damped
oscillation amplitude and a temporal lag. Soil thermal diffu-
sivity is dependent on both the soil texture and water content.
Representation of the soil moisture in GUESS is based
on Haxeltine and Prentice (1996). A two-layer, simple water
balance scheme includes evapotranspiration, snowmelt, per-
colation and runoff. An excess of soil moisture over the
available water holding capacity in the rst and second
layers is attributed to surface runoff and drainage to the
ground water, respectively. The percolation coefcient and
eld capacity are texture-dependent constants.
It has been demonstrated that LPJ (a global version of the
model, sharing all of the components except the vegetation
dynamics component with GUESS; Smith et al., 2001) accu-
rately reproduces the spatial patterns of soil moisture on a
coarse scale (Wagner et al., 2003), as well as the seasonality
of soil moisture for specic sites (Sitch et al., 2003). Further
details about the hydrological evaluation of the model can be
found in Gerten et al. (2004). Sensitivity analysis of the LPJ
model (Zaehle et al., 2005) has revealed that, in water-limited
regions, parameters controlling the evapotranspiration rate
are of considerable importance for the simulated net pri-
mary production. Wolf et al. (in revision) have studied the
uncertainty of GUESS model simulations arising from the
mathematical formulation of the water balance. The simu-
lations for the region in Northern Russia produced by (i) the
standard model, and (ii) GUESS, driven by the soil moisture
estimates produced by the mechanistic model JULES (Cox et
al., 1999), were compared. It was shown that differences in
soil moisture had a minor inuence on the vegetation carbon
uxes simulated with GUESS, whereas the soil carbon uxes
were very sensitive to the soil moisture estimates.
ROMUL (Chertov et al., 2001) is a dynamic soil organic
matter model based on the concept of humus type. The
model was originally called SOMM(Chertov, 1985; Chertov and
Komarov, 1997) and was developed with data from long-term
laboratory experiments on the decomposition of plant litter in
relation to the chemical composition of the material (nitrogen
and ash content), litter/soil temperature and moisture. The
process of decompositionis treated inthe model as sequential
stages of organic matter transformation (humication) per-
formed by different functional groups of soil organisms. The
stages are: L, fresh litter; F, complex of humus substances
with underdecomposed plant debris (CHS); H
O
, organic layer
humus substances (OLH) in the later stages of decomposition
(visually manifested as peat); H
M
, humus of the mineral soil
layer or true humus closely boundto the mineral matrix. The
mass balance for each of these organic matter types (pools) is
described by the system of linear differential equations with
varying coefcients presented below
L
t
= L
O
(k
1
+k
3
)L, (1)
F
t
= k
3
L (k
2
+k
4O
+k
4M
+k
5
)F, (2)
H
O
t
= k
4O
F k
7
H
O
, (3)
H
M
t
= (k
4M
+k
5
)F k
6
H
M
, (4)
where L
O
is a litter input into the soil per unit time, while
L, F, H
O
and H
M
are the current sizes of the respective pools
(kgCm
2
).
Coefcients k
i
(k
1
, . . ., k
7
) (day
1
) account either for com-
plete mineralization or transformation of the specic organic
matter type into another type. Table 1 describes the processes
included in the k
i
coefcients and gives some information on
the groups of soil organisms performing these processes. Each
value of k
i
is presented as a function of the initial N and ash
contents in plant litter, multiplied by the temperature and
moisture response functions:
k
i
= k
i
(N, ash)f
i
(T)g
i
(W), (10)
The functions f
i
(T) and g
i
(W) are empirical and differ for
the different processes, to account for the fact that responses
to environmental parameters are not universal for all types
of organic matter. Here, the dependence of k
i
on nitrogen and
ash contents and soil/litter moisture were taken fromChertov
et al. (2001), while temperature response curves were updated
according to Chertov et al. (2005). Equations parameterizing
the moisture response curves (5)(9) are given in Table 1 as
they are usedfurther inthe discussion. Different k
i
coefcients
were used for the above and belowground litter compartments
(omitted from the above equation system for simplicity). Fur-
thermore, in comparison with a published version of ROMUL,
an additional pool OLH (H
O
) was added to the model (Chertov,
pers. commun., 2004). This accounts for the fact that when
litter quality is poor and/or its acidity is high, the soil Arthro-
478 ecologi cal modelli ng 2 0 4 ( 2 0 0 7 ) 475484
Table 1 Kinetic coefcients k
i
(i =1, . . ., 7) used in the model
Coefcient Process described Group of the soil organisms
performing it
Dependence on the soil moisture, W
(mass%)
k
1
Litter mineralization (L) Fungi and micro-fauna
complex
(5) g
1
=

0 W 7, W > 600
0.0435W0.304 7 < W 30
1 30 < W 300
0.003W+2 300 < W 600
k
2
CHS
a
mineralization (F) Fungi and micro-fauna
complex
(6)
g
2
=

0 W 7, W > 1200
0.0233W0.163 7 < W 50
0.00625W+1.31 50 < W 90
0.00068W+0.811 90 < W 1200
k
3
Litter to CHS transformation (L F) Fungi and micro-fauna
complex
g
3
= g
1
k
4O
CHS to OLH
b
transformation (F H
O
) Bacteria and Arthropoda (7) g
4
=

0.025W W 40
1 40 < W 400
0.0033W+2.333 400 < W 700
0 W > 700
k
4M
CHS to true humus transformation
(F H
M
)
k
5
CHS to true humus transformation
(F H
M
)
Earthworms (8) g
5
=

0 W 2, W > 120
0.0769W0.1538 2 < W 15
1 15 < W 70
0.02W+2.4 70 < W 120
k
6
True humus mineralization (H
M
) Soil bacteria and fungi (9) g
6
=

0.025W W 40
1 W > 40
k
7
OLH mineralization (H
O
) Soil bacteria and fungi g
7
= g
2
a
Complex of humus substances with underdecomposed plant debris (humied forest oor).
b
Organic layer humus substances (peat).
poda do not transport organic material down to the mineral
part of the soil prole, but rather leave it in the organic layer.
The richer is CHS in nitrogen (lower C/N ratio) the higher pro-
portion is going to mineral topsoil (the more active and larger
are Artropoda and Insecta larvae). Therefore, coefcient k
4
describing the transformation of CHS can be divided into two
components: the ow to OLH (k
O
) and the ow to the mineral
soil layer (k
M
):
k
4M
= 0, k
4O
= k
4
,
C
N
10, k
4M
= k
4

1.4 0.02
C
N

,
k
4O
= k
4
k
4M
, 10 <
C
N
35, k
4O
= k
4
, k
4M
= 0,
C
N
> 35,
(11)
The mineralization rate for OLH is described by coefcient
k
7
=0.00015f
2
(T)g
2
(W) (temperature and moisture response
curves are taken as those for CHS).
The model has been evaluated against long-term exper-
imental observations and provides good results for actual
forest data sets (Chertov et al., 1997)
3. Model set-up
In the present study simulations were performed for a geo-
graphical location representative of the Northern Swedish
boreal forest (64.0

N, 19.5

W). Soil texture data were taken

from the FAO data set (Zobler, 1986; FAO, 1991) and average
monthly values for air temperature, precipitation and cloud
cover, for the closest 0.5

0.5

grid square derived from a

meteorological observation network, kindly provided by the
Climate Research Unit (CRU), University of East Anglia (Newet
al., 1999, unpublished), were used to run the model. Although
the simulations were extended into the future, neither climate
predictions nor measured data for the period after 1970 were
used since the potential effects of climate change were not
the focus of this study. The series of inter-annually varying
meteorological parameters, detrended in the case of temper-
ature for the period 19011970, were repeated to represent
uctuations about constant long-term climatic parameters.
Simulations with GUESSROMUL started from bare ground,
but results from the initial period (before the rst equilibrium
withrespect to vegetationandcarbonpool) were not analysed.
Plant species were described explicitly and all model param-
eters were taken from the validated simulations with GUESS
for the Swedish forest (Koca et al., 2006).
Generic disturbance (other than re) was applied with a
prescribed frequency of 100 years. Plant litter C/N ratios in
above and belowground compartments for the given species
were taken from Berg and McClaugherty (2003). According to
observations (T. Persson, pers. commun., 2004) no earthworms
are found in Swedish forests at the latitude simulated here,
therefore coefcient k
5
, characterizing the activity of this type
of soil fauna, was set to zero.
The overall dynamics for a stand were derived by simu-
lating a number (in this case 100) of 1000m
2
patches and
averaging the results. Between-patch variability resulted from
ecologi cal modelli ng 2 0 4 ( 2 0 0 7 ) 475484 479
stochastic variationinestablishment, mortality, re, andother
types of disturbance. Due to averaging over a large number of
patches, stand species and age composition were relatively
stable and well mixed, although dependent to a large degree
on the assumed disturbance interval.
The sumof carbonstoredinall organic soil layers (L +F +H
O
)
was analysed; the humus of the mineral layer (H
M
) was not
included in this study.
Soil moisture in mass% was calculated separately for the
organic and mineral soil layers. Bulk density and wilting point
data for the mineral layer were taken fromthe DAAC database
(Global Soil Data Task Group, 2000). For the organic layer, rough
estimates of bulk density of 0.05gcm
3
were derived fromthe
regression of organic layer depth on relative mass presented
by Chertov (1981).
Twocontrasting soil moisture regimes, mesic andmesic-to-
wet, were considered to investigate the effect of soil moisture
on organic soil layer accumulation. The Swedish National
Forest inventory divides the soils in the country according
to the water table depth (WT) in the following classes: dry
(WT>2m), mesic (1<WT<2), mesic-to-wet, wet and water-
logged (for the last three WT<1m). Beldring et al. (1999) report
a regression linking an average water table depth with soil
moisture, which was developed using data for the Swedish
boreal forest. According to water moisture values produced
by GUESSROMUL the simulation site falls into the category
mesic. That is in accordance with the model soil hydrology
formulation, which assumes free drainage (Section 2). To sim-
ulate mesic-to-wet sites, soil moisture for the mesic site was
simply multiplied by the scaling factor so that WT=0.6m. This
implies that annual variability is the same for sites with dif-
ferent levels of soil moisture, which is not strictly true, but is
realistic as shown by Beldring et al. (1999). The climatic aver-
age of the modelled O soil layer moisture (mass%) is 220% and
440% for the mesic and mesic-to-wet sites, respectively.
It is important to point out that soil moisture inuences not
only the process of decomposition but also the occurrence of
forest res, which in turn inuences vegetation dynamics. To
assess the effect of re frequency on the size of the O layers,
different re return intervals were applied for the mesic and
mesic-to-wet sites. First, the results from a model re rou-
tine were examined but the re return intervals of 360 years
for mesic, and 1000 years for mesic-to-wet sites appeared to
be overestimated. Therefore, estimates found in the literature
for the pre-industrial frequency of re in the simulation area
were used instead. Fire frequency was taken as 50 years for
the mesic, and 300 years for the mesic-to-wet sites, based on
estimates made by examining records of re scarring on trees
in a relatively dry pine forest area (Niklasson and Granstr om,
2000) and a variety of forest types (Engelmark, 1987) in North-
ern Sweden. The data from the latter study support the idea
that the re return interval varies (from 48 to 270 years) and
correlates with forest type, while on mires traces of re are
very rare or absent. A study on Russian forests with natural
re disturbance regimes (Wirthet al., 2002) foundthat, onaver-
age, 50% of the carbon in the O soil layers is consumed by re,
which was integrated into the model. After the year 1870 all
res are assumed to be suppressed (Niklasson and Granstr om,
2000). However, the effects of other forest management prac-
tices were not included in this study.
The analyses presented below are based on the following
model runs:
Run I. Natural re frequency. Simulations continue 2000
years from the time when the C pool in the O layers reaches
a steady state.
RunII. Nore disturbance. Other types of disturbance with
a prescribed frequency of 100 years are included. Simula-
tions continue from 1870 until the C pool in the O layers
reaches a new steady state.
4. Results and discussion
4.1. Comparison of mesic and mesic-to-wet sites
under natural re conditions (model run I)
A mixed stand of spruce, pine and birch was predicted as the
natural vegetation for both the mesic, and for the mesic-to-
wet sites. However, the average contribution of birch biomass
in a stand varied by 7%on the mesic site but only by 3%on the
mesic-to-wet site. Birch is an early successional species so its
contribution increases with increases in re frequency. For the
mesic-to-wet site, the long-term average annual aboveground
litter production was 0.06kgCm
2
year
1
. This is consistent
with an estimate obtained from empirical regressions for
spruce and pine litter fall as a function of actual evapotran-
spiration, in mature forest (Akselsson et al., 2005). Soils in the
more frequently disturbed forest at the mesic site had a much
lower average litter input of 0.045kgCm
2
year
1
due to the
fact that the productivity of the young forest was lower than
that of the mature one. The model results agree with empir-
ical observations (M alk onen, 1974; Berg and McClaugherty,
2003) showing that litter production is a function of tree age,
generally increasing and reaching a steady state as the stand
matures. Fine root litter in the model was assumed to be equal
to the leaf litter.
At the beginning of the simulation, the slowest-developing
O soil layers (H
O
) reached an approximate steady state (with
oscillations due to disturbance) after about 200 years in the
mesic site and 500 years in the mesic-to-wet site. Simulations
from the following 2000 years were used to investigate the
uctuations of the pool sizes around the steady state due to
disturbance of particular patches. We calculated average car-
bon storage at the steady state to be 3.1 and 6.0kgCm
2
for
the mesic and mesic-to-wet stands, respectively (see Fig. 1a, in
which the distribution of the pool sizes within the 2000-year
time series is also shown). This difference in carbon storage
capacity could be explained by:
Organic matter decomposition being suppressed with
increasing soil moisture.
The higher re frequency in the mesic sites causing a sig-
nicant proportion of the soil carbon to be burned.
The overall litter input into the soil being lower at the mesic
site. The higher re frequency would cause young forest,
which is less productive than mature stands, to dominate.
480 ecologi cal modelli ng 2 0 4 ( 2 0 0 7 ) 475484
Fig. 1 Modelled carbon storage in forest soil O layers
under mesic (MES) and mesic-to-wet (MTW) moisture
conditions and a natural re regime. (a) Box plot
characterizing the distribution of the total stand C stored in
soil O layers. The boxes have lines at the lower, median
(bold), and upper quartiles and the whiskers show the
lowest and highest values. Statistics are based on a
2000-year time series with climate and
disturbance-induced uctuations around the steady state.
(b) The relative contribution of peat (H
O
) and the other O
layers (L+F), where L is fresh litter and F is the humied
forest oor.
Model tests have shown that 47% (95% condence inter-
val, CI, 4549%), 16% (CI =1121%) and 37% (CI =3341%) of the
difference can be explained by the rst, second and third rea-
sons, respectively. Therefore, it is suggested that soil moisture
modulates carbon storage in O layers, not only by its direct
effect on organic matter decomposition but also, and equally
importantly, by determining the frequency of the disturbance
events (res).
The relative ratio of the peat pool (H
O
) and other soil Olayer
pools (L +F) in the mesic-to-wet site is about twice as high as
in the mesic site (Fig. 1b).
4.2. Sensitivity of C storage to the frequency of re
disturbance
An estimate of the re return interval in the mesic-to-wet for-
est contains a large level of uncertainty, therefore the model
sensitivity to this parameter was tested. The results, pre-
sented in Fig. 2 show that increasing the re return interval
from 100 to 300 years had a statistically signicant effect on C
storage (the medians of the two groups differ at the 5% signif-
icance level if the inner bounds of the boxes do not overlap),
while the difference in the values between 300 and 500 years
was not signicant.
Fig. 2 Relationship between re return interval and C
storage in O soil layers in the mesic-to-wet site. The boxes
have lines at the lower, median (bold), and upper quartiles
and the whiskers show the lowest and highest values.
Statistics are based on a 2000-year time series with climate
and disturbance-induced uctuations around the steady
state.
There was no direct effect of increased water content on
gross primary production according to the GUESSROMUL
model, presumably because there was no water stress in the
area. Changing the relative contribution of different plant
species in a stand had a negligible effect on the C dynamics of
the O layers.
Increases inthe amount of carbon(or organic matter) insoil
O horizons with increases in stand age have been observed in
various locations within the boreal region (e.g. Bormann et al.,
1995; Yu et al., 2002). Berg and McClaugherty (2003) argue that
such an accumulation can continue for up to a millennium
in undisturbed ecosystems. Furthermore, Wardle et al. (2003)
speculates that forest res are the mainfactors responsible for
reductions in the depth of the organic soil layer in the boreal
region.
The present model study shows that the lower re fre-
quency in temporarily water-logged soils may be one of the
main reasons for the larger C accumulation in the past.
4.3. Comparison of mesic and mesic-to-wet sites
under the conditions of anthropogenic re suppression
(model run II)
The model predicts that O layers have accumulated carbon
from the time re began to be suppressed (in 1870), and will
continue to accumulate it until approximately 2100. However,
whilst this increase is expected to be large for the mesic site,
it should be almost negligible for the mesic-to-wet site. One
explanation for this difference is that the O layers of the soil
at the mesic site are further from steady state than those
at the mesic-to-wet site due to past disturbance. Values for
the prospective equilibrium size of O layers and estimated
sequestration rates are given in Table 2. The sequestration
rate for the mesic site corresponds well with an estimate pro-
vided by the limit value concept (Akselsson et al., 2005) for the
given geographical location. Peltoniemi et al. (2004) developed
a modelling method, validated with empirical data, which
allowed changes in soil organic layer Cstorage following stand
ecologi cal modelli ng 2 0 4 ( 2 0 0 7 ) 475484 481
Table 2 Changes in carbon storage in O soil layers due to re suppression from 1870 to 2100 per year and as a
proportion of the initial storage ((O
1870
O
2100
)/O
1870
)
Site Increase in O layers C for the period 18702100 Steady-state O layer C
storage O
2100
(kgCm
2
)
kgCm
2
year
1
O
2100
O
1870
/O
1870
(%)
Mesic 0.0103 77.8 5.4 (4.76.1)
Mesic-to-wet 0.0022 8.4 6.5 (6.07.2)
The end of the period indicates newsteady state in respect to Osoil layers. The size of a newequilibriumpool is given in the third column. O
1870
:
stand C storage in O soil layers in 1870 representing the pre-industrial steady state. O
2100
: stand C storage in O soil layers in 2100 representing
the steady state under the conditions of anthropogenic re suppression.
disturbance (harvesting) in southern Finland to be predicted.
According to their estimate, an average annual increase of
5.81.0gm
2
year
1
in the O soil layers should be expected
for a stand greater than 20 years of age. However, those results
and the model estimate presented here should not be com-
pared directly, not only because initial C storage and actual
litter production differ, but also because GUESSROMUL was
used to simulate the average response over a number of forest
patches.
4.4. Changes due to precipitation
According to the Swedish National Forest Inventory about 59%
of the forest in the country can be characterised as mesic
and about 31%as mesic-to-wet. Therefore, mesic-to-wet soils,
which differ from mesic soils, both in the amount of car-
bon stored and their ability to sequester carbon in the future,
should not be disregarded. An interesting question there-
fore, is whether the proportion of wet soil in a landscape
remains constant or whether it may be altered by changing
environmental parameters. Analysis of data from the mete-
orological observation network presented in Lindstr om and
Alexandersson(2004) has shownthat there was an8%increase
in precipitation in the boreal region of Northern Sweden from
19011950 to 19502000. At the same time no signicant trend
was observed in temperature.
Toassess roughly howthis affectedwater input intothe soil
could change the relative proportion of wet soils in a land-
scape, a very simplied approach has been applied (Ivanov,
1981). The method was previously used to obtain a dynamic
estimate of the wetland area in river catchments and provided
good results. It is assumed that the elevation y of a free ground
water surface above the impermeable bed at a given point x
along the topography transect ina catchment canbe described
by the parabolic equation (m):
y =

