ICESCM2009/A:11,1246
No  obeci edwi hou p io  e e ence o heau ho 
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Seasonal a iabili yinplank oncommuni ys uc u e,
p oduc i i yand oodweb ans e along hesalini y
g adien o  heBal icSea
by
Lu zPos el1,Ja ie A ís egui2,San iagoHe nández‐Leon2,MayGómez2,
Ca losAlmeida2,Agus inPo illo‐Hahne eld2,Ma iaF.Mon e o2and
Theodo eT.Packa d2
1LeibnizIns i u e o Bal icSeaResea chWa nemünde,Ge many
2Facul addeCienciasdelMa ,Uni e sidaddeLasPalmasdeGC,Spain
Co espondingau ho :lu z.pos el@io‐wa nemuende.de,
elephone:+493815197206, ele ax:+493815197440
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Abs ac 
Theb ackishwa e en i onmen in hesemi‐enclosedBal icSeacauseschangesinplank on
communi ys uc u einclose ela ion ola ge‐scaleci cula ionpa e ns.Inaddi ion,
seasonalchangesinnu ien sle elsands oichiome ic a ios,ligh ands a i ica ion
successi elymodi iedi .Theanalysiso comp ehensi eda ase sonen i onmen al
p ope ies,mic oandmeso‐plank oncommuni ies, hei me abolicac i i yandp oduc i i y
basingonclassicalandenzyma icme hodsallows e i ying hein e playbe ween
communi ys uc u eand oodweb ans e  omKa ega  o heGul o Finlandin he
nine ynine ies.Aspec s,like he a iobe weennewand egene a edp oduc ion, he
pe cen ageo p ima yp oduc i i yu ilizedbyhe e o ophso di e en size,and he
s oichiome yinnu ien  egene a ionbymesozooplank onwe ein es iga ed.Regions
whe enew(p ima y)p oduc ionwasde ec ablewe e es ic ed oa easwi h i e 
discha gea e  hesp ingbloom,bu didne e exceedsomepe cen ageso g oss
p oduc ion.Ni ogenandphospho ouswe eexc e edbymesozooplank oninN/P=6inMay
andN/P=10inAugus in hesame egionbecauseo  hela ge pe cen ageo 
pa henogene ically ep oducingcladoce ansinsumme .He bi o ousplank on<100µ
u ilizedonemagni udemo ema e o au o ophico igin, hanla ge plank on.Al hough
he ewasama ked a iabili yalong hezonalg adien ,o e allseasonali yexceeded
egionalsalini yd i endi e encesin oodweb ans e andin hecommuni ys uc u e.

