JOURNAL OF GEOPHYSICAL RESEARCH, VOL. 101, NO.

D9, PAGES 14,399-14,414, JUNE 20, 1996

Global carbon exchangeand methane emissionsfrom

natural wetlands: Application of a process-basedmodel

Mingkui Cao, StewartMarshall,andKeith Gregson

Departmentof Physiologyand EnvironmentalScience

Universityof Nottingham,Loughborough,England

Abstract. Wetlandsare one of the mostimportantsourcesof atmosphericmethane

(CH4), but the strengthof this sourceis still highly uncertain.To improve estimatesof
CH4 emissionat the regionaland globalscalesandpredictfuturevariationrequiresa
process-based modelintegratingthe controlsof climaticand edaphicfactorsand complex
biological
processes
overCH4 flux rates.Thisstudyuseda methane
emission
model
basedon the hypothesis that plantprimaryproductionand soil organicmatter
decomposition act to controlthe supplyof substrate
neededby methanogens; the rate of
substrate supplyand environmentalfactors,in turn, controlthe rate of CH4 production,
and the balancebetweenCH4 productionand methanotrophic oxidationdeterminesthe
rate of CH4 emissioninto the atmosphere.Coupledto data setsfor climate, vegetation,
soil, and wetlanddistribution,the modelwas usedto calculatespatialand seasonal
distributionsof CH4 emissionsat a resolutionof 1o latitude x 1o longitude.The
calculated net primaryproduction (NPP) of weftandsrangedfrom 45 g C m-2yr-• for
northern bogsto 820 g C m-2yr-1for tropicalswamps.CH4 emission ratesfrom
individualgridcellsrangedfrom0.0 to 661 mg CH4 m-2d-1,with a meanof 40 mg CH4
m-2d-• for northernwetland,150mgCH4 m-2d-1for temperate wetland,and199mgCH4
m-2d-1for tropicalwetland.TotalCH4 emission was92 Tg yr-•. Sensitivity
analysis
showedthat the responseof CH4 emissionto climate changedependsupon the combined
effectsof soil carbonstorage,rate of decomposition,soil moistureand activity of
methanogens.

Introduction tropicalwetlands.However, field measurementsin the 1980s

revealed that northern tundra and peatlandsalso made a
The concentrationof atmosphericCH 4 has increased major contributionto the globalsourcestrength. Matthews
during the last 200 years [Pearman et al., 1986; Steele et and Fung [1987] generateda global distributionof wetlands
al., 1992], and accounts for approximately 15% of from wetland vegetation types, pond soils, and land
anthropogenic greenhouse effects[Rodhe,1990]. Wetlandis inundationdata. Using this data set and CH4 flux rates
one of the most importantterrestrialsourcesof CH 4 because measuredin the early 1980s, theseauthorsestimatedthat the
of its anaerobicconditions,high organicmatter content,and annual emissionof CH 4 from natural wetlandswas 111.1
large area. It is currently estimatedthat wetlandsaccountfor Tg, and that northernwetlandscontributed60% of this total
20% to 30% of the total annual emissionof CH 4 to the emission. Aselmann and Crutzen [1989] developed a
atmosphere[Intergovernmental Panel on Climate Change differentdataset for wetlanddistributionand CH 4 flux rates.
(IPCC), 1990; Fung et al., 1991]. Accuratequantificationof Their estimateof annualCH4 emissionwas in the range40
this source strengthand its spatial and seasonalvariation to 160 Tg. Bartlett and Harris [1993] recentlyreviewedthe
acrossthe globe must play an importantrole in predicting publishedmeasurementsof CH 4 flux rates from wetlands
thetrendin atmospheric CH4 concentration andits impacton globally, andusingaveragedCH 4 fluxesfor variouswetland
climate. types, they calculatedthe total CH 4 emissionto be 109 Tg
There havebeenseveralattemptsto usemeasuredratesof yr-•. Considerable
uncertainty
existsin theestimates
of CH4
emissionsat both the regional and global scales since,
emissionfrom wetlandsto arrive at estimatesfor the global
althoughthere has been a large increasein the number of
total CH4 emissionfrom wetlands.Early studiessuggested
CH4 flux rate measurements,
that tropicalwetlandswere the main sourceof atmospheric their spatialcoverageis still
thattotalemission poor, and most of the measurementswere made over less
CH4;Seilerand Conrad[1987]reported
from than 1 m2 and over periodsof only a few months.Few
was47 Tg yr'•, of which38 Tg yr-• wasreleased
studieshaverecordsspanningmore than 1 year, or a spatial
Copyright1996by theAmericanGeophysical
Union.
scaleof hundredsof squarekilometers.It seemsunlikely that
Papernumber96JD00219. the extrapolationof CH4 rates from point measurements to
0148-0227/96/96JD-00219509.00 the regional or global scale can yield a reliable estimateof
14,399
14,400 CAO ET AL.: CARBON EXCHANGE AND CH4 EMISSIONS FROM WETLANDS

current CH4 emissions.Moreover, climate changeand betweenCH4 emissionand plant primary productionor net
modificationsto hydrologicalconditionsand vegetation CO2 exchange rates has been reported by Whiting and
patternsareoccurring
withinwetlands[Maxwell,1992;Bliss Chanton [1993] and by Sass et al. [1990]. Plant biomass
and Matveyeva,1992], and thereis a needto predictthe enteringthe soil mustfirst be depolymerizedand fermented
resulting variations in CH4 emissions. Existing by a variety of microbial organismsinto low molecular
methodologies are unableto addresstheseconcerns. weight substances,which are then transformedto CH4 by
The rate of CH4 emissionis a reflectionof the complex methanogens [Joneset al., 1987]. The potentialrate of CH4
interaction between the processes of plant primary production is a functionof the amountanddecomposability
production, soil organic carbon decomposition,of soilorganicmatterin the soil [Svensson, 1983; Valentine
methanogenesis, andmethanotrophicoxidation.Improvement et al., 1994], and a linear relationshipbetween CH4
in estimatesof CH4 emissionsunder different climatic production andreadilydecomposable carbonin the soil has
regimesrequires thedevelopment of a process-based model alsobeenreported[Yagiand Minarni, 1990].
to link environmentalfactorsand biologicalprocesses with With a givensupplyof substrate, CH4production depends
CH4 emissionrates. on the control over methanogenesisby environmental
This study concernsthe development of sucha CH4 factors,suchastemperature[Westermann, 1993;Dunfieldet
emission model for natural wetlands, based on substrate al., 1993] and soil moisture [Moore and Knowles, 1989].
supplyby plant primaryproduction
and organicmatter The CH4producedpassesthroughthe oxic-anoxicboundary
decomposition,
regulation
of methanogenesis
by temperature before enteringthe atmosphere.During that passage,it is
andsoilmoisture,andthebalancebetweenCH4production subjectto oxidationby methanotrophic bacteria,where40%
and oxidation.The model was usedto estimatespatialand to 95% of the CH4 produced may be consumedby
seasonalvariationsin CH4 emissionsfrom naturalwetlands methanotrophs in the soil [Kinget al., 1990;Pulliam,1993].
at the global scale. Thus the rate of CH4 productionshouldbe constrained by
availability of substrate, arising from plant primary
The Methane Emission Model
production andorganicmatterdecomposition, andregulated
Emissionof CH4 is the final stepin the mineralizationof by environmental factors.The balancebetweenproduction
organicmatterin anaerobicecosystems andrepresents a part and oxidationof CH4 determinesthe rate of emissioninto
of the cycling of carbon within the plant-soil-atmospherethe atmosphere.The wetland methaneemissionmodel
system.Plant primary productionis the ultimatesourceof (WMEM) was developed usingthesehypotheses as a basis
substratefor CH4 production. A positive relationship (Figure 1).
CO•