p
k
(2ax x
2
) +h
2
, (12)
where p = P E (cmyear
1
) is the climatic average excess of
precipitation over evapotranspiration, k (cmyear
1
) the inl-
tration coefcient, which is dependent on the soil physical
properties, h (m) the depth from the river water surface to the
impermeable bed, and a (m) is the width of the catchment
from the border of the river to the watershed divide. Variables
used in (12) are illustrated in Fig. 3.
Water table depth is calculated as the difference between
the actual topography elevation and the height of the free
ground water surface (y).
The topographical prole used here was derived from a
national topographical map (Lantm ateriet, 2002) of the same
area as the modelled sites. The depth of the impermeable soil
horizon was xed at 3m. Although the ltration coefcient
k has a physical meaning it is rather empirical and here the
value was tuned to produce reasonable values of the water
table depth. The fact that k chosen in that way falls within
the range of observed values shows that the approach was
valid.
Two calculations of the water table depth were made:
the rst with an average precipitation value for the period
19011950 (P
1
, Fig. 3) and the second with an 8% increase
in precipitation (P
2
, Fig. 3). In both cases the same value for
evapotranspiration produced by the GUESS model was used,
assuming no trends in temperature. The areas of the catch-
ment where the water table were below and above 1m were
calculated. With enhanced precipitation the proportion of the
catchment area witha WT<1mincreased from18%to 29%. By
assigning sectors with a WT<1m as mesic-to-wet and those
with a WT>1m as mesic, the data from Table 2, combined
with the initial C storage in O layers estimated from run I,
can be used to calculate an average C sequestration value
for the whole catchment. If the proportion of mesic-to-wet
areas remains constant at 18%, the C sequestration rate will
Fig. 3 Catchment characteristics used for the water table
depth calculations (Eq. (12)). Dashed lines illustrate two free
ground water surface positions (x, y) assuming an average
precipitation for 19011950 (P
1
) and an 8% increase in
precipitation (P
2
). h is the depth from the river water surface
to the impermeable bed and a is the width of the catchment
from the border of the river to the watershed divide.
482 ecologi cal modelli ng 2 0 4 ( 2 0 0 7 ) 475484
be 0.0089kgCm
2
year
1
, while an increase up to 29% of the
area will result in a higher rate of 0.0094kgCm
2
year
1
.
5. Suggestions for the extension of the
model to broader applications
A logical continuation of this work would be to model car-
bon storage in the O layers of the boreal landscape across the
whole gradient, from the dry upland to wet forest. Soil mois-
ture can be estimated by means of hydrological modelling on
a catchment scale, or by indirect observations, for example,
based on ground vegetation type. However, an attempt to test
the model for the whole range of soil moisture conditions has
revealed some limitations. Fig. 4 shows the moisture response
functions g
i
(W) (Eqs. (5)(9) in Table 1) for the kinetic coef-
cients, k
i
. As mentioned in an original SOMM formulation
(Chertov, 1985), no data were found to parameterise g
3
(trans-
formation of plant litter into CHS, regulated by moisture) and
an assumption was made that g
3
=g
1
. Moisture response func-
tions for the mineralization of litter (g
1
) and CHS (g
2
) are both
based on data from long-term laboratory experiments and an
interesting observation is that the range of optimummoisture
differs: with a rather broad interval (30300%) for the litter
and a single peak (50%) for the CHS. The same nding, that
low levels (<100%) of moisture decrease decomposition rates
in the CHS, was also reported by Douglas and Tedrow (1959).
Another important feature, seen in Fig. 4, is that g
1
decreases
to zero as the litter moisture approaches 600% (around com-
plete water saturation). Therefore, for soil that is saturated for
long periods, the GUESSROMUL model erroneously predicts
an innite accumulation of organic matter.
The literature was searched for measured rates of lit-
ter decomposition in the range of soil moisture that would
encompass complete saturation, but with little success. In a
synthesis paper, Walse et al. (1998), reviewing experimental
data on organic matter decomposition with respect to mois-
ture, included no data for moisture levels above eld capacity,
mainly due to a lack of such information in the original pub-
lications. The curves presented in Fig. 5 were derived from
Flanagan and Veum (1974), summarising weight loss and res-
piration experiments performed at tundra eld sites. Large
differences between the curves can be explained by varia-
tions inlitter quality andtemperature. However, inmost cases,
Fig. 4 Moisture response curves for the mineralization of
plant litter (g
1
), and CHS (g
2
), transformation of litter to CHS
(g
3
), and CHS to OLH (g
4
) used in the model. W is litter/soil
moisture expressed in mass%, CHS is a complex of humus
substances with underdecomposed plant debris, and OLH
is organic layer humus substances.
Fig. 5 Observed moisture response curves for plant litter
mineralization derived from various experimental sites at
different temperatures (Flanagan and Veum, 1974). W is
litter/soil moisture expressed in mass%. Complete water
saturation for plant litter is around 600%.
no dramatic reductions in litter decomposition near satura-
tion and even far above it were observed. In comparison with
optimumconditions a relative mineralization rate at W=600%
varied from 0.1 to 1.
Trettin et al. (2001) evaluated 12 commonly used soil
organic matter models and found that they did not describe
anaerobic conditions adequately. Here the question was,
how does decomposition change when the soil approaches
water saturation? None of the models, except CENTURY
(Parton et al., 1987) and WetlandDNDC (Zhang et al., 2002),
extrapolate their simulations across the full range of soil
moisture conditions. However, this was done in the original
SOMMfor the moisture level above eld capacity (30<W<600,
g
1
=g
3
=1) and the results it provided are probably valid rst
approximations. Alternatively a linear function similar to
that presented in the WetlandDNDC model with g
1
=0.75 at
saturation, could be used.
The model test has shown that for two moisture regimes
(mesic and mesic-to-wet) compared in this paper, long-term
carbon accumulation in the O layers is not sensitive to g
1
or g
3
and is determined only by g
2
. The results presented here will
therefore also be valid with a new g
1
parameterisation.
6. Concluding remarks
The importance of soil moisture for the simulation of soil
organic matter dynamics was shown in this study and should
not be neglected when climate change issues are considered.
The GUESSROMUL model is applicable for further investi-
gation in this area and coupling with catchment hydrology
models appears to be promising. However, the model requires
some improvement concerning the dependence of the kinetic
coefcients k
i
on soil moisture levels.
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I
II
I
Variations in net ecosystem exchange of carbon dioxide in a boreal
mire: Modeling mechanisms linked to water table position
Alla Yurova,
1
Annett Wolf,
1,2
Jorgen Sagerfors,
3
and Mats Nilsson
3
Received 5 October 2006; revised 13 February 2007; accepted 7 March 2007; published 19 May 2007.
[1] In mires, which occupy large areas of the boreal region, net ecosystem CO
2
exchange
(NEE) rates vary significantly over various timescales. In order to examine the effect of
one of the most influencing variables, the water table depth, on NEE the general
ecosystem model GUESS-ROMUL was modified to predict mire daily CO
2
exchange
rates. A simulation was conducted for a lawn, the most common microtopographical
feature of boreal oligotrophic minerotrophic mires. The results were validated against
eddy covariance CO
2
flux measurements from Degero Stormyr, northern Sweden,
obtained during the period 20012003. Both measurements and model simulations
revealed that CO
2
uptake was clearly controlled by interactions between water table depth
and temperature. Maximum uptake occurred when the water table level was between
10 and 20 cm and the air temperature was above 15C. When the water table was higher,
the CO
2
uptake rate was lower, owing to reduced rates of photosynthetic carbon fixation.
When the water table was lower, NEE decreased owing to the increased rate of
decomposition of organic matter. When the water table level was between 10 and 20 cm,
the NEE was quite stable and relatively insensitive to both changes within this range
and any air temperature changes above +15C. The optimal water table level range for
NEE corresponds to that characteristic of mire lawn plant communities, indicating
that the annual NEE will not change dramatically if climatic conditions remain within the
optimal range for the current plant community.
Citation: Yurova, A., A. Wolf, J. Sagerfors, and M. Nilsson (2007), Variations in net ecosystem exchange of carbon dioxide in a
boreal mire: Modeling mechanisms linked to water table position, J. Geophys. Res., 112, G02025, doi:10.1029/2006JG000342.
1. Introduction
[2] Boreal mires contain a large carbon pool, accumulated
over millennia, which is significant at both regional and
global scales [e.g., Gorham, 1991; Turunen et al., 2002].
Water balance plays a crucial role among the factors
determining mire growth and development [e.g., Ivanov,
1981; Hilbert et al., 2000]. Therefore climatic variations at
different timescales may affect the rate of CO
2
uptake and
the overall C dynamics of mires [Moore et al., 1998]. In this
context, it is very important to incorporate mire ecosystems
into the framework of dynamic vegetation modeling, a state-
of-the-art approach to carbon balance assessments and
predictions.
[3] The fundamental ecosystem processes (including pho-
tosynthesis, respiration, and soil organic matter dynamics)
are common to mires and other terrestrial ecosystems, so it
should be possible to adapt a general ecosystem model to
mires. However, mires have highly specific self-regulatory
properties that arise from the interactions amongst their
hydrological and biological processes on all scales from the
microtopographical unit to the whole ecosystem [e.g.,
Belyea and Baird, 2006]. Direct measurements of changes
in NEE of individual microforms associated with changes in
the water table level are used to investigate the hydrological
control of mire CO
2
exchange rates. A number of studies
[e.g., Alm et al., 1999; Waddington and Roulet, 2000;
Bubier et al., 2003; Lafleur et al., 2003; Nykanen et al.,
2003; Syed et al., 2006] have demonstrated, using chamber
or eddy covariance measurements over different microtopo-
graphical features, that responses of NEE to changes in the
water table depth can vary widely. Analysis of data using an
ecophysiological model might help to explain the variabil-
ity. However, to date, relatively few process-based general
ecosystem models, particularly ones specific to mires and
run using daily time steps, have been applied to predict NEE
and validated by field measurements from actual mire sites
[Potter et al., 2001; Frolking et al., 2002; Zhang et al.,
2002].
[4] Here we present a modified general ecosystem simu-
lator, GUESS [Smith et al., 2001], which has been previ-
ously used to simulate forest and grassland ecosystem
dynamics, coupled to the soil organic matter dynamics
model ROMUL [Chertov et al., 2001]. The GUESS-
ROMUL model [Yurova and Lankreijer, 2007] was restruc-
JOURNAL OF GEOPHYSICAL RESEARCH, VOL. 112, G02025, doi:10.1029/2006JG000342, 2007
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Article
1
Department of Physical Geography and Ecosystems Analysis, Lund
University, Lund, Sweden.
2
Now at the Department of Forest Ecology, Institute of Terrestrial
Ecosystems, Zurich, Switzerland.
3
Department of Forest Ecology, Swedish University of Agricultural
Sciences, Umea Sweden.
Copyright 2007 by the American Geophysical Union.
0148-0227/07/2006JG000342$09.00
G02025 1 of 13
tured so that it could be applied to Sphagnum-dominated
microtopographical features of mires. We focused on mod-
eling the CO
2
exchange of an area dominated by lawn plant
communities at Degero Stormyr, a minerotrophic mire in
northern Sweden. The results were compared with daily
CO
2
fluxes for 20012003 obtained from eddy-covariance
tower measurements. Furthermore, we derived and analyzed
NEE response functions, from both the model and empirical
data, in relation to the water table depth.
2. Model
[5] The modeling framework used here was the coupled
dynamic vegetation-soil organic matter model GUESS-
ROMUL. A description of the GUESS is given by Smith
et al. [2001], the ROMUL model is summarized by Chertov
et al. [2001], and the combined GUESS-ROMUL system is
described by Yurova and Lankreijer [2007]. Here we only
describe those processes within the model that we revised or
modified in order to simulate mire ecosystems.
[6] The basic unit in the GUESS model is the plant
functional type (PFT), characterized by its physiological,
morphological, and phenological attributes. To simulate
boreal mires, we introduced Sphagnum mosses as a new
PFT within the model. Major changes to the model were as
follows:
[7] 1. A new water balance scheme was introduced.
[8] 2. The photosynthesis model was modified to account
for (1) regulation of Rubisco activity by the tissue N content
and (2) the dependency of moss photosynthetic activity on
water content. Parameters in the equations describing auto-
trophic respiration were also revised.
[9] 3. Specific coefficients were used to describe decom-
position of the Sphagnum litter and peat. Parameterization
of decomposition under anoxic conditions was included in a
soil organic matter dynamics model.
[10] The changes are described in more detail below.
Model parameters and constants are summarized in Table 1.
2.1. Hydrology
[11] An important feature of the mire water regime is that
the water table is so close to the surface that there is a
constant supply (via capillary action) of water to the
unsaturated zone; that is, water can move from the water
table up to the peat surface.
[12] Two layers can be distinguished within the peat
profile: the upper, temporarily aerated layer consisting
largely of living and lightly decomposed plant material
(acrotelm), and the permanently saturated, lower zone,
consisting largely of compact, relatively highly decomposed
plant material (catotelm). Owing to the very strong reduc-
tion in hydraulic conductivity from the acrotelm to the
catotelm [Ivanov, 1981] it can be assumed that the move-
ment of water in the catotelm is negligible compared to its
movement in the acrotelm.
[13] Here we used the set of equations from the Mixed
Water and Heat model (MMWH) [Granberg et al., 1999],
with minor modifications, to describe the water balance in
the acrotelm. The MMWH model was specifically devel-
oped to reconstruct the water table position in a boreal
mixed mire and was validated for the lawn plant community
at Degero Stormyr (the mire used in the present study).
[14] The total volume of water in the profile, from the
peat surface to the depth of the permanently saturated zone
(catotelm, z
cat,,
cm) at the current water table depth (z
wt,,
cm), V
tot
(cm), was defined as
V
tot
f z
cat
z
wt

Z
0
zwt
q
us
z dz; 1
where the first term describes water volume in the saturated
zone (cm) in relation to the thickness of the saturated layer -
(z
cat
z
wt
) and porosity (f). The second term is the water
content in the unsaturated zone (q
us
(z)) integrated from the
current water table depth to the surface. (The sign
convention in this paper is that water table depth is noted
with a negative value when water table is situated below the
peat surface). The function q
us
(z) describes the gradual
change in the capillary water content from its maximum
near the water table (supply) to its lowest value at the
evaporating and transpiring vegetated surface. Here we used
an equation developed by van Genuchten [1980], simplified
by setting the residual water content to zero [Weiss et al.,
1998],
q
us
z f 1 a z
wt
z
n

11=n
: 2
[15] The pressure head (in cm H
2
O) in the van Genuchten
equation was replaced by the distance to the water table
(z
wt
z), assuming that the water profile in the unsatu-
rated zone is in equilibrium. The empirical parameters n and
a were derived from a study of a large number of peat cores
collected in Finland [Weiss et al., 1998],
n 1:49 3:56 r 13:2 r
2
0:0027 ca 0:037 l
1
3
log
10
a 51:8 r
2
0:0057 ca 0:01 sph 0:53 l
1
; 4
where r is peat bulk density (g cm
3
), ca and sph are Carex
and Sphagnum residue contents in the peat (%), respec-
tively, and l
1
has a value of 1 for the peat at a depth of
100 cm and 0 otherwise.
[16] The water balance equation was calculated at daily
intervals, to identify changes in water storage in the profile
(DS, cm d
1
),
DS P E q; 5
where P is rain or snowmelt, E is evapotranspiration and q
is discharge.
[17] The description of saturated flow in the acrotelm is
based on work by Ivanov [1981],
q l
q
i T
r
z
cat
z
wt
; 6
where q is specific discharge (cm d
1
), i is a local slope of
the water table (assumed to be equal to the topographical
slope,
dh
dl
), T
r
(cm d
1
) is transmissivity coefficient, and
l
q
(cm
1
) is a lumped parameter. Zero recharge in the area
and no horizontal flow in the unsaturated zone are assumed.
The peat transmissivity is approximated as an exponential
G02025 YUROVA ET AL.: MODELING CO
2
EXCHANGE IN A BOREAL MIRE
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G02025
function of the water table depth, with coefficients based on
data for the lawn [Ivanov, 1981]: This takes into account the
possibility of a runoff feedback mechanism, for example,
the ability of the mire ecosystem to regulate its water
balance by runoff increasing when the water table is high.
[18] Evapotranspiration is simulated differently here
from the original formulation in the MMWH model.
First, potential evapotranspiration (E
q
cm d
1
) is estimated
using the Priestley-Taylor equation as in the standard
GUESS model [e.g., Gerten et al., 2004]. Then an expo-
nential function is used to adjust actual evapotranspiration
(E, cm d
1
), to correspond to the current water table depth
in the mire.
E E
q
exp l
E
z
wt
: 7
[19] The empirical parameter l
E
(cm
1
) was derived from
evapotranspiration measurements obtained from three Fin-
ish mires, using sample containers with different water
levels [Virta, 1966].
[20] Knowing the change in water storage fromequation (5),
we obtain the new value for the total water volume in the
profile (V
tot
) and solve equation (1) for z
wt
to determine the
new water table depth for the current day.
[21] Acrotelm depth was assumed to be constant (30 cm)
over the simulation period (20012003).
2.2. Moss Production
2.2.1. Photosynthesis
[22] The C
3
photosynthesis in GUESS is formulated
following a simplified version of the Farquhar model
[Farquhar et al., 1980; Collatz et al., 1991; Haxeltine and
Prentice, 1996]. The rate of photosynthesis can be limited
by either the electron transport rate or Rubisco activity. In
the standard version of GUESS, nitrogen supply is assumed
to be always sufficient to provide optimum Rubisco activity
for the given PAR level [Haxeltine and Prentice, 1996]. In
boreal mires, however, nitrogen is often limiting [e.g.,
Malmer and Nihlgard, 1980]. We therefore used an equa-
tion presented by Friend [1995] to calculate maximum
Rubisco activity, V
max
(kgC m
2
d
1
) as a function of nitro-
gen concentration in moss tissues N.
V
max
1036:8 k
c
6:25 8=550 f
N;Rub
N; 8
where k
c
is the temperature-dependent Rubisco carboxylation
turnover rate (molCO
2
mol
1
sites s
1
) and f
N,Rub
is the
fraction of total nitrogen content bound in Rubisco (dimen-
sionless). The nitrogen concentration, in (N, kg m
2
)
recalculated from C:N ratios, and f
N,Rub
, were specified
as model input parameters. The constant 550 has units
(kg mol
1
), and 1036.8 (1210
3
kg mol
1
86400 s d
1
) is a
conversion factor from the original units (mol CO
2
m
2
s
1
)
used by Friend [1995] to kgC m
2
d
1
.
[23] The photosynthetic activity of mosses also depends
on their water content [Titus and Wagner, 1984; Williams
and Flanagan, 1996; Schipperges and Rydin, 1998]. When
the water content of the mosses is low, desiccation adversely
affects their photosynthetic capacity, and when the water
content is too high their photosynthetic rates decrease
because the rate of transport of CO
2
is lower in water than
in air [Silvola, 1990]. The threshold values are species-
Table 1. Conventional GUESS-ROMUL Parameters and Constants Used for the Mire Simulation and New Parameters Included in
Additional Model Equations
Function Symbol Value Units Description Source
Photosynthesis w
min
0 gH
2
0/gC determine the photosynthesis
response curve depending on
the Sphagnum water content
Schipperges and
Rydin [1998]
w
low
12 gH
2
0/gC
w
high
25 gH
2
0/gC
w
max
70 gH
2
0/gC
N
Rub
0.18 proportion of N in Rubisco Friend [1991]
cn 50 C:N ratio in Sphagnum tissues Malmer and Wallen [2005]
Moss respiration b 0.03 shaping parameter in equation (11) simultaneous calibration of b, r
m
and r
g
based on data from
Williams and Flanagan [1998]
r
m
2.0 shaping parameter in equation (12)
r
g
0.5 shaping parameter in equation (10)
Turnover tr 1.0 proportion of the biomass
transferred annually to the litter
Nungesser [2003]
Litter and SOM dynamics
a
k
1
1.610
3
day
1
litter(L) mineralization coefficient Verhoeven and Toth [1995]
k
2
4.010
4
day
1
SOM (F) mineralization coefficient
a
O. G. Chertov
(personal communication, 2004)
k
an
0.50 ratio of aerobic to anaerobic
decomposition rate in the acrotelm
Moore and Dalva [1997]
k
cat
1.010
4
year
1
decay rate in the catotelm Clymo [1984]
r
L
0.02 g cm
3
moss litter density Nungesser [2003,
and references therein]
r
F
; r
H
0.02; 0.05 g cm
3
peat density at various depths Granberg et al. [1999]
Hydrology
f
0.92; 0.98 porosity: acrotelm; catotelm Granberg et al. [1999]
z
cat
30 cm acrotelm depth Granberg et al. [1999]
sph 90 % proportion of Sphagnum and
Carex remains in the peat
M. Nilsson
(personal communication, 2006)
ca 3 %
l
E
0.02 cm
1
shaping parameter in equation (7) based on work by Virta [1966]
a
SOM is soil organic matter.
G02025 YUROVA ET AL.: MODELING CO
2
EXCHANGE IN A BOREAL MIRE
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G02025
dependent and not all mosses show strong reductions in
photosynthesis when water contents increase to high levels.
Here we simulated the potential photosynthetic assimilation
rate further multiplying it by the empirical linear function
f
wc
(dimensionless),
f
wc

w w
min
= w
low
w
min
; w
min
< w w
low
1; w
low
< w w
high
w w
max
= w
high
w
max

; w
high
< w < w
max
0; w ! w
max
; w w
min
8
>
>
>
>
>
>
>
>
>
>
<
>
>
>
>
>
>
>
>
>
>
:
; 9
where w is water content (gH
2
O/gC).
[24] Parameters w
min
, w
low
, w
high
and w
max
(gH
2
O/gC)
were derived from the empirically derived curves published
by Schipperges and Rydin [1998] for the typical lawn
species Sphagnum balticum, which dominates the lawns at
Degero Stormyr. The water content of the apical part of the
Sphagnum shoots was calculated using equation (2), inte-
grated from the peat surface down to a fixed depth (2 cm).
Photosynthesis was assumed to be totally suppressed under
snow cover and when the surface temperature falls below
zero.
2.2.2. Respiration
[25] For Sphagnum, the differentiation between
photosynthetic and nonphotosynthetic tissues is somewhat
obscure. However, Williams and Flanagan [1998] have
postulated that respiration from both compartments
should be specified. We therefore calculate total respiration
R (kg C m
2
d
1
) as follows.
R r
g
A R
nonphot
R
phot

