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Anaerobic decomposition of tropical soils and plant material: Implication for

the CO2 and CH4 budget of the Petit Saut Reservoir

Article  in  Applied Geochemistry · August 2008

DOI: 10.1016/j.apgeochem.2008.04.001

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 Applied Geochemistry 23 (2008) 2272–2283

Contents lists available at ScienceDirect

Applied Geochemistry
journal homepage: www.elsevier.com/locate/apgeochem

Anaerobic decomposition of tropical soils and plant material:

Implication for the CO2 and CH4 budget of the Petit Saut Reservoir
Frédéric Guérin a,b,*, Gwenaël Abril a, Alexis de Junet a,1, Marie-Paule Bonnet c
a
EPOC, UMR CNRS 5805, Université Bordeaux1, Avenue des Facultés, F-33400 Talence, France
b
Laboratoire d’Aérologie, UMR UPS/CNRS 5560, OMP, 14, Avenue Edouard Belin, F-31400 Toulouse, France
c
Laboratoire des Mécanismes et Transferts en Géologie, UMR UPS/CNRS/IRD 5563, OMP, 14, Avenue Edouard Belin, F-31400 Toulouse, France

a r t i c l e i n f o a b s t r a c t

Article history: Tropical hydroelectric reservoirs contribute significantly to atmospheric CH4 and CO2 emis-
Received 22 October 2007 sions. To evaluate the contribution of the mineralization of the flooded soils and biomass to
Accepted 2 April 2008 these atmospheric gas emissions, field and laboratory experiments were conducted. Cores
Available online 14 April 2008
were retrieved inform the littoral zone of the Petit Saut Reservoir (French Guiana), flooded
Editorial handling by G.M. Filippelli
10 a prior to sampling, and different soils and plant material in the tropical forest sur-
rounding the reservoir. All the samples were flooded and incubated in anoxic conditions
in the dark at 30 °C. The potential CH4 and CO2 production rates were determined. Soils
and plant material from the tropical forest were incubated over one year and the produc-
tion measurements were performed at a frequency of 1–5 months. Methane and CO2 pro-
duction rates of soils and littoral sediments were linearly correlated to the organic C (OC)
content of the slurries. The slopes of the relationships were 2:6  102  5:6 
1 1
101 nmol ðCH4 Þ g1OCðdryÞ h and 3:5  102  6:9  101 nmol ðCO2 Þ g1OCðdryÞ h . For plant
material, no relationship between the production rates and the OC content was found
and on average over the year of incubation, production rates were 2:4  103  1
1 1
103 nmol ðCH4 Þ g1OCðdryÞ h and 3:9  103  5  103 nmol ðCO2 Þ g1
OCðdryÞ h , which is one
order of magnitude higher than the mineralization of the soils and sediments. Extrapolated
at the scale of the Petit Saut Reservoir over 10 a, these results show that the mineralization
of the soil and the plant biomass initially flooded contributed to 75–95% of the total C emis-
sions to the atmosphere since the flooding of the reservoir. Methane: CO2 molar ratios were
3 times higher for anaerobic decomposition than atmospheric emissions, quantitatively
consistent with aerobic CH4 oxidation.
Ó 2008 Elsevier Ltd. All rights reserved.

1. Introduction is et al., 2004) and tropical regions (Rosa and Schaeffer,

In recent years, there has been increasing concern about
CO2 and CH4 emissions from artificial hydroelectric reser-
voirs in boreal (Rudd et al., 1993; Duchemin et al., 1995;
Kelly et al., 1997; Huttunen et al., 2002), temperate (Soum-
* Corresponding author. Present address: Max-Planck-Institut für Ter-
restrische Mikrobiologie, Karl-von-Frisch-Strasse, D-35043, Marburg,
Germany. Tel.: +49 6421 178 832; fax: +49 6421 178 809.
E-mail address: guerin@staff.uni-marburg.de (F. Guérin).
1
Present address: Institut de Recherche et Développement (IRD),
Université de La Réunion, BP 172, F-97492 Sainte Clotilde Cedex, France.
1994; Abril et al., 2005; dos Santos et al., 2004; Guérin
et al., 2006; Kemenes et al., 2007). On a global scale, St.
Louis et al. (2000) estimated that reservoirs, and particu-
larly tropical reservoirs, could contribute significantly to
anthropogenic CH4 emissions.
Although the large atmospheric emissions that imme-
diately follow impoundment of hydroelectric reservoirs
are commonly attributed to the degradation of flooded
soil organic matter (OM) and plant biomass (Rudd et al.,
1993; Kelly et al., 1997), the sources of C that drive the
long term emissions is still under debate. In two old bor-
eal reservoirs (>20 a) in Finland (Huttunen et al., 2002)