type

LITD

Temperature
I [ Solar
radiation

Water
EPT
SOMD

CO2 Soil
or other properties
gaseous
carbon
products Precipitation
MPR

MOR CO2

ER• ; Methane
Figure1. Schematic representationof thewetlandMethane emission model(WMEM). Theboldlinesrepresent mass
(carbonand water)exchanges, while the fine linesrepresent interactions
betweendifferentfactors.Four carbon
compartments areusedin themodel:carbon in livingvegetation
(VEGC),carbon in thesoil(SOIC),decomposed carbon
(DCPC),andcarbonin theformof CH4(CH4C).Thecarbonflowsbetween thesecompartments arerepresented
byLITD
(totalcarbon
depositionintothesoilarisingfromvegetation),SOMD(decomposition of organic
carbonin thesoil),MPR
(methaneproduction), MOR (methaneoxidation),and MER (methane emissioninto the atmosphere). EPT is
evapotranspiration.
 CAO ET AL.: CARBON EXCHANGE AND CH4 EMISSIONS FROM WETLANDS 14,401

The WMEM is a compartmentalmodel in which four
carboncompartmentswere used:carbonin living vegetation
(VEGC), carbon in the soil (SOIC), decomposedcarbon
(DCPC), and carbon in the form of CH4 (CH4C). The
carbon fluxes between these pools were assumedto be
controlledby the size of donorpools,by specificparameters
related to characteristicsof the pool involved, and by
environmentalfactors,e.g., solarradiation,temperature,and
soil moisture(see Figure 1). The ratesof changeof the size
of eachof thesepools were calculatedfrom
primary factor associatedwith the variation of CH 4
emissions[Torn and Chapin, 1993] becauseit defines the
anaerobicconditionsand low redox potentialswhich are
essential for methanogenesis.Dise [1991] and Torn and
Chapin [1993] found that water table position and
temperaturetogethercould explain from 75 % to 90% of the
variabilityin CH 4 fluxes, and in thepresentmodelthey were
consideredto be the major environmentalfactorsregulating
the productionof CH 4. ThUS the rate of CH4 production
(MPR) was describedby

dVEGC
= GPPt- RESt- LITDt (1) MPRt= SOMDtPof(WTP)f(TEM) (5)
At

where Pois the proportionof the decomposed organiccarbon

transformedto CH 4 under optimal conditionsof temperature
dSOIC
At
= LITDt _ SOMD t (2) and soil water status for methanogensand the functions
f(WTP) andf(TEM) representproportionalCH4 production
factorsfor soil water table position (WTP) and temperature
(TEM) under suboptimalconditionsrespectively.
dDCPC
= SOMD t _ MPRt _ OCRt (3) Usually, 3-25% of decomposed organic carbon is
At
transformedto CH 4 [Moore and Knowles, 1990; Yavitt et al.
1987], but this proportioncan be as high as 40-60% in an
ideal environment for methanogens. Yavitt et al. [1987]
dCH4C
at
= MPRt_MORt (4) reported approximatelyequimolarproductionof CH4 and
CO2 in anaerobic peatlandsat high temperature (19øC).
Tsutsuki and Ponnamperuma [1987] found that CH4-C
whereGPP is grossprimaryproduction (g C m-2month-i), accountedfor 0.41 to 0.52 of the total decomposedcarbon
RESis plantautotrophic respiration
(g C m-2month-i),LITD (CH4-C + CO2-C), with a mean of 0.47, which was usedas
is the rate of carbon depositioninto the soil arising from the value for Po in the presentmodel.
vegetation (g C m'2month-i),bothaboveandbelowground, CH 4 flux increaseslogarithmicallywith increasingwater
SOMD is the decompositionrate of organic carbon in the tableheight[Moore and Knowles,1989; Christensen,1993],
soil(g C m-2month-i),MPR is therateof CH4formation(g and this positiverelationshipmay be maintaineduntil a 10-
C m'2 month-i),OCR is the rate (g C m-2month -1)at which cm depth of water above the soil surface is reached
carbon is released in the form of other gaseouscarbon [Sebacheret al., 1986]. For inundatedwetlands,we used an
products,suchas CO,_,and MOR is the CH 4 oxidationrate empiricalrelationshipreportedby Rouletet al. [1992]. The
(g C m-2month-i).Thetimestep(at) overwhichcalculations effect of water table positionwas modelledas a multiplier
were performedwas 1 month. acting upon potential CH 4 production,with the maximum
value of 1.0 when the water table was 10 cm or more above

Net Primary Production and Organic Carbon the soil surface, and a smaller value at lower water table
Decomposition positions:

f(SWP) : 0.383 e0'096swP (6)

Net primary production(GPP-RES), depositionof litter
carbon into the soil (LITD), and organic carbon
decomposition (SOMD) were calculated following the where SWP (cm) is water table position,relative to the soil
TerrestrialEcosystemModel, whichis describedin detailby surface. The method by which SWP was calculated for
Raich et al. [ 1991] and McGuire et al. [ 1992]. In the case of noninundatedwetlands is shown in the appendix. In the
inundatedwetland, it was assumedthat the soil is always calculations for the inundated wetlands, the soil was
saturated. considered to be saturated, so that moisture had no effect on
CH 4 production.
Methane Production The observedeffect of temperature,representedby a Q10
value (the rate of changein CH 4 flUX due to a 10øC change
The suitabilityof the soil environmentfor methanogens in temperature),falls within a broad range, from 1.2 to 16
affectsthe proportionof decomposed carbontransformedto [Westermann,1993; Dunfield et al., 1993]. This high degree
CH 4. Moore and Knowles [1989] reported that the molar of uncertainty may be caused by the influence of other
ratio of CH4 to CO2 emissionsfrom wetlands increases factors for example, substrateavailability for methanogens
exponentiallywith water table height. Yavitt et al. [1987] [Whiting and Chanton, 1993; Conrad et al., 1987], since
and Tsutsukiand Ponnamperuma[1987] suggestedthat this temperaturealso influencesplant growth and organicmatter
ratio is also a function of temperature.Field observations decomposition. Laboratory studies have shown that
have demonstratedthat the hydrological regime is the methanogenesis has a Q10value of 1.2-3.5 [Conrad, 1989;
14,402 CAO ET AL.: CARBON EXCHANGE AND CH4EMISSIONS FROM WETLANDS