In oduc ion
Biochemical,especiallyenzyma icme hods o measu ingp oxieso physiological a eswe e
de elopedin henine eense en ies o example o  espi a ion(Elec on anspo sys em
[ETS]ac i i ybyPacka d,1969;OwensandKing,1975),ammoniaexc e ion(Glu ama e
dehyd ogenase[GDH]ac i i ybyBidiga eandKing,1981),andg ow h(e.g.Aspa a 
ansca bamylase[ATC]ac i i ybyBe ge onandBues el,1979).Theuseo  heseme hods
becamemo eandmo ep ac icable(c. .inHa ise al.,2000).Incon as  oclassicalincuba ion
me hods,analyseswe eless imeconsuminganddisloca ed omship olandbased
labo a o ies.The educ iono ini ial olumes o  a iousde e mina ionspe mi eddi e se
measu emen s om hesamesou cema e ial.Thisallowedamul idimensionalmappingo 
a iousphysiological a es(e.g.Packa d,1985)inpa allelwi h hequasi‐synop ic
hyd og aphicalmeasu emen s.
Resul so enzyma icme hodsneed obecalib a edbyclassicalapp oachesmos lybasingon
incuba ions.Bo h ypeso me hodsha e hei special es ic ionsanddiscussionson hei 
accu aciesa eno  ullycomple ed.On heo he hand, hepa allelusehasalsopo en ials o 
es inghypo hesiso  undamen al ele ance, o example hemeaningo allome ic
ela ionshipsinplank ology(Packa dandGómez,2008).
Twen yyea sago,weused heen husiasmdu ing hep ocesso Eu opean euni ica ionand
associa ed unding oexchangeideas, osha eknowledge,and odeepenpa lyexis ing
esea chco‐ope a ions.I  esul edinala ge da ase mainlyonplank on espi a ion,ammonia
exc e ion,andg ow hcollec eddu ingc uiseswhen heLeibnizIns i u eo Bal icSeaResea ch,
Wa nemünde,Ge manyandpa ly heMa ineScienceFacul yo  heUni e si yo LasPalmas,
G.C.,Spain,we ein ol ed.Theac i i iess a edin1989in hecen alpa o  heA lan ic
Ocean(He nández‐Leóne al.,1999),in ensi elycon inuedin heBal icSea(Pos ele al.,1992,
1995)andco e ed egionsinIndianOcean,aNo wegianFjo d,and heAngola‐Benguela‐
F on alRegion(Fig.1).Cu en ly, hema e ialwillbe iewed,da aexpe imen spe o medin
o de  ousei  o complexsolu ions.Thespace‐ empo al esolu iono da aishighes in he
Bal icSea.On heo he hand,combineds udiesonin e ac ionsincommuni ys uc u eand
ma e  ans e in heBal icSeaa e a he  a e(Sandbe g,2007).The e o e,wes a edin his
egionwi hou analysis.La e wewill ocusou e o salsoin hedeepseaa easandon
me hodologicalaspec s.
Me hods
Inp inciple,plank ono  heBal icSeawascollec eddu ingo dina ymoni o ingc uises
be weenBel SeaandGul o Finlandin1990(Fig.2).Phy oplank on ac ionwascollec edby
bo lesampleswhileWP2ne  owso 55and200µmmeshsizeswe eused o zooplank on.
Theme abolicbalanceinoxygenhasbeenmeasu edinmic o‐plank on(<100µm) oms anda d
dep hsdown o20musinganau oma ed e siono  hemic oWinkle me hod(A ís eguiand
Mon e o,2005).Calcula ionso newp oduc ionbaseonaconcep ualmodel.I conside s he
idea ha  hephysiologicalup akeo ni a eby hephy oplank oniscon olledby heni a e
educ ase eac ionwhichwasmeasu ed.Zooplank onsampleswe esie edin osizeclasses,
spli ed o biomassde e mina ions,speciesanalysesandmeasu ingme abolismbyenzyma ic
me hodsandbyclassicalincuba ionme hodsonselec eds a ions(Pos ele al.,1995;
He nández‐LeónandTo es,1997;He nández‐Leóne al.,1995).Phy oplank onda awe eused
omIOWda abase.Me hodsonsamplingandanalysisa edesc ibedinWasmunde al.(2008).
Sea ching o  easonablegeog aphicaldi e encesinphy oplank onandzooplank on
communi ys uc u esin heuppe laye ,weconduc edsimila i yandclus e analysesapplying
B ayCu issimila i yanalysis,squa e oo  ans o ma ionandcomple elinkageas heclus e 
modebyPRIMERso wa e(c .,Cla keandWa wick,1994).