R
nonphot
R
phot
; 10
where the first term describes growth respiration and the
last two maintenance respiration from nonphotosynthetic
R
nonphot
: (kgC m
2
d
1
) and photosynthetic R
phot
:
(kgC m
2
d
1
) tissues, respectively. A is the photosynthetic
CO
2
assimilation rate (kgC m
2
d
1
) and r
g
is a growth
respiration coefficient (1-growth yield).
[26] Maintenance respiration is calculated following
Haxeltine and Prentice [1996] and Sitch et al. [2003].
R
phot
b V
max
11
R
nonphot
r
m
k
r
g T
air
M=cn; 12
where b and r
m
are scaling coefficient, M is total biomass
(kg C m
2
), cn is C:N ratio, g(T
air
) is a temperature
response function described by the modified Arrhenius
equation [Lloyd and Taylor, 1994], and k
r
(day
1
) is the
respiration rate for a 10C baseline.
[27] We were unable to find any studies in which the
respiration rate of Sphagnum species was treated in the
framework of the growth-maintenance paradigm. However,
dark respiration rates of Sphagnum samples have been
measured simultaneously with net assimilation rates by
Williams and Flanagan [1998]. Using the values of envi-
ronmental parameters presented by the cited authors, we
conducted an optimization of three model parameters
(b, r
g
, r
m
) to achieve the same ratio of respiration to net
assimilation as measured by Williams and Flanagan [1998].
Different combinations of b, r
g
and r
m
were potentially
possible, so we selected the one that conformed to the
standard parameter ratio used by GUESS (Table 1).
2.2.3. Growth
[28] At the end of each year, NPP (net primary produc-
tion) was converted to biomass increments after deducting
10% to account for reproduction (converted into the litter
pool). The rest was allocated to new moss biomass. Daily
NPP can be negative for mosses, if respiration exceeds
photosynthesis, for example, on days with low light and
sufficiently high temperatures. Annual litter production was
assumed to equal NPP [Nungesser, 2003].
2.3. Soil Organic Matter Dynamics
[29] It is widely recognized that the decomposition of
Sphagnum moss litter is slow in comparison to that of other
plant genera [Clymo, 1965]. In addition, decomposition also
slows under anoxic conditions.
[30] The ROMUL model used in this study specifies the
following three pools of soil organic matter: fresh litter (L),
complexes of humic substances with recalcitrant remains of
plant debris (F); and humic substances that are in late stages
of decomposition and/or bound to the mineral soil (H). The
rate of mineralization and the flow of matter between
different pools is described by the kinetic coefficients k
i
(i = 1. . .6), which are dependent both on the litter quality
and environmental conditions (temperature and moisture).
To adapt the ROMUL model to the mire sites, we consid-
ered the F pool to be represented by the acrotelm peat. No
transport of organic material to the mineral horizon was
considered. The H pool was interpreted as being the
permanently saturated catotelm peat.
[31] In their comprehensive survey of the decomposabil-
ity of the litter of various plant growth forms (Sphagnum,
evergreen and deciduous shrubs, graminoids and forbs)
from mires in different geographical regions, Dorrepaal et
al. [2005] concluded that the standard chemical variables
(the C:N ratio as used in ROMUL, or the lignin:N ratio)
explain a very small proportion of the variation in decom-
position rates, and that life form per se may be a better
predictor. On the basis of this conclusion, we chose the
specific mineralization rate k
1
= 0.0016 day
1
for the
Sphagnum litter (L pool), derived from data presented by
Verhoeven and Toth [1995] from 73-day-long laboratory
decomposition experiments. The surface peat mineralization
rate k
2
= 0.0004 d
1
for the F pool was obtained from recent
laboratory experiments with a poor fen peat under aerobic
conditions (O. G. Chertov, personal communication, 2004).
The model coefficient k
3
remained as standard for above-
ground tissues [Chertov et al., 2001]. The fixed acrotelm
depth (z
cat
= 0.3 m) determines the annual amount of the
material that submerges into the permanently saturated zone
K (kgC m
2
),
K r
F
z
cat
L=r
L
F=r
F
; 13
where L and F (kgC m
2
) are pool sizes, r
L,
r
F
(kgC m
3
)
are the densities of the corresponding pools and z
cat
is
expressed in meters (m).
G02025 YUROVA ET AL.: MODELING CO
2
EXCHANGE IN A BOREAL MIRE
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G02025
[32] The rate of catotelm decay, k
cat
, was taken from the
literature [e.g., Clymo, 1984; Tolonen et al., 1992; Korhola
et al., 1996; Kuhry and Vitt, 1996]. We also introduced a
new coefficient, k
an
, defined as the ratio of anaerobic to
aerobic decomposition in the acrotelm. To account for the
proportion of the acrotelm peat under the water table at any
given time, the mineralization rate was reduced by multi-
plying it by k
an
. The value of k
an
was taken from Moore and
Dalva [1997].
[33] It should be noted that the suppressed decay rates
found under periodically anoxic conditions in the surface
peat (as described among other things by Moore and Dalva
[1997] and Bergman [1998]) are much higher than the
extremely low decay rates found in deep, permanently
waterlogged peat, which, according to peat core analysis,
could be up to a thousandfold lower [e.g., Clymo, 1984;
Korhola et al., 1996], as hypothesized by Freeman et al.
[2001] owing to the inhibition of phenol oxidases by both
low partial oxygen pressures and high concentrations of
phenolic compounds.
[34] The time steps for the soil organic matter simulations
and the transport of material to the catotelm were 1 day and
1 year, respectively.
2.4. Modeling Protocol
[35] The model was run for the period 20012003,
following a 7000-year spinup: roughly the time since
Degero Stormyr was established [Malmstrom, 1923]. The
climatic data input for the spinup period (monthly air
temperature, precipitation and cloud cover) were repeated
60-year series, using real figures for 19011961 derived
fromthe CRU05 database (provided by the Climate Research
Unit, University of East Anglia; M. New et al., Representing
twentieth century space-time climate variability: Part 2.
Development of 19011996 monthly grids of terrestrial sur-
face climate, 1999, unpublished manuscript) for the coordi-
nates of Degero Stormyr. The daily meteorological data (air
temperature and precipitation) from the station at Degero
Stormyr mire were used for the simulation period 2001
2003. Global solar radiation data were used and PAR was
calculated as 50 percent of the global radiation. For the
current study, we excluded all plant functional types except
mosses from the simulation. The model was run over daily
time intervals.
[36] The following model simulations were conducted:
[37] 1. A run with a simulated water table to assess
overall model performance.
[38] 2. A run with the actual measured water table depths
instead of the modeled values, to estimate the level of
uncertainty added to the model as a result of unknowns
associated with its hydrology section. Data from this run
were also used to analyze the dependency of NEE on the
water table depth.
[39] 3. Two test runs excluding the effect of water depth
on (1) photosynthesis and, subsequently, (2) peat decom-
position rate.
[40] 4. A series of Monte Carlo runs to determine the
models sensitivity to specific parameters and constants
(Table 2).
3. Site Description
[41] The EC measurement tower for monitoring CO
2
exchange at the oligotrophic minerotrophic mire Degero
Stormyr (6411
0
N, 1933
0
E) is part of the Nordic eddy-flux
measurement network, NECC. The mire is located in a
boreal forested zone in a cold temperate humid climate. The
length of the period with air temperatures above 0C is
about 200 days per year. The mean annual temperature at
Kulbacksliden, a meteorological station near the mire, is
+1.2C, the mean July temperature is +14.7C and the
mean January temperature is 12.4C (19611990)
[Alexandersson et al., 1991]. The mean annual precipita-
tion is 523 mm (19611990) [Alexandersson et al., 1991],
34% of which falls as snow.
[42] Degero was initiated approximately 78 kyrs BP and
the current peat depth is mostly between 3 and 4 m, with
maximum depths up to 8 m [Malmstrom, 1923]. Near the
tower the peat depth is 5 m [Granberg et. al., 1999]. The
water table position usually varies from 5 to 19 cm below
the peat surface level (90th percentile to 10th percentile),
but the mire is flooded every spring during snowmelt while
Table 2. Range of Parameters and Constants Used in the Sensitivity Analysis
Parameter Units
Value in
Standard Model Lower Range Higher Range Source and Comments
Production
w
min
gH
2
0/gC 0 0 4 Nungesser [2003, and references therein],
Schipperges and Rydin [1998]
w
low
gH
2
0/gC 12 10 16
w
high
gH
2
0/gC 25 16 30
w
max
gH
2
0/gC 70 28 70
N
Rub
0.18 0.18 0.22 Friend [1991], not specifically for moss
b 0.03 0.01 0.045 assumed
r
m
2.0 1.0 5.0 assumed
r
g
0.5 0.15 0.50 assumed
Decomposition
k
1
day
1
1.610
3
4.010
4
1.010
2
calibration based on work by
Nilsson and Bohlin [2001]
k
2
day
1
4.010
4
9.610
6
6.010
4
Waddington et al. [2001], McKenzie et al. [1998]
k
3
day
1
1.110
2
6.110
3
1.510
2
Chertov [1985], not specifically for moss
k
cat
year
1
1.010
4
810
5
510
4
Clymo [1984], Korhola et al. [1996]
k
an
0.50 0.07 1.0 Bergman [1998], Moore and Dalva [1997]
G02025 YUROVA ET AL.: MODELING CO
2
EXCHANGE IN A BOREAL MIRE
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G02025
the peat is still frozen. Species dominating the lawn plant
community are Sphagnum balticum Russ. C. Jens. with
sparse Sphagnum lindbergii. The vascular plant community
is dominated by Eriophorum vaginatum L., Vaccinium
oxycoccos L., Andromeda polifolia L., with Scheuchzeria
palustris L occurring more sparsely.
3.1. Measurements
[43] The eddy covariance system for measuring CO
2
,
water, and energy fluxes is installed on a tower, and has a
fetch area that is dominated by the lawn microtopographical
feature. The energy balance closure of the system was 96%
over the 3 years, indicating that the data are reliable
[Sagerfors et al., 2007]. The water table relative to the peat
level is monitored automatically using a float and counter-
weight system [Roulet et al., 1991]. In addition, sets of
standard meteorological observations (global and net solar
radiation, air and peat temperature, air humidity and snow
depth) are recorded at the site [Sagerfors et al., 2007]. Data
series for three full years, 20012003, were used for model
validation.
3.2. Evaluation of the Model Performance, Statistical
Data Processing, and Sensitivity Analysis
[44] Simulated water table depth (WT) and net ecosystem
exchange (NEE) data were compared with observed values
on a daily basis over the period 20012003. In accordance
with the recommendations of Willmott [1982] we calculated
the following quantitative measures to evaluate the model
performance: the mean bias error (MBE), the mean square
error (MSE), which was also partitioned into systematic
(MSE
s
) and unsystematic (MSE
u
) components, and the
summary measure the Willmott index of agreement
(d, defined in Appendix A).
[45] To analyze the relationship between the response
variable NEE (y) and the explanatory variable, depth to
the water table in the mire (x), a least squares second-order
regression (y = ax
2
+ bx + c +e) was applied as the best
polynomial model. The t-statistic and corresponding p-value
were used to estimate the significance of the second-order
regression coefficient a (H
o
: a = 0).
[46] Regression lines based on the observed data and
outcomes of the test runs were compared with the standard
model simulation to test the hypothesis of coincidence. A
model, y = a
1
x
2
+ b
1
x + c
1
+ a
2
x
2
z + b
2
x z + c
2
z + e,
was fitted to the data from both of the data sets to be
compared, where the dummy variable z took the value 1
for the first subset (standard simulation) and 0 for the other.
The F-statistic from the ANOVA tables was then used to test
the significance of z, x z, and x
2
z simultaneously.
[47] Quantile regression [e.g., Koenker, 2005] as a rec-
ommended method for estimating effects of limiting factors
in ecology [e.g., Cade et al., 1999], was applied to define
the upper and lower bounds to the limitation posed on NEE
by the water table position. In opposite to classical linear
regression methods, which base their statistics on the
squared deviation from a mean function, quantile regression
methods offer estimating models for median and quantile
functions (Appendix A). The second-order polynomial
model used for the least squares regression above was
linearized by introducing x
2
= x
2
and the regression quan-
tiles {0.05, 0.20, 0.35, 0.5, 0.65, 0.80, 0.95} were calculated
for the resulting model y = ax
2
+ bx + c + e. If we
hypothesize that if other factors confound the influence of
the water table depth on the net CO
2
uptake, then the lowest
quantiles must exhibit the highest rate of change in NEE
with changes in water table depth. (Conventionally, upper
quantiles would be used, but since we are operating with
negative values of NEE, values below the lower regression
quantiles correspond to the maximum net CO
2
uptake rates,
i.e., situations where the effects of other constraints are
lowest). Ninety percent confidence intervals for the quantile
regression coefficients a and b were computed by the rank
inversion method [Koenker, 2005] to estimate if there were
significant differences in the rates of change in NEE to
water table depth calculated for the different quantile
regression lines. All the statistical calculations were imple-
mented in the R programming environment [R Development
Core Team, 2003].
[48] In order to evaluate the model parameter-based
uncertainty concerning the NEE predictions, we performed
a multiparameter sensitivity analysis. Uncertainties in the
key parameters of the LPJ model, which is identical to
GUESS in its representation of physiological and biogeo-
chemical processes, have been determined and investigated
previously using Monte Carlo regression-based analysis
[Zaehle et al., 2005]. In this study we focused only on the
parameters added or modified in GUESS-ROMUL for
modeling the mire ecosystems. The output variable used
in the analysis is NEE. Since NEE is not a monotonous
function of the input parameters, we were not able to use a
regression-based method; hence the variance-based Sobol
sensitivity indices were chosen instead (Appendix A). The
first-order sensitivity coefficient S
j
and the total effect term
TS
j
for the parameter of interest were compared using
Students t-test. Significant differences between the two
were treated as indications of nonlinear interaction between
the model parameters. We used Latin hypercube sampling
(LHS) [McKay et al., 1979] to generate sets of parameters
for the Monte Carlo runs, and where possible the parameter
ranges were derived from the literature (Table 2).
4. Results
4.1. Hydrology
[49] Good agreement between the modeled and measured
water table depths with a mean deviation (MBE) of only
around 1 cm was found for 20012003 (Figure 1), compa-
rable to previously published validation data for 1998
[Granberg et al., 1999]. The Willmott index of agreement
was high (d = 0.92). The root mean square error (RMSE)
was 3.5 cm and the systematic portion accounted for 8% of
this. The difference between the measured and modeled
water table depths was greatest during the spring flood,
reaching up to 10 cm on some days.
4.2. Net Ecosystem Exchange of CO
2
[50] About 4.5 m of the catotelm peat was accumulated
during the 7000-year model initialization period, which
corresponds well with the measured peat depth, 5 m, at
the study site [Granberg et al., 1999].
[51] The model was able to capture the general patterns of
CO
2
exchange observed by the eddy covariance tower at
Degero Stormyr mire (Figure 2). The predicted NEE
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(gCm
2
yr
1
) values for 2001, 2002 and 2003 were 46,
81 and 57, while the measured values were 48, 61,
and 59, respectively. (Net carbon uptake by the mire is
represented by negative NEE values in this paper). The
modeled NEE values were the differences between the
simulated net photosynthesis rates of 162, 244, and
197 gCm
2
yr
1
and simulated heterotrophic respiration
rates of 116, 163, and 140 gCm
2
yr
1
for the three re-
spective years. The deep (catotelm) peat accounted for 10
20% of the total modeled decomposition. The seasonal
switch of the mire between a sink and a source of CO
2
was clearly reproduced by the model (Figure 2a).
[52] Some quantitative measures of the models perfor-
mance at daily time intervals are presented in Table 3. Using
the simulated water table depth instead of the measured
depth did not influence the quality of NEE predictions
significantly (Table 3). In both cases the Willmott index
of agreement (d) was high and the model bias small. The
systematic portion of the mean square error (MSE) was
rather low, but the MSE itself was relatively large. During
the first part of the vegetation period the model simulation
always resulted in a larger net uptake (lower NEE values)
compared to the measured values. During the second part of
the vegetation period there were both positive and negative
deviations (Figure 2b). In general, the simulations for the
Figure 2. (a) Simulated (black line) and measured (crosses) NEE (gC m
2
d
1
) for the lawn at Degero
Stormyr for 20012003. Negative NEE, carbon uptake; positive NEE, carbon release. (b) Deviations
between simulated and measured NEE based on 5-day averages.
Figure 1. Simulated (dash-dotted line) and measured (solid line) water table depth (WT, cm) from the
lawn at Degero Stormyr for 20012003.
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days with negative modeled NEE were poorer than those
for days when no CO
2
fixation took place, but there was
no clear relationship between productivity and model
performance.
4.3. Relationship Between Water Table and NEE
for Modeled and Measured Data
[53] The two least squares regression lines, based on
model simulations and direct measurements for the period
with air temperatures above 5C, are very close to each
other (Figure 3a and Table 4). Both show that the maximum
net CO
2
uptake occurs when the water table is in the range
10 to 20 cm. This range coincides with that for the average
water table depth of this microtopographical unit [e.g.,
Rydin, 1986]. The response function is also relatively
smooth, indicating that within this range the NEE is rather
insensitive to any change in water table depth.
[54] Quantile regression was used to investigate if the
water table depth is an active constraint on NEE across the
whole range of data. The quantile regression lines
(Figures 3b and 3c) run parallel in the statistical sense as
indicated by the 90% confidence intervals (not shown) for
the regression coefficients a and b. Confidence intervals
overlap for all the quantile regression lines based on the
observed values and all but the 95th based on the model
simulations. This means that the effects of the water table
level on NEE were similar across the range of uptake rates
from strongly positive to strongly negative. Only simulated
values comprising the 5% most positive (net release) NEE
values related differently to the water table level; that is,
the water table did not influence NEE within this range
(Figure 3b).
[55] The effects of the interactions between water table
depth and (1) global radiation and (2) air temperature on
NEE were also studied. For this, 2D diagrams of isolines
of NEE versus water table depth and global radiation
(Figures 4a and 4b), and water table depth versus temper-
ature (Figures 4c and 4d) were used. If the water table depth
had no effect on the NEE within a certain temperature or
solar radiation range, the NEE isolines should run parallel to
the x axis (water table depth) within that range. However, at
any given temperature or solar radiation level, the maximum
CO
2
uptake was found to be in the water table depth range
10 to 20 cm; outside this range the CO
2
uptake was reduced
(Figure 4). The same pattern can be seen for both the
measured (Figures 4a and 4c) and modeled (Figures 4b
and 4d) values.
[56] In order to separate the effects of the water table
depth on photosynthesis and peat decomposition, two
additional simulations were performed.
[57] 1. In the first run (Test run 1, psn) it was assumed
that photosynthesis was independent of water content; that
is, the photosynthetic rate was maximal with any water table
depth. This resulted in strong increases in net CO
2
uptake
when the water table was high, but no change in NEE when
the water table was low (Figures 5a and 5c and Table 4).
[58] 2. In the second run (Test run 2, resp) it was assumed
that all the acrotelm litter and peat decayed at a rate
corresponding to the rate of anaerobic decay of Sphagnum
plant material of the same degree of degradation, indepen-
dently of variations in water table depth. This resulted in
Table 3. Quantitative Measures of Model Performance With Respect to Daily NEE Values
a
MBE, gCm
2
d
1
MSE, gCm
2
d
1
MSE
s,
% MSE
u,
% d
Using measured water table 0.057 0.17 9 91 0.80
Using simulated water table 0.075 0.20 15 85 0.79
a
Daily NEE varies between 2.2 gCm
2
d
1
(uptake) and +1.7 gCm
2
d
1
(release). MBE, mean bias error; MSE, mean square error (apportioned into
MSE
s
, systematic mean square error, and MSE
u
, unsystematic mean square error); d, Willmott index of agreement (d = 0, no agreement; d = 1, perfect
match).
Figure 3. Relationship between the NEE (gC m
2
d
1
)
and water table depth (WT, cm) for the lawn at Degero
Stormyr. (a) Measured data (grey diamonds and solid trend
line) and model simulations (crosses and dashed trend line)
analyzed by the least squares second-order polynomial
model. The 95th, 80th, 65th, 50th, 35th, 20th, and 5th
quantile regression lines (b) for measured data and (c) for
model simulations.
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stronger net CO
2
uptake by the mire, due to the decrease in
decomposition (Figures 5b and 5c and Table 4). The most
pronounced differences from those obtained with the orig-
inal parameter settings were observed when the water table
was at a low level, and conditions are normally oxic. Least
squares regression lines based on the results of test runs 1
and 2 are significantly different from the regression line
based on the results from the standard model simulation
(Table 4).
4.4. Sensitivity Analysis
[59] The sensitivity analysis revealed the input parameters
that had strong effects on simulated NEE. The five most
important parameters were: w
high
; w
max
, describing the
inhibition in photosynthesis when the water content of the
moss is too high; r
m
, the maintenance respiration coeffi-
cient; k
an
, describing the degree of reduction in heterotro-
phic respiration under anoxic conditions; and k
2
, the rate of
peat decomposition (Table 5). This is in good agreement
with our analysis of the key factors governing NEE at
changing water table depths (section 4.3). The sensitivity
analysis also showed that the model is nonlinear with
respect to the state variable NEE, since the total effect term
TS
j
and the first-order sensitivity coefficient S
j
for the model
input parameters differ because of the significant contribu-
tion of the interaction terms to the total variance (Table 5).
5. Discussion
[60] We found good agreements between the measured
values and values simulated by GUESS-ROMUL for both
water table depth and NEE (Figures 1 and 2). However, the
MBE of the NEE estimates showed seasonal variability,
which, we believe, reflects model uncertainty in biomass
estimates. Vascular plants (which have strong, seasonal
phenological patterns) were excluded from the simulation,
under the assumption (grounded in the observation that the
net primary production in the tower footprint is almost
completely dominated by the Sphagnum species) that their
contribution to NEE is negligible. The gross primary
productivity (GPP) of vascular plants peaks in July August
when it appears to contribute up to 20% of total GPP (work
Figure 4. Isoline plots for the NEE (gC m
2
d
1
) as functions of water table depth and either air
temperature or global radiation. Each contour connects the points with equal NEE corresponding to a
particular combination of water table depth and (a, b) global radiation or (c, d) air temperature. Figures 4a
and 4c are based on measurements and Figures 4b and 4d are based on model outputs.
Table 4. Estimates of the Regression Coefficients for the Model
y = ax
2
+ bx + c + e
a
Coefficients
p a b c
Measurements 0.00370
b
0.134 0.401 0.12
Simulation, standard 0.00424
b
0.145 0.337 . . .
Test run 1 (psn) 0.00315
b
0.090 0.289 <0.001
Test run 2 (resp) 0.00375
b
0.147 0.165 <0.001
a
Here y is NEE (gCm
2
d
1
) and x is the depth to the water table
(centimeters), and significance P for H
o
: The regression line coincides with
the regression line based on the results from the standard model simulation;
psn denotes photosynthesis and resp denotes respiration.
b
Significance is <0.001.
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in progress), but much lower proportions of GPP are
attributable to the vascular plants during the rest of the
growing season. Even during the driest periods in 2002 and
2003 no sign of Sphagnum desiccation was noted on the
mire.
[61] The importance of autotrophic respiration as the
second most important factor in determining the sensitivity
of NEE provides further evidence of the need to increase
our understanding of plant respiration in ecosystem CO
2
exchange models [e.g., Cannell and Thornley, 2000; Dewar,
2000].
[62] Both the modeled and observed data showed that
NEE varied with changes in the position of the water table.
NEE was minimal (i.e., net CO
2
uptake was highest) when
the water table was 1020 cm below the surface, and it
increased (i.e., net uptake of CO
2
declined) when the water
table either rose or fell beyond this range (Figure 3),
regardless of other constraints (temperature or solar radia-
tion), as shown in the contour plots (Figure 4). The
parallelism of the quantile regression lines in Figures 3b
and 3c also indicates that the role of the water table on NEE
was very similar for all ranges of NEE, and thus that were
no interactions (multiplicative effects) between water table
depth and other variables. However, it does not exclude the
possibility that such variables may have additive effects, as
shown in Figure 4.
[63] Reductions in the mires net uptake of CO
2
when the
water table falls below this range may be due to increases in
temperature stimulating heterotrophic respiration and/or
increases in the proportions of the peat that are decompos-
ing under aerobic conditions [e.g., Silvola et al., 1996].
Reductions in CO
2
uptake when the water table is close to
or above the surface are probably due to the inhibition of
photosynthesis [e.g., Titus et al., 1983; Tuittila et al., 2004],
caused by reductions in rates of CO
2
supply due to its
restricted transport in water and/or reductions in levels of
incoming photosynthetically active radiation.
[64] Using the presented model we attempted to separate
the effects of changes in water table depth on (1) moss
photosynthesis (test run 1, psn), and (2) heterotrophic
respiration (test run 2, resp). When the water table level
was high, less than 10 cm from the peat surface, the
modeled NEE of the lawn was influenced most strongly
by changes in photosynthesis related to the water content of
the Sphagnum shoots (Figures 5a and 5c). When the water
table was low, the increased heterotrophic respiration due to
the proportionally larger aerobic decomposition was respon-
sible for the decrease in NEE (Figures 5b and 5c), while
photosynthesis continued at close to optimal conditions; that
is, despite low water table levels, the capillary action in the
surface peat was sufficient to supply the photosynthetically
active layer with water (Figures 5a and 5c).
[65] The results of this study indicate that there are
unlikely to be any great changes in NEE so long as the
average water table level remains in the wide range from 10
to 20 cm deep. However, if the climate changes there could
be significant long-term changes in the water table position.
If so, the model predicts that the NEE of the lawn commu-
nity will decrease; that is, less carbon will be taken up by
the mire ecosystem. The amplitude of the changes in NEE
Table 5. Sensitivity Indexes for the Five Most Important
Parameters, With Respect to Daily NEE
a
Parameter
Whole
Simulation Period
Days With Air Temperatures
Greater Than 5C
Sj TSj Sj TSj
w
high
0.150 0.264 0.166 0.280
r
m
0.134 0.216 0.055 0.183
k
an
0.089 0.197 0.084 0.184
w
max
0.074 0.160 0.091 0.187
k
2
0.043 0.070 0.041 0.074
a
S
j
, first-order sensitivity coefficient of the parameter j; TS
j
, total effect
term of the parameter j (sum of the first-order sensitivity S
j
and all higher-
order coefficients). Differences between S
j
and TS
j
indicate nonlinear
interactions between the parameters. All the differences presented in the
table are statistically significant (Students t-test) at the P < 0.001 level.
Figure 5. Relationship between the NEE (gC m
2
d
1
)
and water table depth (WT, cm) from the simulation with
the standard parameter settings and two different test runs.
(a) Test run 1: no effect of water table depth on
photosynthesis. (b) Test run 2: decay constant, correspond-
ing to anoxic conditions independent of water table depth.
(c) Trend lines (best-fits by second-order polynomial
regression) based on the data from the standard simulation
(bold solid lines), test run 1 (thin solid line with crosses),
and test run 2 (dash-dotted line).
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would depend on the combination of climatic factors, but
under certain circumstances carbon may even be released
from the lawn. The hydrological feedback mechanism
would mitigate these changes by evaporation decreasing
when the water table is low and runoff increasing (due to
higher peat transmissivity) when it is high. Since long-term
changes in species composition cannot be predicted with the
current version of the GUESS-ROMUL model, long-term
changes in NEE are likely to differ from the projections
based on the 3-year period studied here.
[66] Our future goal is to extend the current homogeneous
model by including the possibility of modeling shifts
between different microcommunities. Other vegetation clas-
ses that may be present in mires have already been param-
eterized for the GUESS model, for example, shrubs (A. Wolf
et al., Future changes in vegetation and ecosystem function
of the Barents Region, submitted to Climate Change, 2007)
and graminoids (R. Wania et al., Integrating peatlands and
permafrost in a dynamic global vegetation model: Evalua-
tion, sensitivity and application, manuscript in preparation,
2007). Following requirements for modeling mires listed by
Belyea and Baird [2006] we can summarize the advantages
and disadvantages of our modeling approach, as follows.
[67] Advantages of the GUESS-ROMUL model are:
(1) process-based representation of the photosynthesis and
transpiration allowing explicit response to climatic forcing;
(2) direct link between the fast ecosystem processes and
other processes as competition for light and water, tissue
turnover and population dynamics; (3) explicit representa-
tion of spatial heterogeneity in vegetation realized by
simulating a number of repetitive patches; and (4) nonlinear
interaction between the net primary production and soil
organic matter dynamics demonstrated in this study by
nonlinear interaction of the key parameters.
[68] Some disadvantages of the model, in comparison
with more advanced features of certain other mire models
(cited in parentheses), are that it does not account for:
(1) localized flows of water and nutrients [e.g., Nungesser,
2003; Swanson and Grigal, 1988; Rietkerk et al., 2004];
(2) plant competition for nutrients [Pastor et al., 2002];
(3) disturbance of the strict division between the acrotelm
and catotelm mediated by oxygen supplied via roots of
sedges and dwarf shrubs [e.g., Frolking et al., 2002; Zhang
et al., 2002]; (4) division of the vertical structure of the peat
into cohorts [e.g., Frolking et al., 2002]; and (5) feedbacks
between the peat accumulation and the hydrological pro-
cesses [Hilbert et al., 2000].
6. Conclusions
[69] In this study the modified GUESS-ROMUL model
of vegetation and soil CO
2
exchange was used to investigate
the effect of the water table depth on net CO
2
uptake rates of
a Sphagnum-dominated mire. NEE data obtained from
eddy-tower measurements over the lawn plant communities
at the oligotrophic minerotrophic mire Degero Stormyr were
used for the validation, and the model was shown to
perform well with daily time steps. The results of the study
indicate the following.
[70] 1. Rates of net CO
2
uptake at Degero Stormyr mire
were relatively constant during the 20012003 growing
seasons when the depth of the water table was between 10
and 20 cm. Model simulations show that any changes in the
water balance that do not cause the water table to shift from
within this normal range are likely to have minor effects on
the annual rates of land-atmosphere CO
2
exchange.
[71] 2. When the water table level was either higher
(>10 cm) or lower (<20 cm) than normal the mire net
CO
2
uptake decreased.
[72] 3. According to model simulations the reductions in
net CO
2
uptake rates that occurred when the water table
level was higher than 10 cm are attributable to reductions
in photosynthetic rates that are probably due to reductions in
the rate of CO
2
assimilation associated with higher than
optimal Sphagnum water contents and increased CO
2
lim-
itations due to the lower transport rate of CO
2
in water.
[73] 4. The reductions in net CO
2
uptake when the lower
water table level was lower than 20 cm resulted from
increases in heterotrophic respiration, rather than reductions
in photosynthesis since no Sphagnum desiccation was noted
on the mire or predicted by the model during 20012003.
[74] These results may be also applied to other mire sites
with similar vegetation, but only under the assumption of
constant plant species composition.
Appendix A
A1. Willmott Index of Agreement d
[75] This dimensionless measure is related to the absolute
size of the difference between the modeled and observed
values.
d 1
X
N
i1
M
i
O
i