Kelly and Chynoweth,1981]. By assuminga Q•ovalue of the soil profile, and the pathwaystaken by CH4 transport
2.0, which is generallyusedfor biochemicalprocesses,and from the soil into the atmosphere[Whalenand Reeburgh,
30øC as the optimal temperaturefor most methanogens 1988; ChantonandDacey, 199i]. The effectof anyof these
[Dunfield et al., 1993; Neue and $charpenseel,1984], the processesis uncertain, and in the present study it was
influenceof temperature, f(TEM), was representedby assumedthat 90% of CH 4 producedin any month was
oxidized in noninundated wetlands [Fechner and Hemond,
f(TEM)
= e0'0693TEM
7.996
(7) 1992].
Finally, the rate of CH• emissioninto the atmosphere
(MER) was calculatedas the differencebetween CH•
whereTEM is the temperature(øC). productionand oxidation:
In the model, CH 4 was producedonly during thaw/wet
seasons,which were definedfollowingFung et al. [1991]. MERt= MPRt_MORt (9)
For a locationthat experiencestemperatureabovefreezing
throughoutthe year, the wet seasonwas considered to be the
periodover which monthlyprecipitationexceededmonthly
potentialevapotranspiration. The thaw seasonwas assumed Application of the Model
to beginwhenmonthlymeantemperature exceeded5øC and
to end when monthlymeantemperaturefell below 0øC. Natural wetlandsare distributedglobally from polar to
tropicalregions,and exist in diverseclimatic, vegetation,
Methane Oxidation and Emission and soil types. Thus the important environmental
characteristicsdeterminingthe rate of CH4 emissionexhibit
CH4 oxidationoccursabovethe water table and in the heterogeneityin both spaceand time. To assessrates of
rhizosphere, whereO2 is likely to be available.It hasbeen methane emission globally with the WMEM, it was
reportedthatfrom 55% to 85% of the CH4 diffusingto the necessary to developa georeferenced databasecontainingall
soil surface may be oxidized [Happell et al., 1993]. In of the independem variablesneededto run the model.The
plant-freepaddylands,CH4 oxidizationmay consume65% databasecontained information on the location, area, soil,
of the CH4production[Holzapfel-Pschorn et al., 1986], and vegetation,andclimateof globalwetlands.This information
in the seedlingstage of rice growth, when rhizospheric was organizedwithin a Geographical InformationSystem
oxidationis not significant,the proportionof the CH4 (GIS) at a resolutionof 0.5 o latitudex 0.5 ø longitude.
oxidizedvaried from 44 % to 59% [Sch'atzet al., 1989]. The The InternationalInstituteof Applied SystemAnalysis
deliveryof O2to the rhizosphere by plantsmay stimulatethe (IIASA) terrestrialclimate data set [Leemansand Cramer,
oxidation of CH4. Positive relationshipsbetween CH4 1991] was used to provide informationon temperature,
oxidationand live root densityhave been reported[Gerard precipitation, and cloudiness. The climaticdata represem
and Chanton,1993; $mitset al., 1990]. The root O2 supply long-termmonthlymeanvaluesat the 0.5ø latitudex 0.5 ø
and thus the oxidative power, shouldincreasewith plant longitudespatialresolution.The globalsolarradiationand
growth [Albera, 1953]. Schatzet al. [1989] foundthat the photosynthetically activeradiationneededto calculateplant
proportionof CH4 beingoxidizedincreased from about50% photosynthesis were estimatedfrom the data on cloudiness
in the early stageto 95% in the late stageof rice growth.It followingthe methodusedby Raich et al. [1991].
seemsthereforethatrhizosphericCH 4 oxidationis correlated The data on wetland location, area, and vegetation
with plant physiologicalactivity. In line with the above information were derivedfromMatthewsandFung[1987].
analysis,it was assumedthat 60% of the CH4 producedin They definednatural wetlandsby integratingthe data of
the anoxic zone was oxidized at the soil-water surface, that vegetation,soil, and land inundation.This resultedin a total
oxidationin the rhizosphereincreasedwith the rate of plant globalwetlandareaof 5.3 x 109km2, occupying 3233 1ø
physiological activity, andthat togethertheymightconsume latitude x 1ø longitudegrid cells. In order to maintainthe
up to 90% of the CH4 produced.Thus,usinggrossprimary resolutionof the climate data set, we extrapolatedthe
production(GPP) to represemphysiological activity,therate wetlanddata to a resolutionof half a degree.The area of
of CH 4 oxidation (MOR) in inundated wetlands was wetlandscalculated by Matthewsand Fung [1987] was for
calculated from inundated soilsonly,butmoist/drytundrais alsobelievedto
be an importantsourceof atmospheric CH4. The area of
(8) moist/drytundracanbe estimated by subtractingthe areaof
MORt = MPRt
0.60
+0.30
GPPt
] GPPma,•
wet tundra(unforestedbogs, 0.884 x 10•2 m2) from the
globalareaof tundra,7.34 x 10•2m2 [Matthews,1983].The
distributionof moist/dry tundra was consideredto be
where GPPmax is the maximumGPP occurringduringany associated proportionallywith thoseof inundatedwetlands.
month. Soil datafor wetlandswere derivedfrom the digitizedFood
When the wetland soil is not inundated,the processof and Agriculture Organization(FAO) map [Zobler, 1986].
oxidationbecomesmore complicated.There shouldbe more The initial-statevaluesfor variousvegetationtypesdefined
oxidationof CH4, but the rate dependson the soil water by Melillo et al. [1993] and McGuire et al. [1992] in their
comere,water table position,O2 and CH4 diffusionwithin globalecosystem modelwere usedin modelcalculations.
 CAO ET AL.: CARBONEXCHANGE AND CH4EMISSIONSFROM WETLANDS 14,403

The WMEM of NPP-C thatwas decomposed

was run for each of the wetland cells until as CH4 andreleasedinto the
a year-to-year
equilibrium
wasreached.Theequilibrium was atmosphere was 2.3% in northern wetland, 4.2% in
considered to have been attained when the flux of net temperatewetland, and 4.8% in tropical wetland. These
primaryproduction,litter deposition,and organicmatter estimates are in line with the results of other studies:
decomposition
differedannually
by lessthan1 g C m-2yr4. Aselmann and Crutzen [1989] calculated that the ratio of
In MatthewsandFung's[1987]wetlanddataset,wetlands CH4-C emissionto NPP-C of wetlandswasfrom2 % to 7 %,
were classifiedinto five types: forestedbog, nonforested and data from simultaneouscarbon exchange and CH4
bog, forestedswamp, nonforestedswamp, and alluvial emission measurement showed that this ratio was 5.5%
formation.In the presentstudy,threeregionalcategories
of [Sebacheret al., 1986], 1.0-14.0% [Moore and Knowles
wetlandswere also used in the analysisof model results:1990], or 2.3% [Whiting and Chanton, 1992]. Pulliarn
northern,temperate,and tropicalwetland.The northern [1993] reportedthat 10% of the decomposed carbonwas
wetland was defined to be north of 50øN, the temperate transformedto CH4, of which 52% was oxidizedand 48%
wetlandwas in the regionsfrom 20øN to 50øN and to the emittedinto the atmosphere.
southof 30øS, and the tropicalwetlandwas between20øN
and 30øS.
Spatial Variation in CH 4 Flux