Resul sandDiscussion
Theac ualsalini ypa e nin heoneme e le el(Figu e3)seems obe he esul o  hela ge‐
scalecoun e ‐clockwiseci cula ionin heBal icp ope .Plank oncommuni ies ollowed his
s uc u e.Fou clus e swe e ounda  he50%simila i yle elusingzooplank onabundanceand
speciescomposi ionin heuppe laye (Figu e4a).Theyshowananalogousdis ibu ionwhen
using he30%simila i yle el(Figu e4b).The esul  ecu ed o phy oplank on.A e  hesame
analysis, hes a ionsg oupedin i edi e en ca ego iesa  he50%le el(Figu e5a)and
showedasequenceo s a ionslikezooplank onwhenusing he30%simila i yle el(Figu e5b).
In hiss ageo  heanalysis,weha en´ looked o  hede ailedin luenceo di e en communi y
s uc u esonma e  ans e .
Thein es iga ionswe edonea e  hephy oplank onsp ingbloom.The e o e,newp ima y
p oduc i i ycalcula ed omni a e educ aseshowedsigni ican highe  aluesina easo 
ex e nalnu ien suppliesonlylikein heOd a i e plumeandin heGul o Finland(Figu e6).
Ingene al,newp oduc ionwasclea lysmalle incompa ison og ossp oduc ion(seeuni s).
Consequen ly, hepe cen ageo newp oduc iondidno exceed5%o g ossp oduc ion,i.e.
egene a edp oduc iondomina eda  hiss ageo seasonalde elopmen al eady.
Con e ing espi a ion oca bonand ela ed op ima yp oduc i i yasdesc ibedinPos ele al.
(1995),up o80%o p ima yp oduc i i ywasu ilizedbyplank on<100µm,andabou 6%by
plank on>100µm(Figu e7).Thispe cen ageexcep ionallyinc easedup o38%insumme 
whenzooplank onwasmo eabundan .
Gene ally,insi u(A heniusco ec ed) espi a iono plank onbe ween100and200µmwasin
u nwi hexc e ionexcep a somes a ions(Figu e8).Thiswasno gene ally hecase,which
migh beo specialin e es .Figu e9 e lec s hesi ua iono Od aRi e inau umn.The
si ua ionwascha ac e izedbyasalini y on inabou 10nau icalmilesdis ance o he i e 
mou h(Fig.9a).Plan pigmen sdec easedd as icallyon heseasideo  heplume.I  i edwi h
hespeci icg ow h a ein e mso ATCac i i yuni s(subs a eu iliza ion),while hespeci ic
espi a ion eachedmaximum aluesin he on ala ea(s ess?).Incon as  oFigu e8,
exc e ionac i i y(GDH)didno spa iallyco ela ewi h espi a ion.I inc easeddowns eam he
on ala eawhen espi a ionwaslow(Figu e9b).
Thes oichiome yinnu ien  egene a ionbymesozooplank onwasno in es iga edinde ail
inou  i s a emp .A e ageso NandP egene a edbyzooplank on<200µmshowed
di e encesinMayandinAugus acco ding oPos ele al.(1995).The a iowassmalle inMay
(N/P=6) haninAugus (N/P=10).I couldbeexplainedbydi e en plank oncomposi ionand
po en ialdi e encesinene gy equi emen s o  ep oduc ion(Gisme ik,1997).InAugus , he
p opo iono cladoce answasmuchhighe  haninMay(Figu e10)whichmaycausela ge P
equi emen s o ene gy ichATPcompoundsdue o hei pa henogene ic ep oduc ionmode.
Byallmeans, hisshouldbecomeclea e whendealingwi h heannual imese ies(Table1).Fo 
he u u e,wealsoseepo en ials o answe ing heques ion:Dochangesincommuni ies
change he ans e o ma e ‐ owhichex en ‐and ice e sa?
Re e ences
A ís egui,J.,Mon e o,M.F.2005.Tempo alandspa ialchangesinplank on espi a ionand
biomassin heCana yIslands egion: hee ec o mesoscale a iabili y.J.Ma .Sys ems
54(1‐4):65‐82
Bidiga e,R.R.andKing,F.D.,1981.Themeasu emen o glu ama edehyd ogenaseac i i yin
P aunus lexuosusandi s olein he egula iono ammoniumexc e ion.Comp.Biochem.
Physiol.,70B,409‐413
Cla ke,K.R.,Wa wick,R.M.,1994.ChangesinMa ineCommuni ies.AnApp oach oS a is ical
AnalysisandIn e p e a ion.Plymou hMa ineLabo a o y,Plymou h.144pp.
Gisme ik,I.1997.S oichiome yo somema ineplank onicc us aceans.Jou nalo Plank on
Resea ch19:279–285.
Ha is,R.P.e al.2000.ICESZooplank onMe hodologyManual.AcademicP ess.684pp.
He nández‐León,S.,Almeida,C.Mon e o,I.,1995.Theuseo aspa a e ansca bamylase(ATC)
ac i i y oes ima eg ow h a esinzooplank on.ICESJou nalo Ma ineSciences,52:377‐
383.
He nández‐León,S.,Pos el,L.,A ís egui,J.e al.,1999.La ge‐scaleandmesoscaledis ibu iono 
plank onbiomassandme abolicac i i yin heno heas e nCen alA lan ic.J.Oceanog .,
55,471–482.
He nández‐León,S.,To es,S.,1997.The ela ionshipbe weenammoniaexc e ionandGDH
ac i i yinma inezooplank on.Jou nalo Plank onResea ch,19:587‐601.
Owens,T.G.,King,F.,1975.Themeasu emen o  espi a o yelec on anspo sys emac i i y
inma inezooplank on.Ma Biol.30:27‐36
Packa d,T.T.,1969.Thees ima iono oxygenu iliza ion a einseawa e  om heac i i yo  he
espi a o yelec on anspo sys eminplank on.PhD hesis,Uni e si yo Washing on,
Sea le
Packa d,T.T.,1985.Oxygenconsump ionin heoceanmeasu ingandmappingwi henzyme
analysis.In:ZinnoA(ed)Ad ancesinchemis y,Se ies209.Mappings a egiesinchemical
oceanog aphy.Ame icanChemicalSocie y,Washing on,DC,p177‐209
Packa d,T.T.,1985.Oxygenconsump ionin heoceanmeasu ingandmappingwi henzyme
analysis.In:ZinnoA(ed)Ad ancesinchemis y,Se ies209.Mappings a egiesinchemical
oceanog aphy.Ame icanChemicalSocie y,Washing on,DC,p177‐209
Packa d,T.T.,andGómez,M.2008.Explo inga i s ‐p inciples‐basedmodel o zooplank on
espi a ion.ICESJou nalo Ma ineScience,65:371–378.
Pos el,L.,He nández‐León,S.,Gómez,M.,To es,S..Mikka ,U.,Po illoHahne eld,A.,1992.
Zooplank onoxygenconsump ionandnu ien  eleasein ela ion ospeciescomposi ion,
animalsizeanden i onmen alcondi ionsin heBal icSeadu ingMayandAugus .In .Coun.
Explo .SeaC.M.1992L:21.1‐16
Pos el,L.,N.Mumm,K ajewska‐Sol ys,A.,1995."Me azooplank ondis ibu ionin he
Pome anianBay(sou he nBal ic)‐speciescomposi ion,biomassand espi a ion."Biule yn
Mo skiegoIns y u uRybackiego/Bulle ino  heSea,RybackiegoIns i u e136(3):61‐73.
Sandbe g,J.,2007.C oss‐ecosys emanalyseso pelagic oodwebs uc u eandp ocessesin he
Bal icSea.EcologicalModeling,201(3‐4):243‐261
Wasmund,N.,Pollehne,F.,Pos el,L.,Siegel,H.,Ze le ,M.L.,2008.Biologische
Zus andseinschä zungde Os seeimJah e2007.Mee eswiss.Be .Wa nemünde74:1‐88.