2
=
X
N
i1
jM
0
i
j jO
0
i
j

2
" #
; A1
where M
i
0
= M
i
O, O
i
0
= O
i
O, M-modeled and O-
observed values.
[76] The Willmott index of agreement varies between 0
and 1, with d = 1 indicating a perfect match.
A2. Quantile Regression
[77] Quantile regression [Koenker and Bassett, 1978] is a
statistical technique that estimates the full range of condi-
tional quantile functions y Xb(t). Estimation of the
coefficients of b(t) is based on minimization of a weighted
sum (with weights depending on the order of the quantiles)
of absolute values of residuals,
min
X
n
i1
r
t
y
i

X
p
j0
b
j
x
ij
! " #
; A2
where n is sample size, p is the number of parameters,
r
t
(e) = e(t I(e < 0)), and I() is the indicator function.
A3. Variance-Based Sobol Sensitivity Indices
[78] The method proposed by Sobol [1990] is based on
decomposition of the function and construction of a model
output into sums of increasing dimensions. The variance D
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of the function can be broken down in a similar manner to
the ANOVA scheme,
D
X
k
j1
D
j

X
1i<jk
D
ij
. . . D
12:::k
; A3
where j indicates one of the k model input parameters ( j =
1,. . ., k). Both the total variance D and each term of this
decomposition can be computed using Monte Carlo
integrals [see Saltelli et al., 2000]. A first-order sensitivity
coefficient, showing the individual effect of the particular
parameter j on the model output, is defined as
S
j

D
j
D
: A4
[79] The total effect term TS
j
[Homma and Saltelli, 1996]
is defined as the sum of the first-order sensitivity coefficient
(S
j
) and all higher-order coefficients containing the param-
eter j, to account for the interaction of j with other
parameters. If the model is nonlinear, then S
j
and TS
j
differ
because of the significant contribution of the interaction
terms to the total variance.
[80] Acknowledgments. We are grateful to three anonymous referees
for their comments on an earlier version of the manuscript. This work has
been supported by the NECC project funded by the Nordic Centre of
Excellence (NCoE) Pilot Programme, the EU project BALANCE, EU grant
EVK2-2002-00169, NCCR-Climate, a research initiative of the Swiss
National Science Foundation and the Swedish Research Council for
Environment, Agricultural Sciences and Spatial Planning (grant 21.4/
2003-0876 to M. N.). The Kempe Foundation is also gratefully acknowl-
edged for grants for the micrometeorological instruments.
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2001GB001838.

M. Nilsson and J. Sagerfors, Department of Forest Ecology, Swedish

University of Agricultural Sciences, Umea, Sweden.
A. Wolf, Department of Forest Ecology, Institute of Terrestrial
Ecosystems, Zurich, Switzerland.
A. Yurova, Department of Physical Geography and Ecosystems Analysis,
Lund University, Solvegatan 12, Lund SE-22362, Sweden. (alla.yurova@
nateko.lu.se)
G02025 YUROVA ET AL.: MODELING CO
2
EXCHANGE IN A BOREAL MIRE
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I
I
III
I
Submitted to Water Resources Research
_____________________________________________________________________
A model of dissolved organic carbon concentrations in peat and stream
waters. I: Development and parameterization
Alla Yurova
1
, Ishi Buffam
2
and Kevin Bishop
3
1
Department of Physical Geography and Ecosystems Analysis, Lund University, Lund, Sweden
2
Department of Forest Ecology, Swedish University of Agricultural Sciences, Ume, Sweden
3
Department of Environmental Assessment, Swedish University of Agricultural Sciences, Uppsala,
Sweden
_____________________________________________________________________
Abstract
In this paper we present a model of the DOC concentrations and fluxes in peatland
water based on the convection-dispersion equation. The dynamics of sorbed soluble
organic carbon (SSOC) in the peat matrix are simulated in parallel with DOC. In a
companion paper (Part II), the model is fully applied to simulate the DOC
concentrations in the outlet of the steam draining a small headwater mire in Northern
Sweden during the period 1993-2001. Here, in Part I, the model is applied solely to
stagnant water conditions, in order to interpret the results of laboratory peat
incubations, with the focus on sorption processes. Some important model parameters
are derived using literature data complemented by information from new incubation
experiments. A dynamic approach is chosen for the representation of sorption kinetics
in the model. Differential sensitivity analysis accompanied by term-by-term
comparison of the processes contributing to changes in DOC are used to demonstrate
the leading role of sorption kinetics in determining DOC concentration during the first
days after the system is reset with a new concentration. Previous models of DOC
have usually been based on static equilibrium representations of sorption, and thus
may only be fully applicable for a rather coarse time resolution. Analysis of the model
highlights the importance of experimental design: when using laboratory peat
incubations to determine DOC production rate, the peat mass to water ratio and initial
SSOC concentration must be standardized or taking into account.
_____________________________________________________________________
Keywords: dissolved organic carbon, peatland, model
* Author for correspondence (alla.yurova@nateko.lu.se)
Introduction
If considered separately, the carbon cycle is not closed by exchange with the
atmosphere in either terrestrial or aquatic ecosystems, since the lateral flux of carbon
from the land to aquatic bodies links the two. In the boreal zone, the flux of dissolved
organic carbon (DOC) is much greater then the particulate organic carbon (POC) and
dissolved inorganic carbon (DIC) fluxes [Cole et al., 2007; Jonsson et al., 2007].
Export of DOC is likely to occur mainly from areas where flowing water bypasses the
adsorbing mineral soils, i.e. in surface and subsurface flows from littoral zones and
wetlands [e.g. McKnight and Aiken, 1998; Qualls, 2000]. Field data collected from
various locations within the boreal region have highlighted the importance of
peatland, a boreal type of wetland, as one of the major sources of terrestrially-derived
dissolved organic carbon (DOC) reaching aquatic ecosystems [e.g. Freeman et al.,
2001a; Tranvik and Jansson, 2002]. The amounts of DOC exported from peatlands
are sufficiently large to be included in regional C balances [e.g. Jonsson et al., 2007]
and to contribute to the peatland C balance [Moore et al., 1998; Fraser et al., 2001;
Billett et al., 2004, Roulet et al., 2007]. Aquatic scientists widely recognize the role of
DOC in providing energy for microbial growth [e.g. Jansson et al., 2000], regulating
UV light attenuation [e.g. Schindler et al., 1996], pH buffering [e.g. Kortelainen et
al., 1989] and metal complexation [e.g. McKnight and Aiken, 1998]. Other effects of
DOC, such as interference with photosynthesis, mild stress on aquatic organisms,
promotion and suppression of microbial growth etc. are less well known but equally
important [Steinberg et al., 2006].
2
The concentration of DOC in peat pore water is a product of inputs (production of
soluble organic matter from living plants and decomposing organic remains, imports
via flowing water, dispersion), outputs (mineralization, hydrological export,
dispersion) and sorptive exchanges with solid organic matter [see, for instance Qualls
and Richardson, 2003]. The complexity of the problem has encouraged the
development of several process-based models of DOC in soils and stream water
[Grieve, 1991; Neff and Asner, 2001; Michalzik et al., 2003; Futter et al., 2007].
Despite the inevitable simplifications when describing chemical and biochemical
DOC transformations, such models have been relatively successful for predicting
DOC concentration and fluxes over daily, weekly or monthly time intervals. Within
the existing models, there is general consensus about which processes to include,
namely microbially-mediated transformations, sorptive exchange and hydrological
transport of the DOC. However, the equations used to describe each process differ
from model to model.
For our study (Parts I and II) we selected a mathematical framework that, to our
knowledge, has not previously been applied to the DOC problem. Solving the
classical convectiondispersion equation [e.g., van Genuchten and Waganet, 1989],
we included two important capabilities that are new to DOC models: (i) accounting
for the non-steady state behavior of organic carbon with respect to sorption and (ii)
dynamically and simultaneously tracing transport and biochemistry in the
heterogeneous peat profile.
The model is built up in two separate stages, the ultimate goal being to simulate DOC
export from boreal mires. In Part I (this paper), the model is fully developed. In
3
addition, the version applicable to a situation with stagnant-water and constant
temperature is used to explore the implications of the model for laboratory incubation
studies. We also performed long-term laboratory incubations with peat samples
collected from the mire studied in Part II to help derive the model parameters. In Part
II (the companion paper), the predictive form of the model is used to simulate DOC
build-up and transport from a small boreal peatland in Northern Sweden during the
period 19932001.
In addition to model parameterization, our main purpose in Part I was to explore the
role of sorption as a major mechanism regulating DOC concentration and release
rates. This was achieved by analyzing the proposed equations and by applying
differential sensitivity analysis to the key model parameters. While laboratory studies
have shown that DOC sorption is time-dependent [e.g. Qualls, 2000; Rasse et al., in
revision], previous DOC models have treated it as an instantaneous process. In the
model developed by Grieve [Grieve, 1991; also applied in Boyer et al., 1996], the
ratio between the store of soluble C and DOC concentration in water is an empirically
fitted constant, while the models of Neff and Asner [2001] and Michalzik et al. [2003]
make use of an equilibrium partition constant, based on laboratory soil incubations, to
relate the amounts of dissolved and sorbed OC. Here we aim to demonstrate the
advantage of the dynamic approach over a static one when modeling DOC sorption.
We use the data from the long-term peat incubations and perform calculations
designed to interpret the data, focusing on sorption action over different time intervals
and under a range of experimental conditions. This approach is particularly important
when trying to use laboratory studies to parameterize models.
4
Model
The model presented here is based on two main premises:
- Soluble organic carbon is present in the mire system in two states: dissolved
(DOC) and sorbed, potentially soluble, but currently solid (SSOC). The balance
between the two phases varies over time, and adsorption and desorption can be
described using first-order kinetics.
- A convectiondispersion equation can provide a suitable model of DOC
transport. Including the terms accounting for the adsorptiondesorption as well
as microbial production and mineralization in this equation, accounts for the full
mass balance for DOC.
The underlying concept of the model is the schematic representation of processes that
generate and consume DOC in a mire ecosystem, as presented in Qualls and
Richardson [2003]. The processes that contribute to in situ changes in pore water
DOC concentration (sorption, microbial transformation) are distinguished from the
various transport processes, including dispersion and advection by both vertical and
horizontal flow (Figure 1). We model sorption and microbially-mediated
transformations of the DOC according to the scheme presented in Figure 2. This part
of the model (shaded in Figure 1) can be used alone to study situations with no water
flow (e.g. systems with stagnant water in laboratory conditions, Part I this paper).
Transport processes are modeled as presented in Figure 3, which is relevant to
application of the model under field conditions (Part II). The hydrological model used
to simulate the flow rate and water content is the Mixed Mire Water and Heat Model
[MMWH, Granberg et al., 1999].
5
Sorption
In our model we use a linear kinetic equation to simulate the adsorption of soluble
organic matter to, and its desorption from, solid organic matter. Sorption plays a key
role in controlling the DOC concentration in pore water. As shown by Qualls [2000],
peat always contains substantial quantities of potentially soluble OC sorbed to solid
organic matter (SSOC); there is always more of this than the amount dissolved during
any single leaching event. In laboratory peat incubations, it is possible to observe the
rapid release of DOC into newly added pore water. This occurs as previously sorbed
DOC is desorbed, and during the period before DOC concentrations in the sorbed and
dissolved phases reach equilibrium [Qualls, 2000]. Equilibration is not achieved
immediately, and the rate of change in the DOC concentration due to sorption is
highest at the beginning of experiment, decreases during incubation, and is negligible
only after 2-4 days [e.g. Rasse et al., in revision]. Equilibrium sorption models do not
describe the dynamics of the process adequately, since they are based on the
assumption that exchange between the sorbed and dissolved phases is so rapid that
there is always a fixed ratio between the two. Hence, a linear kinetic model is a better
when modeling DOC with a relatively high time resolution. Sorption occurs not only
in cases in which the pore water is completely replaced (such as laboratory
experiments), but also in any situation where dilution or advection alter DOC
concentrations in pore water.
Microbially-mediated transformations
Metabolic transformations mediated by microorganisms also strongly affect the DOC
balance. Acting at slower rates than adsorption/desorption processes, such
transformations determine the total amount of soluble OC produced by exoenzymatic
6
dissolution of insoluble organic matter. In addition, they control the rate of
mineralization of DOC and SSOC. In our model we only consider the recalcitrant
fraction of DOC. The DOC present in the pore water of a boreal peatland is almost
exclusively in the form of refractory biomolecules, such as fulvic acids and their
precursors [e.g., McKnight et al., 1985] formed mainly from the decomposition of
Sphagnum. Easily decomposable substances, such as root exudates and simple
products of decomposition, are readily consumed by microorganisms, and have high
turnover rates and very short lifetimes in solution [e.g., Qualls, 2000; Crow and
Wieder, 2005]. They therefore comprise only a small proportion of the DOC in
solution [e.g., McKnight et al., 1985]. Out model does not aim to describe the
dynamics of such short-lived substances, and the parameterization presented below is
based solely on data from the laboratory peat incubations in which the bulk DOC of
the peat solution was measured.
In our simplified model, like many models of soil organic matter dynamics [e.g.,
reviewed by gren et al., 1991], microbial biomass is not explicitly included. Instead,
we simulate microbially-mediated transformations of the DOC (production and
mineralization) that are affected by temperature and the presence of O
2
. We assume
first order kinetics for the mineralization of DOC and SSOC. A zero order production
constant is hypothesized, based on the assumption that substrates for the formation of
DOC are not limited in the peat organic matter.
Bulk DOC concentration is simulated without specifying chemical fractions; we
deliberately avoided separating organic matter into different fractions. Instead, we
believe that a new modeling approach to soil organic matter turnover, based on
7
tracing changes in the continuous distribution of quality of the organic molecules,
known as Q theory [gren and Bosatta, 1998], provides a more promising approach.
However, the application of Q theory to soil solutions has many associated
challenges, such as defining the relationships between quality and solubility, or
between quality and sorption properties, so this work has yet to be undertaken.
Mathematical formulation
The mass balance equation for the concentration of DOC in pore water, c, can be
written in the form of convection-dispersion equation (symbol definition and units in
Table 1):
, c P ) c K s ( ) c ( ) c (
t
) c (
1 D des
+ + - V - V - V =
c
c
u t u
u
T ' [1]
where u is volumetric water content, is peat bulk density, vector q{q
x
, q
y
, q
z
} is
composed of water flux in horizontal and vertical directions, D{D
x
, D
y
, D
z
} is a vector
of dispersion coefficients, t
des
is the kinetic rate of desorption (or adsorption if the
third term in equation [1] becomes negative), K
D
is the partitioning constant,
characterizing an equilibrium between the sorbed and dissolved phases, P is the
microbial production rate of DOC, and
1
is the first-order microbial DOC
mineralization constant. Mathematical operators are:
),
z
c
D (
z
)
y
c
D (
y
)
x
c
D (
x
) c (
z y x
c
c
c
c
+
c
c
c
c
+
c
c
c
c
= - V - V u u u u '
z
) c q (
y
) c q (
x
) c q (
) c (
z
y
x
c
c
+
c
c
+
c
c
= - V T
8
The dispersion coefficient is defined as
,
q
D D
0
u
+ =
where D
0
is the molecular diffusion coefficient, and is dispersivity.
A mass balance for the concentration of the sorbed soluble organic carbon (SSOC), s,
leads to:
, s ) c K s (
t
s
2 D des
=
c
c
t [2]
where
2
is the first-order microbial mineralization constant for the SSOC. We use the
term concentration rather than pool size to describe amounts of SSOC present in
the systems considered in this paper, even though sorbed soluble OC is in a solid state
and thus it is not a wholly appropriate term. The intention is to emphasize that the
term refers to the amounts sorbed per unit mass of peat.
The system (1)-(2) describes DOC concentration at a given time, assuming that the
initial DOC and SSOC concentrations (c
0
and s
0
) are known.
A modified vant Hoff equation is used to account for the temperature dependency of
DOC production and mineralization:
9
) 10 / ) ((
10 1 1
) 10 / ) ((
10
,
Tbasal T
basal
T T
basal
Q
Q P P
basal

=
=

where Q
10
is the relative rate of increase in metabolic rates per 10C increase in
temperature, and T
basal
is a reference temperature (20C was used here) at which basal
rates of microbial DOC production (P
basal
) and mineralization (
1basal
) are estimated.
The constant fractions k
anP
and k
an
are used to relate the anoxic rates of DOC
production and mineralization to the corresponding DOC production and
mineralization rates under aerobic conditions:

an 1 1
anP
k
, k P P conditions anaerobic for
=
=
In our model we made use of separate mineralization rate coefficients for DOC and
sorbed soluble organic carbon (SSOC), to account for the stabilizing effect of sorption
on microbial decomposition [e.g., Kalbitz et al., 2005]. The parameter k
s
is defined as
1
2
s
k

= ,
i.e. the mineralization rate of SSOC (
2
) is k
s
times less then the mineralization rate of
DOC (
1
).
10
Hydrology
Two layers can be distinguished within the peat profile: the upper, periodically
aerated layer consisting largely of living and lightly decomposed plant material
(acrotelm), which is not usually more than 60 cm deep, and the permanently saturated,
lower zone, consisting largely of compacted, relatively highly decomposed plant
material (catotelm). The hydraulic conductivity is high in the upper acrotelm and
declines strongly with depth [Ivanov, 1981]. Consequently, water in the acrotelm
moves rapidly and usually accounts for most of the discharge from the peatland. If the
peatland is formed over minerals soils with low permeability, the water in the
catotelm is mostly stagnant; if, however, the underlying mineral soil is highly
permeable, significant vertical movements can occur [e.g., Reeve et al., 2000]. It has
also been suggested that the acrotelm is the major source of DOC within peatlands
[e.g., McKnight et al., 1985], since decomposition is much slower in the permanently
anaerobic layers, where organic matter is aged and the enzymes regulating organic
matter transformation are suppressed [Freeman et al., 2001a, b], in addition
accumulations of CH
4
and CO
2
further inhibit decomposition [Blodau et al., 2004].
For the field application of our model (Part II) we selected the Mixed Mire Water and
Heat (MMWH) hydrological model [Granberg et al., 1999] to simulate water flux,
water content, water table depth and temperature in the acrotelm. The MMWH model
is designed specifically for peatlands and is based on the method of steady-state
moisture distribution curves [Romanov, 1961]. Utilizing continuous functions for the
moisture profile and hydraulic conductivity, the model gives accurate accounting of
temperature, moisture, and the water table depth in the acrotelm with high vertical
11
resolution. Simplification of the system of equations (1)-(2) for the 1D flow system
described by the MMVH is given below.
Addition: Convectiondispersion equation for the simplified hydrological schemes.
If a 1D hydrological scheme (vertical discreetization) is used and there is no lateral
import of DOC into the peatland from surrounding areas, the finite-difference form of
system (1)-(2) can be written as.
, c P ) c K s (
z
) c q (
z
c q
z
)
z
c
(
D
t
) c (
1 D des
z s
z
+ + =

u t
A
A
A A
A
A
u A
A
u
[1a]
, s ) c K s (
t
s
2 D des
= t
A
A
[2a]
where Az is the depth of a specific layer, q
z
is the vertical water flux and q
s
is the
specific (area) lateral discharge from the layer z.
In this simple model, the DOC export flux from the peatland is the sum of the
products of the concentration and specific discharge from all of the vertical peat
layers.
Model parameterization
The model parameters and initial conditions needed to solve equations (1)-(2) in a
predictive way are summarized in Table 2. Only the parameters relevant when
applying the model to laboratory conditions are specified here in Part I (the other
12
parameters are listed, but their values and ranges are presented in Part II, where the
model is applied to field conditions). Four parameters the sorption equilibrium
partitioning coefficient (K
D
), the initial SSOC concentration (s
0
), and the rates of
microbial DOC production (P), and DOC mineralization (
1
) were estimated using
data obtained from our long-term peat and water incubations (experimental design in
Appendix). In addition, we reviewed the available literature relating to the rates of
DOC release into water during long-term experiments to obtain ranges of estimates
for the two parameters: the DOC production rate (P) and the DOC mineralization rate
(
1
).
Method for obtaining values for model parameters
The required model parameters were determined by a two-step process. First, we had
to accept statements (i) and (ii) presented below. Second, when the numerical values
of the model coefficients had been determined, these assumptions were tested by
means of the analysis of the equations and sensitivity study described under Model
analysis.
We hypothesized that the process of DOC release into the solution could be divided
into two phases, this is conceptually similar to the tea bag approach presented in
Aguilar and Thibodeaux [2005].
(i) During the initial phase, the process is dominated by rapid sorption
desorption kinetics and not detectably influenced by microbially-mediated
rates of production and mineralization [Qualls, 2000]. This phase can take
up to 4 days [e.g. Rasse et al., in revision] until an equilibrium is reached
13
between the DOC in the pore water and the amount of potentially soluble
DOC sorbed on the peat matrix.
(ii) The second phase involves much slower DOC release; this is governed by
microbial DOC production and mineralization, although also mediated by
sorption.
What we refer to here as the first steady state or sorption equilibrium is in fact a
local equilibrium between the DOC solution and the initial SSOC concentration in the
peat matrix; this is reached after the rapid release of DOC in the initial phase of the
experiment. Subsequently, the DOC concentration changes more gradually, as a result
of microbial production and mineralization. It can be shown that a global steady state
is also eventually reached, i.e. the system eventually evolves to a state in which the
production of DOC is balanced by the sum of the mineralization of both DOC and
SSOC. However, this global steady state is mostly of theoretical interest, because it is
only reached after a very long time.
Under laboratory conditions there is no advective water flow and diffusive mixing
within the peat sample can be considered instantaneous. Therefore, equations 1-2 can
be simplified as:
, a M P a
V
M a
K
t
a
c 1 s des
c
D des
c
+ +