Results and Discussion

The spatial variation in CH4 emission rates from
inundated wetlands, calculated with the WMEM, is
Net Primary Productivityand Carbon Exchange presentedin Plate 1. The CH 4 fluxeswithin individualgrid
cellsrangedfrom 0.0 to 118.8 g CH4 111-2
yr-1. The mean
The annualNPP of wetlandscalculatedwith the present rate estimatesfor the five wetland types ranged from 3.6 g
modelrangedfrom0 to 1500g C m-2yr-1(Plate1). For the CH4m-2yr-1for nonforested
bogsto 38.2 g CH4 m-2yr-1for
individualvegetationcategories,themeanvaluesvariedfrom forestedswamps(Table 2). CH 4 fluxes varied widely, even
45 g C m'2 yr-1 for arcticbogsto 820 g C m'2 yr-1 for within the same climatic-ecologicalzone and for a single
tropicalswamps.Generally,this spatialdistributionof NPP wetlandtype. However, they generallyincreasedfrom the
was consistentwith other estimates.Miller et al. [1983] arctic to the tropical region. The latitudinaldistributionof
reportedthat the NPP of variousarctic ecosystems ranged CH4 fluxes showedan increasefrom the north to south,but
from19 to 450 g C m-2yr-1,andthattheaveragevaluewas a decreasebeyond30øS. In terms of wetlandtype, swamps
130g C m-2yr-1for tundraand226 g C m-2yr-1for boreal showeda muchhigher CH 4emissionrate thanbogs;forested
forests. Bartsch and Moore [1985] estimatedthat NPP was wetlandsemitted more CH 4 than nonforested(Table 2).
from50 to 150g C m-2yr-1in thesubarctic regionandfrom The high variability in CH 4 emissionrate is a reflection
100 to 200 g C m-2 yr-1 for boreal bogs. The global of the spatialheterogeneity in climate,vegetation,and soil,
ecosystemmodelpredicteda meanNPP of 120g C m-2yr-1 andtheir complexinteractions.The distributionof CH 4 flUX
for wet/moisttundra,173 g C m'2yr-1for borealwoodland, rate alongthe latitudinalaxis was relatedto temperature,but
and238 g C m'2yr'1for borealforest[Melilloet al., 1993]. more closelyto NPP and SOMD (Figure 2). CH 4 emission
Various field studies have shown that the NPP of marshes rateswere typicallyhigh in the tropics,where temperature,
and swampsin Northern American and Europe is in the NPP, and SOMD are all high. In temperatewetland, NPP
rangefrom 400 to 1000 g C m-2yr4 [McNaughton,1966; andSOMD were not necessarilylower thanthosein tropical
Van der Valk and Bliss, 1971; Baradziej, 1974]. Melillo et wetland,but CH 4 emissionrates were lower becauseof the
al.'s [1993] resultsfor temperateforestsrangedfrom 335 to lowertemperature;87% of bogswerelocatedto the northof
741 g C m-2yr-1.Few field dataareavailablefor the NPP of 50øN, while 83 % of swampswere within the tropicalzone.
tropical wetland. Our resultsin this region were somewhat The meantemperatureover the CH4 productionseasonin
lower, but close to, the estimatesof Warnant et al. [1994] bogswas 9.7øC, while in swampsit was23.8øC, and this
for tropical woodlandsand rain forests,which rangedfrom temperaturedifference,throughits effect on MPR, was
711 to 925 g C m-2yr-1. largelyresponsible for the largedifferencein the calculated
The calculatedNPP of temperateand tropical wetlands CH4 emissionratesbetweenbogsandswamps.The NPP of
was balanced by the rate of soil organic carbon forestedbogs was almostthe sameas that of nonforested
decomposition(SOMD), but the northernwetlandshowedan bogs,but the mean temperatureand SOMD was higher,
excessof NPP over SOMD, resultingin accumulation of soil givingriseto increased
CH4 fluxes(Table1). Similarly,the
organiccarbon(Table 1). Ratesof accumulationwere in the CH4emission ratefromforestedswampwashigherthanthat
range5 to 60 g C m'2 yr-1,with a meanof 33 g C m'2yr'1. from nonforestedbecauseof the higher temperature,NPP,
This was in agreement with the results of Miller et al. and SOMD.
[1983],who reportedthat from 10 to 40 g C m'2 yr'1 was A comparison betweentheresultscalculatedwithWMEM
accumulated in arctic tundra and boreal forests. and measuredCH4 emissionratesreportedin the literature
The proportionof SOMD transformedto CH4-C annually is givenin Table3. In northernwetland,CH4 emission
rates
varied from 2.0% to 40.5% (Table 1), with a mean of calculated with the model were about 2-4 times lower than
14.2% over all wetlands.Of the CH4 produced,from 70% estimates generatedfrom field measurements,but closeto
to 80% was oxidized to CO2 by methanotrophs;the the resultsof field investigations performedat the regional
remainderwas emittedinto the atmosphere.The proportion scale (Table 3). In the Yukon-KuskokwimDelta, Alaska
14,404 CAO ET AL.. CARBON EXCHANGE AND CHnEMISSIONS FROM WETLANDS

Net Primary Productivityof Natural Wetlands

Non
wetland0 50 100 200 300 400 50O 600 700 80O 1500
-1
,,Cm" yr

Methane Emission Rates from Natural Wetlands

Norl , 4 8 12 20 26 32 36 40 46 120
wetlano 0
o CH4m-yr

Plate 1. Net primaryproductivityand methaneemissionratesfor naturalwetlandscalculatedwith the model.

 CAO ET AL.: CARBON EXCHANGE AND CH4 EMISSIONS FROM WETLANDS 14,405

Table 1. Mean Net Primary Productivity(NPP), Rate of gridcellsat 54øNrangedfrom27.4 to 134.7mg CH4m'2d'l,
Decomposition(SOMD), and Carbon Transformationin the with a mean of 59.4 mg CH4 m'2 d'l. The lower CH4
Process of CH4 Emission Calculated with the Wetland emission ratesobtainedin thisstudy,compared with "point"
Methane Emission Model measurements, may arise becausepointmeasurements give
the flux ratesfrom specificsitesat a scaleof 1 m2 or less,
Latitude NPP SOMD RMED RMEE but thosefrom the modelrepresentedthe meanover 50-1500
g C m-2 g C m-2 km2;Fan et al. [1992]andBartlettet al. [1992]showedthat
y-1 y-1 the CH4 flUXrate from enclosuremeasurements was higher
than that from tower and aircraft measurements in the same
80-70øN 183 66 0.06 0.01 area.
70-60øN 172 138 0.12 0.03
In temperateand tropical wetlands,our resultswere in the
60-50øN 243 218 0.16 0.04
samerange as reportedin field measurements.However, the
50-40øN 413 421 0.22 0.06
mean CH 4 emissionrate from temperatewetlandcalculated
40-30 øN 618 620 0.16 0.04
with the modelappearedhigherthanthat suggested by field
30-20øN 615 613 0.25 0.07
experiments.Using data from field campaigns,Bartlett and
20-10øN 461 465 0.28 0.08
Harriss [1993] calculatedthat the averageCH4 flux from
10-0øN 599 604 0.24 0.06
temperate swampswas lower than those from arctic tundra
0-10øS 667 675 0.25 0.06
and boreal forest (Table 3). The temperature,NPP, and
10-20øS 555 548 0.26 0.06
20-30øS 668 667 0.29 0.07
SOMD in temperatewetlandare muchhigherthanthosein
northernwetland, and the CH4 emissionrate would therefore
30-40øS 709 696 0.21 0.05
be expectedto be higher.
40-50øS 622 597 0.12 0.03
For moist/dry tundra, the calculatedCH4 emissionrates
variedfrom0.0 to 55.0 mg CH4m-2d4. Thiswaslowerthan
RMED is the proportionof the decomposed organiccarbon
that of the inundatedwetlandsat the same locationby a
being transformedinto CH4 in methaneogenesis.RMEE is
factorof 5 to 10. The meanratesat variouslatitudesranged
the proportion of the NPP-C being released into the
from 1.0 mg CH4 m-2 d'• to 13.0 mg CH4 m-2 d4. The
atmsophereas CH4.
averageCH 4 emissionrate from all moist/drytundrawas 8.5
mg CH4 m'2 d4. This rate was in line with the resultsof
(61 ø5'N, 162ø92'W), Fan et al. [1992] measuredregional most field measurements. A recent review of 14 field
CH4 fluxesrangingfrom 25 to 30 mg CH4 m 2 d'•, using experimentsby Bartlett and Harriss [1993] showed CH4
eddy correlationtechniqueswith both tower and aircraft. fluxes from moist/dry tundravaried from 0.6 to 29 mg CH 4
The mean CH4 emissionrate calculatedwith our model for m-2d4, with a meanof 7.0 mg CH4 m-2d-1.
gridcellsat 61øN was 25.7 (9.5-85.8) mg CH4 m-2 d-•.
Moore et al. [1990] estimatedCH4 fluxes from a large area SeasonalVariations in CH 4 Emission
(130 km2) in Scheffervilleof Canada(54øN, 66øW). The
CH4 flux rates estimatedfor variouswetland types ranged Figure 3 illustratesthe seasonalpattern in CH4 emission
from7.5 to 71.9 mg CH4m 2 d4, andthe averageflux from rate for different wetlands. For northern wetland, CH 4
the wholeareawas56.3 mg CH4 m-2d'•. Our estimates for emissions in winter were insignificant, while those in