0883-2927/$ - see front matter Ó 2008 Elsevier Ltd. All rights reserved.
doi:10.1016/j.apgeochem.2008.04.001
 F. Guérin et al. / Applied Geochemistry 23 (2008) 2272–2283
and in young experimental reservoirs in Canada (Bodaly
et al., 2004), the CO2 and CH4 emissions were related to
the turnover of autochthonous and allochthonous OM in
the impoundments rather than to the amount of C in
flooded mineral soil and peat. In contrast, in a tropical res-
ervoir, Abril et al. (2005) showed, using a C mass balance,
that 10 a after reservoir filling the mineralization of the
flooded soils and above-ground biomass was still the ma-
jor process contributing to gaseous emissions. The C mass
balance of Abril et al. (2005) based on atmospheric fluxes
did not directly quantify the amount of CO2 and CH4 pro-
duced during organic matter degradation since a large
part of the CH4 produced was oxidized to CO2 in the oxic
epilimnion (Guérin and Abril, 2007) and CO2 was also in-
volved in photosynthesis and respiration processes in the
water column.
In lake sediments, the degradation rates of organic mat-
ter and the resulting CO2 and CH4 benthic fluxes can be ob-
tained by vertical profiles in sediment pore waters or from
benthic chamber experiments. In many reservoirs, these
approaches cannot be performed due to the presence of
tangled trunks and branches that preclude the use of box
corers and benthic chambers.
Taking into account these difficulties, the CO2 and CH4
production rates were investigated during the degradation
of organic matter flooded by an in vitro approach. Soils and
plant samples were taken in the tropical forest around the
Petit Saut Reservoir (PSR) and were incubated with water
in anoxic conditions. The potential production rates of
CO2 and CH4 were then followed over one year. Cores were
also retrieved from the littoral zone of the reservoir in
2003 and 2004 (10 a after the reservoir filling), and were
incubated in the same conditions for comparison. These re-
sults were then extrapolated to estimate the evolution of
CO2 and CH4 production within the reservoir over 10 a
and to compare it with the total atmospheric emissions
from the reservoir (Abril et al., 2005) as well as the mass
balance of pelagic aerobic CH4 oxidation (Guérin and Abril,
2007).
2. Materials and methods
2.1. Study site
The Petit Saut dam was constructed in the tropical for-
est of French Guiana on the Sinnamary River, 80 km up-
stream from its mouth to the Atlantic Ocean. The PSR
started to be filled in January 1994 and it covers 100 km
of the Sinnamary River course. At its maximum level of
35 m, first reached in July 1995, 365 km2 of un-cleared
tropical forest were flooded. The terrestrial OM flooded at
PSR was composed of soil, litter, tree trunks as well as
leaves that fell into the water a few months after flooding.
Tree trunks remained standing, apparently unaffected by
the flooding. After flooding, the organic matter pool under-
went mineralization under high temperature. This led to a
chemical stratification of the lake water body with an oxic
epilimnion and an anoxic CO2- and CH4-rich hypolimnion
and high CH4 and CO2 emissions (Galy-Lacaux et al.,
1997). This situation has persisted to the present (Abril
et al., 2005; Guérin and Abril, 2007).
2273
2.2. Sampling methodology
In March 2003, soil and plant samples were taken in the
primary forest surrounding the PSR. The soil samples were,
from the top to the deeper layers, (1) the litter (Lit) (5 cm
thick), (2) the organic-rich soil (OS) which was 10 cm thick
and had a brown–black colour and (3) the first 20 cm of the
red tropical lateritic soil (LS). A suspended soil (SS) was
also sampled on a tree 2-m above the ground. Suspended
soils develop on the branches of trees due to both deposi-
tion of dead leaves from the tree itself and due to the accu-
mulation of the decaying material of epiphytes which grow
in those suspended soils. Leaves of Ficus (Trigona Linnaeus),
palm tree (Astrocaryum sp.) and epiphyte (Anthurium sp.)
were also collected. Taking into account the natural biodi-
versity of forest in French Guiana (more than 3000 tree
species), it is impossible to determine if these plants are
representative of the total flooded forest biomass. They
were chosen based on the different structures and tex-
tures, which presumably lead to different CO2 and CH4 pro-
duction rates.
In addition to this material from the forest (see above),
sediments (flooded soils + recently sedimented OM) were
also collected in the littoral zone of the PSR, 20 km up-
stream of the dam. Three 20 cm-long sediment cores were
collected under 50-cm of water during the dry season
(December 2003) and three cores during the wet (May
2004) season. The sediments retrieved during the dry sea-
son were always submerged as sampling was performed
when the lake was at its minimum level. In contrast, the
sediment collected during the wet season undergoes sea-
sonal drying. The sediment cores were cut in 2 cm slices
within 2 h of sampling. The samples from one core were
used for organic C (OC), isotopic composition and C/N ratio
analyses. In this case, the sediments were immediately
dried at 60 °C and stored at ambient temperature until
analysis. A sub-sample of each slice was immediately
sealed in a pre-weighed vial for determination of porosity.
The two other cores were used for determination of CH4
concentration and subsequent incubation for CO2 and
CH4 production. Each slice of these two cores was trans-
ferred into pre-weighed glass vials containing 10 mL of
deionised water that were quickly closed with rubber stop-
pers, vigorously shaken to break up the sediment and en-
sure equilibration between and liquid phase before
analysis (Abril and Iversen, 2002).
2.3. CO2 and CH4 production rates
Soil samples (200 g) were ground and homogenized
and plant leaves were finely chopped. Five sub-samples
of soil and plant material of each type were then redistrib-
uted to five replicate vials for the preparation of slurries.
The composition of the slurries is given in Table 1. It was
expected that most of methanogenic micro-organisms
present in the PSR originated from the forest soil that
was flooded and plant material was probably free of meth-
anogens. Therefore, the latter material was incubated with
a mixture of lateritic soil and organic soil (1:1 w/w) in or-
der to obtain environmental conditions as close as possible
to that occurring in situ during and after impoundment.
2274 F. Guérin et al. / Applied Geochemistry 23 (2008) 2272–2283
Table 1
Composition of the slurries for determination of CO2 and CH4 production
rates
Incubation Composition of the slurry
Suspended soil (SS) 10 g of soil + 10 mL DW
Litter (Lit) 10 g of soil + 10 mL DW
Lateritic (LS) 10 g of soil + 10 mL DW
Organic soil (OS) 10 g of soil + 10 mL DW
Lateritic and organic soil 5 g of LS + 5 g of OS + 10 mL DW
mixture (LS–OS)
Ficus leaves (Ficus) 9 g of LO + 1 g of leaves + 10 mL DW
Epiphyte leaves (Epi) 9 g of LO + 1 g of leaves + 10 mL DW
Palm tree leaves (Palm) 9 g of LO + 1 g of leaves + 10 mL DW
Abbreviation: DW, deionized water.
For incubations, the slurries were placed into five serum
bottles (160 mL), closed with butyl stoppers and alumin-
ium crimps and flushed with N2 over 30 min. The produc-
tion rates of CO2 and CH4 were determined by anoxic
incubation of the slurries at 30 °C in the dark. Incubations
were performed without agitation to avoid the destruction
of syntrophic microbial associations involved in methano-
genesis (Dannenberg et al., 1997). Before each analysis,
serum bottles were vigorously shaken for about 30 s to en-
sure equilibration between the liquid and gas phase. From
April 2003 to March 2004 this operation was repeated 7
times at a frequency of 1–5 months with the same serum
bottles in order to evaluate the evolution of the production
rates with time. One week before the determination of the
production rates, the serum bottles were flushed with N2
over 30 min to eliminate the overpressure, accumulated
CH4 and CO2 and any volatile compounds inhibiting meth-
anogenesis (Williams and Crawford, 1984) and anaerobi-
cally produced CO2 (Magnusson, 1993) that accumulated
in the headspace. When no measurements were performed
for more than one month, serum bottles were also flushed
with N2. Total CH4 and CO2 concentrations in the vials were
measured in the headspace of the serum bottles 3-times a
week to calculate the production rate, as the slope of the
concentration versus time. For the incubations of soils
and soils + plant leaves, production rates are the average
of the production rates of the five replicates (±SD). For
soils, CO2 and CH4 production rates reported hereafter
1
are production rates per gram of soil ðnmol g1 soil h Þ
whereas for the plant material, the production rates are
those attributed to the mineralization of plant material
only. Therefore, the CO2 and CH4 production of the plant
material was calculated by subtracting the contribution
of the LS–OS mixture to the LS–OS–vegetal mixture. At
times when production rates were not measured, they
were calculated by linear interpolations of existing data.
Hereafter, yearly average production rates refer to the
average of measured and calculated production rates.
Production rates were also determined by incubation in
anaerobic conditions of the core slices. After the determi-
nation of CH4 concentration in pore water of the flooded
soils within 4 h of sampling, the headspace of the vials con-
taining a 2 cm-thick slice with 10 mL of deionised water
was flushed with N2. Total CO2 and CH4 concentrations in
the vials (110 mL) were determined by taking gas samples
(duplicates at each depth) in the headspace every two days
over a week in order to determine the production rates.
Incubations were performed at 30 °C in the dark and with-
out agitation. As the results obtained from duplicate cores
were different by one order of magnitude in some cases,
they are described separately.
2.4. CO2 and CH4 analysis
Methane and CO2 concentrations in the cores and dur-
ing incubations were obtained with a gas chromatograph
equipped with a flame ionization detector and a thermal
conductivity detector, respectively. Calibration of the
detectors was performed using Air Liquid gas standards
(pCH4 = 10 and 1000 ppmv; pCO2 = 40,000 ppmv). Meth-
ane and CO2 concentrations together with the porosity,
the volume of water added and the specific gas solubility
for CH4 (Yamamoto et al., 1976) and CO2 (Weiss, 1974)
were used for calculation of total CH4 and CO2 content in
the vials and the CH4 concentrations in interstitial waters.
2.5. Organic carbon, C/N and d13C analyses
For OC analysis, dried soil samples were ground and
homogenized. Dry samples were acidified with HCl 2 N to
remove carbonates and dried at 50 °C overnight before
analysis. A Carbon LECO CS-125 analyzer was used based
on direct combustion in an induction furnace and infrared
absorption determination of the CO2 produced. Analyses
were done in duplicate.
Carbon/N weight ratios on sediments were measured
by a CN Elemental Analyzer (Flash Elemental) after Nie-
uwenhuize et al. (1994). Basically, it relies on a catalytic
combustion (1020 °C) of OM with O2 and quantification
of each gas formed by combustion (CO2 and N2) with a
catharometer. Analyses were done in triplicate.
Sub-samples of sediments, soils and plant material
(1 g), were acidified (Nieuwenhuize et al., 1994) before
the determination of the d13C of OM. Measurements were
performed with a 2500 CARLO-ERBA CN Elemental Ana-
lyzer coupled to an Isotope Ratio Mass Spectrometer. Car-
bon isotope ratios are expressed in the delta notation
(d13C) relative to Pee Dee Belemnite. Calibration of the
mass spectrometer was done with two standards at 10‰
and 35‰.
3. Results
3.1. Sediments in the reservoir
The vertical profiles of OC, C/N and d13C obtained in the
sediment core collected at the littoral station are shown in
Fig. 1. Results obtained for the litter and organic soil sam-
ples are also reported in Fig. 1 for comparison. In the first
8 cm of the core, OC decreased from 14% to 1% and the
C/N ratio decreased from 31% to 17% and remained con-
stantly low (Fig. 1a and b). The d13C values decreased from
29‰ to 30.5‰ in the first 4 cm and then increased pro-
gressively to 25.7‰ at 20 cm depth. At the sediment sur-
face, OC was lower than in the Lit and OS (Fig. 1). The C/N
ratios of the surface sediments were the same as in the Lit
whereas the C/N ratio in the OS was the same as in the
 F. Guérin et al. / Applied Geochemistry 23 (2008) 2272–2283 2275

a OC (%, w/w) b C/N c δ 13C (%)

3 6 9 12 15 18 21 14 18 22 26 30 -30.5 -29.5 -28.5 -27.5 -26.5 -25.5
Depth (cm) 0

-5
c
-10

-15

-20

Fig. 1. Depth profiles of (a) organic C (OC), (b) C:N ratio, and (c) d13C–OC and in littoral cores retrieved in December 2003 at the PSR.

a CH4 (mmol.L -1) b CH4 production (nmol.g -1.h-1)

