Plant and Soil 251: 303317, 2003.

2003 Kluwer Academic Publishers. Printed in the Netherlands.

303

Soil and total ecosystem respiration in agricultural fields: effect of soil and
crop type
Annalea Lohila1,3 , Mika Aurela1 , Kristiina Regina2 & Tuomas Laurila1
1 Finnish

Meteorological Institute, Air Quality Research, Sahaajankatu 20 E, 00810 Helsinki, Finland. 2 Agrifood
Research Finland, Soils and Environment, MTT, 31600 Jokioinen, Finland. 3 Corresponding author

Received 29 April 2002. Accepted in revised form 20 October 2002

Key words: mineral soil, peat soil, soil carbon content, soil respiration, total ecosystem respiration

Abstract
A study was made of the effect of soil and crop type on the soil and total ecosystem respiration rates in agricultural
soils in southern Finland. The main interest was to compare the soil respiration rates in peat and two different
mineral soils growing barley, grass and potato. Respiration measurements were conducted during the growing
season with (1) a closed-dynamic ecosystem respiration chamber, in which combined plant and soil respiration
was measured and (2) a closed-dynamic soil respiration chamber which measured only the soil and root-derived
respiration. A semi-empirical model including separate functions for the soil and plant respiration components was
used for the total ecosystem respiration (TER), and the resulting soil respiration parameters for different soil and
crop types were compared. Both methods showed that the soil respiration in the peat soil was 23 times as high
as that in the mineral soils, varying from 0.11 to 0.36 mg (CO2 ) m2 s1 in the peat soil and from 0.02 to 0.17
mg (CO2 ) m2 s1 in the mineral soils. The difference between the soil types was mainly attributed to the soil
organic C content, which in the uppermost 20 cm of the peat soil was 24 kg m2 , being about 4 times as high as
that in the mineral soils. Depending on the measurement method, the soil respiration in the sandy soil was slightly
higher than or similar to that in the clay soil. In each soil type, the soil respiration was highest on the grass plots.
Higher soil respiration parameter values (Rs0 , describing the soil respiration at a soil temperature of 10 C, and
obtained by modelling) were found on the barley than on the potato plots. The difference was explained by the
different cultivation history of the plots, as the potato plots had lain fallow during the preceding summer. The total
ecosystem respiration followed the seasonal evolution in the leaf area and measured photosynthetic flux rates. The
23-fold peat soil respiration term as compared to mineral soil indicates that the cultivated peat soil ecosystem is a
strong net CO2 source.

Introduction
The soil is a major biospheric reservoir for carbon,
containing globally twice as much C as the atmosphere
and three times as much as vegetation. About one-third
of the carbon stored in soils is present in the northern peatlands (Gorham, 1991). In Finland, more than
half of the original peatland area has been drained for
forestry or farming purposes. Nowadays about 2000
km2of peat soils are under cultivation, accounting for
FAX No: +358-9-1929 5403.

Email: Annalea.Lohila@fmi.fi

about 10% of the total area of arable soils (Myllys,

1996). Drainage and agricultural use increase the aeration of the peat, and lead to enhanced microbial
degradation of organic material, turning the peatland
from a carbon dioxide sink into a source (Nyknen
et al., 1995; Silvola et al., 1996a). During the last
few years, an increasing number of studies on CO2
dynamics on unaltered, natural wetlands (e.g., Alm
et al., 1999a, b; Aurela et al., 1998, Bubier et al.,
1998; Suyker et al., 1997) or on managed cutover or
restored peatlands (e.g., Petrone et al., 2001; Sundh
et al., 2000; Tuittila et al., 1999; Waddington and
Warner, 2001; Waddington et al., 2002) have been

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made. However, on agricultural peat soils, where
regular soil fertilization and mechanical disturbance
further stimulate the mineralisation rate of the old peat,
measurements of CO2 emission rates are scarce (Koizumi et al., 1999; Maljanen et al., 2001; Nyknen et
al., 1995).
The net carbon dioxide balance of an agricultural
ecosystem depends on the magnitude of two fluxes
having opposite signs: CO2 uptake by gross photosynthesis and CO2 release by shoot, root and soil
respiration. As the seasonal variation in shoot and
root respiration rates can be assumed to be roughly
equal for similar crops experiencing the same growth
conditions, the major difference between the net CO2
balances of agricultural ecosystems established on different soil types arises from the differences in soil
heterotrophic respiration. To make accurate estimates of ecosystem CO2 dynamics, it is necessary to
quantify all the separate fluxes comprising the net
balance of carbon dioxide. When measurements of
the total ecosystem respiration are used, a modelling
approach is necessary in order to partition the autotrophic and heterotrophic components. This is often
quite a complicated task, and the process models used
to estimate ecosystem respiration do not often differentiate between these two, even though different
driving variables control plant and soil derived respiration. The most important factors affecting the soil
respiration rate are soil temperature (Buyanovsky et
al., 1986; Kutsch and Kappen, 1997; Rochette et al.,
1995), soil moisture (Akinremi et al., 1999; Buyanovsky et al., 1986; Kutsch and Kappen, 1997), soil
organic carbon quantity and quality (La Scala et al.,
2000), and soil texture, which also affects the gaseous
diffusion in the soil (Bouma and Bryla, 2000). Primary
production has been shown to be one of the main contributors not only for plant and root respiration, but
also for soil rhizosphere respiration, both in laboratory experiments as well as in many different crop,
grass and forest ecosystems in both the shorter and
the longer term (Hgberg et al., 2001; Kuzyakov
and Cheng, 2001; Raich and Tufekcioglu, 2000). In
soil respiration models, however, primary production
has not been commonly used to explain the daily or
seasonal variation in the soil respiration.
This study is a part of a 3-year (19992002) national global change research program (FIGARE). A
subproject of this program, AGROGAS, aims to produce estimates of the yearly balances of CO2 , N2 O and
CH4 from typical Finnish agricultural soils. The results will be utilised in determining the greenhouse gas

emissions from agriculture. This paper presents the

results from chamber-based measurements of the ecosystem and soil respiration fluxes for one growing season. The objectives of this study were (1) to quantify
and compare the soil and ecosystem respiration rates
in different agricultural ecosystems consisting of organic and mineral soils and three crop species and
(2) to utilise and adjust a total ecosystem respiration
model for future CO2 balance calculations.