Table 2. MethaneEmissionRatesof VariousWellandTypes,CalculatedUsingtheWetlandMethane

Emission Model

Ratesof CH4 Emission,mg CH4m-2d4

Total
Latitude Forested Non Forested Non Alluvials Mean Emission,
Bogs forested Swamps forested Tg yr-•
Bogs Swamps

80-70 øN 18.0 12.2 6.9 0.0 0.0 12.1 0.3

70-60øN 35.7 22.4 29.6 31.6 10.8
60-50 øN 57.3 30.6 87.8 103.5 51.2 10.7
50-40øN 90.8 35.8 177.5 164.4 120.5 7.4
40-30øN 168.2 126.4 125.5 122.5 143.5 3.2
30-20øN 119.4 242.5 318.7 334.5 235.1 3.6
20-10øN 239.7 170.8 192.7 8.2
10-0øN 206.6 161.2 203.9 150.4 195.7 182.1 10.2
0-10øS 205.5 205.9 152.3 200.9 198.5 14.2
10-20øS 221.3 164.5 154.5 184.3 8.2
20-30 øS 206.3 318.3 249.8 10.8
30-40øS 145.1 213.3 180.5 3.1
40-50øS 89.1 89.1 0.1
14,406 CAO ET AL.' CARBON EXCHANGE AND CH4EMISSIONS FROM WETLANDS

900 30

800-
'25
.o 700-
._

E
ß 600- -2O

.c 500'
E
'• 400'

-lO
'• 300'
a-
n 200'
z

100-

O• ' ' 0
90 •0 3'o
North Latitude South

Temperature [] NPP -•-- Methaneemission

Figure2. Latitudinal
distributions
of temperature
(øC),netprimary
productivity
(NPP,g C m-2yr-•),andmethane
emission
(10-•g CH4m-2yr-•).Thetemperature
shownisthemean
temperam.re
inthemethane
production
season.

summeraveragedfrom 40 to 60 mg CH4 m'2 d-1. In temperature,NPP, and SOMD. Tropical wetlandshoweda

temperatewedand, CH4 emissionrates in summerranged different seasonalpattern in CH4 emission.BecauseCH 4
from 100to 400 mg ca 4 m'2d-1,whilein theotherseasons productionceasedduringthe dry season,the shift between
the rate varied from 20 to 150 mg ca 4 m'2 d'l. These dry andwet seasons playedan importantrole in determining
seasonalvariationswere closelycorrelatedwith changesin the seasonalvariation in CH 4 fluxes. To the north of the

Table 3. A ComparisonBetweenthe CalculatedCH4 EmissionRatesWith the Model and Values

Reportedin the Literature,DerivedFrom Field Measurements
i ,

CH4 EmissionRatesFrom CalculatedCH4 EmissionRates

Field Measurements, with the Present Model,
mg CH4 m'2d'l mg CH4 m'2d'l
Range Meana Range Mean
Northern wetland 39.7
Wet tundra 20-150b(25-85)d 96 6.4-50.6
Boreal 30-200c(19-50)d 87 12.1-132.6
Temperatewetland 149.6
Bogs 20-300 e 135 50.6-278.2
Forestedswamps i0-250f 75 35.4-301.4
Nonforestedswamps 5-150 s 70 12.3-286.1

Tropicalwetland 199.0
Forestedswamps 75-350 h 165 127.9-360.3
Nonforestedswamps 30-390 h 233 120.3-474.7

a Bartlett and Harriss [1993]

bSebacheret al. 1986; WhalenandReeburgh[1990], Bartlettet al. [1992], andMorrissey
and Livingston;[1992].
c Crill et al. [1988]; Vitt et al. [1990]; andMoore and Knowles1989).
a ratesobtained
at regionalscale[Bartlettet al., 1992;Fanet al., 1992;Whalen
and Reeburgh,1992; Christensen,1993].
e Yavittet al. [1990]; Routletet al. [1992a]; and Harffss et al. [1985].
f Harffssand Sebacher [1981], andWilson et al. [1989].
gBarber et al. [1988], Burkeet al. [1988]; Harffss et al. [1988].
hBartlettet al. [1990], Devol et al. [1988, 1990]; Tathyet al. [1992], Keller [1990].
 CAO ET AL.' CARBONEXCHANGEAND CH4EMISSIONSFROM WETLANDS 14,407

180! 1990; Christensen,1993], while in tropicalwetland,it may

16o.;L arisefrom the transitionbetweenwet anddry seasons.In the
Amazonfloodplain,Devol et al. [1990] observedthat the
12o
averageCH4 flux duringthe periodof low waterlevel was
aboutone third of that for the period of high water levels.
.•1oo-
60-
Total CH 4 Emission Into the Atmosphere
40

