Seasonal variability in plankton community structure, productivity and food web transfer along the salinity gradient of the Baltic Sea
Abstract
The brackish water environment in the semi‐enclosed Baltic Sea causes changes in plankton community structure in close relation to large‐scale circulation patterns. In addition, seasonal changes in nutrients levels and stoichiometric ratios, light and stratification successively modified it. The analysis of comprehensive data sets on environmental properties, micro and meso‐plankton communities, their metabolic activity and productivity basing on classical and enzymatic methods allows verifying the interplay between community structure and food web transfer from Kattegat to the Gulf of Finland in the ninety nineties. Aspects, like the ratio between new and regenerated production, the percentage of primary productivity utilized by heterotrophs of different size, and the stoichiometry in nutrient regeneration by mesozooplankton were investigated. Regions where new (primary) production was detectable were restricted to areas with river discharge after the spring bloom, but did never exceed some percentages of gross production. Nitrogen and phosphorous were excreted by mesozooplankton in N/P=6 in May and N/P=10 in August in the same region because of the larger percentage of parthenogenetically reproducing cladocerans in summer. Herbivorous plankton
Full text
ICESCM2009/A:11,1246
No obeci edwi hou p io e e ence o heau ho
Seasonal a iabili yinplank oncommuni ys uc u e,
p oduc i i yand oodweb ans e along hesalini y
g adien o heBal icSea
by
Lu zPos el1,Ja ie A ís egui2,San iagoHe nández‐Leon2,MayGómez2,
Ca losAlmeida2,Agus inPo illo‐Hahne eld2,Ma iaF.Mon e o2and
Theodo eT.Packa d2
1LeibnizIns i u e o Bal icSeaResea chWa nemünde,Ge many
2Facul addeCienciasdelMa ,Uni e sidaddeLasPalmasdeGC,Spain
Co espondingau ho :lu z.pos el@io‐wa nemuende.de,
elephone:+493815197206, ele ax:+493815197440
Abs ac
Theb ackishwa e en i onmen in hesemi‐enclosedBal icSeacauseschangesinplank on
communi ys uc u einclose ela ion ola ge‐scaleci cula ionpa e ns.Inaddi ion,
seasonalchangesinnu ien sle elsands oichiome ic a ios,ligh ands a i ica ion
successi elymodi iedi .Theanalysiso comp ehensi eda ase sonen i onmen al
p ope ies,mic oandmeso‐plank oncommuni ies, hei me abolicac i i yandp oduc i i y
basingonclassicalandenzyma icme hodsallows e i ying hein e playbe ween
communi ys uc u eand oodweb ans e omKa ega o heGul o Finlandin he
nine ynine ies.Aspec s,like he a iobe weennewand egene a edp oduc ion, he
pe cen ageo p ima yp oduc i i yu ilizedbyhe e o ophso di e en size,and he
s oichiome yinnu ien egene a ionbymesozooplank onwe ein es iga ed.Regions
whe enew(p ima y)p oduc ionwasde ec ablewe e es ic ed oa easwi h i e
discha gea e hesp ingbloom,bu didne e exceedsomepe cen ageso g oss
p oduc ion.Ni ogenandphospho ouswe eexc e edbymesozooplank oninN/P=6inMay
andN/P=10inAugus in hesame egionbecauseo hela ge pe cen ageo
pa henogene ically ep oducingcladoce ansinsumme .He bi o ousplank on<100µ
u ilizedonemagni udemo ema e o au o ophico igin, hanla ge plank on.Al hough
he ewasama ked a iabili yalong hezonalg adien ,o e allseasonali yexceeded
egionalsalini yd i endi e encesin oodweb ans e andin hecommuni ys uc u e.
In oduc ion
Biochemical,especiallyenzyma icme hods o measu ingp oxieso physiological a eswe e
de elopedin henine eense en ies o example o espi a ion(Elec on anspo sys em
[ETS]ac i i ybyPacka d,1969;OwensandKing,1975),ammoniaexc e ion(Glu ama e
dehyd ogenase[GDH]ac i i ybyBidiga eandKing,1981),andg ow h(e.g.Aspa a
ansca bamylase[ATC]ac i i ybyBe ge onandBues el,1979).Theuseo heseme hods
becamemo eandmo ep ac icable(c. .inHa ise al.,2000).Incon as oclassicalincuba ion
me hods,analyseswe eless imeconsuminganddisloca ed omship olandbased
labo a o ies.The educ iono ini ial olumes o a iousde e mina ionspe mi eddi e se
measu emen s om hesamesou cema e ial.Thisallowedamul idimensionalmappingo
a iousphysiological a es(e.g.Packa d,1985)inpa allelwi h hequasi‐synop ic
hyd og aphicalmeasu emen s.