Materials and methods

The experimental sites
The study was carried out at Jokioinen, in Southern
Finland. In 1999, an experimental plot (46 30 m)
with nine subplots (10 10 m) was established on
organic (Terric histosol, 6054 N, 23 31 E), sandy
(Eutric cambisol, 60 51 N, 2328 E) and clay (Vertic
cambisol, 6048 N, 2328 E) fields, located no more
than 15 km from each other. The soil types have been
described in more detail by Yli-Halla and Mokma
(2001). In the growing season of 2000, three separate
subplots of barley (Hordeum vulgare L. Saana), perennial ley (a mixture of Phleum pratense and Festuca
pratensis, hereafter referred to as grass) and potato
(Solanum tuberosum L. Van Gogh) were grown on
the experimental plots. The distribution of the subplots
obeyed the latin square design. In the preceding year,
barley and grass had been grown on the same subplots
as in summer 2000, while the potato plots had been left
fallow. The barley plots were ploughed in the autumn
and sown again the following spring. The organic peat
field has been in agricultural use for at least a century.
The mean summer water table depth was about 80 to
100 cm in each soil type.
Barley was sown at a rate of 256 kg ha1 on 8 9
May, 2000 in the mineral soils and on 11 May in the
peat soil. It was harvested on 28 August on the peat
and clay soils and on 5 September on the clay soil.
The grass was cut for the first time on 13, 14 and 19
June on the clay, sandy and peat soils, respectively. In
the clay soil, the grass was re-sown after the first cut.
The second grass cutting took place on 7 August on
the clay soil and on 8 August on the peat and sandy
soils. Potato was sown on 23, 24 and 25 May on the
clay, sandy and peat soils, respectively. The stems of
the potatoes were removed on 11, 15 and 25 August on
the sandy, peat and clay soils, respectively. The potato
was harvested on 18 and 21 August on the peat and

305
sandy soils, respectively, and on 5 September on the
clay soil.
In 2000, the mean annual temperature and precipitation at Jokioinen were 6.0 C and 652 mm, respectively. According to the long-term meteorological data
for Jokioinen weather station (60 49 N, 23 30 E), the
mean annual temperature was 3.9 C and mean annual
precipitation was 581 mm during the normal period
(19611990) (Finnish Meteorological Institute, 1991).
Analysis of vegetation and soil physical and chemical
characteristics
The leaf area index (LAI) was measured with an LAI2000 Plant Canopy Analyzer (LI-COR, Inc, NE) 12
times a week on every subplot, inside and beside the
collars. Plant biomass was determined at the time of
harvest from oven-dried (40 C) samples from three
replicate total ecosystem respiration measurement collars. The yields of the barley and potato crops were
determined with a threshing machine and a potato
harvester from about 20 m2 area of each subplot.
The soil moisture was measured continuously at a
depth of 15 cm in the peat soil growing barley and
at a depth of 7 cm in the clay soil growing barley
with a ThetaProbe Soil Moisture Sensor type ML2x
(Delta-T Devices Ltd., U.K.). The soil moisture was
also determined biweekly from oven-dried (105 C)
soil samples from three pooled 25-cm cores with a
diameter of 3 cm. The soil bulk density was determined three times during the growing season from three
replicate oven-dried 15-cm soil cores (diameter 5 cm)
from each plot. The particle density of the soil was
determined from five replicate samples as described in
Blake and Hartge (1986). The soil pH was determined
from suspensions of 30 ml of soil and 50 ml of water.
Analyses of total C and N were made from ground,
air-dried samples using a Leco-analyser CN-2000.
The basic soil characteristics are presented in Table
1. The soil organic carbon and nitrogen content, as
well as the C/N-ratio, were highest in the peat soil.
The mean water-filled pore space during summer 2000
was lower in the sandy soil than in the peat or clay
soils. The soil moisture determined manually by soil
sampling varied between 0.20 and 0.55 m3 m3 in
the peat, between 0.13 and 0.23 m3 m3 in the sandy
soil and between 0.29 and 0.44 m3 m3 in the clay.
These values supported the results obtained with the
automatic device, according to which the soil moisture
varied in the peat soil from 0.15 to 0.35 m3 m3 , and
in the clay soil from 0.27 to 0.45 m3 m3 . In each