Globally, total CH 4 emissionfrom natural wetlandswas

0::
1 2 3 4 5 6 7 8 9 10 11 12 estimatedto be 92 Tg yr-•, of whichinundatedwetlandand
Month moist/drytundraemitted85 Tg and7 Tg, respectively.The
latitudinal distribution of fluxes from the various wetland
= Northern [] Temperate ,•, Tropical typesis presentedin Figure 4. The large fluctuationsin CH4
emission shown in this figure representthe combined
Figure3. Seasonal
variations
inmethane
emission
rates
(mgCH4m'2 variationsin flux and wetlandarea along the latitudeaxis.
d'l) in thenorthern,temperate,
andtropicalwetland.
High peaksappearedin bothnorthernandtropicalwetlands.
In northernwetland, CH4 flux rates were low, but the area
equator, the rainy season was generally from May to of wetlandwas large. The inundatedwetlandcovers2.7 x
October, when the mean CH4 flux rates rangedfrom 150.0 10•2m2, accounting for 53% of the totalareaglobally,and
to 300.0mg CH4m-2d-1,butin dry months,CH4flux rates thereis 6.46 x 1022m2 moist/drytundra.Northernwetland
were 50-120 mg CH4 m-2d-•. The highestCH4 flux rates emitted16.7 Tg yr-•, andmoist/drytundraemitted6.6 Tg
occurredin May or June. To the southof the equator,the yr-•; togetherthey contributed
25% to the globalemission.
high temperatureand rainy seasonsoccurbetweenOctober The mean CH4 flux rate from temperatewetlandwas much
and March. Accordingly, high CH4 emission rates, of higher thanthat of the northernwetland,but the area, 5.8 x
150-300mg CH4 m'2 d-1, appearedin theseseasons.In 10• m2, is smaller.Its totalemissionwas 17.2 Tg yr-•. The
contrast,the averageCH4 flux rate in otherseasonswasonly CH4 emissionfrom tropical wetland was 51.4 Tg yr-•,
10-100mg CH• m-2d-•. accountingfor 56% of the global emission.Althoughthe
Field experiments
havealsoshowna significantseasonal area of tropical wetland was less than that of northern
variation in CH4 flux rates. Whalen and Reeburgh [1992] wetland, the mean CH 4 flUX rate was much higher. CH 4
reported that CH4 flux rates increasedby 4 orders of emissions from different wetland types are shown in
magnitudefrom winter to midsummerin their 4-year Table 3.

observationsin tundra. In northernand temperatewetland, The calculatedtotal CH 4 emissionfrom moist/dry tundra

seasonalvariation was found to be driven largely by wascloseto theestimates
of from4.0 Tg yr-• to 4.8 Tg yr-•
temperature[Bartlettet al., 1992], soil moisture[Mooreet [Bartlett and Harriss, 1993; Bartlett et al., 1992; Fung et
al., 1994; Roulet et al., 1992], or both [Moore and Knowles, al., 1991], but it is much smaller than the result of

2.5
Forested
Bog
Nonforested
Bog
+ Moist/dry
tundra
ForestedSwamp
_

• Nonforested
Swamp
1 Alluvial
førmatiøns

0.5-

0
L_
ii iii ii iii ii iii Ii ii iii ii iii ii Iiiii Iiiii iiiii iiiii ii I

90 60 30 0 30 60 90
North Latitude South

Figure4. Latitudinaldistribution
of totalannualmethaneemissions
fromvariouswetlandtypes,predictedwith the
WMEM.
14,408 CAO ET AL.: CARBON EXCHANGE AND CH4 EMISSIONS FROM WETLANDS

have measuredCH4 fluxes in Siberia and the European

Arctic to discoverthatfluxesthereare considerably lessthan
thosein Alaska. The Alaskanvalueshave previouslybeen
assumedto apply throughoutthe northernhemisphere,and
so overestimates of the total flux are to be expected.
Seasonalvariation in total CH 4 emissionis illustratedin
Figure 5. The global emissionof CH4 in July (12 Tg) was
abouttwice as large as in Januaryor December.In northern
wetland,CH 4 emissionsin summer(June-Augus0accounted
for more than 70% of the annual emission, while almost no
CH4 was releasedin winter. In temperatewetland,summer
o
emission still dominated(47 %), but there was some CH4
Month
emittedin winter (12%). In tropicalwetland,mostCH4 was
emittedduring the wet season;only 28% of the CH4 was
Figure 5. Seasonalvariation in methaneemissionfrom wetlands releasedin the dry season.
predictedwith the WMEM.
Our results are comparedwith other estimatesof the
globalCH 4 emissionin Figure 6. Matthewsand Fung [1987]
Christensen [1993], 12.1-22.9Tg yr4. Accordingto hisfield used the limited databaseof CH 4 fluxes then availablefor
measurements,Christensenassumedthe mean CH 4 flUXwas northernwetlandto estimateglobalemissions.The CH4 flUX
25 mg CH4 m-2 d'l, which is twice the rate usedin this rateusedfor bogswas about2 timeshigherthanfor swamps
study. However, more recently, Christensenet al. [1995] and 7 times higher than for alluvial formations.
4O

35-

5-

0
60 30 0
North Latitude South

250-

c
.o 150-
.e_.
E
ß 100-

50-

0
90 go o
North Latitude South

(b)
Bartlett and • Asclmann and [] Matthews and
Hatriss(1993) Crutzen(1989) Fung(1987) Modelsimulation

Figure 6. Comparisons
betweenthe CH4 emissionscalculatedwith the WMEM and publishedestimates:(a) total CH4
emissions
(Tg yr4), (b) meanCH4 emission
rates(mg CH4 m'2
 CAO ET AL.: CARBON EXCHANGE AND CH4 EMISSIONS FROM WETLANDS 14,409

Consequently,
northernwetlandwasestimated
to contribute
asmuchas58.6%
ofthetotal
CH4emission,while
the • 0.9 "=-Q,o = 1.$
tropical
wetland
only
contributed
28.4%
ß It seems
that
the • o.s. -,a- Q,o = 2.0
role of northern wetland was overestimated.Aselmann and '• o.?. -•- Q,o = 2.5
• 0.6'
Crutzen[1989] classifiedwetlandsinto bog, fen, swamp, -•- Q,o = 3.0
•' 0.5-
marsh,floodplain,andshallowlake,andgaveeachof them
a ell4 fluxrateandanactiveell4 producing
period.Asthe • 0.4.
flux ratesgivento swamps,
marshes,
andfloodplains
were E o.a,
higher
andtheperiods
of ell4 production
werelonger,
they =•0.2-
o

estimated
thattropicalwetlandaccounted
for 55% of the o

total ell4 emission,while the northernand temperate o oo

g. 5 10 15 20 25 30

wetlands contributed 31% and 14%, respectively. By Temperature

(øC)
reviewingrecentfield measurements of ell4 emissionrates, o
(a)
30
Bartlett and Hatriss [1993] defined CH4 flux rates for -• 1.2
variouswetland types in differentregions,and calculated
25
thatnorthern,temperate,
andtropicalwetlandscontributed ß 0.8-

28.9% 8.5% and62.6% respectively,to the totalglobal c- {3.8

20 •
CH4 emission.The total CH4 emissionestimatedusingthe
WMEMwaswithin
therange
ofexisting
estimates
andthe E 0.4
•5 ß
contributions
of the three latitudinalregionswere quite -Z o.a
similar to the results of Aselmannand Crutzen [1989] and
-•0 •
Bartlett and Harriss [1993], although the estimate for o -0.2