=
c
c
t t [3]
, a a
V
M a
K
t
a
s 2 s des
c
D des
s

=
c
c
t t [4]
14
where M is the mass of the peat sample, V the volume of water added, while a
c
and a
s
are the amounts of DOC and SSOC, respectively, defined as a
c
=cV and a
s
=sM.
We assumed that, in a continuous laboratory incubation of peat with stagnant water,
after the first steady state between DOC and SSOC is reached the sorption reaction
system is in quasi-equilibrium if examined at a daily time step. That is because the
change in total soluble OC is so small that the system continuously adjusts to the state
when s(t)=K
D
c(t). The analytical solution of equations (3)-(4) can then be obtained:
,
V
M
K k 1
M P
)) t t (
V
M
K 1
V
M
K k 1
exp( )
V
M
K k 1
M P
V c ( V ) t ( c
D s 1
1
D
D s
1
D s 1
1
|
.
|

\
|
+

+
|
|
|
|
.
|

\
|
+
+

|
.
|

\
|
+

[5]
where c
1
is the DOC concentration at time t
1
, the end of the initial phase of incubation
when the first steady-state is established between the sorbed and dissolved phases.
Assuming that the steady-state has been reached, c
1
can be calculated from the mass
conservation as follows:
,
1
V
M
K
c
V
M
s
c
D
0 0
1
+
+
= [6]
where c
0
and s
0
are the initial DOC and SSOC concentrations, respectively.
The initial DOC concentration, c
0
, and the pore water DOC concentration in the first
sorption steady state, c
1
, can be measured in a laboratory peat incubation. Our peat
15
incubation experiments were performed with two types of water: water from the
stream at the outlet of the mire studied in Part II (c
0
=44 mg L
-1
DOC), and tap water
(deep groundwater) from a research station near the site (c
0
=0.3 mg L
-1
DOC) (details
in Appendix). The stream water (44 mg L
-1
DOC) was also incubated on its own to
measure DOC mineralization in the water.
K
D
, s
0
: data from laboratory incubations
The time t
1
was determined from the DOC vs. time curve as the critical point at which
the first local steady state is reached. Using two measurements of the DOC
concentration (c
1
) at time t
1
, one for each of the two different initial DOC
concentrations (c
0
), we used equation (6) to estimate, simultaneously, the partitioning
coefficient K
D
and the initial SSOC concentration, s
0
.
P.
1
: data from laboratory peat and water incubations
DOC concentrations from the readings starting from t
1
were inserted into equation (5)
and the values of P and
1
, optimized using the least squares method. All experiments
were performed at 20C, so that we could estimate the basal rates P
basal
and
1basal
.
The values of parameters t
des
and k
s
were as presented in Table 2. The data from the
incubations of water without peat were used to calculate the DOC mineralization rate
(first order decay coefficient) in water alone (Table 2). Our original intention was to
compare the resulting value with the corresponding DOC mineralization coefficient,

1basal
, obtained by optimization, using the data from peat incubations to determine
whether the presence of a peat matrix changes the DOC solution mineralization rate.
16
However we discovered that the optimization procedure was insufficiently sensitive to
measure
1basal
, and the lower range of this parameter was not detectable, presumably
because the mineralization rate is an order of magnitude lower than the rate of
production determined at the same time during the optimization. Therefore, only the
upper range of
1bsal
was determined by optimization. The range of the DOC
production rate, presented in Table 2, reflects experimental uncertainty.
P,
1
: literature data
The least square optimization for equation (5) was also utilized to process the data
relating to DOC release rates from the long-term laboratory peat incubations found in
the literature [Moore and Dalva, 2001; Aguilar and Thibodeaux, 2005; Chow et al.,
2006]. The listed studies did not contain information about the values of model
coefficients K
D
(sorption partitioning coefficient) or k
s
(degree of reduction in the
microbial mineralization when the soluble OC is sorbed) required to calculate the
DOC production and mineralization rates in accordance with equation (5). The
original concentration of SSOC (amount present sorbed on the peat matrix, s
0
) was
not reported either. To overcome this problem we used a range of parameter values:
K
D
=0.019-0.091 Lg
-1
(minimum taken from Qualls [2000], maximum from Rasse et
al. [in revision]); k
s
=1/6-1/3 [Kalbitz et al., 2005]; s
0
=1-10 mgg
-1
(assumed)). The
results of the calculations therefore present possible ranges for the parameters of
interest (Table 3). When appropriate, the mineralization rate was estimated from the
optimization, otherwise the literature range of the decay constant of the slowly
decaying DOC fraction in throughfall, soil and stream water (
1
=1 10
-5
-4 10
-5
h
-1
,
[Qualls 2000]) was used. The range of the calculated production rate here reflects
17
both experimental uncertainty and uncertainty included in the estimates of the other
parameters that were used.
Model analysis
To test the validity of our hypothesis that long-term laboratory incubations could be
considered as two discrete phases, we examined the partitioning between net sorption
(adsorption minus desorption) and net microbial production (production minus
mineralization) for the day-to-day DOC concentrations during the incubations .This
was done by numerically estimating each term in equation (3). Hourly simulated
outputs were accumulated within daily time intervals and two starting DOC
concentrations (0.3 and 44 mgL-1) were considered, in accordance with our
experimental design.
The local sensitivity coefficients for the model parameters K
D
, t
des
, P and
1
were
calculated by differential analysis [e.g., Saltelli, 2000]. Differential analysis is often
used in studies of chemical kinetics [e.g. Saltelli, 2000]. Equations are derived from
the original equation system to simulate the partial derivatives (local sensitivity
coefficients) of the variable of interest (e.g. DOC concentration) with respect to the
model parameters The direct method, utilizing the simultaneous numerical integration
of a combined system of equations (3)-(4) and forward sensitivity equations, was then
applied here using the CVODES programming package [Cohen and Hindmarsh,
1996; program codes and documentation available from
http://www.llnl.gov/casc/sundials/].
18
Two important characteristics: (i) the DOC during the first steady-state (c
1
), and (ii)
the rate of DOC release into the water during the second stage of the incubation, were
further evaluated. First, we examined the dependency of c
1
on the initial SSOC
concentration (s
0
) and the peat-to-water ratio (M/V) given by equation (6). In addition,
the net DOC release per unit peat mass per unit time,
M t
V c
A
A
, was derived by
applying equation (5) to the time interval At. By changing the input parameters that
are used in the resulting equation, we analysed the influence of such factors as the
initial SSOC concentration (s
0
) and peat-to-water ratio (M/V) on the rate of DOC
release observed during the laboratory incubations.
Results
Parameterization results
Our estimate of the sorption partitioning coefficient K
D
was 0.033 Lg
-1
; lower than
that estimated by Rasse et al. [in revision] based on a surface peat sample from a mire
in Norway (K
D
=0.091 Lg
-1
) and slightly higher than 0.019 Lg
-1
, the value we
calculated using the data by Qualls [2000], who analysed a surface peat sample from a
marsh in the Everglades (Florida, USA). Our estimate of the initial SSOC
concentration s
0
=2.0 mgg
-1
is close to that in a Lamnela peat sample from a drained
black spruce forest soils in Alaska (s
0
=2.7 mgg
-1
[Qualls and Richardson, 2003]).
As summarized in Table 3, the rate of microbial production can be substantially
higher than the rate of release of DOC from peat into water. As interpreted by our
model, the rate of release is less than the rate of production because soluble OC
produced by microbial transformations of the peat is distributed by sorption between
19
the solid and dissolved phases. In experiments, measurements of DOC in solution will
only record the increases in the dissolved phase.
The estimated rate of microbial production we found in our incubation of surface
Sphagnum peat is of the same order as that from the sapric Sphagnum peat sample
incubations of Moore and Dalva [2001] (Table 3). A fibric peat sample [Moore and
Dalva, 2001] and a peat sample from a river delta [Aguilar and Thibodeaux, 2005]
had higher rates of microbial production, while a different river delta sample [Chow et
al., 2006] showed much lower production, resulting in net mineralization. The upper
limit for the mineralization rate in our sample, defined by the optimization, was 1.0
10
-4
h
-1
. This is of the same order as the upper range of the decay constant of the
slowly decaying DOC fraction in throughfall, soil and stream water [0.410
-4
h
-1
,
Qualls, 2000].
The role of sorption as a regulator of DOC concentrations at different time
intervals during peat incubations
The role of sorption can be illustrated by comparing modeled net sorption and net
microbial production (Figure 4 a). For the chosen parameter range, sorption strongly
dominated over production during the first 39 h of incubation. It was difficult to
estimate the partitioning later in the experiment as here we compared very small
numbers so that numerical errors became an important issue. To overcome this
problem we compared the weighted local sensitivity coefficients (
i
i
p
c
p
c
c
) calculated
for the main model parameters (p
i
) with hourly time resolution and high precision
(Figure 4b). The kinetic constant of sorption, t
des
, was the most important parameter
for the first 14 hours of incubation, after that the sorption partitioning coefficient K
D
20
was most important. After 39h the sensitivity coefficient for the production rate P
became higher than that for the kinetic desorption t
des
coefficient.
The two coefficients related to sorption have distinctly different meanings. While t
des
is indicative of the time-dependent kinetics of the process, K
D
characterizes the
partitioning between the solid and dissolved phase when the system is in a steady state
in relation to sorption. Both methods that we used to distinguish sorption from
microbial processes (Figure 4a,b) indicated that partitioning the DOC release into two
phases was valid. There was a clear division between the first phase when sorption
kinetics dominated and the second phase when the observed DOC concentration was
in equilibrium, with adsorptive partitioning of microbially-produced DOC.
Abiotic factors affecting the first (local) steady-state DOC concentration and the
rate of DOC release during the second phase of the long-term peat incubation
As expected according to the model, the amount of DOC that was rapidly released
during the first phase of incubation (c
1
) exhibited a linear relationship with the
amounts of soluble OC originally sorbed on the peat matrix (Figure 5a). Increases in
the ratio of peat mass to water volume resulted in non-linear increases in the amount
of DOC released (Figure 5b). There was also a notable difference in the DOC release
rate during the second phase of incubation depending on the experimental set-up:
- The DOC release rate has a negative linear relationship to the initial SSOC
concentration, s
0
(Figure 5c). The initial SSOC concentration presumably varies
naturally among peat samples of different origins and is also affected by the
rinsing or washing of the sample before the start of the experiment. Within the
21
expected natural range of SSOC concentrations the effect on DOC release rate is
presumably not high (up to 10%), but it is significant.
- The DOC release rate is non-linearly negatively related to the peat mass-to-water
ratio in the experiment, M/V (Figure 5d). For example, DOC release per mass
unit is much smaller for an unsaturated sample than for a saturated or flooded
sample. This is noteworthy, since here we only deal with the
adsorption/desorption processes affected by the relative proportion of the solid
phase and the effect of moisture on microbial activities is not considered.
Summary and discussion
The convectiondispersion equation used in this paper was developed to describe the
transport and transformations of DOC in a peatland system. To our knowledge this
mathematical approach to the problem of DOC dynamics is new, although in
analogous fields of hydrology, such as pesticide transport in soil, solving the
convection-dispersion equation has become almost standard practice [Lapidus and
Amundson, 1952; van Genuchten and Wagenet, 1989; Baskaran et al., 1996].
The dynamic, rather than static, representation of sorption kinetics is one of the
strengths of the proposed model. In this paper we evaluated mathematically the
mechanism of DOC regulation by sorption kinetics studied in the laboratory
experiments [e.g., Qualls 2000, Rasse et al., in revision]. It is known and often cited
in relation to laboratory DOC studies that (i) the initially rapid release of DOC from
the peat matrix into newly added water is mainly due to abiotic processes [e.g.,
Qualls, 2000, Moore and Dalva, 2001; Aguilar and Thibodeaux, 2005; Chow et al.,
22
2006] and (ii) depending on the initial concentration, DOC can be either added to or
removed from the solution as a result of sorption [Qualls, 2000]. In the current study
we provide equations to explicitly describe these mechanisms.
By comparing the terms in the differential equation for DOC concentration and
performing differential sensitivity analysis we have been able to establish that there is
a clear division between the two phases of DOC release into water during long-term
peat incubations. The first phase was short (39 h for the model fitted to our laboratory
study) and governed solely by sorption, while further, much slower, increases in DOC
concentration during the second phase were due to microbial activity. In addition, the
derived equation (6) illustrated that, depending on the initial DOC and SSOC
concentrations, DOC concentration can either increase or decrease during the first
phase of incubation.
The model also suggests that, during the second phase, the additional soluble OC
produced slowly due to microbial transformations of the peat is distributed by
sorption between the solid and dissolved phases. Only the DOC contribution is
recorded when the change in solution concentration is measured, which means that the
true production rate is likely to be higher, and includes the fraction that becomes
sorbed (illustrated by comparing columns three and four in Table 3).
Using the model equations we were able to show how the release of DOC into water
during the long-term incubations can vary substantially according to the experimental
set-up. The DOC release rate into solution was affected by the peat-to-water ratio
(Figure 5d) and the initial amounts of soluble DOC sorbed on the peat matrix (Figure
23
5c). Both effects persisted for the whole time period we considered in our simulations
(five years). It was not only the amounts of DOC released initially, but also the
subsequent release rate that was affected by the initial SSOC concentrations and the
peat mass to water volume ratio. Different peat-to-water ratios resulted in different
rates of DOC release because the same net production was partitioned differently
between the sorbed and solid phases. This consideration is of great importance when
conclusions about microbial processes are drawn by comparing the results of different
laboratory incubations. Usually only changes in DOC concentration are measured and
directly converted to production rates, without taking into account the peat-to-water
ratio. The effect of sorption, rather than other factors such as, for example, the effect
of moisture on microbial activity, could account for the different DOC release rates if
data from two experiments with mixtures of peat and water in different ratios are
compared. The effect of sorption would be even stronger if water was flowing through
the peat, significantly changing the total amount of SSOC; in this paper, however, we
deal only with stationary water.
In Part I we only apply the model to laboratory conditions with stagnant water.
However, the general model formulation we presented here is sufficiently broad to use
in field applications. In the accompanying paper the model is applied to simulate the
variations in DOC concentrations observed during 1993-2001 in a stream draining a
small headwater mire in Northern Sweden. The sensitivity of the results to the
parameter choice is also discussed.
24
Appendix
Method for the long-term (165 day) incubation of a peat-water slurry with two
different initial DOC concentrations.
Two peat samples were collected, using a 5 5 70 cm peat corer, from
Kallkllsmyren, an 8 ha peat mire in northern Sweden (see Sirin et al. 1998 for a
detailed description) on 21 September 2005. Core 1 was taken from an area
dominated by cottongrass and sedge, while core 2 was taken from an area with
cottongrass, sedge and Sphagnum. A 10 cm
3
plug of peat was taken from depths of 40
cm and 60 cm and combined to give a 20 cm
3
plug (approximately 35 g wet wt, 3 g
dry wt). Peat plugs were then incubated in 300 mL pre-acid washed high-density
polyethelene bottles filled with water. Two types of water, with different DOC
concentrations, were used: tap water (GW = deep well groundwater) from a research
station near the site (0.3 mg L
-1
DOC) and stream water (S) from the mire outlet (44
mg L
-1
DOC). Each combination of water type (S and GW) and core (1 and 2) were
incubated in triplicate in the dark at 20C.
Periodic measurements (after 1h, 2d, 43d, and 165d) of absorbance at 254 nm were
made in order to calculate DOC concentration in the aqueous phase of the waterpeat
slurry. At the time of each measurement, sample bottles were shaken vigorously for 1
minute, then allowed to stand for 30 minutes, after which 5 mL of the sample was
filtered (disposable Millipore 0.45 m MCE membrane filters) and analyzed for
absorbance using an HP 8452-UV Spectrophotometer. Incubation bottles were topped
up with sample water, leaving no headspace during the subsequent incubation. In
addition, bottles of water without peat were incubated as control treatments under the
25
same conditions. The stream water incubation (44 mg L
-1
DOC) was used to measure
the rate of DOC mineralization in water alone. Absorbance at 254 nm was used to
estimate DOC concentration using a well-established rating curve for samples from
the stream at the Kallkllsmyren mire outlet:
254
23 A c = N = 83; R
2
= 0.85; RMSE = 4 mg L
-1
(13%) (1.1A)
Where c is DOC concentration (mg L
-1
) and A
254
= absorbance at 254 nm (AU)
measured in a 1 cm quartz cuvette. Measurements of DOC (Combustion and
measurement as CO
2
using a Shimadzu TOC-V
CPH
analyzer) for selected samples
from the incubation experiment gave values always within 20% of the value estimated
from eq. 1.1A.
26
Acknowledgements
We thank Daniel Rasse for insightful comments and suggestions, as well as for
sharing the data from his on-going research. This work has been supported by the
NECC project funded by the Nordic Centre of Excellence (NCoE) Pilot Programme
and by the Swedish Research Council for Environment, Agricultural Sciences and
Spatial Planning.
27
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30
Figure captions:
Figure 1. Schematic diagram of the model formulation and structure.
Figure 2. Schematic diagram of the modeled pools of soluble organic matter and
processes that control microbially-mediated transformation (production and
mineralization) and abiotic exchanges (adsorption and desorption) between the solid
and dissolved phases. Parameters are described in detail in Table 1.
Figure 3. Schematic diagram of DOC transport within the peat profile, based on the
convection-dispersion framework.
Figure 4. (a) Modeled estimates of partitioning of the DOC release rate observed in
laboratory peat incubations into net sorption (desorption minus adsorption) and net
production (microbial production minus mineralization) components in incubations
with two starting DOC concentrations (c
0
) in water. (b) Normalized first-order local
sensitivity coefficients computed by the forward direct method for the main
parameters in the model applied to laboratory incubations (calculated for an
experiment with c
0
=0.0 mgL
-1
).
Figure 5. Simulated effect of the initial soluble solid organic carbon (SSOC)
concentration, and peat-to-water ratio (M/V) on the first steady state DOC
concentration (a, b); and on the DOC release rate during the second phase of a long-
term peat incubation (c, d) (K
D
=0.033 Lg
-1
, c
0
=0.0 mg L
-1
, P=1.0 10
-3
mgg
-1
h
-1
,
1
=4
10
-5
h
-1
, k
s
=0.3 ; (a, c) M/V=10.0 gL
-1
; (b, d) s
0
=2.0 mg g
-1
). A Sphagnum peat sample
with density of 0.02 gcm
-3
and porosity 0.98 is saturated at M/V=20gL
-1
and
31
oversaturated when M/V is below 20gL
-1
. Lines 20 days and 60 days in subplot
(d) coincide.
32
Table 1. Model variables: symbol definitions and units
Symbol Definition Dimension Units
c DOC solution concentration M L
-3
g cm
-3
(eq.1,1a);
mg L
-1
(eq. 3-6)
s Sorbed soluble organic carbon
(SSOC) concentration
M
0
g g
-1
(eq2, 2a);
mg g
-1
(eq.3-6)
a
c
Amount of DOC in the solution M mg
a
s
Amount of soluble organic carbon
sorbed
M mg
u
Volumetric water content L
0
-

Peat bulk density M L

-3
g cm
-3
q{q
x
, q
y
,q
z
}
Water flux (Darcian velocity) L T
-1
cm h
-1
D{D
x
, D
y
,D
z
}
Dispersion coefficient L
2
T
-1
cm
2
h
-1
M Mass of peat M g
V Volume of water L
3
L
q
s
Specific discharge LT
-1
cm h
-1
33
Table 2. Model parameters. The list includes all model parameters, but only the
parameters relevant to the model application for laboratory conditions are specified.
The other parameters are specified in Part II (Table 1) where the model is applied to
field conditions.
Parameter Symbol Estimated/ taken from Units Value/ range
Sorption
Desorption kinetic constant t
des
Rasse et al., in revision
Surface peat from the
mire in Norway
h
-1
0.078
Partitioning coefficient
describing the sorption
equilibrium
K
D
Estimated from eq. 6 L g
-1
0.033
Initial DOC concentration in
water
c
0
Measured mg L
-1
Tap water: 0.3
Stream water: 44
Initial SSOC concentration s
0
Estimated from eq. 6 mg g
-1
2.0
Microbial
The rate of microbial DOC
production at 20C
P
basal
Estimated from least
square optimization
of eq. 5
mggh
-1
0.4-2.410
-3
The rate of the microbial
mineralization of DOC
at 20C

1basal
Estimated from
- water incubation
- least square
optimization of eq. 5
h
-1
0.410
-4
-1.010
-4
The constant reducing the
microbial mineralization
rate when the soluble OC
is sorbed
k
s
Kalbitz et al. [2005]
organic horizon of a
podzol soil
- 1/6-1/3
Modifier to account for the
effect of temperature
on P and
1
Q
10
Not used here: constant
temperature 20
0
C
- -
Modifier to account for the
effect of oxygen supply on P
k
anP
Not used here: aerobic
conditions only
- -
Modifier to account for the
effect of oxygen supply
on
1
k
an
Not used here: aerobic
conditions only
- -
Hydrological
Molecular diffusion
coefficient
D
0
Not used here: assumed
that the sample is
homogeneous
cm
2
h
-1
-
Dispersivity Not used here: assumed
that the sample is
homogeneous
cm -
34
Table 3. The rates of DOC production, P
basal
, and mineralization,
1basal,
derived from
the long-term laboratory peat incubations at approximately 20C.
P
basal
, Sample/location Peat-to-
water
ratio,
gL
-1
Net DOC
release
measured,
10
-2
mgg
-1
d
-1
10
-2
mgg
-1
d
-1
mgg
-1
h
-1

1basal
,
h
-1
Source and
comments
Surface
Sphagnum peat /
small headwater
mire
10-14 1.2-2.3 1.8-5.8 7.510
-4
-
2.410
-3
a
-
1.010
-4
This study
(c
0
=0.3 mg L
-1
)
Surface
Sphagnum peat /
small headwater
mire
14 0.7 1.0-1.7 4.210
-4
-
7.110
-4
a
- This study
(c
0
=44 mg L
-1
)
Fibric
Sphagnum peat /
bog
10-100
b
4.8 7.4-43.3 3.110
-4
-
1.810
-2
a
- Moore and Dalva,
[2001]
Sapric
Sphagnum peat /
bog
10-100
b
3.0 5.0-26.4 2.110
-3
-
1.110
-2
a
- Moore and Dalva
[2001]
Surface peaty
soil with 19%
OC/ river delta
140 5.3 18.2-56.2 7.610
-3
-
2.310
-2
a
- Aguilar and
Thibodeaux [2005]
Subsurface
fibrous peat /
river delta
3.310
3
-6.610
-4 c
1.410
-3
-
8.610
-3
3.310
-6
-
1.910
-5
1.910
-4
-
6.710
-4
Chow et al. [2006]
d
Subsurface
fibrous peat /
river delta
1.410
3
-7.010
-3 c
7.910
-3
-
4.610
-2
1.410
-6
-
6.710
-6
1.110
-3
-
3.710
-3
Chow et al. [2006]
d
Subsurface
fibrous peat /
river delta
500 -3.110
-2 c
3.410
-3
-
1.610
-2
4.610
-4
-
1.410
-3
Chow et al. [2006]
d
a
Not detectable by the parameter optimization
b
Presented as a range in the source publication [Moore and Dalva, 2001].
c
From day 7 to day 60
d
this study uses different techniques to determine the DOC release rate, so the data on
total soluble OC (SSOC+DOC) were used for the calculations here
35
Processes Sorption Microbial Hydrological
Time scale Short/hours Long/weeks-years Any, dependent on
flow rate
How
modelled
Linear kinetics Zero-order
production, first-
order
mineralization
A gradient of the the
DOC flux density
Parameters K
D
,,
des
P,
1
k
s
Q
10
, k
anP ,
k
an
D
0
,
MMWH model
Paper I. Implementation: numerical
or analytical solution of CDE
Paper II. Implementation: numerical
solution of CDE
W
a
t
e
r

f
l
u
x
T, water content
Figure 1. Schematic diagram of the model formulation and structure.
36
Solid OC
M
s
M>>s+c
Soluble OC sorbed
on solid OC
SOC(s)
Dissolved OC
DOC(c)
Sorption
-
des
K
D
c
Desorption

des
s
Production
P
Mineralization

1
k
s
s
Mineralization

1
c
Figure 2. Schematic diagram of the modeled pools of soluble organic matter
and processes that control microbially-mediated transformation (production and
mineralization) and abiotic exchanges (adsorption and desorption) between the
solid and dissolved phases. Parameters are described in detail in Table 1.
37
0.4 0.6 0.8 1
Water content,
z
q
z
c
q
x
c q
x
c
-Ddc/dz
dilution
advective
flux
dispersive
flux
Water table
Depth to the
catotelm
Figure 3. Schematic diagram of DOC transport within the peat profile,
based on the convection-dispersion framework.
38
0
0.1
0.2
0
0.1
0.2
0 24 48 72 96