soil type the soil moisture was generally lowest in the

grass plots and highest in the potato plots (data not
shown). There was a similar seasonal trend in the soil
moisture measured by both methods: in each soil type
the water content decreased until the end of June and
increased again at the end of July. The amounts of
applied fertilizers are shown in Table 2.
The total dark respiration and the soil respiration
measurements
The total ecosystem respiration (TER) is the sum of
the root and soil heterotrophic respiration, added to
the shoot respiration. The TER was measured weekly
between 08:00 and 17:00 on each subplot from May
16 to September 21, except during some rainy periods.
At the beginning of the growing season, a 60 60-cm
steel or aluminum collar (height 15 cm), with a water
groove for making the chamber airtight, was inserted
into the soil on each subplot to a depth of about 10 cm.
The TER was measured with an opaque aluminium
chamber (height 20 cm) equipped with a long pressure equilibrium tube and a fan inside the chamber.
The change in the chamber CO2 concentration was
analysed by an infra-red gas analyser (LCA-2, Analytical Development Company Ltd., England) with an air
flow of about 0.2 l min1 . Each measurement took 12
min. The TER flux rate was calculated from the linear
increase in the chamber headspace CO2 concentration.
The chamber air temperature was monitored simultaneously. Additional steel collars (20 or 40 cm) were
used later in the growing season due to higher vegetation, and to ensure adequate air mixing an extra fan
was installed inside the chamber. Soil temperature at a
depth of 4 cm outside the collar was measured at the
same time as the respiration measurement. In the barley and potato plots, the collars were reinstalled after
sowing or planting, respectively. In the grass plots, the
collars were moved to new places after both harvests.
Soil respiration measurements (SR) were conducted with a dynamic, closed-chamber infra-red gas analyser system (EGM-2, SRC-1, PP-Systems, Hitchin,
U.K.). Small PVC collars (diameter 10 cm, height 5
cm) were inserted into the soil (to a depth of 3 cm)
at least 24 h before the measurements. On each experimental field, two to four collars were inserted on
two replicate subplots growing barley or grass. Half
of the collars were placed between the rows, while the
other half were inserted in the rows. To eliminate the
plant respiration, the above-ground parts of the plants
were cut off before inserting the collars and were kept

306
Table 1. Soil characteristics and particle size fractions of the experimental fields
Peat soil
Physical and chemical properties
Org C g kg1 (SE)
250 40
N (Leco) g kg1 (SE)
12 2
440 70
Humus g kg1 (SE)
Amount of C at 0-20 cm (kg m2 )
24
Bulk density (g cm3 )
0.48
1.93
Particle density (g cm3 )
Soil pH
5.8
Total porosity (%)
81
Soil moisture (m3 m3 )a
Average (min,
max)
0.36 (0.200.55)
Water-filled pore space (%)
61

Sandy soil

Clay soil

24 5
1.7 0.3
42 8
5.8
1.20
2.59
6.0
56

30 2
2.3 0.1
51 4
6.3
1.06
2.65
6.5
57

0.18 (0.130.23)
31

0.38 (0.290.44)
55

96
135
769

573
334
93

Soil particle size fractions

Clay (g kg1 )
Silt (g kg1 )
Sand (g kg1 )

nd
nd
nd

a Determined by soil sampling, mean of the values from 2 May 28 September, 2000.

nd not determined.

short by regularly clipping them. Soil temperature was

measured simultaneously at a depth of 4 cm. The calculation of the SR flux was done automatically by
fitting a quadratic equation to the relationship between
the increasing CO2 concentration and elapsed time.
Each measurement took 12 min.
Two periods of SR measurements were conducted:
one before barley sowing on 34 May, representing
spring conditions, and another in the autumn between
August 3 and 29. The collars were reinstalled into the
soil at the beginning of the autumn campaign. During
the spring period, a total of two to three measurements was conducted on each collar. In the autumn,
the measurements were made 12 times per week. The
bare soil measurements were done close to the barley
plots in a place kept free of vegetation. In the autumn,
no bare soil measurements were made in the sandy
soil. The SR measurements were always done between
08:00 and 17:00.
The fan in the SRC-1 designed for mixing purposes
has been found to cause an overestimation of the soil
carbon dioxide efflux by pumping CO2 storages out of
the soil pores (LeDantec et al., 1999). It is possible
that the degree of the overestimation varies depending on the soil texture, and may be higher for porous
soils like peat. To ensure that a systematic error in
the SR estimate between the soil types was not being

made, at least two consecutive measurements on each

collar were carried out. If the difference was more
than 15%, further measurements from the same collar
were conducted until a constant flux level was reached.
Occasionally a decreasing trend in the flux rate was
observed, but this was independent of the soil type.
The SR data are presented as means of seven to 26
replicates standard error (SE). Only measurements
in the soil temperature range 815 C were included
so that the averages were as close to 10 C as possible. Comparison among means was carried out using
the Tukeys honestly significant difference (HSD) test
(=0.05).
Modelling the total ecosystem respiration
In order to quantify the relationships between the environmental and phenological variables and the total
ecosystem respiration, as well as to divide the total respiration into its soil (R s ) and plant (Rd ) components,
an equation combining Rd and Rs was fitted to the
TER chamber data (Aurela et al., 1998, 2001a; Kim
et al., 1992). The equation for the soil respiration used
here was developed by Lloyd and Taylor (1994). To
make the model work better in heavily-managed agricultural ecosystems, we added a term (SA) to the soil
respiration part, reflecting the effect of plant activity

307
Table 2. The amount of applied fertilizers in the experimental fields

Barleya
Grassb
Potatoa

N-P-K (kg ha1 )

Peat soil

Mineral soils

60 12 21
60 4.6 7
100 50 116

100 20 35
100 8 12
100 50 116

a Fertilizers were applied once, at sowing/planting.

b Fertilizers were applied twice: on 611 May and on 19 June.