0.5 1.0 1.5 2.0 2.5 3.0 10 20 30 40 50
0 0
-2 -2
-4 -4
Depth (cm)

-6 -6
-8 -8
-10 -10
-12 -12
-14 -14
-16 -16
-18 Dec. 2003 -18
-20 -20

c 0
0.1 0.2 0.3 0.4 0.5 0.6 d 0
10 20 30 40 50

-2 -2
-4 -4
Depth (cm)

-6 -6
-8 -8
-10 -10
-12 -12
-14 -14
-16 -16
-18 Dec. 2003 -18
-20 -20
CO2 production (nmol.g -1.h-1)
e 0
0.1 0.2 0.3 0.4 f 0
50 100 150 200 g 0
10 20 30 40

-2 -2 -2
-4 -4 -4
Depth (cm)

-6 -6 -6
-8 -8 -8
-10 -10 -10
-12 -12 -12
-14 -14 -14
-16 -16 -16
-18 May 2004 -18 -18
-20 -20 -20

h 0
0.01 0.02 0.03 0.04 0.05 0.06 i 0
1 2 3 4 j 0
10 20 30 40
-2 -2 -2
-4 -4 -4
Depth (cm)

-6 -6 -6
-8 -8 -8
-10 -10 -10
-12 -12 -12
-14 -14 -14
-16 -16 -16
-18 May 2004 -18 -18
-20 -20 -20

Fig. 2. Depth profiles of CH4 concentration (a, c, e and h), CH4 (b, d, f and i) and CO2 (g and j) production rates in littoral cores retrieved in December 2003
and May 2004 at the PSR.

2–8 cm interval of the core. The d13C of Lit (28.7‰) and zone of the PSR (2 profiles for each season) (Fig. 2). In
OS (29‰) were similar to those in the first centimetres December (Fig. 2a), CH4 concentrations reached
of the core (Fig. 1c). 2.86 mmol L1 at 5 cm. At these depths and temperature,
No common features were found for the depth profiles the maximum solubility of CH4 is around 1.5 mmol L1,
of CH4 concentrations in pore water obtained in the littoral which indicates the existence of CH4-rich bubbles. For all
2276 F. Guérin et al. / Applied Geochemistry 23 (2008) 2272–2283

other profiles, concentrations were 1–2 orders of magni- The evolution of the monthly CO2 and CH4 production
tude lower (Fig. 2b–d). rates over one year after flooding the soils and the leaves
The vertical profiles of CH4 (Fig. 2b, d, f and i) produc- in anoxic conditions is shown in Fig. 3. During the 1-a incu-
tion rates revealed that CH4 production occurred only in bation period, the CO2 production rates of the LS were 1–2
the first 8 cm of the flooded soils of the littoral zone. Dee- orders of magnitude lower than those of SS, OS, LS–OS and
per, production rates were below detection limit. Carbon Lit (Fig. 3). During the first 6 months of incubation, CO2
dioxide production (Fig. 2g and j) occurred in the first production rates of all the soils followed the same pattern
12 cm and was below detection limit below this. From (Fig. 3). The CO2 production rates decreased sharply (by a
one core to another, the maximum CH4 production rates factor of 2–4) after 3 months of incubation. Afterwards,
1
varied between 3.6 and 200 nmol g1 sed h without any sea- CO2 production rates decreased by a factor of 2–3 for OS,
sonal trend (Fig. 2b, d, f and i). By contrast, for CO2, the ver- LS and LS–OS whereas they increased by a factor of 2–3
tical profiles of production rates obtained with the for Lit and SS (Fig. 3). The yearly averages of CO2 produc-
sediment of the two replicate cores in December were of tion rates ranged from 4 ± 4 (LS) to 92  58 nmol g1 soil (SS)
the same order of magnitude (Fig. 2g and j). and were intermediate for the Lit, OS and LS–OS (Table 2).
Methane concentrations and production rates of CO2 For the plant material, CO2 production rates were 2 or-
and CH4 varied widely from one core to another. This re- ders of magnitude higher than the production rates of soils
flects the large spatial heterogeneity of the factors control- (Fig. 3). The CO2 production rates by plants were maximum
ling anaerobic processes in the flooded soils of the littoral on the second month of incubation and then decreased for
zone of the PSR. all plant material (Fig. 3). After the second month, the CO2
production rates decreased and stabilized to a rate of
1
3.2. Soils and plants from the forest  50 nmol g1 leaf h for the Palm and Epi after one year of
incubation. For the Ficus, CO2 production rates were at a
The OC content of the different soils used for incubation minimum 3–4 months after the beginning of the incuba-
was different (Table 2), the suspended soil (SS) having by tion and the increased again to reach a value up to
1
far the highest OC value (w/w) (42%). From the top to the 2600  67 nmol g1 leaf h (Fig. 3). Yearly averages of CO2
bottom of the soil sequence, the OC content decreased production rates for plant material ranged from
from 20% to 0.37% for the litter (Lit) and the lateritic soil 344 ± 551 (Epi) to 1889  3882 nmol g1 leaf (Ficus), that is
(LS), respectively (Table 2). The organic soil (OS) which cor- 1–2 orders of magnitude higher than the production rates
responds to an intermediate state in the pedogenesis pro- of the soils (Table 2).
cess contained 17% of OC. The LS–OS mixture had an For all soils (OS, LS, LS–OS and Lit) and during the whole
intermediate value (8.4%). The organic content of the plant experiment, CH4 production rates ranged from
1
material was higher than in the soils and was almost con- 0:016  0:017 nmol g1 soil h (LS) to 195  31 nmol g1 soil
stant around 46% for all plants. After the addition of 1 g of (Lit). Note that the CH4 production rate of the SS was below
plant leaves to 9 g of LS–OS, the slurries containing plant detection limit during the first week of incubation. Whereas
material had an organic content around 12% (Table 2). CH4 production rates of OS and LS–OS were maximum after
1 month of incubation, the highest CH4 production rates
over the experiment were reached after 3 and 5 months
Table 2 for the LS–OS and the Lit, respectively. After these periods
Organic carbon (OC) content, average of CO2 and CH4 production rates of high methanogenic activity, CH4 production rates de-
obtained during the year of incubation, molar ratio of production rates of creased for all the soils. Yearly averages of CH4 production
CH4 to CO2 and OC (%) mineralized during one year of incubation
rates were an order of magnitude higher for the Lit, OS
Slurry Initial OC % Average production CH4:CO2 C and LS–OS than they were for the LS and the SS (Table 2).
(w/w) rate nmol G1 h1 loss%a The CH4 production rates of the three different types of
CO2 CH4 plant leaves were up to an order of magnitude higher than
Soilsb the production rates of soils (Fig. 3). The temporal patterns
SS 42.0 87 ± 63 6±4 0.001 1 of the CH4 production rates show a contrast for the three
Lit 20.5 79 ± 43 49 ± 70 0.071 7 different types of plant leaves (Fig. 3). The maximum CH4
OS 17.0 59 ± 87 28 ± 39 0.333 6 production rate of Palm was reached during the second
LS 0.4 4±3 0.17 ± 0.18 0.003 14
month and then, the methanogenic activity decreased by
LS–OS 8.4 48 ± 72 20 ± 24 0.015 9
2 orders of magnitude. For Epi, the maximum CH4 produc-
Vegetal leavesc
tion rate was obtained during the first week of incubation.
Ficus 45.0 1889 ± 3882 719 ± 1249 0.050 79.0
Palm 44.5 1057 ± 3223 525 ± 781 0.250 40.0 Afterwards, the methanogenic activity decreased and
Epi 47.5 344 ± 551 870 ± 1153 1.667 49.0 reached a minimum 5 months after the beginning of the
incubation. During the two final months, CH4 production
The composition of the slurries is given in Table 1.
a
Calculated by the difference between the initial OC content and the rates were 10–20 times higher than the minimum CH4 pro-
cumulated production of CO2 and CH4 and the weight of soil in the slurry duction rates of Epi. For Ficus, two peaks of high methano-
(10 g). genic activity were obtained, one after 1 month and one
b 1
Production rates for soils are expressed in nmol g1
soil h .
c
after 5 months of incubation. Afterwards, the CH4 produc-
Calculated by subtracting the contribution of 9 g of the LS–OS to CO2
and CH4 production to the total CO2 and CH4 production by the LS–
tion rates decreased by 2 orders of magnitude. Yearly aver-
OS + vegetal leaf mixture. The difference is attributed to 1g of plant ages were 1–3 orders of magnitude higher than the
1
material and the production rates are expressed in nmol g1 leaf h . production rates of soils (Table 2).
 F. Guérin et al. / Applied Geochemistry 23 (2008) 2272–2283 2277