0
2
4
6
8
Incubation time, h
K
D
t
des
P

1
net production
net sorption
a
b
c
0
=0 mgL
-1
Figure 4. (a) Modeled estimates of partitioning of the DOC release rate observed
in laboratory peat incubations into net sorption (desorption minus adsorption) and
net production (microbial production minus mineralization) components in
incubations with two starting DOC concentrations (c
0
) in water. (b) Normalized
first-order local sensitivity coefficients computed by the forward direct method
for the main parameters in the model applied to laboratory incubations (calculated
for an experiment with c
0
=0.0 mgL
-1
).
c
0
=44 mgL
-1
p
i
d
c
/
d
p
i
,

m
g
L
-
1
h
-
1
D
O
C

r
e
l
e
a
s
e
,

m
g
g
-
1
h
-
1
39
0
20
40
60
80
0 5 10
0.0
0.5
1.0
1.5
2.0
2.5
0 50 100 150 200
after 20 days
after 60 days
after 5 years
after 20 days
after 60 days
after 5 years
a b
c
d
Figure 5. Simulated effect of the initial soluble solid organic carbon (SSOC)
concentration, and peat-to-water ratio (M/V) on the first steady state DOC
concentration (a, b); and on the DOC release rate during the second phase
of a long-term peat incubation (c, d) (K
D
=0.033 Lg
-1
, c
0
=0.0 mgL
-1
,
P
basal
=1.0
.
10
-3
mgg
-1
h
-1
,
1basal
=4
.
10
-5
h
-1
, k
s
=0.3 ; (a, c) M/V=10.0 gL
-1
;
(b, d) s
0
=2.0 mgg
-1
). A Sphagnum peat sample with density of 0.02 gcm
-3
and porosity 0.98 is saturated at M/V=20gL
-1
and oversaturated when M/V is
below. Lines after 20 days and after 60 days in subplot (d) coincide.
Initial SSOC concentration, s
0
, mgg
-1
M/V, gL
-1
D
O
C

r
e
l
e
a
s
e
,

1
0
-
2
m
g
g
-
1
d
a
y
-
1
F
i
r
s
t

s
t
e
a
d
y

s
t
a
t
e

[
D
O
C
]
,

c
1
,

m
g
L
-
1
40
I
IV
I
Submitted to Water Resources Research
_____________________________________________________________________
A model of dissolved organic carbon concentrations in peat and stream waters.
II: Application and sensitivity analysis
Alla Yurova
1
, Andrey Sirin
2
, Kevin Bishop
3
and Hjalmar Laudon
4
1
Department of Physical Geography and Ecosystems Analysis, Lund University, Lund, Sweden
2
Laboratory of Peatland Forestry and Hydrology, Institute of Forest Science RAS, Uspennskoye,
Russia
3
Department of Environmental Assessment, Swedish University of Agricultural Sciences, Uppsala,
Sweden
4
Department of Ecology and Environmental Sciences, Ume University, Ume, Sweden
_____________________________________________________________________
Abstract:
A new model of DOC concentrations and fluxes in peatland water was introduced in
Part I of this study. Here, in Part II, the model is used to simulate DOC concentrations
in the outlet of the steam draining a small headwater mire in Northern Sweden during
the period 19932001. A relatively good model fit (MBE=-0.5-2.5 mgL
-1
, Willmott
index of agreement d>0.7) was found for all the categories of stream discharge,
except periods with very low flow (q<0.3 mm day
-1
). Heterogeneity within the
vertical peat profile simulated by the model was sufficiently strong to introduce
significant fluxes of DOC due to sorption and dispersion. Previous models of DOC
rely on bucket representations of the water movement with only a few vertical layers,
so it is worth examining whether such simplifications could result in the loss of some
episodic variations in DOC concentrations. When seeking explanations for the
interannual variability in DOC concentrations we, like previous authors, could find
the influence of temperature, flow path and intensity. However, the model has helped
to demonstrate that the system may have some memory: the store of sorbed soluble
organic carbon (SSOC) in one year affects the DOC concentrations and fluxes in the
following year.
_____________________________________________________________________
Keywords: dissolved organic carbon, peatland, model, uncertainty
* Author for correspondence (alla.yurova@nateko.lu.se)
Introduction
Peatlands are, collectively, one of the major sources of dissolved organic carbon
(DOC) in boreal watercourses [Urban, 1988; Dillon and Molot, 1997; Hope et al.,
1997; Gergel et al., 1999; Laudon, 2004; gren et al., 2007]. Changes in DOC
outputs from peatlands are of interest because (inter alia) they affect both regional
carbon balances [e.g., Jonsson et al., 2007] and aquatic biota [e.g., Laudon et al.,
2005; Steinberg et al., 2006]. Recent analyses have also emphasized the importance of
including DOC fluxes when resolving the total C balance of a peatland [Roulet et al.,
2007; Sagerfors et al., 2007]. Not only average DOC fluxes are significant, but also
interannual variation is comparable to that of vertical net carbon exchange [Roulet et
al., 2007]. Models are needed to test hypotheses regarding the causes of the seasonal
and interannual variability that has been recorded, and to predict DOC fluxes from
peatland catchments under different climate change scenarios and other human
impacts [e.g. Futter et. al, 2007].
Given the strong variations of DOC concentrations from peatlands, modeling them is
challenging. It was found that DOC is negatively correlated with water flow through a
system for a number of sites within the boreal region [Laudon et al., 2004; Fraser et
al., 2001]. The relationship is, however, demonstrable only if data from the major
hydrological event of the year, the spring flood, are included in the calculations
[Fraser et al., 2001]. Across the range of flow intensities the DOC concentration and
its relationship to flow can vary both seasonally and annually. The role of particular
environmental factors, including temperature and precipitation, in determining the
variability of the DOC concentration and fluxes is still open to debate [e.g., Freeman
et al., 2001a; Tranvik and Jansson, 2002; Worrall et al., 2004; Erlandsson et al., in
2
review]. It is widely appreciated that both in situ microbial DOC transformation
(production and mineralization) and hydrological transport are important influences
on DOC concentrations [e.g., Tranvik and Jansson, 2002], but the relative
contribution of each factor remains largely unknown. Experimental evidence supports
the view that the sorption equilibrium, based only on physico-chemical properties, is
likely to be as important for the export of the DOC from the wetland as biotic and
hydrological processes [Qualls and Richardson, 2003].
In Part I of this study we introduced a novel (to our knowledge) application of the
convectiondispersion equation to model DOC concentration and transport in a
peatland. The main processes included in the proposed model were: microbially
mediated DOC production and mineralization, hydrological transport and adsorptive
exchanges between two components of the system, namely the solution DOC and
sorbed soluble organic carbon (SSOC) of the peat matrix. In Part I we also discussed
the importance of sorption as a major process regulating the DOC concentration in
laboratory incubations. It was shown that kinetics equations describe the behavior of
such systems during the first 1-2 days of incubation better than steady state
approximations. An examination of the sorption kinetics associated with the water
flow in a vertically heterogeneous peat profile was left for Part II, this paper. This is a
key study to determine whether the model is valid for field situations. Here we apply
the convectiondispersion equation, at a relatively fine vertical resolution, to the
processes within the acrotelm. The model output should describe a degree of
heterogeneity that current DOC models do not consider, since they rely on bucket
[Neff and Asner, 2001; Michalzik et al., 2003] or semi-distributed [Boyer et al., 1996;
Futter et. al., 2007] hydrological approaches, incorporating very few vertical layers.
3
We tested the proposed model using long-term discharge and DOC concentration data
from the stream draining a small headwater mire, Kallkllsmyren, in Northern
Sweden [Sirin et al., 1998] collected during the period 19932001. The main
objective of Part II was to assess the applicability of the proposed model and
determine the main driving parameters by means of sensitivity and uncertainty
analyses. We addressed three specific questions when studying the mire:
(i) What are the components (production, storage and output) in the annual DOC
balance of the mire for the period 19932001?
(ii) How important is sorbed soluble organic carbon as a regulator of DOC
concentration over different time scales?
(iii) How significant is the vertical heterogeneity and exchange between the different
peat layers?
Methods
Study site
The Kallkllsmyren mire is situated in the Vindelns Forest Research Park, 60 km
northwest of Ume, Northern Sweden (6414'N, 1946'E). The mire catchment is one
of the headwaters in the 68 km Krycklan catchment [e.g., Buffam et al., 2007]. It is
situated upstream of the Nynget research catchment, which has been monitored since
1980 [Grip et al., 1990]. The site is located in the boreal zone with a cold temperate
humid climate. The mean annual temperature for the period 19812001, measured at
the nearby Svartberget Forest Research Station, was 1.5
o
C [Ottosson-Lfvenius et al.,
2003]. The average annual precipitation for the same period was 660 mm, with a
4
runoff of around 330 mm. About 45% of the precipitation falls as snow, and up to
60% of the annual runoff from the studied catchment can be snowmelt [Bishop et al.,
1995].
Kallkllsmyren is a complex mire occupying a topographic depression. The mire is 2
to 5 m deep and the underlying sediments are blocky glacial tills. On the eastern and
western sides the mire is surrounded by rather steep slopes (Figure 1). The slope to
the north is very gentle, and the southern part of the depression opens into the valley.
The slopes are afforested with spruce and pine and the soils are podzols. The mire is
drained by a single stream, which originates from the mire southern edge. A ditch was
dug for the stream in the 1920s, but the mire edge was left untouched.
The main section of the mire is raised, sloping down towards the edges; it has distinct
ombrotrophic features with vegetation composed of a sparse Scots pine stand with
dwarf-shrubs and Sphagnum mosses, together with Carex at the edges (Figure 1).
Mineratrophic fen vegetation occupies the southern area of the mire adjacent to the
stream, and there are also two belts of this vegetation situated on the eastern and
western sides of the raised section (Figure 1). Surface peat in both sections of the mire
is formed from Sphagnum remains with some Carex in the mineratrophic parts. The
degree of decomposition of the surface peat is less than 27% in the main section and
about 5% at the downslope edge.
The hydrological response at the outlet of the mire is very rapid, with peaks delayed
by no more than a few hours from the maximum precipitation or snowmelt inputs.
Isotope hydrograph separation has shown that 70-90% of the runoff during rain-driven
5
events is pre-event water [Bishop, 1991], while during spring floods, the pre-event
water component is 5070%, including 515% derived from the deep ground water
[Laudon et al., 2007]. Sirin et al. [1998] and Petrone et al. [2007] provide more
detailed descriptions of the Kallkllsmyren mire.
The measurements
The stream water samples and discharge measurements were obtained from the stream
draining the mire about 50 m below its origin. The catchment area draining into the
sampling point is 18 ha, of which 9ha is occupied by the mire. During snow-free
periods of the year, discharge was computed hourly, based on water level
measurements made using a pressure transducer. Rating curves were derived using
timed measurements of discharge volume over the weir. Water sampling was
conducted weekly to biweekly during periods with low flow and up to daily during
snowmelt and a number of rainfall episodes since 1993. Samples were collected
monthly during 1991-1993 in acid-washed, 250 ml HDPE bottles and frozen until
analyzed. TOC was measured with a Shimadzu TOC-5000 total organic compounds
analyzer using catalytic combustion. As shown by Ivarsson and Jansson [1994] and
Gadmar et al. [2002] the particulate fraction of TOC is negligible in the type of water
studied here. Thus, while TOC was measured and is reported here, the results actually
reflect the DOC. The site has been the subject of intensive fieldwork [Bishop, et al.,
1990; Bishop and Pettersson, 1996; Laudon et al., 1999; Hruska et al., 2001] and
additional parameters, such as O
18
isotopes and major ion concentrations have been
periodically measured in water samples collected at the stream origin as well as from
piezometers installed at various depths in the mire [Sirin et al., 1998].
6
To calculate annual DOC fluxes from measured DOC concentrations we used daily
average values of stream discharge and linearly interpolated the DOC concentrations.
The model
The main model has three key components: Heat flux, Hydrology and DOC Mass
Balance (Figure 2). We use the Mixed Mire Water and Heat model [MMWH;
Granberg et al., 1999] as modified by Yurova et al. [2007] to describe water fluxes,
vertical distribution of water content in the acrotelm, snow dynamics and heat transfer
in the peat profile. The system of two mass balance equations, one for the sorbed and
one for the dissolved OC(the latter in the form of a convectiondispersion equation),
is solved to estimate DOC concentration in the pore water and the flux entering the
stream, as described in Part I of this paper. Numerical values of the model parameters
were measured or calibrated (Table 1).
The model was developed in C++ code (Microsoft Visual C++) and the numerical
solution of the system of DOC and SSOC mass balance equations was calculated
using the CVODE program package [Cohen and Hindmarsh, 1996; program codes
and documentation available at http://www.llnl.gov/casc/sundials/]. The vertical
resolution of the model is 5 cm in the acrotelm and 10 cm in the catotelm. The time
steps within the model are hourly, and the results were analyzed using daily
averaging. Below are important details for the particular application of the model.
Hydrology and heat transfer
The MMWH model was designed to simulate the flow and moisture dynamics in the
upper, active, intermittently aerated zone of a peatland, the acrotelm. A zero flux
7
(impermeable) lower boundary is placed at the level of constant saturation, and it is
assumed that no flow occurs below this, i.e. in the catotelm [see Reeve et al., 2000 for
an examination of the validity of this assumption]. The model is based on steady-state
vertical moisture distribution curves [Romanov, 1961], which, as we discuss below,
was found to be applicable to the parameter values and time step used in this study.
The model considers variable lateral flow at different levels, to account for reduction
in hydraulic conductivity with depth [e.g., Ivanov, 1981]. The MMWH model was
first developed for the hydrology of an area with prescribed geometry and a single
vegetation type. Here it was applied to the whole mire and model parameters
characterizing the geometry, hydraulic conductivity and surface roughness of the mire
were combined into the calibrated parameters. The vertical peat profile was
discreetized into 5 cm deep layers, to partition the lateral flow. Water content was
integrated within each layer. It was assumed that vertical drainage is not restricted by
ice, but when the ice content of a layer is exceeds 0.7 lateral flow ceases.
Model equations and parameters are given in Granberg et al. [1999] and recent
changes are documented in Yurova et al. [2007]. Here, the model was applied to a
number of layers, rather than the whole active layer. Hence, layer-specific, rather than
integral hydraulic conductivity is used. The general form of the equation used to
calculate the specific discharge then becomes:
)
=
z
zcat
z
dz k q o , [1]
where z (cm) is depth to the water table (negative when below the peat surface), z
cat
(cm) is the acrotelm depth, o (cm
-1
) is a lumped parameter and the hydraulic
8
conductivity, k
z
(cmh
-1
), is approximated as an exponential function of the depth
(k
z
=exp(|z), where | is a lumped parameter based on data presented in Ivanov [1981],
and the linear scaling coefficient before the exponent is included in o).
Evapotranspiration is assumed to decrease exponentially with depth (E=E
PT
exp(l
E
z),
where E
PT
(cmday
-1
) is potential evapotranspiration and l
E
is a lumped parameter.
During rain or snowmelt events it was assumed that drainage is vertical and that the
vertical water flux, q
v
, equals precipitation intensity, corrected for the change in water
content due to a rise in the water table. In addition, a generalized form of the Manning
equation [e.g., Beven, 2001] was applied here to describe the overland flow:
67 . 1
s SO
h q o = , [2]
where q
S0
(cm h
-1
) is specific (area) discharge via surface overland flow, h (cm) is
water table depth above the peat surface and o
s
is a lumped parameter.
The Heat balance equation was solved in a slightly different way than in the original
MMWH, mainly for practical reasons. We used a solution presented by Wania et al.
[in preparation] and implemented it using a C++ program. One important difference
from the formulation of Granberg et al. [1999] is that the snow layer and meltwater
pooled on the mire surface are included as additional layers in the vertical profile
when simulating the heat balance.
9
DOC concentration
To simulate DOC and sorbed soluble organic carbon (SSOC) concentrations in the
acrotelm, we applied equations (1a)(2a), presented in Part I. The initial SSOC
concentration in the acrotelm peat was determined utilizing data from laboratory
incubations of surface peat from the studied mire, as described in Part I. For the
catotelm a more simple approach was taken. We assumed that a very slow production
rate [e.g., Freeman, 2001b] does not change the total DOC amount significantly on
the decadal time scale that we are studying here. We, therefore, assumed a constant
SSOC concentration in equilibrium with the dissolved phase, affected only by
diffusive flux to and from the acrotelm. Data from the sampling wells in the mire
were used to estimate SSOC concentration in the catotelm. A few model parameters
relevant to the field application are discussed in some detail below, the rest of the
parameters remained unchanged from those derived from the laboratory experiments
or the literature, as described in Part I of this study.
In our model we use a single value of Q
10
[Moore and Dalva, 2001] as a modifier of
the model parameters P
basal
,
1basal
and
2basal
(
2
is included in the model as

2
=
1
k
s
). This approach should only be regarded as a first approximation. Chow et al.
[2006] demonstrated that Q
10
is different for the mineralization of soil organic matter
(thought to be one step in DOC production) and mineralization of DOC. Similarly, it
is likely that Q
10
also differs for the mineralization of DOC and SSOC. Unfortunately
no published data could be found that specifies Q
10
for DOC production, DOC
mineralization or SSOC mineralization.
10
Two important model parameters are the ratios of anaerobic to aerobic DOC
production rate (k
anP
) and of anaerobic to aerobic mineralization rate (k
an
). Anaerobic
to aerobic mineralization rate ratio may also differ between the organic matter, which
is a source of the DOC, and the DOC itself. Some studies have indicated that the net
DOC production rate is the same in oxic and anoxic conditions [Moore and Dalva,
2001], while others have demonstrated the effect of anoxia on DOC production and
mineralization [e.g., Freeman et al., 2001b]. In the absence of validated data from the
site, with which to quantify the parameters, we defined them based on the range of
values available within the literature. The upper limit was set at 1.0 (no effect of
anoxia) and the lower limit at 0.07 [Bergman, 1998; Moore and Dalva, 1997:
decomposition of surface peat in anoxic vs. oxic conditions in laboratory experiments]
for both coefficients (k
anP
and k
an
).
In a current study, living vegetation is not modeled explicitly as a separate layer.
Instead DOC microbial production in the uppermost peat layer is increased by
multiplying the basal production rate, P, by f
Ps
. f
Ps
is a constant ratio between DOC
production from living tissues to that from peat. Is does not depend on the current
NPP, but is calculated assuming an average NPP (taken from eddy-tower estimates at
the near-by Deger mire [Yurova et al., 2007]) and the ratio of DOC release from
Sphagnum to that from peat presented in Moore and Dalva, [2001]. Here, we ignore
the release of easily decomposable substances, such as root exudates, that have a short
lifetime in solution [e. g., Crow and Wieder, 2005]. The model was formulated for the
bulk of relatively refractory DOC reaching the stream, and not for the labile fractions
with short turnover times. Fractionation of DOC in water samples from the mire
stream and peat extractions (laboratory study in Part I) using the UV spectral
11
deconvolution method [Hassouna et al., 2007] demonstrated that 50% of the DOC is
hydrophobic, 18% transic and 22% hydrophilic. The proportion of these fractions is
the same for the stream and the peat sample extracts [M. Hassouna, unpublished
data]. Therefore, approximately 10% of the DOC is non-humic; this corresponds well
with earlier estimates from the study site [Bishop and Pettersson, 1996]. As shown by
McKnight et al. [1985] the hydrophobic acids in Sphagnum bog waters are mainly
fulvic; the hydrophilic acids present are intermediate compounds in the formation of
fulvic acids. Both are known to be relatively recalcitrant.
When water floods over the saturated peat profile, there is DOC exchange between
the surface water and the pore water in the uppermost peat layer. This is described in
the model using a formulation similar to the dispersion term in eq. (1), Part I, but
with a specific dispersion coefficient, D
s
.
Modeling protocol
The model was run for the Kallkllsmyren mire site using a series of climate data for
temperature, precipitation and potential evapotranspiration recorded at the nearby
Svartberget research station [Ottosson-Lfvenius et al., 2003]. The parameters and
constants used are defined and listed in Table 1. The parameters o, |, o
s
and l
E
, used
in the hydrology submodel, were optimized by minimizing the mean squared
difference between the modeled and observed specific discharge. The only three
parameters that were adjusted in the DOC concentration submodel were: the
dispersion coefficient for the overland flow, D
s
, the ratio of anoxic to oxic DOC
production rate, k
an
., and the ratio of anoxic to oxic DOC mineralization rate, k
an
..
12
The following model simulations were conducted:
- Model initialization was made using climate data from 1986 to 1993, the start of
the actual simulation period. Initial SSOC concentrations were varied from 1 to 5
mg g
-1
in order to study the sensitivity of the system to initial conditions
- A model run for the whole simulation period (19932001)
- A series of Monte Carlo runs (1000) to determine the model sensitivity to
specific parameters and constants (Table 2).
The model results were analyzed on the basis of daily averages. Observed annual
DOC fluxes were estimated on the basis of interpolated DOC concentrations
measured at the stream outlet. To evaluate the annual DOC balance predicted by the
model, the total production and mineralization in the acrotelm were simulated
separately and the net production was estimated as the difference between the two
summed over a year. Annual net production in the acrotelm was then compared with
the stream export (flux) of DOC and the total amounts of SSOC and DOC stored in
the acrotelm as predicted by the model. In addition, the annual volume-weighted DOC
concentrations were calculated, based on the observed data and model simulations, to
eliminate the dominance of flow in the DOC fluxes. This reveals how the
concentrations vary between years. Volume-weighted concentration (both observed
and modeled) was then correlated to the modeled amounts of SSOC and DOC stored
in the acrotelm before the spring flood event, to determine whether the latter is related
to the former. Simulated concentrations weighted for different seasons were correlated
to the simulated stored amounts of SSOC and DOC in the preceding seasons. The
seasons were defined arbitrarily as: winter, the period when the surface temperature
13
of the peat was at or below 0C; spring, the period from the start of snow melt till
the leveling off of the spring flood flow rate; and summer as the rest of the year.
To test the applicability of the steady-state vertical moisture distribution curves, the
1D Richards equations were solved for the observed range of precipitation amounts
and water table depths. The code of Bolster and Raffensperger [1996] based on
Hornberger et al. [1998] was used. van Genuchten soil moisture functions [van
Genuchten, 1980] were simplified, as they are in the current version of the MMWH
that is based on parameterization [Weiss et al., 1998]. The range for the saturated
hydraulic conductivity of the surface peat was derived from the literature [Romanov,
1961; Letts et al., 2000; Reeve et al., 2000].
To investigate the dynamics of the stream DOC concentration during hydrological
episodes when DOC concentration was high, two rain-driven flow events were
chosen to characterize the system behavior. The episodes differed in the intensity of
the rain event. In both cases the stream DOC concentration increased with the first
peak in the discharge. After the second (larger) runoff peak, the concentration
stabilized at approximately the same level as prior to the first event (12 July30
August 1993, Figure 4a) and dropped about 10 mgL
-1
during the second event (27
August28 September 2001, Figure 4b). There was more rainfall during the second
event than the first. To illustrate the effect of vertical heterogeneity within the peat
profile during the second rainfall event, we partitioned the six layers within the
acrotelm with respect to: (i) net microbial production (mineralization) of DOC; (ii) net
release (removal) of DOC due to sorption; (iii) stream export; (iv) vertical advective
fluxes from (and to) adjacent layers; and (v) vertical diffusive fluxes. The sum of the
14
listed components for the whole of the 27 August28 September 2001 event was
compared within each layer and between the layers.
Evaluation of the model performance, uncertainty, and sensitivity
Following the recommendations of Willmott [1982], we calculated the following
quantitative measures to evaluate the model performance: the mean bias error (MBE),
the root mean square error (RMSE) and the Willmott index of agreement, d. The
Willmott index of agreement is a dimensionless measure that varies between 0 and 1,
with d =1 indicating a perfect match. It is calculated as follows:
, |) ' | | ' (| / ) ( 1
1 1
2 2
(