on soil respiration. The model takes the form

TER = R d + R s




1
1

= PI Rd0 exp b d
T0 + 273 T1 + 273




1
1

+ SAR s0 exp 308.56

,
56.02 T s + 46.02

where PI is an empirically-determined dimensionless effective phytomass index describing seasonal

changes in the photosynthetically active vegetation.
The concept of PI was originally introduced by Aurela
et al. (2001a). PI has been derived from the net ecosystem CO2 exchange data (measured with a transparent
chamber) by determining the magnitude of the gross
photosynthesis at a photosynthetic photon flux density (PPFD) level of 1200 mol m2 s1 , denoted as
GP1200. PI was normalised to unity at its maximum
measured on barley at the end of June.
Rd0 is the rate of plant dark respiration (in mg
(CO2 ) m2 s1 ) at the reference temperature T0 =
10 C. The value of Rd0 , 0.14 mg (CO2 ) m2 s1 ,
was selected on the basis of some trial fitting procedures and other studies. The same parameter value was
used for all three crops. Kim et al. (1992) estimated
the highest Rd0 to be 0.08 mg (CO2 ) m2 s1 for
warm season grasses with a maximum LAI of about
3. On a wheat field (LAI 3) at air temperatures of
about 1015 C the highest night-time total ecosystem respiration values were about 0.20 mg (CO2 ) m2
s1 (Baldocchi 1994). The peak LAI in our data was
higher than in these studies. Tl is the leaf temperature ( C), instead of which we used the chamber air
temperature. bd is a physiological constant (K) often
represented as Ea /R, where Ea is the activation energy quantifying the sensitivity of canopy respiration
rates to temperature. R is the universal gas constant
(8.314 K1 mol1 ). We selected a constant value of
bd = 5000 to describe the temperature dependence of
the plant dark respiration of barley, grass and potato.

This value is somewhat smaller than that measured by

Baldocchi (1994) for wheat (bd = 6990), higher than
that Gent and Kiyomoto (1992) measured for wheat
(Q10 = 1.3, corresponding to bd of about 2500), but
close to the value measured by Kim et al. (1992) for a
temperate grassland ecosystem, bd =5410.
In the soil respiration term, Rs0 is a parameter describing the soil respiration (in mg (CO2 ) m2 s1 ) at
the reference soil temperature of 10 C. In this work,
Rs0 is thought to mainly represent the heterotrophic
oxidation of parent soil organic matter, possibly augmented with some part of the root respiration. Ts is the
soil temperature ( C) at a depth of 4 cm. Due to the relatively small amount of data and consequently limited
temperature range, it was not possible to experimentally determine the accurate temperature response of
the soils used in this study. In the model, the temperature response in the range from 10 to 20 C corresponds
to a Q10 of about 2.3. This is close to the values Koizumi et al. (1999) observed (1.72.3) in soil types and
at a location nearly identical to ours. In addition, our
soil respiration measurements support these values.
SA, the effective soil activity, is a dimensionless
semi-empirical coefficient that we added to the original soil respiration model in an attempt to better
reproduce the TER. By doubling as a function of time
in the course of the growing season, SA describes the
level of soil activity and thus soil respiration mainly
due to the input of easily-degradable carbon derived
from plant activity. Root exudates and plant and root
litter allocated into the soil during the growing season
enhance the soil respiration by stimulating microbial
growth and activity. The idea of using SA in the modelling arose from the TER results shown in this paper;
these were still quite high late in the summer after
barley ripening, when plant and root respiration had
ceased. These flux rates could not be explained by
higher soil temperatures, either. Throughout the data, a
better fit in the modelling was obtained by using SA as
a multiplier in the temperature-dependent respiration
model.
In barley and potato, SA starts to increase soon
after sowing (Figure 1). The maximum is reached
at about the time of the reproductive growth stage
in potato and barley. For simplicity, the increase is
assumed to be linear. About 2 weeks after the harvest, when most of the plant-derived carbon has been
incorporated into the soil, SA begins to decrease exponentially, being 1 again at the beginning of the next
growing season. In grass, the increase begins when the
daily effective temperature sum (ETS, with base tem-

308

Figure 1. Annual variation of the semi-empirically-determined effective soil activity (SA) used in the modelling.

perature 5 C) begins to accumulate. The level of 2 is

reached when the ETS does not increase any further.
This is the point at which assimilation is assumed to
cease.
The use of the chamber and soil temperatures as
driving variables for the plant and soil respiration, respectively, may produce a bias in the interpretation of
Rs0 and Rd0 , as these two temperatures presumably
correlate. Since no separate measurements exist for
the soil and plant respiration (the SR measurements
cannot be used directly to calibrate the soil respiration
level due to the small number of measurements and the
discontinuous time series in the SR data), we do not
know the proportions of these two components. However, this is not crucial in this work, as the idea was
to compare different treatments. Seeing that the combined above-ground plant dark respiration is a fixed
value in the modelling (Rd0 = 0.14 mg (CO2 ) m2 s1
when PI = 1), and is additionally normalised to the
other treatments by PI, we claim that the Rs0 parameter values from different soilcrop combinations
can be compared reliably.
In the modelling, Rs0 was the only unknown parameter, and was solved separately for each of the nine
data sets consisting of three different soil and plant
types. Air and soil temperatures, calculated PI values
and SA were used as input parameters. All curve fittings were performed with the MarquardtLevenberg
algorithm.