SS Lit
350 250 350 250

CO2 (nmol gsoil h-1)

CO2 CO2

CH4 (nmol gsoil h-1)

CO2 (nmol gsoil h-1)
300 300

CH4 (nmol gsoil h-1)

CH4 200 CH4 200
250 250
200
150 200 150

150 100 150 100

100 100
50 50
50 50
0 0 0
0
0 1 2 3 4 5 6 7 8 9 10 11 12 13 0 1 2 3 4 5 6 7 8 9 10 11 12 13

OS Ficus
350 250 20000 5000

CO2 (nmol gleaf h-1)

CO2 CO2

CH4 (nmol gleaf h-1)

CO2 (nmol gsoil h-1)

CH4 (nmol gsoil h-1)

300 CH4
CH4 200 4000
15000
250
200 150 3000
10000
150 100 2000
100 5000
50 1000
50
0 0 0 0
0 1 2 3 4 5 6 7 8 9 10 11 12 13 0 1 2 3 4 5 6 7 8 9 10 11 12 13

LS Palm
350 350 CO2 (nmol gleaf h-1)
20000 5000
CO2 CO2
CH4 (nmol gsoil h-1)

300

CH4 (nmol gleaf h-1)

300
CO2 (nmol gsoil h-1)

CH4 250 CH4 4000

250 15000
200
200
150 3000
150 1.25 10000
15 1.00 2000
10 0.75
0.50 5000
5 1000
0.25
0 0.00
0 0
0 1 2 3 4 5 6 7 8 9 10 11 12 13 0 1 2 3 4 5 6 7 8 9 10 11 12 13

LS-OS Epi
350 250 20000 5000
CO2 (nmol gleaf h-1)

CO2 CO2

CH4 (nmol gleaf h-1)

CH4 (nmol gsoil h-1)

300
CO2 (nmol gsoil h-1)

CH4 200 CH4 4000

250 15000

200 150 3000

10000
150 100 2000
100
50 5000
1000
50
0 0 0 0
0 1 2 3 4 5 6 7 8 9 10 11 12 13 0 1 2 3 4 5 6 7 8 9 10 11 12 13
Month Month

Fig. 3. Evolution over one year of monthly production rates of CO2 and CH4 during the incubation of flooded soils (SS, LS, OS, Lit, LS–OS) and plant material
(Ficus, Palm, Epi). For vegetal material, the production rates are those attributed to the mineralization of plant material only (see text). Note the different
scales for the different panels.

The temporal patterns of the CH4:CO2 molar ratios transiently, the major end product. From one soil to another,
(Fig. 4) were used to evaluate the relative contribution of the yearly averages of the CH4:CO2 ratios varied by 2 orders
CO2 and CH4 production during the incubations. A CH4:CO2 of magnitude (Table 2). The low CH4:CO2 ratio of the SS was
molar ratio <1 indicates that, during the mineralization pro- due to its very low capacity to produce CH4 and to its high
cess, more than 50% of the C is converted to CO2 and vice ver- CO2 production rates (Table 2). The degradation of Epi pro-
sa. For all incubations during the whole period, the CH4:CO2 duced more CH4 than CO2 as the yearly average CH4:CO2 ra-
molar ratios ranged from 4  104 to 50. This large range of tio of this slurry is >1 (Table 2).
values reveals (1) large temporal variations for a given sub-
strate and (2) important differences from one substrate to 4. Discussion
another (Fig. 4). Globally, except for Epi, the fraction of the
mineralized C that was transformed in CH4 increased signif- 4.1. Controlling factors of CH4 and CO2 production by soils
icantly during the first 3–4 months. Afterwards, the fraction and vegetation
of C mineralized to CO2 increased for Lit, SS, Ficus and LS or
remained constant for OS, LS–OS and Palm. For LS–OS, Lit The high CO2 production rates at the beginning of the
and all incubations with plant material, CH4 was, at least incubation probably resulted from the oxidation of OM
2278 F. Guérin et al. / Applied Geochemistry 23 (2008) 2272–2283
Ficus Epi SS
Palm
Lit OS
45
35
25
15
5 consisted of 0.1–35% CH4. Assuming that the production of
2.0
CH4/CO2
1.5
1.0
0.5
0.0
1 2 3 4 5 6 7 8 9 10
Month
Fig. 4. Evolution over one year of monthly CH4:CO2 molar ratios during
the incubation of flooded soils (SS, LS, OS, Lit, LS–OS) and vegetal material
(Ficus, Palm, Epi).
2
by bacteria using NO 3 , Fe(III), Mn(IV) and SO4 as electron
acceptors (Conrad, 1989; Froelich et al., 1979). After 2
months of incubation, the net decrease of the CO2 produc-
tion rates could be attributed to the depletion of the elec-
tron acceptors and the occurrence of methanogenic
processes. When environmental conditions are favorable
to methanogenesis (anoxia with depletion of the alterna-
tive electron acceptors), the methanogenic population in-
creases and becomes more and more active (e.g., Le Mer
and Roger, 2001; Conrad, 1989). However, methanogenesis
can occur simultaneously with SO2 4 -reduction (Oremland
et al., 1982) or Fe(III) reduction (Yao et al., 1999). In terres-
trial aquatic ecosystems, CH4 is mainly produced by two
pathways: via the reduction of CO2 by H2 or via the reduc-
tion of acetate which produces CH4 and CO2 (e.g., Krüger
et al., 2002). Krüger et al. (2002) showed that following
the flooding of rice field soils, CH4 is mainly produced by
reduction of CO2 before being mainly produced by acetate
reduction at the end of the growing season. The concomi-
tant occurrence of the consumption of alternative electron
acceptors and the prevalence of methanogenesis via CO2
reduction during the first months of the present incubation
could explain the apparent decrease of CO2 production
(Fig. 3). For SS and Ficus, the CO2 production rates in-
creased again at the end of the experiment for a reason
which cannot be explained. Between 2 and 6 months after
the beginning of the experiments, CH4 production rates
were maximum and then decreased to low values as previ-
ously observed by Yao et al. (1999) during long term incu-
bation of 16 rice field soils. In the experiment of Yao et al.
(1999), the methanogenic activity also reached a steady
state as observed for most of the present incubations (ex-
cept Ficus and Epi).
The maximum and steady state production rates of CO2
and CH4 found by Yao et al. (1999) for rice fields incubated
at 25 °C were higher than the production rates by the LS
and in the same range of the production rates obtained
with Lit, the OS, the SS and LS–OS throughout the incuba-
LS
tion period. The CH4 and CO2 production rates obtained by
LS-OS
Magnusson (1993) during anoxic incubation at 16 °C of
forest and peat soils from a boreal environment over
120–140 days were similar to the steady state production
rates obtained with Lit, OS, LS, SS and LS–OS. Yao et al.
(1999) found that the gaseous release of their incubations
non-CH4 hydrocarbons were negligible, it can be estimated
that during the experiment of Magnusson (1993), the gas-
eous release consisted of <0.1–28% of CH4 after 120–140
days of incubation. Based on the same assumption, during
the present soil incubations the proportion of CH4 in the
gaseous release ranged from <0.1% to 55%, which is up to
twice as high as for boreal forest and peat soils and 1.6
times higher than for rice field soils. These facts reveal that
whether the degradability of the OM from tropical forest
11 12 soils is similar to that of rice field soils, the mineralization
of tropical soils produces proportionally more CH4 than
rice field soil. The lower CO2 and CH4 production rates
and the lower proportion of CH4 in the total gaseous re-
lease obtained by Magnusson (1993) than those obtained
by Yao et al. (1999) and during this study are probably
due to the lower incubation temperature.
As previously observed during anoxic incubation of rice
field soils (Garcia et al., 1974; Wassmann et al., 1998; Yao
et al., 1999), peat soils (Magnusson, 1993; Moore and Dal-
va, 1993) and river sediments (Gebert et al., 2006), poten-
tial CH4 and CO2 production rates are linearly correlated to
the OC content of the incubated material (Fig. 5). Note that
even if the correlation between production rates and the
OC were significant for all these studies, the production
rates correlated best with Loss by Ignition and the ratio
of soil N to electron acceptors in the studies of Magnusson
(1993) and Yao et al. (1999), respectively. The slope of the
1
relationship between production rates (nmol g1 soil h ) and
the OC content (gOC/gsoil) are the production rates by mass
1
unit of OC (nmol g1 OC h ). The slopes of these relationships
strongly depend on the duration of the incubation as
shown by the experiments of Yao et al. (1999) and Mag-
nusson (1993) and this study (Table 3). For the present
experiments, the best fits between the production rates
and OC were found when considering the yearly averages
of CO2 and CH4 production rates (Table 2). The slopes of
these relationships are not significantly different from
the slope of the relationships determined by the incubation
of the littoral sediments. Therefore, a unique relationship
can be established between the CO2 production rates and
the OC of the soils and the littoral sediments of the PSR
(r2 = 0.78, p = 0.007 and n = 10 for CO2) and a similar rela-
tionship for CH4 (r2 = 0.72, p = 0.0018 and n = 10 for CH4)
(Fig. 5). The slopes of the relationships are 2:6  102 
1
5:6  101 nmol ðCH4 Þ g1 OC h and 3:5  102  6:9  101
1
nmol ðCO2 Þ g1OC h . These relationships show that CH4
and CO2 production depends on the amount of available
substrate for the bacteria involved in the mineralization
of the OM and explain most of the differences of produc-
tion rates between one soil and another (see Fig. 5).
However, the OC content of the substrate is not the un-
ique controlling factor of CH4 and CO2 production rates.
The SS was not considered for these relationships as the
 F. Guérin et al. / Applied Geochemistry 23 (2008) 2272–2283 2279