+ =
_ _
= =
N
i
N
i
i i i i
O M O M d [3]
where O M M
i i
= ' , O O O
i i
= ' , M-modeled and O-observed values.
In order to evaluate the model parameter-based uncertainty, we performed a global
multi-parameter sensitivity analysis [e.g., Saltelli, 2000]. Latin hypercube sampling
[LHS, McKay et al., 1979] was used to generate sets of parameters for use in the
Monte Carlo runs. Where possible, the parameter range was derived from the
literature (Table 2). Uncertainty and sensitivity analysis was performed on the
simulated DOC concentrations. Uncertainty bounds (lower and upper 10% quantiles)
were calculated based on the results of the Monte Carlo runs for each day during the
simulation period, following the GLUE methodology [Beven, 2001]. The Willmott
index of agreement was chosen as a likelihood measure. In the sensitivity analysis, the
15
rank partial correlation coefficient (RPCC) was chosen as a measure of the parameter
importance, indicating the extent to which a particular model parameter determines
the model output (range for the parameter vs. range in the output).
Results
General agreement between simulated and measured discharge, stream DOC
concentrations and fluxes: Error assessment
The simulated discharge corresponded well with the measurements at the mire outlet
(MBE=-0.11 mm, RMSE=1.5 mm d=0.85 mean=1.1 mm) and no systematic bias was
found for particular periods or discharge categories. The largest discrepancies (both
negative and positive) between the modeled and measured discharge occurred during
the spring flood.
Seven years after the start of the simulation (1986) the model performance was no
longer influenced by the initial conditions. The simulated values for DOC
concentration from the runs with different initial SSOC concentrations converged
after about five years.
We found relatively good agreement between the simulated and measured DOC
concentrations both overall (Figure 3) and for particular events (Figure 4). However,
the model performance varied greatly between the categories for total discharge
(Table 3). The model performed very poorly when the discharge was below 0.3 mm
day
-1
(Table 3 and cross symbols on Figure 3). Above this threshold, the model
performance increased sharply and the best fit between the modeled and measured
16
DOC concentrations was found for the highest discharge category (Table 3). As a
result of the high correlation between discharge and DOC fluxes, when averaged over
a year, the agreement between the measured (interpolated) and simulated annual DOC
fluxes was very high (d=0.95; MBE=1.1 gm
-2
year
-1
RMSE=2.0 gm
-2
year
-1
).
The parameter associated with DOC microbial production, k
anP
, achieved optimal
model performance when set to a value of 0.07. This is the lower limit of the
anaerobic-to-aerobic decomposition rates for organic matter in the surface peat [e.g.,
Bergman, 1998; Moore and Dalva, 1997]. In contrast, the model performed best when
k
an
, the parameter associated with DOC mineralization, was allocated a value of one,
i.e. no reduction in DOC mineralization rate due to anoxia.
Uncertainty and sensitivity analysis
Prediction bounds determined by the uncertainly analysis covered most of the
observed DOC concentrations. The most common exceptions were the very low
concentrations recorded during periods of very low flow, below 0.3 mmday
-1
(cross
symbols on Figure 3), which the model overestimated.
Sensitivity analysis (Table 2) revealed that the most influential model parameters are
the DOC production rate (P
basal
), Q
10
, and the parameter accounting for the decease in
the DOC production rate under anaerobic conditions (k
anP.
). Also important, but less
so, are the DOC mineralization rate (
1basal
) and the sorption constants (t
des
and K
D
).
The other parameters exerted less influence over the model performance.
17
Model study of the vertical peat profile of the acrotelm
Estimates produced using Richards equations (not shown) indicated that the steady-
state moisture profile within the unsaturated zone is always reached within an hour of
the end of a rainfall event, even during high-intensity events. This is because of the
relatively shallow unsaturated layer, but, more importantly, the high hydraulic
conductivity of the surface peat. The result is promising as it indicates that the
MMWH model based on steady-state moisture distribution curves is likely to be a
valid approximation for the time scale studied here.
Figure 5 illustrates the simulated fluxes that contribute to the change in DOC
concentration in pore water; the values for the individual layers are summed over the
duration of the event being investigated, i.e. 27 August28 September 2001 (Figure
4b). The figure does not show the entire summed DOC since it does not include
SSOC mineralization; it represents the total solute DOC concentration only. The first
interesting feature to note is a strong vertical heterogeneity between the 5cm layers.
Microbial DOC production decreases with depth under anoxic conditions and as the
temperature decreases. The stream export term also decreases, due to reduction in
hydraulic conductivity and therefore flow intensity with depth. The DOC is released
into the solution as a result of desorption in the three uppermost layers, but it is
removed from the solution by sorption in the layers below. Therefore, by sorption, the
layers more than 15 cm below the surface accumulate soluble OC carbon produced in
the layers above. Due to vertical advective transport, DOC is lost from the uppermost
layer and transferred to the layers below that were permanently saturated during the
events studied. Dispersion operates throughout the whole acrotelm and its input to the
18
total DOC balance is not insignificant. Another important feature to note from Figure
5 is that, for all the layers except the uppermost, the contribution of sorptive release or
removal of DOC is similar to or even exceeds the net microbial production.
Components of the annual DOC-SSOC balance
Two main simulated components of the annual DOCSSOC balance, net microbial
production within the acrotelm and stream export, are of a similar order of magnitude
but are not equal for the year under consideration; in fact, the difference between the
two can be large (Figure 6). Interannual variation in the volume-weighted DOC
concentration reflects the variation in the amounts of DOC and SSOC stored within
the acrotelm in the previous year (Figure 7a, the DOC and SSOC are the model
estimates at the start of the current hydrological year, and are also equal to those at
the end of the preceding year). The correlation between the volume-weighted stream
DOC concentration in any given season and the store of DOC and SSOC in the
preceding season is high for the winter (r=0.75, P=5.8 10
-5
) and summer (r=0.87,
P=1.910
-5
), but lower for the spring (r=0.40, P=0.023) (Figure 7b).
Discussion
The relatively good overall results of the simulations indicate that the model is able to
describe the main mechanisms responsible for the variability in DOC concentration.
Very generally, concentrations rise to their highest levels (e.g. the episode in
September 1996 on Figure 3) when conditions favorable for DOC production (good
aeration and suitable temperatures) are combined with and/or shift the flow depth to
the layers that were not previously exposed to water flow. Some precipitation is,
therefore, always needed to mobilize the DOC present in the unsaturated layer and to
19
shift the sorption equilibrium by adding low DOC water so that additional amounts of
SSOC are released into the water being discharged. According to the model, this
explains the correspondence in timing of the initial increase in stream DOC
concentration and the increase in stream discharge following precipitation events
(e.g., Figure 4 a, b). Strong and prolonged events may, however, reduce the DOC
concentrations (e.g., the 27 August28 September event shown in Figure 4b) through
the removal of significant amounts of SSOC from a particular layer. After the flow
ceases, there is a build up of new DOC in the pore water; simultaneously, SSOC may
accumulate on the peat matrix as a result of both in situ production and dispersive and
advective exchange with the adjacent layers. The model shows that the vertical peat
profile is strongly heterogeneous (Figure 5) and the role of dispersion is important
down to the bottom of the acrotelm. Sorption may release additional DOC in some
layers and remove it in others (Figure 5), depending on the sorption equilibrium.
Sorption makes a large contribution to DOC in a particular layer similar to that of
microbial production (Figure 5). The lowest DOC concentrations were observed in the
stream during the spring flood (Figure 3). The flow is so intensive during this period
that significant amounts of SSOC are removed from the acrotelm while the
concentrations remain low. This could also explain the relatively slow recovery of
concentrations after the spring flood (up to one month).
No deep flow mechanism is considered in the current model, which describes only the
water balance in the acrotelm. We hypothesize that this is the main reason for the poor
model performance during periods of very low flow (Table 3 and cross symbols in
Figure 3). The water discharged during such periods appears to be groundwater from
the podzols surrounding the mire, which has bypassed the peat soils of the mire. In
20
terms of the total DOC budget, the ground water contribution is rather low, since the
flow itself is low; this input is, however, persistent. During periods of low flow, when
flow out of the mire virtually ceases, the ground water bypass can strongly influence
the DOC concentrations at the outlet. This bypass may be related to the
hydrochemical anomaly observed by Sirin et al. [1998] at a depth of 2 m in the
catotelm.
Results of the sensitivity study show that only a few parameters strongly influence the
model output. They are the DOC production rate and its modifiers that account for the
effects of temperature and anoxic conditions (Table 2). In addition, the model
predictions are also largely dependent on the chosen values of DOC mineralization
rate and the two sorption constants. These parameters require further investigation to
improve model generalization.
The balance between the net production of DOC and SSOC and the DOC
hydrological export, as predicted by the model (Figure 6), may go some way to
explaining the interannual variability in the total amounts of SSOC and DOC stored in
the acrotelm. Although the amount of stored SSOC and DOC in the acrotelm is
several times larger than the annual export, the variation in the amount stored as a
result of the dominance of either production or export in a given year is significant
and may persist in subsequent years. The system therefore has a memory, as
demonstrated by the good correspondence between the simulated storage and the
observed annual volume-weighted stream DOC concentration (Figure 7a). An
example of system memory has been described by Worrall et al. [2006] discussing
the role of drought in a mire in the UK. Unlike the example in Worrall et al. [2006],
21
the mire we studied exhibits a link between the summer concentrations and the
intensity and duration of the spring flood event, as shown by the very high correlation
between the storage of DOC and SSOC after the spring flood and the volume-
weighted summer stream DOC concentration [Figure 7b: summer]. The opposite is
also true: winter concentrations may be determined by the changes in SSOC amounts
during the previous summer. The situation during the spring flood is more
complicated and the amount of DOC and SSOC stored in the acrotelm before the
event is not such a large determinant of the stream DOC concentration (Figure 7b:
spring).
Net microbially mediated DOC production is influenced first by temperature, but also
by the proportion of the acrotelm in which aerobic conditions favor the exoenzymatic
hydrolysis of organic matter, a critical step in DOC formation. The model is
parameterized so that the DOC production rate is higher in aerobic than in anaerobic
conditions, but DOC mineralization is not. That resulted purely from the model
calibration. We can, however, speculate that the increased DOC production rate in
aerobic (vs. anaerobic) conditions is associated with enrichment of the solution with
phenolic compounds that may impair DOC mineralization [Freeman et al., 2001b].
The suppressed DOC production leads to the low net production simulated by the
model in the wet years (e.g., 1993, Figure 6). At this stage we cannot justify how valid
the assumption is, but it certainly deserves further attention.
Conclusions
In Part I of this study we described the model of DOC concentration and export from
a peatland. We highlighted the importance of modeling sorptive exchange as a
22
dynamic process, rather than a simple steady state, when considering data from
laboratory peat incubations. We also demonstrated how some abiotic factors may,
through the action of sorption, strongly affect the rate of DOC release into water, as
observed during long-term laboratory incubations.
In this paper the DOC concentrations recorded during 19932001 in an outlet of the
steam draining a small headwater mire in Northern Sweden were used to validate the
model. Relatively good agreement between the measured and simulated DOC
concentrations was found both for the whole simulation period and for particular
events (Figures 3 and 4). The results of this study indicate the following:
- Observed concentrations at the outlet of the stream draining the studied mire
when the flow of intensity exceeds 0.3 mm may, largely, be modeled by the
combination of production, sorption and transport processes within the layered
acrotelm, i.e. by the convectiondispersion equation.
- It is important to include vertical heterogeneity in the model in order to
adequately describe the processes that are either strongly influenced by depth
(production, horizontal transport) or driven by the vertical heterogeneity
(sorption, dispersion).
- Sorption is an important process in determining DOC concentration and fluxes. It
may cause a release of DOC into the pore water in some layers and removal in
others, that is similar to the amounts of microbially produced DOC. During each
event, part of the DOC in the stream export originates from that which was stored
in the layer prior to the event, part is released into the solution from the peat
matrix due to desorption and the remainder is newly produced.
23
- Interannual variability in the SSOC and DOC stores within the acrotelm depends
on the conditions for microbially mediated DOC production and mineralization
during the current year (temperature, aeration) and flow intensity. Some
memory is, however, characteristic of the system, meaning that the store in one
year affects the concentrations of fluxes in the following year. DOC export in the
studied mire is very closely linked to the total annual discharge
Though more detailed formulation of the subsurface flow and the ground water flow
may be needed to account fully for DOC transport within the mire, we contend that
even the relatively simple MMWH model can serve well, when the water movement
in the acrotelm dominates the total discharge. We also hope that the current model
formulation will be suitable for ecosystem models designed to examine the C balance
of peatlands.
Acknowledgements
This work has been supported by the NECC project funded by the Nordic Center of
Excellence (NCoE) Pilot Programme and by the Swedish Research Council for
Environment, Agricultural Sciences and Spatial Planning. Ishi Buffam is
acknowledged for preparing the map used in Figure 1.
24
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10.1029/2006JG000342.
28
Figure captions:
Figure 1. (a) Map of Sweden with study site location. (b) Upper portion of the
Nynget drainage basin at Svartberget near Vindeln, Vsterbotten in Northern
Sweden, showing the location of the Kallkllsmyren mire and the measurement site at
the stream origin. (c) Vegetation map of the Kallkllsmyren mire.
Figure 2. Schematic diagram of the main model components
Figure 3. Simulated (line) vs. measured (symbols) DOC concentrations at the outlet of
Kallkllsmyren mire. () DOC measurements corresponding to stream discharge
values less than 0.3 mm, (+) DOC measurements corresponding to stream discharge
value greater than 0.3 mm. The gray area indicates the upper and lower 10%
prediction bounds obtained from 1000 Monte-Carlo runs processed using the GLUE
methodology. Panel above: measured stream discharge plotted with y axis inverted
(i.e. with peak flows pointing downwards).
Figure 4. Simulated and measured DOC concentrations for two episodes: (a) 12 July-
30 August, 1993, and (b) 27 August-28 September, 2001. Upper panels illustrate the
measured total stream discharges plotted with y axis inverted (i.e. with peak flows
pointing downwards).
Figure 5. Modeled estimates of partitioning between the processes contributing to
changes in pore water DOC concentrations during the event from 27 August to 28
29
September 2001 in each of the six vertical peat layers of the acrotelm. Layer 1 is the
uppermost layer.
Figure 6. Simulated components of the annual DOC-SSOC balance: microbially
mediated net production and DOC stream export. Changes from year to year in the
amounts of SSOC stored in the acrotelm are also shown (estimated for the dates
before the first autumn frost).
Figure 7. (a) Interannual variations in the amounts of SSOC and DOC stored before
the start of the hydrological year (spring flood) simulated by the model. Interannual
variation in the volume-weighted annual stream DOC concentration, simulated by the
model and calculated from the interpolated observations. (b) Correlation between the
amounts of SSOC and DOC stored in the preceding season (winter and spring,
respectively), and the volume-weighted seasonal stream DOC concentrations in spring
and summer simulated by the model.
30
Table 1. Model parameters and constants
Function Symbol Value Units Description Source
Hydrology o 0.045 cm
-1
lumped discharge parameter,
eq. 1
Calibrated
| 0.20 cm
-1
lumped transmitivity parameter Calibrated from the data
range in Ivanov [1981]
o
s
0.11 cm
-0.67
h
-1
lumped surface discharge
parameter, eq.2
Calibrated
l
E
0.03 - lumped evapotranspiration
parameter
Calibrated from the data
range in Virta [1966],
Romanov [1961]
| 0.98
0.92;
- porosity: acrotelm; catotelm By analogy, e.g.
Granberg et al [1999]
z
cat
-30 cm acrotelm depth By analogy, as average
sph 90 % proportion of Sphagnum remains
in the peat
Measured
ca 5 % proportion of Carex remains in
the peat
Measured
0.02;
0.06
g cm
-3
peat density, acrotelm; catotelm By analogy, e.g.
Nungesser [2003]
DOC
dynamics
K
D
0.033 Lg
-1
sorption partitioning constant Part I, Table 2
t
des
0.078 h
-1
desorption kinetic constant Rasse et al., in revision:
surface peat from the mire
in Norway
P
basal 1.410
-3
mgg
-1
h
-1
microbial DOC production rate
in the acrotelm at 20
o
C
Part I, average Table 2
f
Ps
1.8 - correction for the DOC
production by vegetation
Moore and Dalva [2001],
comp. Sphagnum and
fibric peat

1basal
0.410
-4
h
-1
DOC and SSOC mineralization
rate in the acrotelm at 20
o
C
Part I, water incubation
Table 2
k
s
1/6 - The constant reducing the
microbial mineralization rate
when the soluble OC is sorbed
Kalbitz et al. [2005]:
podzol organic horizon
Q
10
1.7 - Modifier to account for the
effect of temperature on
P and
1
Moore and Dalva [2001]:
fibric and sapric peat
k
anP
1 - ratio of anaerobic to aerobic
DOC production
Calibrated from the rage
in Moore and Dalva
[1997]
k
an
0.07 - ratio of anaerobic to aerobic
DOC mineralization
Calibrated from the rage
in Moore and Dalva
[1997]
D
0
4.3 10
-2
cm
2
h
-1
molecular diffusion coefficient
Karlstrm [1995]
10 cm dispersivity Reeve et al. [2001]
D
s
0.19 cm
2
h
-1
surface dispersion coefficient Calibrated
31
Table 2. Range of parameters and constants used in the sensitivity analysis.
Parameter range* Parameter RPCC
min max
Source and comments
P
basal
0.84 0.410
-3
2.410
-3
Table 2 Part I
Q
10
-0.78 1.6 4 Chow et al. [2006],
Moore and Dalva
[1997]
k
anP
0.64 0.07 1 Bergman [1998};
Moore and Dalva
[1997]

1basal

-0.54 0.410
-4
1.010
-4
Table 2 Part I
t
des

0.53 -50% 50% -
K
D
-0.51 0.019 0.091 Qualls [2000], Rasse et
al., in revision
k
an
-0.33 0.07 1 Moore and Dalva
[1997]
|
0.21 12 34 From optimization
k
s
-0.19 1/6 1/3 Kalbitz et al. [2005]:
l
E
0.13 0.01 0.03 From optimization
o
o

0.09 0.09 0.6 From optimization
o
-0.07 0.0113 0.0693 From optimization
D
0
-0.06 1.2 10
-2
7.2 10
-2
Karlstrm [1995]
D
s
0.05 0.01 1 -

-0.04 0.2 100 Reeve et al. [2001],
Ours et al. [1997]
f
Ps
0.01 1 3 -
* units as in the Table 1
32
Table 3. Quantitative measures of model performance with respect to DOC
concentrations. MBE, mean bias error; RMSE, root mean square error; d, Willmott
index of agreement (d=0, no agreement; d=1, perfect match) calculated for the whole
simulation period and for particular discharge categories.
Daily discharge
interval (N=657),
mm
MBE RMSE d
0-0.15 12.8 16.3 -0.65
0.16-0.3 3.8 11.9 0.32
0.3-0.49 2.5 7.4 0.70
0.5-1.4 1.1 8.9 0.74
1.5-23 -0.5 7.2 0.89
All 3.4 10.6 0.68
33
Figure 1. (a) Map of Sweden with study site location. (b) Upper portion of the
Nynget drainage basin at Svartberget near Vindeln, Vsterbotten in Northern
Sweden, showing the location of the Kallkllsmyren mire and the measurement
site at the stream origin. (c) Vegetation map of the Kallkllsmyren mire.
21 E
69 N
13 E
55 N
a
1 2 3 4
5 6 7 8
100m
Vegetation types:
1. Pine (H 7-8m), shrub, sphagnum
2. Pine (H 3-4m), shrub, cotton grass, sphagnum
3. Cotton grass, sphagnum, some pines
4. Pine and spruce (H 4-5m), sedge in the hollows,
shrub and sphagnum in the hummocks
5. Cotton grass and sedge
6. Sedge, swampy vegetation
7. Hollows with open peat and water
8. Cane (reed)-sedge, swampy vegetation
c
Catchment boundary
Stream
Stream sampling site
Topo lines
Peat mire
Lake
100m
b
300
290
280
34
...........................
Snow dynamics
Soil temperature
Freezing and
thawing
.............................
Evapotranspiration
Runoff in saturated
zone
and overland flow
Water content in
unsaturated zone
................................
Convection and
dispersion
Sorption/desorption
Metabolic production
and degradation
Mixing in stream
water
Heat transfer Hydrology
DOC mass balance
air temperature, precipitation, potential evapotranspiration,
peat type, composition and physical properties, acrotelm depth
runoff,
DOC concentration in peat pore and stream water,
DOC flux out of the mire
Inputs:
Outputs:
Figure 2. Schematic diagram of the main model components
35
20
10
0
d
i
s
c
h
a
r
g
e
,

m
m
Apr93 Apr94 Apr95 Apr96 Apr97 Apr98 Apr99 Apr00 Apr01
0
10
20
30
40
D
O
C

c
o
n
c
e
n
t
r
a
t
i
o
n
,

m
g
L
-
1
lower 10% upper 10% obs(q>0.3mm) obs(q<0.3mm) stand sim
E
p
i
s
o
d
e
E
p
i
s
o
d
e
Figure 3. Simulated (line) vs. measured (symbols) DOC concentrations at the outlet of
Kallkllsmyren mire. (
x
) DOC measurements corresponding to stream discharge values less
than 0.3 mm, ( ) DOC measurements corresponding to stream discharge value greater than
0.3 mm. The gray area indicates the upper and lower 10% prediction bounds obtained from
1000 Monte-Carlo runs processed using the GLUE methodology. Panel above: measured
stream discharge plotted with y axis inverted (i.e. with peak flows pointing downwards).