Resul so enzyma icme hodsneed obecalib a edbyclassicalapp oachesmos lybasingon
incuba ions.Bo h ypeso me hodsha e hei special es ic ionsanddiscussionson hei
accu aciesa eno ullycomple ed.On heo he hand, hepa allelusehasalsopo en ials o
es inghypo hesiso undamen al ele ance, o example hemeaningo allome ic
ela ionshipsinplank ology(Packa dandGómez,2008).
Twen yyea sago,weused heen husiasmdu ing hep ocesso Eu opean euni ica ionand
associa ed unding oexchangeideas, osha eknowledge,and odeepenpa lyexis ing
esea chco‐ope a ions.I esul edinala ge da ase mainlyonplank on espi a ion,ammonia
exc e ion,andg ow hcollec eddu ingc uiseswhen heLeibnizIns i u eo Bal icSeaResea ch,
Wa nemünde,Ge manyandpa ly heMa ineScienceFacul yo heUni e si yo LasPalmas,
G.C.,Spain,we ein ol ed.Theac i i iess a edin1989in hecen alpa o heA lan ic
Ocean(He nández‐Leóne al.,1999),in ensi elycon inuedin heBal icSea(Pos ele al.,1992,
1995)andco e ed egionsinIndianOcean,aNo wegianFjo d,and heAngola‐Benguela‐
F on alRegion(Fig.1).Cu en ly, hema e ialwillbe iewed,da aexpe imen spe o medin
o de ousei o complexsolu ions.Thespace‐ empo al esolu iono da aishighes in he
Bal icSea.On heo he hand,combineds udiesonin e ac ionsincommuni ys uc u eand
ma e ans e in heBal icSeaa e a he a e(Sandbe g,2007).The e o e,wes a edin his
egionwi hou analysis.La e wewill ocusou e o salsoin hedeepseaa easandon
me hodologicalaspec s.
Me hods
Inp inciple,plank ono heBal icSeawascollec eddu ingo dina ymoni o ingc uises
be weenBel SeaandGul o Finlandin1990(Fig.2).Phy oplank on ac ionwascollec edby
bo lesampleswhileWP2ne owso 55and200µmmeshsizeswe eused o zooplank on.
Theme abolicbalanceinoxygenhasbeenmeasu edinmic o‐plank on(<100µm) oms anda d
dep hsdown o20musinganau oma ed e siono hemic oWinkle me hod(A ís eguiand
Mon e o,2005).Calcula ionso newp oduc ionbaseonaconcep ualmodel.I conside s he
idea ha hephysiologicalup akeo ni a eby hephy oplank oniscon olledby heni a e
educ ase eac ionwhichwasmeasu ed.Zooplank onsampleswe esie edin osizeclasses,
spli ed o biomassde e mina ions,speciesanalysesandmeasu ingme abolismbyenzyma ic
me hodsandbyclassicalincuba ionme hodsonselec eds a ions(Pos ele al.,1995;
He nández‐LeónandTo es,1997;He nández‐Leóne al.,1995).Phy oplank onda awe eused
omIOWda abase.Me hodsonsamplingandanalysisa edesc ibedinWasmunde al.(2008).
Sea ching o easonablegeog aphicaldi e encesinphy oplank onandzooplank on
communi ys uc u esin heuppe laye ,weconduc edsimila i yandclus e analysesapplying
B ayCu issimila i yanalysis,squa e oo ans o ma ionandcomple elinkageas heclus e
modebyPRIMERso wa e(c .,Cla keandWa wick,1994).
Resul sandDiscussion
Theac ualsalini ypa e nin heoneme e le el(Figu e3)seems obe he esul o hela ge‐
scalecoun e ‐clockwiseci cula ionin heBal icp ope .Plank oncommuni ies ollowed his
s uc u e.Fou clus e swe e ounda he50%simila i yle elusingzooplank onabundanceand
speciescomposi ionin heuppe laye (Figu e4a).Theyshowananalogousdis ibu ionwhen
using he30%simila i yle el(Figu e4b).The esul ecu ed o phy oplank on.A e hesame
analysis, hes a ionsg oupedin i edi e en ca ego iesa he50%le el(Figu e5a)and
showedasequenceo s a ionslikezooplank onwhenusing he30%simila i yle el(Figu e5b).