results of plant material do not fit with the relationships

a 1900 OS between the OC content of soil material and the CO2 and
1500 LS
LS -OS
CH4 production rates (Fig. 5). This reveals that the quantity
1100
Li t of OC is not the only controlling factor for CO2 and CH4 pro-
700
Cores duction. For each measurement of the production rates, the
300
CO2 (nmol g -1 h-1)

SS
remaining OC in the slurry was calculated assuming that
Ficus
Pa lm OC content decreased only because of the release of CH4
100 Ep i and CO2. Then, the CO2 and CH4 production rates were
1
80 computed by mass unit of organic C ðnmol g1 OC h Þ for

60 each measurement. No relationships were found between

the production rates and the OC content in individual slur-
40
ries. Thus the average over the incubation period was con-
1
20
sidered. They were 2:4  103  1  103 nmol ðCH4 Þ g1 OC h
3 3 1 1
0 and 3:9  10  5  10 nmol ðCO2 Þ gOC h , which is an
order of magnitude higher than for soils. Thus, the plant
0.0 0.1 0.2 0.3 0.4 0.5 material contains OC much more bio-available for bacteria
involved in the degradation of OM and for the methano-
b 900 genic communities than the C from soils. The different
700 plant materials exhibited different production rates, which
are probably related to the lignin and fiber content of the
500
leaves (Miyajima et al., 1997).
CH4 (nmol g-1 h-1)

300 In a reservoir, the flooded OM consists of soil and plant

material. Nevertheless, as revealed by the common rela-
100 tionships for the incubation of the different soils and the
flooded soils from the littoral station, the OC of soil OM
80
is alone a reliable index to predict the production rates in
60
the flooded OM of a tropical reservoir flooded 10 a ago.
40 However, in a reservoir, the sedimentation of autochtho-
20 nous and allochthonous OM fuels the CO2 and CH4 produc-
tion as shown for methanogenesis in natural lakes (Kelly
0
and Chynoweth, 1981). Therefore, the prediction of CH4
0.0 0.1 0.2 0.3 0.4 0.5 and CO2 production at the bottom of reservoirs by incuba-
OC (w/w) tion of soils similar to those flooded during reservoir
impoundment should be considered as the minimum po-
Fig. 5. Yearly average of CO2 (a) and CH4 (b) production rates of the tential production. Also, if one wants to predict CO2 and
incubations and the sediments from the littoral zone (Cores) versus the
CH4 production in a tropical reservoir by this in vitro ap-
initial organic C content (OC). Production rates are linearly correlated to
OC for OS, LS, LO and Lit and Cores, only. proach, the incubation must be done at a temperature
close to in situ since the temperature significantly affects
gas production rates (e.g., Kelly and Chynoweth, 1981).
Table 3
Specific production rates ðnmol g1
1
OC h Þ of CO2 and CH4 of rice field and
4.2. Flooded soils and plant biomass in the Petit Saut
tropical forest soils obtained during short and long term incubations in Reservoir
anaerobic conditions

Environment Incubation Specific production Reference The kinetics of CO2 and CH4 production obtained by the
1
duration days rate nmol g1
OC h incubations of flooded soils and plant material in anoxic
Rice field CH4 14 9.4  102 Wassmann conditions can be used to compute the potential produc-
soils 56 1.3  103 et al. (1998) tion rates by reservoir surface area (similar to a benthic
Rice field CH4 6–78 4.4  102 Yao et al. flux) of the OM flooded in the PSR over 10a. The CO2 and
soils 120 1.2  101 (1999) CH4 production during soil and plant biomass mineraliza-
CO2 19–75 1.4  103 tion at the reservoir scale was calculated at monthly inter-
120 1.2  101
vals using the C density (MgC km2) of the different pool of
Tropical CH4 60–120 3.8  102 This study OM in the PSR and the CO2 and CH4 production rates by
forest 365 3.9  101 1
mass unit of organic C ðnmol g1 OC h Þ. It was assumed that
soils CO2 30–60 8.3  102
365 2.6  102 C of the different pools is only lost in the form of CO2 and
CH4. At each interval, the total amount of C (CO2 + CH4) lost
during the previous interval was subtracted from the C
density of each C pool.
temporal pattern and the intensity of the production rates The initial pool of OM flooded in the PSR was around
differed significantly from the other soils, reflecting a to- 9000 GgC of which 58% originated from plant biomass
tally different bacterial assemblage due to its formation (Table 4). The C from the plant material mainly consisted
without any contact with the soil on the forest floor. The of ligneous biomass (wood), and the leaf biomass and
2280 F. Guérin et al. / Applied Geochemistry 23 (2008) 2272–2283
Table 4
Carbon density of the pool of organic matter (OC) flooded in the Petit Saut Reservoir
OM pool Thickness Volumetric weight
(cm) (Mg m3)
Organic 10 1a
Lateritic 30 1.4b
Total soil 40
Tree trunksc
Above-ground
biomassc
Total
a
Cleavland et al. (2003).
b
Centurion et al. (2004).
c
Galy-Lacaux (1997).
bushes represented only 2% of the total initial pool (Table
4). Although Campo and Sancholuz (1998) showed that,
over 46 a, 40% of the tree trunk density was lost in a reser-
voir in Uruguay, it is still unknown if tree trunks release C
into the water column of reservoirs and the fate of this C is
not documented. The potential for CO2 and CH4 production
from the ligneous material was not measured and is as-
sumed to be negligible for the calculation; only the non-
ligneous pool was considered. Half of the C pool of the
plant biomass was assumed to be present for the first year
and the second half was added at the beginning of the sec-
ond year, as the leaves of the trees fell about a year after
the start of filling. The second C pool in the PSR is the soil,
which contains 46% of the total initial pool (Table 4). The C
density of a 40 cm-thick soil calculated from the field data
(Table 4) is very similar to the C density of 12.3 GgC km2
given by Houghton et al. (2001) for tropical forest soils. For
calculation purposes, the organic and lateritic layers were
considered as two individual C pools with their own C den-
sities (Table 4). The litter was neglected since its C density
is only 0.02 GgC km2.
Over 10 a, the computed production rates of the soil
material decreased from 460 ± 80 to 262 ± 20 MgC–
CO2 km2 a1 (Fig. 6a) and from 347 ± 70 to 197 ±
20 MgC–CH4 km2 a1 (Fig. 6b). The computed production
rates due to the mineralization of plant material are max-
imum during the second year (140 MgC–CO2 km2 a1 and
85 MgC–CH4 km2 a1) and decrease to negligible values
6 a after flooding (Fig. 6a and b). An interesting point to
note is that the plant material is only 5% of the total C con-
sidered in these calculations but it was estimated to con-
tribute up to 10% of the total of both CO2 and CH4
production. Assuming that the C can only be lost in the
form of CO2 and CH4, it corresponds to the mineralization
of 54% of the OC from the soils and 100% of the C originat-
ing from plant material, which is 20% of the total C pool
including tree trunks.
These results are consistent with the data from the core
retrieved in December 2003 in the littoral zone of the PSR.
Ten years after reservoir filling, at the littoral station, the
first 2 cm of the sediment matched well with the litter
d13C and C/N signature. The 2–6 cm depth slices are very
close to the organic soil signature. The terrestrial origin
of the OM of these littoral sediments was also confirmed
by OC/pigment ratios (de Junet et al., in press) in the first
6 cm of the core. Deeper in the core, lateritic soil was found
Water content Soil density OC Carbon density Total %
(%) (Gg km2) (%) (GgC km2) (GgC)
40 60 17. 11.0
20 336 0.4 1.2
396 6.0 12.2 3690 42
16.5 4950 56
0.6 180 2
8820
with OC <1%, a constant C/N ratio at 16, an absence of
pigment and a net increase in d13C (Fig. 1). Such increase
in d13C with depth is typical of that found in soils as a re-
sult of complex biological and chemical processes occur-
ring over long time scales (Garten et al., 2000; Powers
and Schlesinger, 2002). It thus appears that the sediment
core sampled here is composed of the soil flooded when
the PSR was impounded 10 a previously. However, due to
the intense mineralization of OM at the bottom of the
PSR, the top 5 cm is depleted in OC by 20–30% in compar-
ison with the original OS and Lit (Fig. 1a) showing that C
escaped from the soils after flooding as hypothesized. Nev-
ertheless, the residual C density in the littoral cores cannot
be compared directly to the C density of the initial soil
since the OC-rich layer in the core was restricted to the
top 5 cm whereas this layer extended to 10 cm depth in
the forest soil. This fact suggests that, in the littoral zone,
two processes modified the depth profiles of OC: (1) the
overlying water probably contributed to the compaction
of the flooded soils and (2) part of the OM had also been
eroded and was probably mineralized in the water column
and at the bottom of deeper zones of the reservoir.
In Fig. 6a and b, depth-integrated production rates of CO2
and CH4 from the littoral cores are compared to the benthic
fluxes derived from the incubations. Benthic fluxes from the
cores are assumed to be representative of the physical and
biogeochemical state of the OC pool 10 a after flooding. Ben-
thic fluxes of CO2 from the two cores sampled in May were
very similar and are nearly twice as high as the benthic
fluxes extrapolated from the incubations (470 versus
260 MgC–CO2 km2 a1) (Fig. 6a). For CH4, benthic fluxes
obtained from the cores showed a large spatial heterogene-
ity (Fig. 6b) consistent with the depth profiles of CH4 con-
centrations (Fig. 2). Benthic fluxes ranged from 90 to
470 MgC–CH4 km2 a1 for the December period and from
45 to 1530 MgC–CH4 km2 a1 for the May period
(Fig. 6b). The average of these four fluxes is more than twice
as high as the benthic flux from the incubation. Even if some
discrepancies exist between these two estimates, the ben-
thic fluxes from the incubations are nevertheless on the
same order of magnitude as the benthic fluxes from the lit-
toral cores. The CO2 and CH4 benthic fluxes obtained by
these two approaches are one order of magnitude higher
than the benthic fluxes determined by Fick’s first law of dif-
fusion in a 70 a-old tropical reservoir (Abe et al., 2005). This
comparison of the benthic fluxes in two tropical reservoirs is
 F. Guérin et al. / Applied Geochemistry 23 (2008) 2272–2283 2281