temperatewetlandappearedsomewhathigh. c- -0.4

-o.6
o -0.8 o
Sensitivity Analysis •- 90 •0 o 3'0 6'o 90
North Latitude South
Sensitivityto Model Parameters
= 010 = 1.5 [] 010 = 2.5 A 010 = 3.0 .............
Temperature
Temperatureand the positionof the soil water table are
important factors modulating CH4 production, but the (b)
reportedquantitativerelationshipsvary between different
studiesandare ratheruncertain.ThereforetheconsequencesFigure 7. Sensitivityanalysis. (a) The relationshipsbetween
of usingdifferentrelationshipsin the modelwere studied. temperatureandtherateof methanogenesis withdifferentQ10values.
The relationshipwith Q10= 2.0 wasusedin the presentmodel. (b)
In the presentmodel, a Q10valueof 2 was usedto define
Thechangein CH4emissions withdifferenttemperaturerelationships,
the directinfluenceof temperatureon methanogenesis, but relative to the resultsof the presentmodel.
it mayrangefrom 1.2 to 3.5 [Conradet al., 1987;Kellyand
Chynoweth,1981]. Thusdifferenttemperature relationships
with Q10valuesof 1.5, 2.5, and 3.0 (Figure 7a) were used
in themodelto showtheeffectonCH4emissions. Compared watertablepositionon CH 4productionwere considered.The
to the resultspresentedabove, a Q10value of 1.5 gave an linear relationship was obtained by assuming that the
increasein the rate of CH4 emission,while valuesof 2.5 and proportionalCH 4 productionfactor is 0.0 when the water
3.0 led to a reduction.The scaleof the changeswaslargein table position is 30 cm below soil surface and increased
northernand temperatewetlands(30% to 100%), and small linearly with the increasein soil water tableheightto 10 cm
in tropical wetland (10-30%) (Figure 7b). As Figure 7a above the soil surface. The exponentialrelationshipwas
shows,the larger the Q10value, the larger the reductionin derived from the data of Moore and Knowles [1989]. The
CH4 productionrate with decreasing temperature;the lower calculatedCH 4 emissionswith thesetwo relationshipswere
the temperature,the larger the differencebetweenCH4 higherby 9.5 % and 20.7 %, respectively,thanthe estimates
productionrateswith differentQ10values.The estimateof predictedby the presentmodel.
total CH4 emissionfrom inundatedwetlandswith a Q,0 of
1.5 was 108.4 Tg yr'l, whilethosewith Q,0of 2.5 and3.0 Responseto Changesin Climatic Conditions
were73 Tg yr4 and65 Tg yr4, respectively. Thususing
different (reasonable) temperature relationships for General circulation model (GCM) projectionsindicate
methanogenesis mightcausevariationsin the resultingtotal that, with a doubling of the current concentrationof
CH4 emissionof from -24 % to 27 %. atmospheric CO2,an averageannualglobalwarmingof from
Various relationshipsbetweenwater table positionand 1.9 to 5.2øC [Lashofand Ahuja, 1990] and a globalannual
CH4 emissionhave been reported[Vourlitiset al., 1993; precipitation increaseof 15% may occur[Boeret al., 1990].
Moore andKnowles,1989]. Linearandexponentialfunctions The temperatureand precipitationchangenot only affects
(Figure 8) were tested in the model to calculate CH4 methanogenesis directly,but alsothe equilibriumof carbon
emissionsfrom moist/drytundra,where the effectsof soil exchange in wetland ecosystems. The analysisdescribed
14,410 CAO ET AL.' CARBONEXCHANGE AND CH4EMISSIONSFROM WETLANDS

1
water table as a result of increasedevapotranspiration.
The
+ The relationshipusedin
balancebetweentheseeffectsdeterminesthe changein CH 4
the presentmodel
e-
0
0.8-
• A linear relationship ., emission. At constanttemperature,increasingprecipitation
of between 5 % and 15 % causedincreasesin SOMC, NPP,
SOMD, and YMER. For example, when precipitationwas
0.6-
increasedby 10%, SOMC wasincreasedby 6.8%, NPP and
0.5-
SOMD were increasedby 5.8% and 11.8%, respectively,
0.4- and YMER was enhancedby 35 %.
0.3-
0.2- (a) NorthernWetland
0.7

o.1
e- 0.6'
o
-30 -25 -20 -15 -10 -5 0 5 10
e- 0.5.
water table position(cm) 0

o 0.4'
Figure8. The threerelationships
betweensoilwatertableposition ._=

andmethanogenesisusedin the sensitivityanalysis. e- 0.3-

• 0.2-
¸
below wasperformedto testthe sensitivityof the model,and
,_

00.0.1
it is not intendedthat the resultsbe interpretedas a realistic p
predictionof future trends. 0
o 1 2 3 4
For inundated wetlands, the sensitivity to temperature Temperatureincrease
increaseacrossthe range from 1øC to 4øC was studied
basedon the assumption that soil water conditionsremained 0.5
(b) Temperate Wetland
unchanged(Figure 9). In northernwetland,meanNPP, soil
carbonstorage(SOMC), SOMD, and annualCH4 emission to 0.4-
rates (YMER) all increased. In temperate wetland, the
positive effect on NPP, SOMD, and YMER remained, 0
• 0.3-
althoughthe increase was smaller than that in northern
._•
wetland,anda negativeeffecton SOMC wasobserved.This • 0.2-

was due to a larger increasein the rate of decomposition

relative to the rate of carbon input into the soil from to 0.1

vegetation.In tropical wetland, CH4 emissionwas still ¸

1• 0
enhanced, but SOMC, NPP, and SOMD declined with o

increasedtemperature.The decreasein NPP wascausedby

-0.1
an excessof plantrespirationoverthe rateof photosynthesis. 0 • • • 4
The low rate of increase in carbon input into the soil Ternperature increase

comparedwith the increasein the rate of decomposition (c) Tropical Wetland

resultedin a decreaseof soil carbonstorage,which in turn,
reducedthe rate of decomposition. It was no surprisethat
x o.1
northernwetlandshoweda largerandmorepositiveresponse
to temperature increase. Increased temperaturewould o
• o
improveconditionsfor plant growthand microbialactivity o

and so enhancethe rate of carbon exchangeand CH4 ß -o.1-

emission.In the caseof a 2øC increase,the CH4 emission
rate was increasedby 36% for northernwetland,29% for J:
o
-0.2-
temperatewetland,and 12% for tropicalwetland.The total
CH4 emissionfrom all inundatedwetlandscouldbe increased o -0.3-

by 19%. o

For moist/drytundra,the responses bothto temperature ,o -0.4

0 • • • 4
and precipitationchangeswere studied.When temperature Temperature increase
was increasedby between IøC and 5øC, both NPP and
SOMD were enhanced;soil carbon storageand methane
= SOMC '- NPP .*: SOMD [] YMER
emissionwere increasedwith the 1øC and2øC temperature
increasesbut decreased with a greatertemperature increase Figure 9. Changesin net primaryproductivity(NPP), soilorganic
(Figure 10). Temperature increase can, on one hand, carbondecomposition (SOMD), soilorganiccarbonstorage(SOMC),
stimulateCHn productionby increasingcarbon substrate andyearlymethaneemission(YMER) in inundatedwetlandsarising
supply and the activity of methanogens but, on the other fromtemperature increase( øC), relativeto thevaluesobtainedunder
hand, may attenuateCH4 productionby loweringthe soil current conditions.
 CAOETAL.'CARBON
EXCHANGE
ANDCH4EMISSIONS
FROMWETLANDS 14,411