TableandFigu es

Table1:Re iewonzooplank onda acollec ion(P ojec ,when,whe e,pa ame e measu ed,
ela edaspec s)


Figu e1:Loca ionso zooplank ons udiesoncommuni ys uc u e,me abolismandg ow hby
enzyma icandclassicalme hodsbyLeibnizIns i u eo Bal icSeaResea ch,
Wa nemünde,Ge many,andpa lybyMa ineScienceFacul yo  heUni e si yo Las
Palmas,G.C.,Spain
Figu e2:Samplingloca ionsin heBal icSeainMay1990
Figu e3:Salini ypa e nin heoneme e le eldu ingMay1990
Figu e4:(a)Simila i yplo basingonabundanceandspeciescomposi iono zooplank on>
100µmin heuppe laye inMay1990and(b)dis ibu iono s a ions ela edon he
30%simila i yle el
Figu e5:(a)Indi idualdominance( ela i eabundance)wi hin i ephy oplank onclus e s
du ingMay1990in heuppe 20mand(b)dis ibu iono s a ion ela edon he30%
simila i yle el
Figu e6:New e sus egene a edp oduc iono plank on<100µmin heuppe 20minMay
1990:G ossp oduc ion,newp oduc ioncalcula ed omni a e educ aseac i i y,and
hepe cen ageo newp oduc iona g ossp oduc ion
Figu e7:Pe cen ageo p ima yp oduc i i yu ilizedbyplank on<100µmandbyplank on>
100µm(PBPome anianBay,ASA konaSea,BSBo nholmSea,S&CGSSou he nand
cen alGo landSea,NBPNo he nBal icp ope ,GoFGul o Finland,WGSWes e n
Go landSea,BS2Bo nholmSea2ndsu ey,AS2A konaSea2ndsu ey)
Figu e8:Compa isono  hecou seo insi uspeci icETS‐andGDHac i i yo  he ac ion100
o200µmbe ween hewes e nBal icSea(s a ion12)and heGul o Finland(s a ion
305)and e u n oA konaSea(s a ion113)inMay,1990
Figu e9:Cou seo  heinsi uspeci icme abolic a esandg ow ho  henOd aRi e mou hin
all1993:(a)salini yandplan pigmen s,(b)speci ic a eso ETS,GDH,andATC
Figu e10:Compa isono a e age axomomiccomposi iono zooplank on>200µminMayand
Augus in ela ion o hea e ageN:P a ioin heBal icSea(acco ding oPos ele al.,
1992)