CO2 and CH4. Total CO2 production ranges from 167 ± 35

a 600
to 79 ± 8 GgC–CO2 a1 and is lower than CO2 emissions
Plant
500
by 35% on average over 10 a. Total CH4 production ranges
soils
from 122 ± 31 to 59 ± 8 GgC–CH4 a1. Methane production
CO2 (MgC km-2 a-1)

cores is systematically higher than emissions. The difference be-

400
tween production and emissions is small during the first
300 3 a and becomes very significant afterwards. Over 10 a,
total gaseous production consisted of 60% of C–CO2 and
200
40% of C–CH4 whereas total atmospheric emissions
100
consisted of 20% of C-CH4 just after impoundment to 7%
10 a after (calculated from Abril et al., 2005). In addition,
0 cumulated production of C–CH4 plus C–CO2 over 10 a
1 2 3 4 5 6 7 8 9 10 (1.9 ± 0.2 TgC) is about 73–95% of emissions (2.2 TgC; Abril
et al., 2005).
b 1500 Plant
The shift in the C balance between produced and emit-
1250 ted CH4 and CO2 can be explained by the intensive aerobic
soil
1000 CH4 oxidation in the water column of the PSR and in the
CH4 (MgC km-2 a-1)

cores
750 river downstream (Guérin and Abril, 2007). The yearly
500 average of aerobic CH4 oxidation estimated by difference
400
between the extrapolation of the incubation and the emis-
sions range between 20 and 81 GgC–CH4 a1 over 10 a
300
(Fig. 6c), and is estimated to range from 43 to 59 GgC–
200 CH4 a1 for the year 2003 (Table 5). For the year 2003 at
100 the PSR, Guérin and Abril (2007) estimated from direct
0 measurements of CH4 oxidation that total aerobic CH4 oxi-
1 2 3 4 5 6 7 8 9 10 dation was on average 67 ± 64 GgC–CH4 a1, with impor-
tant seasonal variation (5–264 GgC–CH4 a1). These two
c 350
CH emission
independent estimates of CH4 oxidation are very similar
300 and confirm that the methanotrophic activity in the PSR
CO emission
is up to one order of magnitude higher than what is com-
250 CH production
monly found in natural lakes (Guérin and Abril, 2007). The
GgC a-1

200 CO production seasonal variation of CH4 oxidation in the PSR could not be
reproduced by in vitro experiments since they were related
150
to variation in CH4 concentrations in the water column due
100 to strong seasonal variation in the residence time of water
and to the physics of the reservoir (stratification–destrati-
50
fication) (Guérin and Abril, 2007).
0 Assuming a methanotrophic bacterial growth efficiency
1 2 3 4 5 6 7 8 9 10 (BGE) of 50%, aerobic CH4 oxidation corresponds to the
Year
production of 26 ± 6 GgC–CO2 a1 (Table 5) and the same
Fig. 6. Ten-year evolution of the benthic fluxes of CO2 (a) and CH4 (b) amount of bacterial biomass (Guérin and Abril, 2007).
computed from the production rates of the incubations and the C densi- BGE varies between 5% and 80% among different lakes
ties of the different pools of OM flooded in the PSR (plant material and and seasons in the boreal environment (Bastviken et al.,
soils). Vertically integrated production of CO2 and CH4 from the littoral 2003). This estimate is therefore highly dependent on the
cores are added for comparison with the benthic fluxes 10 a after flood-
ing. Panel (c): Comparison of total CO2 (GgC–CO2 a1) and CH4 (GgC–
assumption on BGE that is not documented at the PSR.
CH4 a1) production and total emission (GgC a1) from the PSR over 10 a. Nevertheless, the difference between CO2 production and
Emissions are from Abril et al. (2005). The average surface area of the PSR CO2 emission is 43 ± 8 GgC–CO2 a1, that is 50% higher
considered for calculation is 300 km2.