In hiss ageo heanalysis,weha en´ looked o hede ailedin luenceo di e en communi y
s uc u esonma e ans e .
Thein es iga ionswe edonea e hephy oplank onsp ingbloom.The e o e,newp ima y
p oduc i i ycalcula ed omni a e educ aseshowedsigni ican highe aluesina easo
ex e nalnu ien suppliesonlylikein heOd a i e plumeandin heGul o Finland(Figu e6).
Ingene al,newp oduc ionwasclea lysmalle incompa ison og ossp oduc ion(seeuni s).
Consequen ly, hepe cen ageo newp oduc iondidno exceed5%o g ossp oduc ion,i.e.
egene a edp oduc iondomina eda hiss ageo seasonalde elopmen al eady.
Con e ing espi a ion oca bonand ela ed op ima yp oduc i i yasdesc ibedinPos ele al.
(1995),up o80%o p ima yp oduc i i ywasu ilizedbyplank on<100µm,andabou 6%by
plank on>100µm(Figu e7).Thispe cen ageexcep ionallyinc easedup o38%insumme
whenzooplank onwasmo eabundan .
Gene ally,insi u(A heniusco ec ed) espi a iono plank onbe ween100and200µmwasin
u nwi hexc e ionexcep a somes a ions(Figu e8).Thiswasno gene ally hecase,which
migh beo specialin e es .Figu e9 e lec s hesi ua iono Od aRi e inau umn.The
si ua ionwascha ac e izedbyasalini y on inabou 10nau icalmilesdis ance o he i e
mou h(Fig.9a).Plan pigmen sdec easedd as icallyon heseasideo heplume.I i edwi h
hespeci icg ow h a ein e mso ATCac i i yuni s(subs a eu iliza ion),while hespeci ic
espi a ion eachedmaximum aluesin he on ala ea(s ess?).Incon as oFigu e8,
exc e ionac i i y(GDH)didno spa iallyco ela ewi h espi a ion.I inc easeddowns eam he
on ala eawhen espi a ionwaslow(Figu e9b).
Thes oichiome yinnu ien egene a ionbymesozooplank onwasno in es iga edinde ail
inou i s a emp .A e ageso NandP egene a edbyzooplank on<200µmshowed
di e encesinMayandinAugus acco ding oPos ele al.(1995).The a iowassmalle inMay
(N/P=6) haninAugus (N/P=10).I couldbeexplainedbydi e en plank oncomposi ionand
po en ialdi e encesinene gy equi emen s o ep oduc ion(Gisme ik,1997).InAugus , he
p opo iono cladoce answasmuchhighe haninMay(Figu e10)whichmaycausela ge P
equi emen s o ene gy ichATPcompoundsdue o hei pa henogene ic ep oduc ionmode.
Byallmeans, hisshouldbecomeclea e whendealingwi h heannual imese ies(Table1).Fo
he u u e,wealsoseepo en ials o answe ing heques ion:Dochangesincommuni ies
change he ans e o ma e ‐ owhichex en ‐and ice e sa?
Re e ences
A ís egui,J.,Mon e o,M.F.2005.Tempo alandspa ialchangesinplank on espi a ionand
biomassin heCana yIslands egion: hee ec o mesoscale a iabili y.J.Ma .Sys ems
54(1‐4):65‐82
Bidiga e,R.R.andKing,F.D.,1981.Themeasu emen o glu ama edehyd ogenaseac i i yin
P aunus lexuosusandi s olein he egula iono ammoniumexc e ion.Comp.Biochem.
Physiol.,70B,409‐413
Cla ke,K.R.,Wa wick,R.M.,1994.ChangesinMa ineCommuni ies.AnApp oach oS a is ical
AnalysisandIn e p e a ion.Plymou hMa ineLabo a o y,Plymou h.144pp.
Gisme ik,I.1997.S oichiome yo somema ineplank onicc us aceans.Jou nalo Plank on
Resea ch19:279–285.
Ha is,R.P.e al.2000.ICESZooplank onMe hodologyManual.AcademicP ess.684pp.
He nández‐León,S.,Almeida,C.Mon e o,I.,1995.Theuseo aspa a e ansca bamylase(ATC)
ac i i y oes ima eg ow h a esinzooplank on.ICESJou nalo Ma ineSciences,52:377‐
383.