36
0
10
20
30
q
,

m
m
07/12 07/26 08/09 08/23
20
40
60
[
D
O
C
]
,

m
g
L

1
0
10
20
30
q
,

m
m
08/28 09/04 09/11 09/18 09/25
20
40
60
[
D
O
C
]
,

m
g
L

1
modelled DOC obs. DOC
a: 12July30August 1993
b: 27August28September 2001
Figure 4. Simulated and measured DOC concentrations for two episodes:
(a) 12 July-30 August, 1993, and (b) 27 August-28 September, 2001. Upper
panels illustrate the measured total stream discharges plotted with y axis
inverted (i.e. with peak flows pointing downwards).
37
layer1 layer2 layer3 layer4 layer5 layer6
c
o
m
p
o
n
e
n
t
s

o
f

t
h
e

D
O
C

b
a
l
a
n
c
e
,

g
m

1
0
1
2
3
net production
sorptive release
stream export
advective flux
dispersive flux
Figure 5. Modeled estimates of partitioning between the processes contributing to changes
in pore water DOC concentrations during the event from 27 August to 28 September 2001
in each of the six vertical peat layers of the acrotelm. Layer 1 is the uppermost layer.
38
1993 1994 1995 1996 1997 1998 1999 2000 2001
0
10
20
30
40
50
60

net production stream export SSOC&DOC stored
c
o
m
p
o
n
e
n
t
s

o
f

t
h
e

b
a
l
a
n
c
e
,

g
m
-
2
y
e
a
r
-
1
;
s
t
o
r
e

g
m
-
2
Figure 6. Simulated components of the annual DOC-SSOC balance: microbially
mediated net production and DOC stream export. Changes from year to year in
the amounts of SSOC stored in the acrotelm are also shown (estimated for the
dates before the first autumn frost).
39
1993 1995 1997 1999 2001
20
40
60
80
S
S
O
C
&
D
O
C

s
t
o
r
e
d
,

g
m
-
2

D
O
C
,

m
g
L
-
1
mod.SSOC&DOCstored before spr.food
mod. vol.weighted annual stream DOC
obs. vol.weighted annual stream DOC
10
15
20
25
30
30 40 50 60
30
40
50
60
mod. SSOC&DOC stored season before, gm
-2
m
o
d
.

v
o
l
.

w
e
i
g
h
t
e
d

s
e
a
s
o
n
a
l

s
t
r
e
a
m

D
O
C
,

m
g
L
-
1
a
b: spring
b:summer
r=0.40
r=0.87
Figure 7. (a) Interannual variations in the amounts of SSOC and DOC stored
before the start of the hydrological year (spring flood) simulated by the model.
Interannual variation in the volume-weighted annual stream DOC concentration,
simulated by the model and calculated from the interpolated observations.
(b) Correlation between the amounts of SSOC and DOC stored in the preceding
season (winter and spring, respectively), and the volume-weighted seasonal
stream DOC concentrations in spring and summer simulated by the model.
40
MEDDELANDEN FRN LUNDS UNIVERSITETS
GEOGRAFISKA INSTITUTION
Serie Avnandlingar
I. Herman Richter: Sknes karta frn mitten av 1500-talet till omkring 1700:
bidrag till en historisk-kartografisk underskning. (1929)
II. Josef Westin: Kulturgeografiska studier inom Ntra-, Nske- och
Utbyarnas flodomrden samt angrnsande kusttrakter. (1930)
III. Herman Richter och Wilhelm Norlind: Orbis Arctoi Nova et Accurata
Delineatio Auctore Andrea Bureo Sueco 1626. (1936)
IV. Sven Bjrnsson: Sommen-sundenomrdet : en geomorfologisk studie.
(1937)
V. Arne Sandell: Tektonik och morfologi inom dalformationen med
omgivande urbergsterrng. (1941)
VI. Sven Dahl: Torna och Bara : studier i Sknes bebyggelse- och
nringsgeografi fre 1860. (1942)
VII. Karl Erik Bergsten: Islvsflt kring norra Vttern : fysisk-geografiska
studier. (1943)
VIII. Carl Erik Nordenskjld: Morfologiska studier inom vergngsomrdet
mellan Kalmarsltten och Tjust. (1944)
IX. Sven Bjrnsson: Blekinge : en studie av det blekingska kulturlandskapet.
(1946)
X. Karl Erik Bergsten: stergtlands bergslag : en geografisk studie. (1946)
XI. Tor Holmquist: Den hallndska vinterfiskehamnsfrgan. (1947)
XII. Olof ngeby: Landformerna i nordvstra Jmtland och angrnsande delar
av Nord-Trndelag. (1947)
XIII. Axel Wennberg: Lantbebyggelsen i nordstra stergtland 1600-1875.
(1947)
XIV. Lars Bjerning: Sknes jord- och stenindustri : dess utveckling,
lokalisering och betydelse ur nringsgeografisk synvinkel. (1947)
XV. Allan Weinhagen: Norbergs bergslag samt Gunnilbo och Ramns till
omkring 1820 : studier i omrdets nrings- och bebyggelsegeografi.
(1947)
XVI. Helge Stlberg: Smlands skogs- och trfrdlingsindustrier : en nrings-
geografisk studie. (1947)
XVII. Folke Lgnert: Veteodlingen i sdra och mellersta Sverige. (1949)
XVIII. Yngve Nilsson: Bygd och nringsliv i norra Vrmland : en
kulturgeografisk studie. (1950)
XIX. Olof ngeby: Evorsionen i recenta vattenfall. (1951)
XX. Karl Erik Bergsten: Sydsvenska fdelseortsflt. (1951)
XXI. Folke Lgnert: Valmanskren p sknes landsbygd 1911-1948. (1952)
XXII. Olof Nordstrm: Relationer mellan bruk och omland i stra Smland
1750-1900. (1952)
XXIII. Arvid Bergdahl: Israndsbildningar i stra Syd- och Mellansverige med
srskild hnsyn till sarna. (1953)
XXIV. Sven E Behrens: Morfometriska, morfogenetiska och tektoniska studier
av de nordvstsknska urbergssarna, srskilt Kullaberg. (1953)
XXV. Torsten Hgerstrand: Innovationsfrloppet ur korologisk synpunkt.
(1953)
XXVI. Gunhild Weimarck: Studier ver landskapets frndring inom Lnsboda,
rkeneds socken, nordstra Skne. (1953)
XXVII. Ingemar Larsson: Structure and landscape in western Blekinge, southeast
Sweden. (1954)
XXVIII. Sven Godlund: Busstrafikens framvxt och funktion i de urbana
influensflten. (1954)
XXIX. Folke Lgnert: Syd- och mellansvenska vxtfljder. Del I : de ldre
bruknings-systemens upplsning under 1800-talet. (1955)
XXX. Olof ngeby: Toppkonstans, erosionsytor och passdalar i Jmtland och
Trndelag. (1955)
XXXI. Gunnar Johnsson: Glacialmorfologiska studier i sdra Sverige. (1956)
XXXII. Folke Lgnert: Syd- och mellansvenska vxtfljder. Del II : 1900-talet.
(1956)
XXXIII. Olof Nordstrm: Befolkningsutveckling och arbetskraftsproblem i stra
Smland 1800-1955. (1957)
XXXIV. Sven Godlund: Befolkning - regionsjukhus - resmjligheter - regioner.
(1958)
XXXV. Elis Plsson: Gymnasiers rekrytering och lokalisering. (1958)
XXXVI. Harald Svensson: Glaciation och morfologi : en glacialgeografisk studie i
ett tvrsnitt genom Skanderna mellan sdra Helgelandskusten och
Kultsjdalen. (1959)
XXXVII. Erik Ljungner: Nahuel Huapi : ein geographischer Querschnitt durch die
Anden in Patagonien. (1959)
XXXVIII. Nils Lewan: Om pendling mellan bostad och arbetsplats : en
underskning med material frn sydvstra Skne. (1960)
XXXIX. ke Mattsson: Morphologische Studien in Sdschweden und auf
Bornholm ber die nichtglaziale Formenwelt der Felsenskulptur. (1962)
XL. Stig Nordbeck: Framstllning av kartor med hjlp av siffermaskiner.
(1964)
XLI. Olof Nordstrm: Svensk glasindustri 1550-1960. (1962)
XLII. Jan Davidsson: Littoral processes and morphology on scandian flatcoasts.
(1963)
XLIII. Martin Markgren: Detaljmorfologiska studier i fast berg och
blockmaterial : geomorfologisk studie inom Fennoskandia med Skne.
(1962-1963)
XLIV. Martin Markgren: Geomorphological studies in Fennoscandia. II : chute
slopes in northern Fennoscandia. A : regional studies. (1964)
XLV. Martin Markgren: Geomorphological studies in Fennoscandia. II : chute
slopes in northern Fennoscandia. B : systematic studies. (1964)
XLVI. Lennart Amen: Stadsbebyggelse och domnstruktur. (1964)
XLVII. Arvid Bergdahl: Det glaciala landskapet. (1961)
XLVIII. Olof Nordstrm - Solveig Mrtensson: Turism p land (1966)
XLIX. Jan O Mattsson: The temperature climate of potato crops. (1966)
L. Nils Lewan: Landsbebyggelse i frvandling. (1967)
LI. Gsta Nordholm: Sknes ldre ekonomiska geografi. (1967)
LII. Sven Godlund - Torsten Hgerstrand - Bengt Svanstrm: Samhlls-
utvecklingen och samhllsplaneringen. (1967)
LIII. Tor Fr Rasmussen: Storbyutvikling og arbeidsreiser. (1966)
LIV. Erik Fagerlund - Harald Svensson - Sven Lindqvist m fl:
Infrardtermografi : principer och naturgeografiska tillmpningar. (1967)
LV. Lars Eldblom: Structure foncire, organisation et structure sociale. (1968)
LVI. Knut Norborg: Jordbruksbefolkningen i Sverige. (1968)
LVII. Gunhild Weimarck: Ulfshult (1968)
LVIII. Rune Jrvenstedt - Sven Lindqvist - Jan O Mattsson m fl:
Televisionssystem i naturgeografisk forskning. (1968)
LIX. Arne Jakobsson: Omflyttningen i Sverige 1950-1960. (1969)
LX. ke Hillefors: Vstsveriges glaciala historia och morfologi. (1969)
LXI. Sven Lindqvist: Bebyggelseklimatiska studier. (1970)
LXII. Torsten Hgerstrand, Gunnar Trnqvist m fl: Urbaniseringen i Sverige.
(SOU 1970:14)
LXIII. Bengt Sahlberg: Interregionala kontaktmnster : personkontakter inom
svenskt nringsliv en flygpassagerarstudie. (1970)
LXIV. Bjrn Hedberg: Kontaktsystem inom svenskt nringsliv : en studie av
organisationers externa personkontakter. (1970)
LXV. Mats G Engstrm: Regional arbetsfrdelning : nya drag i frvrvsarbetets
geografiska organisation i Sverige. (1970)
LXVI. Torsten Persson: Geomorphological studies in the south-Swedish
highlands. (1972)
LXVII. Dewitt Davis Jr: A factoral ecology of Malm 1965 : the social
geography of a city. (1972)
LXVIII. Zoltan Petery: Studier i svensk regionplanering : regionplanering enligt
byggnadslagen i mindre regioner. (1972)
LXIX. Tommy Book: Stadsplan och jrnvg i Norden. (1974)
LXX. Hans Abrahamson: Studier i jordbrukets omstrukturering. (1974)
LXXI. Christer Persson: Kontaktarbete och framtida lokaliseringsfrndringar.
(1974)
LXXII. Ulf Helldn: Karst : en studie av Artfjllets karstomrde samt jmfrande
korrosionsanalyser frn Vstspetsbergen och Tjeckoslovakien. (1974)
LXXIII. Jnos Szeg: Befolkningstthet, markanvndning, planering (vol 1 o 2).
(1974)
LXXIV. Raul Nino Guerrero: Rural to urban drift of the unemployed in Colombia.
(1975)
LXXV. Ulf Erlandsson: Fretagsutveckling och utrymmesbehov. (1975)
LXXVI. Sture berg: Methods of describing physical access to supply points.
(1976)
LXXVII. Bo Lenntorp: Paths in space-time environments : a time-geographic study
of movement possibilities of individuals. (1976)
LXXVIII. Richard hman: Palsar i Nordnorge : en studie av palsars morfologi,
utbredning och klimatiska frutsttningar i Finnmarks och Troms fylke.
(1977)
LXXIX. Bjrn Gyllstrm: The organization of production as a space-modelling
mechanism in underdeveloped countries. (1977)
LXXX. Anders Jrnegren - Fosco Ventura: Tre samhllens frndringshistoria :
exploateringen av den fysiska miljn i historisk belysning. (1977)
LXXXI. Tommy Book: Stadsplan och jrnvg i Storbritannien och Tyskland.
(1978)
LXXXII. Jan O Mattsson - Leif Brjesson: Lokalklimatiska temperaturstudier inom
ett sknskt fruktodlingsdistrikt med srskilt beaktande av frostlntheten.
(1978)
LXXXIII. Bjrn Terje Asheim: Regionale ulikheter i levekr. (1979)
LXXXIV. Solveig Mrtensson: On the formation of biographies in space-time
environments. (1979)
LXXXV. Erik Wallin: Vardagslivets generativa grammatik - vid grnsen mellan
natur och kultur. (1980)
LXXXVI. Reinhold Castensson: Vlja fr framtid - om markanvndningsval och
frtroendemannainflytande i kommunal planering. (1980)
LXXXVII. Kerstin Montal: Industri och vatten : den vattenfrorenande industrins
lokaliseringsproblem i Malmhus ln. (1980)
LXXXVIII. Tommy Carlstein: Time resources, society and eecology : on the capacity
for human interaction in space and time in preindustrial societies. (1980)
LXXXIX. Jonas kerman: Studies on periglacial geomorphology in west
Spitsbergen. (1980)
XC. Leif Engh: Karstomrdet vid Lummelunds bruk, Gotland, med speciell
hnsyn till Lummelundagrottan. (1980)
XCI. Karna Lidmar-Bergstrm: Pre-quaternary geomorphological evolution in
southern Fennoscandia. (1982)
XCII. Lars-Olof Olander: Staten, kommunerna och servicen : tiden kring
kommun-reformen i ett ekonomiskt geografiskt perspektiv. (1984)
XCIII. Bo Malmstrm och Owe Palmr: Glacial och periglacial geomorfologi p
Varangerhalvn, Nordnorge : geomorfologisk kartering med analys av
glaciala former och blockhav. (1984)
XCIV. Franz-Michael Rundquist: Hybrid maize diffusion in Kenya : policies,
diffusion patterns and consequences. (1984)
XCV. Girma Yadeta: Dynamic processes of development in marginal areas : a
case study from the pokot of north west Kenya. (1985)
XCVI. Anders Sporrek: Food marketing and urban growth in Dar Es Salaam.
(1985)
XCVII. Rolf Nyberg: Debris flows and slush avalanches in northern Swedish
Lappland distribution and geomorphological significance. (1985)
XCVIII. Lennart Olsson: An integrated study of desertification - applications of
remote sensing, GIS and spatial models in semi-arid Sudan. (1985)
XCIX. Mikael Stern: Census from heaven? : population estimates with remote
sensing techniques. (1985)
C. Katarina Olsson: Remote sensing for fuelwood resources and land
degradation studies in Kordofan, the Sudan. (1985)
CI. Gran Loman: The climate of a sugar beet stand - dynamics, impact on
the crop and possibilities of improvement. (1986)
CII. Eric Clark: The rent gap and urban change : case studies in Malm 1860-
1985. (1987)
CIII. Karin Hall-Knyves: Remote sensing of cultivated lands in the south of
Sweden. (1988)
CIV. Eva Ahlcrona: The impact of climate and man on land transformation in
central Sudan : applications of remote sensing. (1988)
CV. Kerstin Cederlund: Offentlig verksamhet : sysselsttning territoriellt och
funktionellt. (1988)
CVI. Per Olof Hallin: Tid fr omstllning : om hushlls anpassningsstrategier
vid en frndrad energisituation. (1989)
CVII. Jens Mller: Godsen och den agrara revolutionen : arbetsorganisation,
domn-struktur och kulturlandskap p sknska gods under 1800-talet.
(1989)
CVIII. Juha Uitto: The Kenyan conundrum : a regional analysis of population
growth and primary education in Kenya. (1989)
CIX. Ola Jonsson: Informationsteknologi och arbete : fallstudier inom svensk
sjukvrd. (1989)
CX. Tora Friberg: Kvinnors vardag. Kvinnors arbete och liv : anpassnings-
strategier i tid och rum. (1990)
CXI. Tomas Nihln: Eolian processes in southern Scandinavia and the
Mediterra-nean area. (1990)
CXII. Anders Lfgren: Att flytta hemifrn : boendets roll i ungdomars
vuxenblivande ur ett situationsanalytiskt perspektiv. (1990)
CXIII. Irma Guilln: Cuidad Guayana en stad, tv vrldar : en studie av ett
regionalt utvecklingsprojekts lokala effekter. (1991)
CXIV. Hans Holmn: Building organizations for rural development : state and
cooperatives in Egypt. (1991)
CXV. Petter Pilesj: GIS and remote sensing for soil erosion studies in semi-
arid environments : estimation of soil erosion parameters at different
scales. (1992)
CXVI. Ann-Cathrine quist: Tidsgeografi i samspel med samhllsteori. (1992)
CXVII. Jos da Cruz: Disaster and society : the 1985 Mexican earthquakes.
(1993)
CXVIII. Tomas Germundsson: Landsbygdens egnahem : egnahemsrrelsen,
smbruket och landskapet i sydsvenskt perspektiv. (1993)
CXIX. Ann-Katrin Bcklund: JUST-IN-TIME : hur industriella
rationaliseringsstrate-gier formar arbetsdelning och kompetens. (1994)
CXX. Jon Knudsen: Kulturspredning i et strukturelt perspektiv : eksemplifisert
ved politisk og religis endring under moderniseringen av det norske
samfunn. (1994)
CXXI. Tone Haraldsen: Teknologi, konomi og rom : en teoretisk analyse av
relasjoner mellom industrielle og territorielle endringsprosesser. (1994)
CXXII. Peter Jnsson: Wind climate during the instumental period and recent
wind erosion in southern Scandinavia. (1994)
CXXIII. Peter Schlyter. Palaeo-wind abrasion in southern Scandinavia : field and
laboratory studies. (1995)
CXXIV. Jun Yamashita: Spatial interaction and spatial structure : a study of public
facility location. (1995)
CXXV. Mats Riddersporre: Bymarker i backspegel : odlingslandskapet fre
kartornas tid. (1995)
CXXVI. Anders Schrstrm: Pathogenic paths? : a time geographical approach in
medical geography. (1996)
CXXVII. Lars Eklundh: AVHRR NDVI for monitoring and mapping of vegetation
and drought in east African environments. (1996)
CXXVIII. Magnus Jirstrm: In the wake of the green revolution : environmental and
socio-economic consequences of intensive rice agriculture the problem
of weeds in Muda, Malaysia. (1996)
CXXIX. Stefan Anderberg: Fldesanalys i den hllbara utvecklingens tjnst :
reflektioner kring en metabolism studie av Rhenomrdets utveckling.
(1996)
CXXX. Karl-Johan Lundquist: Fretag, regioner och internationell konkurrens :
om regionala resursers betydelse. (1996)
CXXXI. Badr-Eldin Taha Osman: GIS-hydrological modelling in aridlands : a
geographical synthesis of surface waters for the African Red Sea region in
the Sudan. (1996)
CXXXII. Marie Stenseke: Bonden och landskapet : gares och brukares relationer
till markerna och frutsttningarna fr en uthllig markanvnding. (1997)
CXXXIII. Kristina Blennow: Spatial variation in near-ground radiation and low
temperature interactions with forest vegetation. (1997)
CXXXIV. Lennart Runesson: Tomtrtt : ett markpolitiskt instrument i upplsning.
(1997)
CXXXV. Johan Hultman: The eco-ghost in the machine : reflexions on space, place
and time in environmental geography. (1998)
CXXXVI. Jonas Ard: Remote sensing of forest decline in the Czech Republic.
(1998)
CXXXVII. Per Hillbur: The knowledge arena : approaching agroforestry and
competing knowledge systems a challenge for agricultural extensionl
(1998)
CXXXVIII. Tom Mels: Wild landscapes : the cultural nature of Swedish national
parks. (1999)
CXXXIX. Carolyn Hannan-Andersson: Promoting equality between women and
men in bilateral development cooperation : concepts, goals, rationales and
institutional arrangements. (2000)
CXL. Nikolaus Solakius: The parnassus zone, central Greece. (2000)
CXLI. Jonathan Seaquist: Mapping primary production for the west African
Sahel using satellite data. (2001)
CXLII. Karin Book och Lena Eskilsson: Stadens struktur : varfr och hur? (2001)
CXLIII. Maria Wikhall: Universiteten och kompetenslandskapet : effekter av den
hgre utbildningens tillvxt och regionala spridning i Sverige. (2001)
CXLIV. Rannveig Olafsdottir: Land degradation and climate in Iceland : a spatial
and temporal assessment. (2002)
CXLV. Marie Ekstrm: Relationships between atmospheric circulation and wind
erosion in southern Sweden and Australia. (2002)
CXLVI. Maj-Lena Finnander Linderson: The spatial distribution of precipitation
in Scania, southern Sweden : observations, model simulations and
statistical downscaling. (2002)
CXLVII. Richard Ek: resundsregion - bli till! : de geografiska visionernas
diskursiva rytm. (2003)
CXLVIII. Olivia Louw: Exploring the culture of non-payment in the post-apartheid
South Africa. (2003)
CXLIX. Cecilia Kjellman: Ta plats eller f plats? : studier av marginaliserade
mnniskors frndrade vardagsliv (2003)
CL. Christina Scholten: Kvinnors frsrjningsrum : hegemonins frvaltare
och murbrckor (2003)
CLI. Michael Runnstrm: Land degradation and mitigation in northern China :
evaluated from the biological production (2003)
CLII. Sara Brogaard: Recent changes in land use and productivity in agro-
pastoral Inner Mongolia, China (2003)
CLIII. Jan_Henrik Nilsson: stersjomrdet : studier av interaktion och barrirer
(2003)
CLIV. Thomas Hickler: Towards an integrated ecology through mechanistic
modelling of ecosystem structure and functioning (2004)
CLV. Andreas Persson: Hydrological modelling, topographical influence and
yield mapping in precision agriculture (2004)
CLVI. Maria Olsrud: Mechanisms of below-ground carbon cycling in subarctic
ecosystems (2004)
CLVII. Sandra C. Fernndez: Farewell to the peasantry? : (Post)modernising rural
Mexico : the case of the ejido peasants in the Isthmus of Tehuantepec
(2004)
CLVIII. Andrs P. Lesk: To own the phone : spatial diffusion, ownership and
regulation of telephone service in Argentina, 1878-1990 (2004)
CLIX. Henrik Svensson: ppna och slutna rum : enskiftet och de utsattas
geografi : husmn, bnder och gods p den sknska landsbygden under
1800-talets frsta hlft (2005)
CLX. Pablo Morales: Modeling carbon and water fluxes in European terrestrial
ecosystems (2006)
CLXI. Emmanuel S. Gritti: Global changes and European terrestrial ecosystems
(2006)
CLXII. Ola Thufvesson: Kreativitetens yttre villkor : miljer, rrlighet och
nobelpristagare (2006)
CLXIII. Deniz Koca: Impacts of regional climate change on Swedish forests : an
evaluation using process-based regional ecosystem modelling approach
(2006)
CLXIV. Bodil Elmqvist: Livelihood diversification and land use change in the
Sahel : an interdisciplinary analysis of gum arabic in Sudan (2006)
CLXV. Jan Vang-Lauridsen: Industrial dynamics and the spatial organization of
industries (2006)
CLXVI. Heidi Wiig Aslesen: Innovation in an urban context (2006)
CLXVII. Torbjrn Johansson: Temporal and spatial variability of carbon cycling in
a subarctic landscape (2006)
CLXVIII. Anders Lund Hansen: Space wars and the new urban imperialism (2006)
CLXIX. Lars Coenen: Faraway, so close! : the changing geographies of regional
innovation (2006)
CLXX. Pontus Olofsson: Remote sensing of carbon balance across Scandinavian
forests (2007)
CLXXI. Margareta Rmgrd: The power of place : existential crises and place
security in the context of pregnancy (2006)
CLXXII. Helena Eriksson: Leaf area index of Scandinavian forests : methods using
in situ and remotely sensed data (2007)
CLXXIII. Ransom Lekunze: Corporate social responsibility and development : the
case of the Chad Cameroon oil pipeline project (2007)
CLXXIV. Alla Yurova: Hydrological aspects of the carbon balance in a boreal
catchment : a model study (2007)
I