Table 5
consistent with the decrease by one order of magnitude of CH4 and CO2 balance (GgC a1) in the PSR for the year 2003
CH4 and CO2 emissions from reservoirs between young res-
CO2 CH4 Reference
ervoirs (<10-a) and 40–70-a old reservoirs (St. Louis et al.,
2000; dos Santos et al., 2004). However, more studies dedi- Production 79 ± 8 59 ± 8 This study
Emission 122 9 Abril et al. (2005)
cated to determination of benthic fluxes in tropical reser-
Production–emission 43 ± 8 +50 ± 11 This study
voirs are therefore needed to definitely assess this question. CH4 oxidationa +26 ± 6 50 ± 11 This study
CH4 oxidationb +34 ± 32 67 ± 64 Guérin and Abril (2007)
4.3. A tentative budget a
Calculated from the difference between production and emission
assuming a bacterial growth efficiency of 50% for CO2 production by
In Fig. 6c, extrapolated CO2 and CH4 production over aerobic CH4 oxidation.
b
10 a are compared to the total atmospheric emissions of Calculated from a kinetic approach.
2282 F. Guérin et al. / Applied Geochemistry 23 (2008) 2272–2283
than the calculated CO2 production by CH4 oxidation
(Table 5). The internal recycling of CH4, CO2 and autochtho-
nous OM in the lake water column and in the river down-
stream changes the partitioning between those two gases
and cannot be estimated by the anoxic microcosm experi-
ments used for this study. Part of the CO2 produced by the
decomposition of the flooded OM is used by phytoplankton
to produce labile OM. A fraction of this freshly produced
OM is respired in the water column and returned to the
system as CO2. The remaining OM sinks and fuels CO2
and CH4 production in the anoxic hypolimnion and con-
tributes to the C trapping in reservoirs. Integrated over
the PSR, the sedimentation of OM in the reservoir was esti-
mated to be 1.90 ± 0.07 GgC a1 from sediment trap data;
most of this material was of planktonic origin as shown
by pigment analyses (de Junet et al., in press). However,
the internal recycling does not change the amount of C in-
volved in CO2 and CH4 production in the system. It thus
indicates that an additional source of C representing 5–
30% of emissions is missing in this C balance. Supply of
allochthonous material from the rivers upstream of the
PSR is 35 GgC a1 but (1) part of this C is refractory and
thus it would contribute only partially to the gap between
gaseous production from the initial pool of flooded OM and
the emissions from the PSR and (2) the supply of allochth-
onous C is balanced by the export to the ocean
(50 GgC a1; Abril et al., 2005). Lateral supply by runoff
of allochthonous OM, CO2, dissolved inorganic C and CH4
from the forest surrounding the PSR to the lake have not
estimated until now despite such lateral C transfer is
known to fuel significant CO2 and CH4 emissions from nat-
ural aquatic ecosystems (Cole et al., 2007). The potential
supply of OM from the decomposition of wood (standing
and tangled trees) was also not estimated despite the fact
that wood is the largest C pool in the reservoir.
5. Summary and conclusions
The potential CO2 and CH4 production rates following
the flooding of soils and in the sediments are mainly con-
trolled by the organic C content of the soils. The common
relationship between production rates and OC for the dif-
ferent soil incubations and the cores indicate that the OC
in soils is a reliable index to predict the long term produc-
tion rates of flooded soils in tropical reservoirs. In contrast,
for the same amount of organic matter, the production
rates obtained during the mineralization of plant material
are an order of magnitude higher. This point has important
implications: despite the low C density of plant material in
the PSR (2% of the total C flooded), the plant material con-
tributes up to 10% of the total gaseous production.
When extrapolated to the reservoir scale, CH4 and CO2
production during the decomposition of the flooded OM
contributes up to about 75–95% of the total atmospheric
emissions, with a C–CH4:C–CO2 ratio around 0.75 whereas
the C–CH4:C–CO2 ratio of atmospheric emissions is around
0.16 (Abril et al., 2005). This difference is mainly due to the
intensive aerobic CH4 oxidation in the water column that
significantly reduces atmospheric CH4 emissions and con-
tributes to CO2 emissions (Guérin and Abril, 2007). In this
mass balance 5–30% of C is missing. The supply of allochth-
onous C from the three tributaries of the PSR is balanced by
the export of C downstream of the dam. For future studies,
two additional sources must be considered. In natural
lakes without tributaries, allochthonous C represents
around 50% of the C in their water column (Cole et al.,
2000). That means that direct lateral inputs of C from the
terrestrial ecosystems surrounding the lakes by runoff
are significant and can contribute to CO2 and CH4 emis-
sions. The contribution of this C pathway has never been
quantified for a reservoir. The wood density of tree trunks
decreases with time when they are standing in reservoirs
(Campo and Sancholuz, 1998). It is still unknown whether
tree trunks release C into the water column and the fate of
this C is still not documented.
The C mass balance of the PSR is one of the most com-
prehensive estimated for a hydroelectric reservoir. Meth-
ane and CO2 production (this study), aerobic CH4
oxidation (Guérin and Abril, 2007) and atmospheric CH4
and CO2 emissions (Abril et al., 2005) were all deter-
mined by several independent methods which all con-
verge to similar estimates. The main conclusion of this
study and the study of Abril et al. (2005) is that 10 a after
impoundment, the mineralization of the flooded soil and
biomass is still, by far, the major contributor to the
greenhouse gas emissions from the PSR. It appears thus
to be robust. However it contrasts with the conclusion
of Huttunen et al. (2002) and Bodaly et al. (2004) regard-
ing the origin of OM driving long and short term emis-
sions in boreal reservoirs. These differences between
tropical and boreal reservoirs are probably related to
the temperature (e.g., Kelly and Chynoweth, 1981) and
OM quality (e.g., Miyajima et al., 1997) differences. In
boreal reservoirs flooding peatland, the low temperature
and the high content of slowly decomposable OM lead
to lower CO2 and CH4 production rates than at the bot-
tom of tropical reservoirs.
From calculations, 54% of the soil OC degassed as CH4
and CO2 in 10 a. This is in the upper range since allochtho-
nous OM also contributed to atmospheric emissions. The
contribution of the turnover of allochthonous OM to emis-
sions from reservoirs increases with time as the initial pool
of OM flooded is more and more exhausted. A better
understanding of ‘‘carbon spiraling” (Newbold et al.,
1982) in reservoirs appears thus to be a key component
to predict emissions from reservoirs over long time scales
(>10–20 a). As the majority of the world’s hydropower po-
tential remains in the tropics, future research on this topic
must focus on these climatic zones.
Acknowledgements
The authors thank R. Delmas (LA, Toulouse) who was
the initiator of the Petit-Saut study, H. Etcheber for OC
analysis and for his encouragement, and A. Grégoire
(EDF) for his continuous confidence. The authors thank
two anonymous reviewers for their constructive com-
ments. This research was funded by Electricité de France
(EDF) and the CNRS national programs (PNCA and ECCO).
F.G. benefited from a Ph.D. Grant by EDF.
 F. Guérin et al. / Applied Geochemistry 23 (2008) 2272–2283 2283