Results
PI, LAI, biomass and yield
The highest individual value of GP1200 , 2.47 mg
(CO2 ) m2 s1 , which was used to normalise the PI
scale, was measured on the peat soil growing barley
at the end of June (Figure 2). If the three highest PI
values measured on the peat soil growing barley are
excluded, the PI maxima in barley and grass were
fairly similar, despite the higher LAI maxima in grass
(Table 3). As expected, PI and LAI seemed to be
tightly coupled to each other during the period of vegetative growth (results not shown). After the start of
senescence, LAI is no longer a reliable measure of the
green leaf area. Thus PI may be used throughout the
growing season instead of LAI to describe the seasonal
evolution of the green phytomass activity.
In barley plots, the total above-ground biomass,
expressed on a dry weight basis, varied between 0.8
and 0.9 kg m2 . In potato plots, the combined above
and below-ground biomass was close to 1.0 kg m2
in the peat and sandy soils, but was lower in the clay
soil. The potato yield was higher in the sandy than in
the peat soil. This may be explained by the fact that
the potato on the peat soil received too much N, which
may possibly have led to increased investment in leaf
and stem growth and to less-than-normal investment
in the tubers. Additionally, the harvest loss is greater
in the clay soil, as there the potato tubers are stuck in
the stiff clay lumps. The biomass of grass is a sum of

309
Table 3. LAI maxima, above-ground biomass, and average yield for barley, grass and potato. Also shown is the date when the highest
LAI was measured. The biomass was determined from the collar area, while the yield was determined from the subplot. Biomass and
yield are an average of three subplots. Grass biomass was determined on 13 June and 7 August, and potato and barley biomass from
the harvest

LAImax
Date (LAImax )
Biomass (kg m2 )a
Yield (kg m2 )b

Peat

Barley
Sandy

Clay

Peat

Grass
Sandy

Clay

Peat

Potato
Sandy

Clay

6.9
12/7
0.90
0.49

4.7
7/7
0.80
0.47

4.6
26/7
0.87
0.48

7.4
7/8
1.23
nd

7.9
7/8
0.88
nd

6.9
7/8
0.96
nd

5.1
26/7
0.96
1.08

4.7
20/7
1.06
1.72

3.4
2/8
0.65
0.89

a Collar area harvested, average of the three subplots. In grass, the two harvests are summed. In potato the tuber biomass is also
included. All values are expressed on a dry weight basis.
b Whole subplot harvested, average of the three subplots. In potato, only the fresh weight of the tuber is shown.
nd not determined.

two harvests on 12 June and 7 August. There was less

grass phytomass on the sandy and clay soils than on
peat soil for the first harvest. This can also be seen in
PI, which is lower on the sandy and clay soils than on
the peat soil before the first grass cutting (Figure 2).
Seasonal trend in the total ecosystem respiration rates

Figure 2. Phytomass index (PI), derived from the measured photosynthetic CO2 uptake rates, together with the highest measured
LAI for each treatment. PI was normalised to unity at its maximum
measured on barley on 30 June.

The highest carbon dioxide fluxes from the bare barley

fields, before crop emergence, were measured on the
peat soil, averaging 0.24 mg (CO2 ) m2 s1 (at mean
soil temperature 18.1 C). The corresponding fluxes
on the clay and sandy soils growing barley averaged
0.07 (17.5 C) and 0.14 (20.0 C) mg (CO2 ) m2 s1 ,
respectively. On all soil types, the highest TER fluxes
were measured between 28 June and 8 July, practically simultaneously with the barley flowering (Figure
3a). The peak TER values (average of the three plots)
measured on each soil type growing barley were 0.89,
0.66 and 0.67 mg (CO2 ) m2 s1 on the peat, clay,
and sandy soils, respectively. In general, the evolution
of the total ecosystem respiration closely followed the
development of LAI and PI until flowering. After that,
the TER was higher than what could be expected on
the basis of PI, probably reflecting the stronger role of
soil respiration towards the end of the summer.
As with barley, the TER fluxes on the grass increased with PI, dropping after each cut (Figure 3b).
The highest TER values (average of the three plots)
were measured on the peat and sandy soils at the beginning of August (1.1 and 0.57 mg (CO2 ) m2 s1 ,
respectively) and on the clay soil at the end of August
(0.51 mg (CO2 ) m2 s1 ).
For the potato, the average TER fluxes from the
bare soil were again greatest on the peat soil, 0.11

310

Figures 3a-b. Measured total ecosystem respiration () (average of three subplots, SE) in summer 2000 on different soil types for (a) barley
(b) grass and (c) potato. Also shown are the chamber air temperature () and the soil temperature (
) at a depth of 4 cm. The time of the
harvest is indicated by the broken line. For potato, the dotted line indicates the time of tuber harvest.

mg (CO2 ) m2 s1 (mean soil temperature 16.1 C).

On the bare clay and sandy soils, the CO2 fluxes were
0.05 (16.1 C) and 0.06 mg (CO2 ) m2 s1 (17.7 C),
respectively (Figure 3c). The magnitude of the TER
followed the same pattern as was seen for barley and
grass, the total respiration rates increasing with PI.
The peak TER values (average of two to three plots)
measured on each soil type growing potato were 0.84,
0.26 and 0.39 mg (CO2 ) m2 s1 in the peat, clay,
and sandy soils, respectively. The rapid decline in TER
and PI (Figure 2) soon after the potato flowering was
caused by an infection of potato blight.