arenchymoussystemof plantscan also serveas an effective

pathway for the transport of ca 4 from the soil to the
x 0.4-
atmosphere.Thusthe presenceof plantsmay alsoreducethe
0 0,2- exposure of ca 4 to potential oxidation [Whalen and
Reeburgh,1988]. In addition, environmentalfactors, suchas
.c: O•
temperatureandpH, alsoinfluenceca 4 oxidation[Dunfield
c -0.2-
et al., 1993], but theseprocessesand relationships
are also
_ -0.4- poorly understood.Hence the treatmentof this important
• -o.6-
processwithin the current model is entirely empirical and
rather crude.
o -0.8
Further improvementsto the model depend on an
Temperature increase improvementin the understandingof importantprocesses
and relationshipsinvolved.For progressto be madein this
= SOMC •, NPP '" SOMD • YMER area, the focusof field studiesshouldmove away from flux
measurementper se, toward investigationof the mechanistic
Figure10. Changes in netprimaryproductivity (NPP), soilorganic
carbondecomposition (SOMD), soilorganiccarbonstorage (SOMC),
processesand quantitative relationshipsconcerning the
andyearlymethaneemission(YMER) of moist/drytundraarising impact of environmentalfactors on CH 4 productionand
from increasedtemperature(øC), relativeto the valuesobtained oxidation in soil. Methanotrophic oxidation especially
under current conditions. warrants further research. Since oxidation consumes much
of the ca 4 produced,it is probablya major determinantof
the magnitudeof CH 4 emissionsinto the atmosphere.The
Conclusion and Discussion resultsof the present study were comparedwith estimates
based on measurementsmade over a small spatial scale.
The presentstudyrepresentsthe first publishedattemptto Furthervalidationwith more dataat the regionalscalewould
developand use a process-based model to estimatespatial allow development of the model and a more reliable
and seasonal variations in CH4 emissions from natural parameterization.
wetlands at the global scale. The CH4 emission rates As with any model, the resultof the WMEM dependson
calculated using the model for various wetland types the qualityof the datausedto drive it. The spatialresolution
were in the same range as those suggestedby field of soil, vegetation,and wetland,and the temporalresolution
measurementsreported in the literature. The total ell4 of the climate were limiting factors. Data on the seasonal
emission,92 Tg yr-•, is in the middleof the existingrange variationin wetland area and soil water table positionwith
of estimates:higher than the results of Aselmann and fine spatialresolutionare not currentlyavailable,but these
Crutzen[1989], but lower than others[Matthewsand Fung, are importantfactorsin determiningCH 4 emissions.While
1987; Bartlett et al., 1990; Fung et al., 1991; Bartlett andthe developmentof a more detailed databasefor soil,
Harriss, 1993]. Sensitivityanalysisshowedthat the response vegetation, and climate is in progress, remote-sensing
of CH 4 emission to climate change depends upon the imagery may also provide useful informationon vegetation
combined effects of soil carbon storage, rate of phenology, seasonalvariations in soil temperatureand
decomposition, soil moisture,and activityof methanogens. moisture, river water level, and wetland area. When the
The presentmodelstill containssomeuncertainties. It is presentmodel is coupledto thesedatabases,its ability to
clear that the model is sensitive to the values of the simulateand interpretthe globalpatternsof CH 4 emissions
parameters describingthe response of CH4 productionrate shouldbe greatly enhanced. Nevertheless,the "realistic"
to water table position and temperature.However, the resultsobtainedfrom the model demonstratethe ability to
relationshipsbetweenCH4 productionand soil water table simulateCH 4 emissionswithin the framework of climate-
positionand temperaturereportedin the literaturevary vegetation-soilinteractions at the global scale. This
widely and are uncertain[Svensson and Rosswall,1984; represents a major stepforwardin the quantification of CH4
Sebacheret al., 1985]. Even if theseparametervaluescould emission from global wetlands, and when linked with
be determinedreliably, the scaleat which the model must scenariosof climate and land utilization change,the model
operateimposesthe need for a "smoothing" of the true will providea basisfor predictionof futureCH4 emissions.
spatialand temporalheterogeneityin soil moistureand
temperaturevalues.
The model's treatmentof CH 4 oxidationprovides an Appendix Calculation of Soil Moisture and
exampleof an aspectof the carbon flux that is poorly Water Table Position for Dry/Moist Tundra
understood.The availabilityof 02 in the soil may be a chief
factor controllingthe oxidationrate. It is relatedto soil Monthly variations in soil moisture and water table
moisture,02 diffusionthroughthe soil-waterbody or plant position in the moist/dry tundra were simulatedwith a
aerenchymous system,and competitionfor 02 by other simpleone-dimensional(vertical) model as follows.
processes[King et al., 1990; Gerardand Chanton,1993]. For the moist/dry tundra, water inputswould be simply
Plants may enhanceca n oxidationby providing 02 to precipitationand snowmelt.Accordingto Abet and Federer
methanotrophs associatedwith the rhizosphere,but the [1992], a constant fraction of precipitation (IFRA) was
14,412 CAO ET AL.: CARBON EXCHANGE AND CH4EMISSIONS FROM WETLANDS

considered to be intercepted
andevaporated beforeentering Wt = [z'SAT
SW(Z)
dZ (16)
intothe soil (12%). The remainingprecipitation
wasdivided j0

betweensnowand rain. The partitionwas calculatedas

where SAT is the volumetric water content at the saturation
point. Equation(16) can be integratedand invertedto give
a quadraticfunctionfor the water table depthas a function
of total water content, SWC. After the water balance for
SNFC = 0.0 > 5oc each time step was solved, the water table depth was
TEM t _
calculated from this quadratic equation, and the water
contentof the unsaturatedzone was determinedfrom (16).

SNFC
=ITEM
t-21 -7
2øC < TEMt < 5øC (10)

SNFC = 1.0 TEMt _<2øC References

(PRCP, cm) Abet, J.D., and C.A. Federer,A generalized,lumped-parameter

whereSNFC is the fractionof theprecipitation
as snow. TEM is monthlymeantemperature(øC).
Snowmelt(SNME) (cm) was given as
SNME = 4.5TEMt
Total water input into the soil was
WATI = PRCP(1 - IFRA - SNFC)+ SNME (12)
In Frolking and Crill's
evapotranspiration
(AEP) was givenas
AEP= PE
model of photosynthesis,evapotranspirationand net primary
productionin temperateand borealecosystems,Oecologia,92,
463-474, 1992.
Albera, T., Growth and root developmentof lowlandrice and its
TEMt > l oC (11) relationto oxygensupply,Plant Soil, 5, 1-18, 1953.
Aselmann, I., and D.J. Crutzen, Global distribution of natural
freshwaterwetlands and rice paddies, their net primary
productivity, seasonalityand possiblemethaneemissions,J.
Atmos. Chem., 8, 307-385, 1989.
Baradziej,E., Net primary productionof two marshcommunities
nearIspiniain the Niepolomiceforest(southPoland),Ekol. Pol.,
[1994] model, soil water 22, 145-172, 1974.
Barber, T.R., R.A. Burke and W.M. Sackett, Diffusive flux of
methanefromwarmwetlands,GlobalBiogochem.Cycles,2, 411-
425, 1988.
Zw < Zs Bartlett, K.B., and R. C. Harriss, Review and assessmentof methane
(13) emissionsfrom wetlands,Chemosphere, 26, 261-320, 1993.

AEP
=[PE(Zw-
Zm)]
Z>ZS ,

Zf- Zm w
•

Bartlett, K.B., P.M. Crill, J.A. Bonassi,J.E. Richey, and R.C.

Harriss, Methane flux from the Amazon River floodplain:
Emissionsduring rising water, J. Geophys.Res., 95, 16,772-
wherePE is the potentialevapotranspiration
calculatedwith 16,788, 1990.
Bartlett, K.B., P.M. Crill, R.L. Sass,R.C. Harriss, and N.B. Dise,
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