ETS
GDH
ATC
EggP od
Remi
Respi
Abundance
Aspec
BALTEX 05,1990OpenBal ic
Sea,
Pome anian
Bay
xxxxxxx
sp ing,summe ,
(au umn)
Salini yg adien
HELCOM 08,1990 x x x
BALTEX 05,1991 x x x x x x
BALTEX 08,1991 x x x x x x
BALTEX 11,1991 x x x x x x
BALTEX 05,1992 x x x x x x
PLANKTON 08,1992xxx xxx
PLANKTON 10,1993 Nea coas al
&openBal ic
Sea
xxxxxxx
PLANKTON 09,1995 x x x x x x x
PLANKTON 04,1996 x x x x x x x
STORE‐
FJORD
06,1993 No way x x x x x x x
Ve icalp o ile
o400(?)m
Annual
coas al
S a ion
1999
2000
Wa ne‐
münde,
RugenIsland
xxx xxx
Seasonalcycle,
weeklysampling
TRUMP 09,1993 Nea coas al
(Pome anian
Bay)
xxx xxx
Salini yg adien
TRUMP 06,1994 x x x x x x
TRUMP 01,1995 x x x x x x
TRUMP 04,1995 x x x x x x
TRUMP 07,1995 x x x x x x
TRUMP 09,1995 x x x x x x
TRUMP 10,1995 x x x x x x
TRUMP 09,1993 x x x x x x
ABFZ97 1997 ABFZ x x x x x x
F on alzone,
Zona ion
ABFZ2000
(M48‐3)
2000 ABFZ x x x x
Indik 1995 Me idional
sec ion
xxx xxx
Openocean
ATLEX89 1989 La i udional
sec ion
xxx xxx
Openocean

Table1:Re iewonzooplank onda acollec ion(P ojec ,when,whe e,pa ame e measu ed, ela ed
aspec s)

STOREFJORD93 BALTEX90‐92
PLANKTON92‐95
TRUMP
ABFZ97,2000
INDIAN
OCEAN95
ATLEX89
Figu e1:
Loca ionso zooplank ons udiesoncommuni ys uc u e,me abolismandg ow hbyenzyma icand
classicalme hodsbyLeibnizIns i u eo Bal icSeaResea ch,Wa nemünde,Ge many,andpa lybyMa ine
ScienceFacul yo  heUni e si yo LasPalmas,G.C.,Spain
BALTEX90‐
92
Figu e2:
Samplingloca ionsin heBal icSeainMay1990
<100µm(BALTEX5_90)
>100µm(Pos ele al.1995)
Ina e age:abou 6%
Æsumme ,whenmeso‐zooplank onisabundan –up o38%!
P ima yp oduc i i yu ilizedbyhe e o ophs
25‐
80%
Figu e7:
Pe cen ageo p ima yp oduc i i yu ilizedbyplank on<100µmandbyplank on>100µm

Figu e8:
Compa isono  hecou seo insi uspeci icETS‐
andGDHac i i yo  he ac ion100 o200µm
be ween hewes e nBal icSea(s a ion12)and heGul o Finland(s a ion305)and e u n oA konaSea
(s a ion113)inMay,1990
lu z.pos el@io‐wa nemuende.de
Pome anianBay,
Bal icSea, all1993
Pos ele al.(1995)
Salini yg adien
Eu ophica iong adien
Figu e9:
Cou seo  heinsi uspeci icme abolic a esandg ow ho  henOd aRi e mou hin all1993:
(a)salini yandplan pigmen s
lu z.pos el@io‐wa nemuende.de
Pome anianBay,
Bal icSea, all1993
Speci ic espi a ion a e Speci icg ow h a e
Speci icexc e ion a e
Figu e9:
Cou seo  heinsi uspeci icme abolic a es
andg ow ho  henOd aRi e mou hin all
1993:(b)speci ic a eso ETS,GDH,andATC
Cladoce ans
Pa henogene ic
ep oduc ionmode–
highsho  e mene gy equi emen s(ATP)
Copepods
A e ages1988,1990,1991,>200µm
S oichiome y
innu ien  ecycling
N:PÆ6(inMay)
N:PÆ10(inAugus )
Figu e10:
Compa isono a e age axomomic
composi iono zooplank on>200µminMayandAugus in ela ion o
hea e ageN:P a ioin heBal icSea(acco ding oPos el
e al.,1992)