He nández‐León,S.,Pos el,L.,A ís egui,J.e al.,1999.La ge‐scaleandmesoscaledis ibu iono
plank onbiomassandme abolicac i i yin heno heas e nCen alA lan ic.J.Oceanog .,
55,471–482.
He nández‐León,S.,To es,S.,1997.The ela ionshipbe weenammoniaexc e ionandGDH
ac i i yinma inezooplank on.Jou nalo Plank onResea ch,19:587‐601.
Owens,T.G.,King,F.,1975.Themeasu emen o espi a o yelec on anspo sys emac i i y
inma inezooplank on.Ma Biol.30:27‐36
Packa d,T.T.,1969.Thees ima iono oxygenu iliza ion a einseawa e om heac i i yo he
espi a o yelec on anspo sys eminplank on.PhD hesis,Uni e si yo Washing on,
Sea le
Packa d,T.T.,1985.Oxygenconsump ionin heoceanmeasu ingandmappingwi henzyme
analysis.In:ZinnoA(ed)Ad ancesinchemis y,Se ies209.Mappings a egiesinchemical
oceanog aphy.Ame icanChemicalSocie y,Washing on,DC,p177‐209
Packa d,T.T.,1985.Oxygenconsump ionin heoceanmeasu ingandmappingwi henzyme
analysis.In:ZinnoA(ed)Ad ancesinchemis y,Se ies209.Mappings a egiesinchemical
oceanog aphy.Ame icanChemicalSocie y,Washing on,DC,p177‐209
Packa d,T.T.,andGómez,M.2008.Explo inga i s ‐p inciples‐basedmodel o zooplank on
espi a ion.ICESJou nalo Ma ineScience,65:371–378.
Pos el,L.,He nández‐León,S.,Gómez,M.,To es,S..Mikka ,U.,Po illoHahne eld,A.,1992.
Zooplank onoxygenconsump ionandnu ien eleasein ela ion ospeciescomposi ion,
animalsizeanden i onmen alcondi ionsin heBal icSeadu ingMayandAugus .In .Coun.
Explo .SeaC.M.1992L:21.1‐16
Pos el,L.,N.Mumm,K ajewska‐Sol ys,A.,1995."Me azooplank ondis ibu ionin he
Pome anianBay(sou he nBal ic)‐speciescomposi ion,biomassand espi a ion."Biule yn
Mo skiegoIns y u uRybackiego/Bulle ino heSea,RybackiegoIns i u e136(3):61‐73.
Sandbe g,J.,2007.C oss‐ecosys emanalyseso pelagic oodwebs uc u eandp ocessesin he
Bal icSea.EcologicalModeling,201(3‐4):243‐261
Wasmund,N.,Pollehne,F.,Pos el,L.,Siegel,H.,Ze le ,M.L.,2008.Biologische
Zus andseinschä zungde Os seeimJah e2007.Mee eswiss.Be .Wa nemünde74:1‐88.
TableandFigu es
Table1:Re iewonzooplank onda acollec ion(P ojec ,when,whe e,pa ame e measu ed,
ela edaspec s)
Figu e1:Loca ionso zooplank ons udiesoncommuni ys uc u e,me abolismandg ow hby
enzyma icandclassicalme hodsbyLeibnizIns i u eo Bal icSeaResea ch,
Wa nemünde,Ge many,andpa lybyMa ineScienceFacul yo heUni e si yo Las
Palmas,G.C.,Spain
Figu e2:Samplingloca ionsin heBal icSeainMay1990
Figu e3:Salini ypa e nin heoneme e le eldu ingMay1990
Figu e4:(a)Simila i yplo basingonabundanceandspeciescomposi iono zooplank on>
100µmin heuppe laye inMay1990and(b)dis ibu iono s a ions ela edon he
30%simila i yle el
Figu e5:(a)Indi idualdominance( ela i eabundance)wi hin i ephy oplank onclus e s
du ingMay1990in heuppe 20mand(b)dis ibu iono s a ion ela edon he30%
simila i yle el
Figu e6:New e sus egene a edp oduc iono plank on<100µmin heuppe 20minMay
1990:G ossp oduc ion,newp oduc ioncalcula ed omni a e educ aseac i i y,and
hepe cen ageo newp oduc iona g ossp oduc ion
Figu e7:Pe cen ageo p ima yp oduc i i yu ilizedbyplank on<100µmandbyplank on>
100µm(PBPome anianBay,ASA konaSea,BSBo nholmSea,S&CGSSou he nand
