Table Of ContentRespirationinAquaticEcosystems
This page intentionally left blank
Respiration in Aquatic
Ecosystems
EDITED BY
PaulA. del Giorgio
UniversitéduQuébecàMontréal,Canada
Peter J. le B.Williams
UniversityofWales,Bangor,UK
1
3
GreatClarendonStreet,OxfordOX26DP
OxfordUniversityPressisadepartmentoftheUniversityofOxford.
ItfurtherstheUniversity’sobjectiveofexcellenceinresearch,scholarship,
andeducationbypublishingworldwidein
Oxford NewYork
Auckland Bangkok BuenosAires CapeTown Chennai
DaresSalaam Delhi HongKong Istanbul Karachi Kolkata
KualaLumpur Madrid Melbourne MexicoCity Mumbai Nairobi
SãoPaulo Shanghai Taipei Tokyo Toronto
OxfordisaregisteredtrademarkofOxfordUniversityPress
intheUKandincertainothercountries
PublishedintheUnitedStates
byOxfordUniversityPressInc.,NewYork
©OxfordUniversityPress2005
Themoralrightsoftheauthorhavebeenasserted
DatabaserightOxfordUniversityPress(maker)
Firstpublished2005
Allrightsreserved.Nopartofthispublicationmaybereproduced,
storedinaretrievalsystem,ortransmitted,inanyformorbyanymeans,
withoutthepriorpermissioninwritingofOxfordUniversityPress,
orasexpresslypermittedbylaw,orundertermsagreedwiththeappropriate
reprographicsrightsorganization.Enquiriesconcerningreproduction
outsidethescopeoftheaboveshouldbesenttotheRightsDepartment,
OxfordUniversityPress,attheaddressabove
Youmustnotcirculatethisbookinanyotherbindingorcover
andyoumustimposethissameconditiononanyacquirer
AcataloguerecordforthistitleisavailablefromtheBritishLibrary
LibraryofCongressCataloginginPublicationData
(Dataavailable)
ISBN 0198527098(hbk)
ISBN 019852708X(pbk)
10 9 8 7 6 5 4 3 2 1
TypesetbyNewgenImagingSystems(P)Ltd.,Chennai,India
PrintedinGreatBritain
onacid-freepaperbyAntonyRoweLtd.,Chippenham
Preface
Respiration,initsvariousbiochemicalmanifestations,istheprocessbywhichallorganismsobtainvital
energyfromavarietyofreducedcompounds,andrepresentsthelargestsinkoforganicmatterinthebio-
sphere.Althoughrespirationisatthecenterofthefunctioningofecosystems,muchofcontemporaryecology
haschosentofocusattentionandresearchontheproductiveprocessesratherthanoncatabolism.Almost
certainly,respirationrepresentsamajorareaofignoranceinoutunderstandingtheglobalcarboncycle.We
decidedtoembarkinthisprojectduringthe2000ASLOmeetinginCopenhagen, rightafterasessionon
aquaticrespirationorganizedbyErikSmithandoneofus.Toourknowledge,itwasamongthefirstsessions
entirelydevotedtoaquaticrespirationataninternationalmeeting, andalthoughwehadhopedtoattract
someinterest,weneverexpectedtofillthelargestroomintheconferencecenter!Itbecomeclearthatthere
wasatremendousinterestinthetopicwithintheaquaticscientificcommunity,andthattherewasaneedfor
synthesisanddirectioninthisareaofresearch.Notextbookthathadreviewedandsynthesizedtheextant
informationonrespirationinnaturalaquaticsystems,independentofproductionorothertraditionalareasof
focus.Inspiteofitsobviousecologicalandbiogeochemicalimportance,mostoceanographicandlimnologi-
caltextbooksusuallydealwithrespirationsuperficiallyandonlyasanextensionofproduction.Wesetout
tofillthisgapandprovidethefirstcomprehensivereviewofrespirationinaquaticsystems,withthehelpof
anoutstandinganddiversegroupofresearcherswholaiddownabiochemicalbasis,examinedthepatterns
andscalesofrespirationindiverseaquaticecosystems.Theresultisasynthesisthatspansthemajoraquatic
ecosystemsofthebiosphere,andanindepthanalysisofthecurrentstateofunderstandingofrespirationin
aquaticsystems.Itisourhopethatthiscollectiveeffortwillhelpestablishthemainscientificquestionsand
challengesthatfacethisparticularfield,andmoreimportant,helpbringrespirationintofocusasapriority
areaoffutureresearch.
Thebookismainlydirectedtorespirationoccurringinthewatercolumnofaquaticsystems,becausethere
areotherexcellenttextbooksthatextensivelydiscusssedimentmetabolism.Sedimentrespirationisdiscussed
inthebroadercontextinindividualchapters, aspartofthespecificaimofintegratingwatercolumnand
benthicmetabolicprocesses.Wefirst(Chapters1-5)laydownabasistounderstandrespirationwithinmajor
categoriesoforganisms,thephotoautotrophs,thebacteria,theprotozoaandtheplanktonicmetazoa.Wethen
(Chapters6-12)considertheprocessinaseriesofaquaticecosystemsandthecurrentstatusofattemptsto
modelrespirationinvariousplanktongroups(Chapter13).Asthebookprogressesweshallseektoassessthe
currentstateofknowledgeconcerningrespirationinaquaticecosystems,andtoaddressaseriesofquestions
thatarekeytounderstandingthisprocessattheecosystemandbiosphericlevel.