Measured and modelled soil respiration

The measured total ecosystem respiration (TER) represents a combination of soil, root and shoot dark
respiration. In order to compare the soil respiration
rates in different agricultural ecosystems, the soil respiration was separated from the TER by modelling.
Irrespective of crop type, the soil respiration parameter, Rs0 , was always highest for the peat soil and
smallest for the clay soil (Figure 4). Furthermore, for
each soil type Rs0 was always highest for grass and
lowest for potato. The ratio of Rs0 between the peat
and the mineral soils varied between 1.9 and 3.0, being highest for the potato plots and lowest for the
grass plots. However, the difference between them was

311

Figure 3c.

rather constant for different crop types, ranging from

0.053 to 0.071 mg (CO2 ) m2 s1 .
The averages of the soil respiration (SR) measurements conducted in the spring and autumn periods
are shown for each soilcrop-combination in Figure
5. As the above-ground vegetation was kept short
by clipping, no movement of photosynthates into the
rhizosphere occurred. Thus the measured respiration
mainly reflects soil heterotrophic respiration and root
respiration, except in the bare soil having no vegetation, where the flux originates from the soil only.
Within each treatment the highest CO2 effluxes were
measured on the peat soil, averaging 23 times as high
as that on the mineral soils. However, most of the
differences between the soil types were not significant in the spring period. The soil respiration ranged
from 0.11 to 0.35 mg (CO2 ) m2 s1 and from 0.02 to

0.17 mg (CO2 ) m2 s1 on the peat and mineral soils,

respectively. No significant differences were found
between the SR values for the mineral soils. Similarly to the parameter Rs0 , SR was generally highest
on grass plots. Distinctly the lowest SR rates were observed on bare soil. In the grass plots, SR was higher
in the collars inserted in the rows than in those located
between the rows (results not shown). The average of
these collars was used in the further analysis.
The performance of the TER and SR models are
shown in Figures 6 and 7, respectively. The model
was better able to reproduce the TER data from barley and potato than from grass (Figure 6). In peat soil
grass, where the fit between the data and the model
was poorest (r 2 =0.10), the model considerably overestimated TER after the harvest, suggesting a general
overestimation of soil respiration in the model. On the
whole, owing to the more radical variation of the plant
respiration in grass due to the harvests taking place
at the maximum LAI, the modelled estimates of the
total ecosystem respiration from grass do not follow
the measured data very well. However, the fit between
the observed and modelled soil respiration was relatively good (r 2 =0.66), particularly when taking into
account the fact that the parameters for modelling the
soil respiration were acquired from the TER data by
using the Equation (1).
The ratio of the soil and the total ecosystem respiration during the peak growth was studied by using the
measured input variables from the TER measurements
together with the modelled parameter values (Equation (1), Figure 4), and reproducing the soil and plant
dark respiration values. Only the TER measurements
conducted between the end of June and the beginning
of August, when PI was > 50% of its maximum or
LAI was > 3, were included. The results are presented
in Figure 8 as averages of 1017 calculations. According to these estimations, the soil respiration in the peat
soil was on average responsible for 62% of the TER at
the height of the summer (Figure 8). In sandy and clay
soils the average proportion of the soil respiration was
46 and 49%, respectively.

Discussion
Seasonal trend in the total ecosystem respiration
In both annual crops, barley and potato, the TER level
increased quite rapidly during the vegetative growth
and declined when the crop was maturing. Slightly

312

Figure 4. Values of Rs0 ( 95% confidence limits), the modelled parameter describing soil respiration (mg (CO2 ) m2 s1 ) at 10 C in
different plots. All relationships were highly significant (p < 0.0001).

lower TER rates were measured for the potato as

compared to the other crops. This was at least partly
explained by the smallest LAI for potato.
The increase in the TER rates during the vegetative growth stage was not only due to the increases
in shoot respiration but also to that in rhizosphere
respiration. During intense growth, the rhizosphere
respiration is enhanced by the photosynthetic activity
due to the allocation of assimilates into the roots and
soil (Kuzyakov and Cheng, 2001). This was demonstrated in field conditions by Rochette et al. (1992)
who found the soil respiration to correlate with the
daily net assimilation rates in a barley field. Buyanovsky et al. (1986) observed the highest root biomass
in wheat to occur at about the time of flowering.
The highest root respiration has been shown to correlate with the maximum amount of plant root matter
(Schssler et al., 2000). Thus the peak level in the
root and above-ground plant respiration are reached
at about same time. By using isotope techniques,
Rochette and Flanagan (1997) found that in a corn
field the contribution of the rhizosphere to the soil surface CO2 flux increased to a maximum of 60% at the
height of summer. At the beginning and the end of the
growing season its role was markedly less.
The decline in the TER rates started during the
reproductive growth stage, as the above-ground plant
and root respiration gradually ceased. The seasonal
changes in the plant dark respiration are known to
correlate with the changes in photosynthesis. According to McCree (1974), the dark respiration may be

divided into maintenance and growth respiration, the

latter being directly proportional to daily photosynthesis. For example, on 3 August, the PI of the sandy
soil barley had decreased to about 30% of its maximum, indicating that the plant respiration level had
probably decreased. This was actually observed in the
TER values. In the peat soil barley on 34 August, the
PI was still about 60% of the seasonal maximum and
the TER level was still quite high. In cereals, the translocation of the photosynthates is typically shifted from
the roots to the grain after flowering, and the supply
into the soil of easily degradable C gradually diminishes. The rhizosphere respiration thus also decreases,
as less new substrates are available. For similar PI values, a greater part of the TER probably comes from
soil respiration in the reproductive growth stage than
during the vegetative growth stage. In a winter wheat
field Buyanovsky et al. (1986) measured the highest
soil respiration rates after the harvest. The result was
attributed to the soil heterotrophic activity associated
with the decomposition of fresh roots, but also to high
soil temperatures, as the soil surface was no longer
protected by a plant canopy.
The highest average TER values obtained in this
study, ranging from 0.66 to 0.89 for barley and from
0.51 to 1.1 mg (CO2 ) m2 s1 for grass, were higher
than those measured by Gent and Kiyomoto (1992)
with the chamber method for wheat growing in fine
sandy loam, about 0.33 mg (CO2 ) m2 s1 . Nocturnal TER rates of about 0.2 mg (CO2 ) m2 s1 have
been measured with the eddy covariance method for