cen alGo landSea,NBPNo he nBal icp ope ,GoFGul o Finland,WGSWes e n
Go landSea,BS2Bo nholmSea2ndsu ey,AS2A konaSea2ndsu ey)
Figu e8:Compa isono hecou seo insi uspeci icETS‐andGDHac i i yo he ac ion100
o200µmbe ween hewes e nBal icSea(s a ion12)and heGul o Finland(s a ion
305)and e u n oA konaSea(s a ion113)inMay,1990
Figu e9:Cou seo heinsi uspeci icme abolic a esandg ow ho henOd aRi e mou hin
all1993:(a)salini yandplan pigmen s,(b)speci ic a eso ETS,GDH,andATC
Figu e10:Compa isono a e age axomomiccomposi iono zooplank on>200µminMayand
Augus in ela ion o hea e ageN:P a ioin heBal icSea(acco ding oPos ele al.,
1992)
ETS
GDH
ATC
EggP od
Remi
Respi
Abundance
Aspec
BALTEX 05,1990OpenBal ic
Sea,
Pome anian
Bay
xxxxxxx
sp ing,summe ,
(au umn)
Salini yg 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 08,1992xxx xxx
PLANKTON 10,1993 Nea coas al
&openBal 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 icalp o ile
o400(?)m
Annual
coas al
S a ion
1999
2000
Wa ne‐
münde,
RugenIsland
xxx xxx
Seasonalcycle,
weeklysampling
TRUMP 09,1993 Nea coas al
(Pome anian
Bay)
xxx xxx
Salini yg 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
ABFZ97 1997 ABFZ x x x x x x
F on alzone,
Zona ion
ABFZ2000
(M48‐3)
2000 ABFZ x x x x
Indik 1995 Me idional
sec ion
xxx xxx
Openocean
ATLEX89 1989 La i udional
sec ion
xxx xxx
Openocean
Table1:Re iewonzooplank onda acollec ion(P ojec ,when,whe e,pa ame e measu ed, ela ed
aspec s)
STOREFJORD93 BALTEX90‐92
PLANKTON92‐95
TRUMP
ABFZ97,2000
INDIAN
OCEAN95
ATLEX89
Figu e1:
Loca ionso zooplank ons udiesoncommuni ys uc u e,me abolismandg ow hbyenzyma icand
classicalme hodsbyLeibnizIns i u eo Bal icSeaResea ch,Wa nemünde,Ge many,andpa lybyMa ine
ScienceFacul yo heUni e si yo LasPalmas,G.C.,Spain
BALTEX90‐
92
Figu e2:
Samplingloca ionsin heBal icSeainMay1990
<100µm(BALTEX5_90)
>100µm(Pos ele al.1995)
Ina e age:abou 6%
Æsumme ,whenmeso‐zooplank onisabundan –up o38%!
P ima yp oduc i i yu ilizedbyhe e o ophs
25‐
80%
Figu e7:
Pe cen ageo p ima yp oduc i i yu ilizedbyplank on<100µmandbyplank on>100µm
Figu e8:
Compa isono hecou seo insi uspeci icETS‐
andGDHac i i yo he ac ion100 o200µm
be ween hewes e nBal icSea(s a ion12)and heGul o Finland(s a ion305)and e u n oA konaSea
(s a ion113)inMay,1990
lu z.pos el@io‐wa nemuende.de
Pome anianBay,
Bal icSea, all1993
Pos ele al.(1995)
Salini yg adien
Eu ophica iong adien
Figu e9:
Cou seo heinsi uspeci icme abolic a esandg ow ho henOd aRi e mou hin all1993:
(a)salini yandplan pigmen s
lu z.pos el@io‐wa nemuende.de
Pome anianBay,
Bal icSea, all1993
Speci ic espi a ion a e Speci icg ow h a e
Speci icexc e ion a e
Figu e9:
Cou seo heinsi uspeci icme abolic a es
andg ow ho henOd aRi e mou hin all
1993:(b)speci ic a eso ETS,GDH,andATC
Cladoce ans
Pa henogene ic
ep oduc ionmode–
highsho e mene gy equi emen s(ATP)
Copepods
A e ages1988,1990,1991,>200µm
S oichiome y
innu ien ecycling
N:PÆ6(inMay)
N:PÆ10(inAugus )
Figu e10:
Compa isono a e age axomomic
composi iono zooplank on>200µminMayandAugus in ela ion o
hea e ageN:P a ioin heBal icSea(acco ding oPos el
e al.,1992)