Finally,wewishtothankHughDucklow,JonathanCole,MikePace,IanJointandMortenSøndergaard
forreviewingvariousportionofthisbook.WealsothankIanShermanandAnitaPetrieofOxfordUniversity
Pressfortheirpatienceandencouragementforthisproject,AnnMitchelforprovidingacoverdesignand
PaulaCondeforhelpineditingmanuscripts. PauldelGiorgioacknowledgesthefinancialsupportofthe
NationalScienceandEngineeringCouncilofCanada.Finally,wewouldliketodedicatethisbooktoLawrence
PomeroyandtothelateRobertPeters,twopioneersandvisionarieswhounderstoodearlyontheimportance
ofrespirationinaquaticecosystems.
This page intentionally left blank
Contents
Listofcontributors ix
1 Respirationinaquaticecosystems:historyandbackground 1
PeterJ.leB.WilliamsandPaulA.delGiorgio
2 Ecophysiologyofmicrobialrespiration 18
GaryM.King
3 Respirationinaquaticphotolithotrophs 36
JohnA.RavenandJohnBeardall
4 Respirationinaquaticprotists 47
TomFenchel
5 Zooplanktonrespiration 57
SantiagoHernández-LeónandTsutomuIkeda
6 Respirationinwetlandecosystems 83
CharlotteL.Roehm
7 Respirationinlakes 103
MichaelL.PaceandYvesT.Prairie
8 Estuarinerespiration:anoverviewofbenthic,pelagic,andwholesystemrespiration 122
CharlesS.Hopkinson,JrandErikM.Smith
9 Respirationanditsmeasurementinsurfacemarinewaters 147
CarolRobinsonandPeterJ.leB.Williams
10 Respirationinthemesopelagicandbathypelagiczonesoftheoceans 181
JavierArístegui,SusanaAgustí,JackJ.Middelburg,andCarlosM.Duarte
11 Respirationincoastalbenthiccommunities 206
JackJ.Middelburg,CarlosM.Duarte,andJean-PierreGattuso
12 Suboxicrespirationintheoceanicwatercolumn 225
LouisA.Codispoti,TadashiYoshinari,andAllanH.Devol
13 Incorporatingplanktonrespirationinmodelsofaquaticecosystemfunction 248
KevinJ.Flynn
14 Theglobalsignificanceofrespirationinaquaticecosystems:fromsinglecells
tothebiosphere 267
PaulA.delGiorgioandPeterJ.leB.Williams
This page intentionally left blank
List of contributors
SusanaAgustí AllanH.Devol
IMEDEA(CSIC-UIB) SchoolofOceanography
GrupodeOceanografíaInterdisciplinar(GOI) UniversityofWashington
InstitutoMediterráneodeEstudios Seattle,WA98195
Avanzados,C USA
MiquelMarqués,21
CarlosM.Duarte
07190Esporles(Mallorca)
IMEDEA(CSIC-UIB)
Spain
GrupodeOceanografíaInterdisciplinar(GOI)
InstitutoMediterráneodeEstudiosAvanzados,C
JavierArístegui
MiquelMarqués,21
FacultaddeCienciasdelMar
07190Esporles(Mallorca)
CampusUniversitariodeTafira
Spain
UniversidaddelasPalmasde
GranCanaria TomFenchel
35017LasPalmasdeGranCanaria MarineBiologicalLaboratory
Spain UniversityofCopenhagen
DK-3000Helsingør
JohnBeardall Denmark
SchoolofBiologicalSciences
MonashUniversity KevinJ.Flynn
Clayton EcologyResearchUnit
VIC3800 UniversityofWalesSwansea
Australia SwanseaSA28PP
UK
LouisA.Codispoti
Jean-PierreGattuso
UniversityofMarylandCenterfor
Laboratoired’OcéanographiedeVillefranche
EnvironmentalScience
CNRSandUniversityofParis6
HornPointLaboratory
B.P.28
P.O.Box775
06234Villefranche-sur-MerCedex
Cambridge
France
MD21613
USA SantiagoHernández-León
BiologicalOceanographyLaboratory
PaulA.delGiorgio FacultaddeCienciasdelMar
Départementdessciencesbiologiques UniversidaddeLasPalmasdeGranCanaria
UniversitéduQuébecàMontréal CampusUniversitariodeTafira
CP8888,Succ.CentreVille 35017LasPalmasdeGC
Montréal,Québec CanaryIslands
CanadaH3C3P8 Spain
Description:Respiration represents the major area of ignorance in our understanding of the global carbon cycle. In spite of its obvious ecological and biogeochemical importance, most oceanographic and limnological textbooks invariably deal with respiration only superficially and as an extension of production an