313

Figure 5. Averages of the measured soil respiration (SE) for each soil type in spring and autumn 2000. Average soil temperatures are shown
above the vertical bars. Soil temperatures varied between 8 and 15 C. Different letters denote significantly different SR rates (Tukey HSD test,
=0.05) (e.g., c differs significantly from a, b and d, but not from bc or cd).

wheat (Baldocchi, 1994). The reported respiration

rates were lower because these studies were conducted at night-time, when soil and air temperatures are
usually moderate, while our measurements were made
during the daytime with relatively high air (30 C)
and soil (20 C) temperatures. Additionally, the low
TER value in the study of Baldocchi (1994) may
be explained by the negligible amount of parent soil
respiration, as the soil contained only 0.5% organic
matter. Soil respiration was, however, not measured
in his study. In peat soil, typical daytime peak TER
values for barley measured using nearly the same
chamber construction as us were about 0.6 mg (CO2 )
m2 s1 , the maximum values being as high as 1.11
mg (CO2 ) m2 s1 (Maljanen et al., 2001). These are
in good agreement with our results for peat soil, the
highest average TER being 0.89 and 1.1 for barley and
grass, respectively.
The effect of the crop type on the soil respiration
In the case of each soil type, the parameter value for
the soil respiration, Rs0 , was highest in the grass and
lowest in the potato plots (Figure 4). Also in direct soil
respiration measurements (SR), the highest respiration
values were found in the grass plots (Figure 5). In the
clay soil, in autumn, however, the highest SR was observed in barley, which was probably due to the 4 C
higher average soil temperature. The root biomass was
not studied here, but the root biomass, rootshoot ratio and annual carbon input to the soil is normally
higher in perennial leys than in barley. The more living

roots there are in the soil, the greater is the amount of

CO2 produced by the roots. Consequently, the amount
of microorganisms utilizing the root exudates in the
rhizosphere also increases, enhancing the soil respiratory activity. In a carbon budget study performed in
an agroecosystem, twice as much carbon was allocated below ground in the case of grass as compared to
barley (Paustian et al., 1990). As a consequence, the
soil respiration was also higher in the grass than in the
barley. Tufekcioglu et al. (2001) found higher soil respiration rates in riparian buffers covered with different
grasses than in the adjacent crop fields. The difference
was attributed to soil organic carbon content, greater
fine root biomass and higher soil moisture content.
There are two possible explanations for the low Rs0
observed for potato. First, the potato plots were left
fallow the preceding summer; the barley plots were,
however, growing barley the year before. Fertilizers
and fresh carbon in the form of root exudates and straw
litter had thus been added to the soil in the barley
plots. Secondly, the PI, LAI and the duration of intense
growth were lower in potato than in grass or barley
during the growing season 2000. As can be seen in
Figure 2, the photosynthetically active period lasted
only about 50 days for potato, while for barley it lasted
over 80 days and in grass even longer. Thus the amount
of assimilates allocated into the soil was presumably
greatest in the grass and lowest in the potato plots. The
TER measurements conducted at nearly similar soil
temperatures on the bare barley and potato plots before crop emergence showed the lowest CO2 effluxes
on the potato plots (Figure 3c). This indicates that the

314

Figure 6. Measured and modelled (Equation (1)) TER values for each soil and plant type combinations (n = 1). The solid line represents the
1:1 relationship.

preceding years history may account for the observed

difference. No significant differences in the amount of
soil C, soil N, pH, bulk density or any other factor
could be found between the barley and potato plots on
any of the soil types that could explain the differences
in the parameter values describing soil respiration. It
is probable that these kinds of changes in the soil respiration equivalent to the SA used in the modelling
that are probably induced by the recent plant activity
do not affect the total soil organic C content, being
rather a qualitative change in carbon composition or
an induction of microbial activity than a noticeable
change in the amount of organic C.

The effect of the soil type on the soil respiration

According to both the soil respiration parameter (Rs0 )
derived from the modelled TER data, and the measured soil respiration (SR), the combined soil and
rhizosphere respiration in the agricultural peat soil in
the growing season was about 23 times as high as that
in the mineral sandy and clay soils. A similar pattern
was observed for each crop type. The higher peat soil
respiration is most likely to be due to the higher soil
organic carbon content. The total amount of organic
carbon in the uppermost 20 cm of the soil was about
4 times higher in the peat than in the mineral soils.
Soil respiration has been shown to correlate positively
with the soil organic carbon content (Franzluebbers

315

Figure 7. Measured and modelled (Rs in Equation (1)) SR values

for each soil type growing barley and grass (n = 2 4). The solid
line represents the 1:1 relationship.

Figure 8. Averages of the modelled soil and plant dark respirations

for all soil and crop combinations during the period of about 1 month
when the peak TER values were measured (n = 9 17). The calculations have been performed using the measured environmental
variables during the TER measurements.