References Guérin, F., Abril, G., 2007. Significance of pelagic aerobic methane
oxidation in the methane and carbon budget of a tropical reservoir.
J. Geophys. Res. 112, G03006. doi:10.1029/2006JG000393.
Abe, D.S., Adams, D.D., Sidagis Galli, C.V., Sikar, E., Tundisi, J.G., 2005.
Guérin, F., Abril, G., Richard, S., Burban, B., Reynouard, C., Seyler, P.,
Sediment greenhouse gases (methane and carbon dioxide) in the
Delmas, R., 2006. Methane and carbon dioxide emissions from
Lobo-Broa Reservoir, São Paulo State, Brazil: concentrations and
tropical reservoirs: significance of downstream rivers. Geophys. Res.
diffuse emission fluxes for carbon budget considerations. Lakes
Lett. 33, L21407. doi:10.1029/2006GL027929.
Reservoirs Res. Manage. 10, 201–209.
Houghton, J.T., Ding, Y., Griggs, D.J., Noguer, M., van der Linden, P.J.,
Abril, G., Iversen, N., 2002. Methane dynamics in a shallow non-tidal
Xiaosu, D. (Eds.), 2001. Intergovernmental Panel on Climate Change.
estuary (Randers Fjord, Denmark). Mar. Ecol. Prog. Ser. 230, 171–181.
Climate Change 2001: The Scientific Basis. Contribution of Working
Abril, G., Guérin, F., Richard, S., Delmas, R., Galy-Lacaux, C., Gosse, P.,
Group I to the Third Assessment Report of the Intergovernmental
Tremblay, A., Varfalvy, L., dos Santos, M.A., Matvienko, B., 2005.
Panel on Climate Change. Cambridge University Press, Cambridge,
Carbon dioxide and methane emissions and the carbon budget of a
United Kingdom and New York, NY, USA.
10-year old tropical reservoir (Petit Saut, French Guiana). Glob.
Huttunen, J.T., Väisänen, T.S., Hellsten, S.K., Heikkikinen, M., Nykänen, H.,
Biogeochem. Cycles 19, GB4007. doi:10.1029/2005GB002457.
Jungner, H., Niskanen, A., Virtanen, M.O., Lindqvist, O.V., Nenonen, O.,
Bastviken, D., Ejlertsson, J., Sundh, I., Tranvik, L., 2003. Methane as a
Martikainen, P.J., 2002. Fluxes of CH4, CO2, and N2O in hydroelectric
source of carbon and energy for lake pelagic food webs. Ecology 84,
reservoirs Lokka and Porttipahta in the northern boreal zone in
969–981.
Finland. Global Biogeochem. Cycles 16, 1–17.
Bodaly, R.A., Beaty, K.G., Hendzel, L.H., Majewski, A.R., Paterson, M.J.,
Kelly, C.A., Chynoweth, D.P., 1981. The contribution of temperature and
Rolfhus, K.R., Penn, A.F., St Louis, V.L., Hall, B.D., Matthews, C.J.D.,
the input of organic matter in controlling rates of sediment
Cherewyk, K.A., Mailman, M., Hurley, J.P., Schiff, S.L., Venkiteswaran,
methanogenesis. Limnol. Oceanogr. 26, 891–897.
J.J., 2004. Experimenting with hydroelectric reservoirs. Environ. Sci.
Kelly, C.A., Rudd, J.W.M., Bodaly, R.A., Roulet, N.P., St. Louis, V.L., Heyes, A.,
Technol. 38, 346A–352A.
Moore, T.R., Shiff, S., Aravena, R., Scott, K.J., Dyck, B., Harris, R., Warner,
Campo, J., Sancholuz, L., 1998. Biogeochemical impacts of submerging
B., Edwards, G., 1997. Increases in fluxes of greenhouse gases and
forests through large dams in the Rio Negro, Uruguay. J. Environ.
methyl mercury following flooding of an experimental reservoir.
Manage. 54, 59–66.
Environ. Sci. Technol. 31, 1334–1344.
Centurion, J.F., Beutler, A.N., de Souza, Z.M., 2004. Physical attributes of
Kemenes, A., Forsberg, B.R., Melack, J.M., 2007. Methane release below a
kaolinitic and oxidic oxisoils resulting from different usage systems.
tropical hydroelectric dam. Geophys. Res. Lett. 34, L12809.
Brazilian Arch. Biol. Technol. 47, 725–732.
doi:10.1029/2007GL029479.
Cleavland, C.C., Townsend, A.R., Schmidt, S.K., Constance, B.C., 2003. Soil
Krüger, M., Eller, G., Conrad, R., Frenzel, P., 2002. Seasonal variation in
microbial dynamics and biogeochemistry in tropical forests and
pathways of CH4 production and in CH4 oxidation in rice fields
pastures, southwestern Costa Rica. Ecol. Appl. 13, 314–326.
determined by stable carbon isotopes and specific inhibitors. Global
Cole, J.J., Pace, M.L., Carpenter, S.R., Kitchell, J.F., 2000. Persistence of net
Change Biol. 8, 265–280.
heterotrophy in lakes during nutrient addition and food web
Le Mer, J., Roger, P., 2001. Production, oxidation, emissions and
manipulations. Limnol. Oceanog. 45, 1718–1730.
consumption of methane in soils: a review. Eur. J. Soil Biol. 37, 25–50.
Cole, J.J., Prairie, Y.T., Caraco, N.F., McDowell, W.H., Tranvik, L.J., Striegl,
Magnusson, T., 1993. Carbon dioxide and methane formation in forest
R.G., Duarte, C.M., Kortelainen, P., Downing, J.A., Middelburg, J.J.,
mineral and peat soils during aerobic and anaerobic incubations. Soil
Melack, J., 2007. Plumbing the global carbon cycle: integrating inland
Biol. Biochem. 25, 877–883.
waters into the terrestrial carbon budget. Ecosystems 10, 171–184.
Miyajima, T., Wada, E., Ba, Y., Vijarnsorn, P., 1997. Anaerobic
Conrad, R., 1989. Control of methane production in terrestrial ecosystems.
mineralization of indigenous organic matters and methanogenesis
In: Andrea, M.O., Schimel, D.S. (Eds.), Exchange of Trace Gases
in tropical wetland soils. Geochim. Cosmochim. Acta 61, 3739–3751.
Between Terrestrial Ecosystems and the Atmosphere. John Wiley &
Moore, T.R., Dalva, M., 1993. The influence of temperature and water table
Sons, Chichester, pp. 39–58.
position on carbon dioxide and methane emissions from laboratory
Dannenberg, S., Wulderl, J., Conrad, R., 1997. Agitation of anoxic paddy
columns of peatland soils. J. Soil Sci. 44, 651–664.
soil slurries affects the performance of the methanogenic microbial
Newbold, J.D., Mulholland, P.J., Elwood, J.W., O’Neil, R.V., 1982. Organic
community. FEMS Microbiol. Ecol. 22, 257–263.
carbon spiraling in stream ecosystems. Oikos 38, 266–272.
de Junet, A., Abril, G., Guérin, F., Billy, I., De Wit, R., in press. A multi-
Nieuwenhuize, J., Maas, Y.E.M., Middelburg, J.J., 1994. Rapid analysis of
tracers analysis of sources and transfers of particulate organic matter
organic carbon and nitrogen in particulate materials. Mar. Chem. 45,
in a tropical reservoir (Petit Saut, French Guiana). River Res. Appl.,
217–224.
doi:10.1002/rra.1152.
Oremland, RS., Marsh, LM., Polcin, S., 1982. Methane production and
dos Santos, M.A., Matvienko, B., Rosa, L.P., Sikar, E., dos Santos, E.D., 2004.
simultaneous sulphate reduction in anoxic, salt marsh sediments.
Gross greenhouse gas emissions from Brazilian hydro reservoirs. In:
Nature 296, 143–145.
Tremblay, A., Varfalvy, L., Roehm, C., Garneau, M. (Eds.), Greenhouse
Powers, J.S., Schlesinger, W.H., 2002. Geographic and vertical patterns of
Gas Emissions-Fluxes and Processes: hydroelectric Reservoirs and
stable carbon isotopes in tropical rain forest soils of Costa Rica.
Natural Environments, Environ. Sci. Ser.. Springer, New York, pp. 67–
Geoderma 109, 141–160.
291.
Rosa, L.P., Schaeffer, R., 1994. Greenhouse gas emissions from
Duchemin, E., Lucotte, M., Camuel, R., Chamberland, A., 1995. Production
hydroelectric reservoirs. Ambio 23, 164–165.
of the greenhouse gases CH4 and CO2 by hydroelectric reservoirs in
Rudd, J.W.M., Harris, R., Kelly, C.A., Hecky, R.E., 1993. Are hydroelectric
the boreal region. Global Biogeochem. Cycles 9, 529–540.
reservoirs significant sources of greenhouse gases? Ambio 22, 246–248.
Froelich, P.N., Klinkhammer, G.P., Bender, M.L., Luedtke, N.A., Heath, G.R.,
Soumis, N., Duchemin, E., Canuel, R., Lucotte, M., 2004. Greenhouse gas
Cullen, D., Dauphin, P., 1979. Early oxidation of organic matter in
emissions from reservoirs of the Western United States. Global
pelagic sediments of eastern equatorial Atlantic: suboxic diagenesis.
Biogeochem. Cycles 18. doi:10.1029/2003GB002197.
Geochim. Cosmochim. Acta 43, 1075–1090.
St. Louis, V., Kelly, C., Duchemin, E., Rudd, J.W.M., Rosenberg, D.M., 2000.
Galy-Lacaux, C., 1997. Modification des échanges de constituants mineurs
Reservoir surface as sources of greenhouse gases to the atmosphere: a
atmosphériques liées à la creation d’une retenue hydroélectrique.
global estimate. Bioscience 20, 766–775.
Impact des barrages sur le bilan du CH4 dans l’atmosphère. Ph.D.
Wassmann, R., Neue, H.U., Bueno, C., Lantin, R.S., Alberto, M.C.R., Buendia,
Thesis, Univ. Paul Sabatier (Toulouse III).
L.V., Bronson, K., Papen, H., Rennenberg, H., 1998. Methane
Galy-Lacaux, C., Delmas, R., Jambert, C., Dumestre, J.F., Labroue, L.,
production capacities of different rice soils derived from inherent
Richard, S., Gosse, P., 1997. Gaseous emissions and oxygen
and exogenous substrates. Plant Soil 203, 227–237.
consumption in hydroelectric dams: a case study in French Guiana.
Weiss, R.F., 1974. Carbon dioxide in water and seawater: the solubility of
Global Biogeochem. Cycles 11, 471–483.
a non-ideal gas. Mar. Chem. 2, 203–215.
Garcia, J.L., Raimbault, M., Jacq, V., Rinaudo, G., Roger, P.A., 1974. Activités
Williams, R.J., Crawford, R.L., 1984. Methane production in Minnesota
microbiennes dans les sols de rizière du Sénégal: relations avec les
peatlands. Appl. Environ. Microbiol. 47, 1266–1271.
propriétés physico-chimiques et influence de la rhizosphère. Rev.
Yamamoto, S., Alcauskas, J.B., Crozier, T.E., 1976. Solubility of methane in
Ecol. Biol. Sol 11, 169–185.
distilled water and seawater. J. Chem. Eng. Data 21, 78–80.
Garten, C.T., Cooper, L.W., Post III, W.M., Hanson, P.J., 2000. Climate
Yao, H., Conrad, R., Wassmann, R., Neue, H.U., 1999. Effect of soil
controls on forest soil C isotope ratios in the southern Appalachian
characteristics on sequential reduction and methane production in
Mountains. Ecology 81, 1108–1119.
sixteen rice paddy soils from China, the Philippines, and Italy.
Gebert, J., Köthe, H., Gröngröft, A., 2006. Prognosis of methane formation
Biogeochemistry 47, 269–295.
by river sediments. J. Soils Sediments 6, 75–83.

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