et al., 2002; La Scala et al., 2000; Tufekcioglu et

al., 2001), but peat soils were not included in these
studies. There are also other factors influencing CO2
production and release from the soil. Soil texture affects the movement of water and gases in the soil and
may have profound effects on the soil CO2 efflux rates.
Peat soils are much more porous than mineral soils:
peat had the smallest bulk and particle density and the
largest porosity (Table 1). As a result, the diffusion of
CO2 and O2 is not very easily restricted, resulting in
better aeration and more favourable conditions for aerobic decay of soil organic C. On the other hand, peat
has a very good water retention capacity that offers
an excellent protection against soil drying (Pivnen,
1982), although in wet summers a good water retention capacity may lead to insufficient aeration in the

root zone. In the peak growing season our peat soil did
not overwet, but, owing to the high evapotranspiration,
stayed rather dry at a depth of 15 cm for a long period
despite days of relatively high rainfall.
The difference in Rs0 between clay and sandy soils
was small, but, however, the smallest Rs0 was always
observed in the clay soil. Nevertheless, the soil C content was slightly higher in the clay than in the sandy
soil. The lower soil respiration rate may be attributed
to the mineral structure of clay, which is known to
interact with soil organic C, thus protecting it from
decomposition. Clay also commonly has a lower hydraulic conductivity than sandy soil, which in the case
of high precipitation may lead to reduced gaseous diffusion in the clay. Soil respiration is suppressed by
decreased oxygen and high concentrations of accumulated CO2 that may to some extent inhibit the root and
microbial respiration rates (Bouma and Bryla, 2000).
In the present study, the percentage of water-filled pore
space was higher in the clay soil than in the sandy soil
in summer 2000 (Table 1). However, differences in the
soil respiration (SR) between the clay and sandy soils
were not found in this study.
The SR measurements were made at soil temperatures close to 10 C. The smallest CO2 effluxes were
measured on bare clay and sandy soils in the spring,
and the highest efflux rates on the peat soil growing grass in the autumn. Low soil respiration rates
on bare soil are explained by the limited substrate
supply into the soil. The contribution of plant roots
to the CO2 fluxes from soils has been found to vary
between 35 and 60% (Andrews et al., 1999; Rochette
and Flanagan, 1997; Silvola et al., 1996b). The difference in the SR values between the bare peat and
mineral soils (about 0.07 mg (CO2 ) m2 s1 ) (Figure
5) was nearly equal to the corresponding difference in
the Rs0 (0.0580.067 mg (CO2 ) m2 s1 ). We suggest
that this difference roughly represents the oxidation of
older peat material, assuming that the turnover of fresh
C derived from agricultural activities is of a similar
magnitude regardless of soil type. The assumption is,
of course, rather rough, and other factors may lie behind the observed differences in Rs0 between the soil
types. One such factor is the inter-site difference in
PI. As plant growth strongly controls the rhizosphere
respiration, differences in the growth variables should
influence the soil respiration as well.
According to the model calculations, during the
peak of the growing season about half of the total ecosystem respiration originates from the soil respiration,
the proportion being highest on the peat soil (62%).

316
Especially on the mineral soils, where the proportion
of Rs to TER averaged 46 and 49% for sandy and
clay soils, respectively, the estimates agree quite well
with the results of Franzluebbers et al. (2002), who
recently found this ratio to be 40% during the peak
growing season on a tallgrass prairie. This consistency in the results is encouraging and suggests that the
model used in this paper works fairly well. The calculated shoot respiration was always highest in barley
and lowest in potato, probably owing to the differences
in PI between the crops.
In other studies, peak summer soil CO2 effluxes
ranging from 0.14 to 0.22 mg (CO2 ) m2 s1 on
bare peat soil and from 0.28 to 0.44 mg (CO2 ) m2
s1 on peat soil growing grass have been measured
(Nyknen et al., 1995). These values, measured during higher soil temperatures than the SR in our study,
were similar to or higher than our SR results from peat
soil, which were about 0.11 mg (CO2 ) m2 s1 on
bare peat soil and 0.210.36 mg (CO2 ) m2 s1 on
peat soil growing grass. Rochette et al. (1992) measured higher soil CO2 efflux rates for fallow than for
barley. In their study the CO2 effluxes in clay loam
with a soil C content of < 2.4% varied between about
0.05 and 0.35 mg (CO2 ) m2 s1 , but the soil temperatures were almost always higher than in our SR
measurements. Taking into account the different soil
temperatures, these results are consistent with our SR
results from the mineral soils, ranging from 0.02 to
0.17 mg (CO2 ) m2 s1 in different treatments. In the
study of Rochette et al. (1992), the higher soil respiration for fallow than for barley resulted mainly from the
lower summer-time soil moisture and temperature for
barley. Rather small differences in soil CO2 effluxes
between different soil types were found by Koizumi
et al. (1999), who included in their study peat, sandy
and clay soils similar to ours. The soil respiration at a
soil temperature of 10 C was close to 0.07 mg (CO2 )
m2 s1 for each soil type. However, due to different
temperature and soil moisture responses in the soil carbon dioxide efflux, the 5-month cumulative respiration
fluxes estimated with a statistical model were 1.67,
0.98 and 1.31 kg CO2 m2 5 months1 in the peat,
sandy and clay soils, respectively; the peat soil thus
released the largest amount of CO2 , at least during the
growing season.
In conclusion, the CO2 efflux from a peat soil
which had been in agricultural use for at least a century was more than twice as great as that from mineral
soils. This result presumably has implications for the
estimates of the net CO2 balances of these different

ecosystems, which will be quantified in the near future. Knowing the CO2 balance of cultivated peat soils
is important, as new ways of mitigating global climate
change through enhancing the land carbon sink are
being continuously sought. In countries having a high
proportion of agricultural peat soils, the improved soil
carbon accumulation obtained through changes in cultivation practices and land use may be small compared
to the total net CO2 emissions from agricultural soils.

Acknowledgements
We thank Johanna Haapala and Mikko Leimu for field
assistance and Risto Tanni for the maintenance of the
experimental fields and for supporting us with the
background data. We are also grateful to Agrifood
Research Finland for providing the facilities for the
work and to the Universities of Joensuu and Kuopio
for providing the technical equipment. This work was
a part of the Finnish Global Change Research Program (FIGARE), and was funded by the Academy of
Finland.

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Section editor: R. Merckx

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