i1.nd67 67 gi112.indd Tel.: +31(0)534874507 University of Twente P.O. Box 6,7500AAEnschede, The Netherlands; Fax: +74959382054 Kosygin Street 19,Moscow, 119991RussianFederation; Tel.: +74959393810, 1 Alexandr D. Shalabodov the rise ofthemassindividualorganismsthe rise biodiversity,of plankton community and of dominatingcomplexes, increasing index occurs, byreplacement which isconfirmed species recolonizationlakes withnorthern stage ofdecrease ofthetoxic pollution,the formed plankton communities. Onthe inecosystem,turnover dominated inthe (r-strategists), providing more rapidbiomass biogenic inflow. Smallforms oforganisms intoxicconnections conditionsand were increased dueto lackofcompetitive speciesabundanceandbiomass Eurybiontic were biodiversity community decreased. of degradation, speciesabundanceand the decrease ofthetoxic stress. Onthestage trendsand degradation after andrecovery background parameters ecosystem quality: and nutrients. There are three stagesof ofpollutionwithtoxicimpact substances The ecosystemstransfo andfauna. their origin, water chemistry past were oligotrophic andsimilarin isgiven. and Imandra, inthe The lakes Ladoga,Onega,the Russianlarge lakes: aquatic ecosystem changesin long-term 2 * Tatyana I.Moiseenko 4 3 Tel. [email protected] +78122307838,e-mail: Academy ofSciences, 50,185030,, A.Nevsky RussianFederation; Federation; Tel.: (3452)461131,Fax: [email protected] (3452)41-00-59,e-mail: ABSTRACT. EUROPEAN LAKES THE CASE OF THE NORTH-EAST LAKE ECOSYSTEMS UNDER POLLUTION: LONG-TERM CHANGES IN THE LARGE Northern Northern Water Problems Institute, Research Karelian Centre oftheRussian ; e-mail: [email protected] Corresponding author;e-mail: Tyumen State University ofRussia;Semakova Street 10,625003 Tyumen, Russian Faculty of Geo-Information Science and Earth Observation (ITC) Observation Faculty andEarth Science ofGeo-Information V.I. ofRAS andAnalyticalChemistry Vernadsky ofGeochemistry Institute

A retrospective analysisof 1* , Andrey N.Sharov rmed underthe 4 2 , Alexey A. Voinov mid-19 negative environmental changesinthe century, dramaticallymanifested itselfin period of industrial revolution ofindustrial period inthe18 environment, whichdates backto the Prolonged anthropogenic pollutionofthe recovery. ecosystem, reference condition,disturbance, one. modification, whichdiffers from anatural formation ofitsmature andmore stable decreaseafter ofthetoxic indicates impact upper trophic level. The ecosystem state in ecosystems are efficientlyutilized atthe of thecommunities. Accumulated nutrients substances. of environmental pollutionbytoxic stress andrevealed thesevere hazard biologic systems to anthropogenic transformations and theresponses of of theanthropogenic environmental have given insightinto theregularities INTRODUCTION KEY WORDS: th century. Numerous investigations long-term pollution,aquatic long-term 3 , 10.0210:05:34 10:05:34 21.03.2012 th

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6 8 1999; Palmer processesof ecosystem [Depledge, recovery can coordinate aimedatacceleration actions of increasing anddecreasing pressure, one communities andecosystems inconditions ofsuccessions trajectory Knowing mechanism.successions andrecovery role inunderstandingofanthropogenic incalculably. plays akey The ecologytheory proceed completely, nonlinearly, andoften such processes asdegradations andrecoveries andthatiswhycontaminating disturbance Water ecosystems, as arule, suffer multi- a newconfiguration? recover toxic after disturbance, ortheyattain question: isitpossiblefor ecosystems to [Cairns, 2005].Many scientistsraisea toxictheir successionsafter disturbance ofecosystems recovery,scenarios including have beenattempting to predictallthe disturbance. However, onlyrecently scientists thecontaminating after ecosystems recovery Science hasbeencompilinginformation on stream into environment, includingwater. towards thedeclineofdangerous pollutants’ years, theretendency hasbeenadistinct 2006; Palmer in thescientificliterature [Cairns, 2005;Harris, toxic pollutants, hasbeenwell documented decreasing anthropogenic inputs, including ofaquaticecosystems duetorecovery processes ofecosystems recovery. The it ispossibleto efficientlyaccelerate the decreasing anthropogenic loadsare known, under theconditionsofincreasing and successions ofcommunitiesandecosystems of regularities ofrecovery. thetrajectories If the anthropogenic successionsandthe role plays inunderstanding akey theory way. unpredictable often The ecological develop inacomplicated, non-linear, and stress. Hence, theirdegradation andrecovery amulti-contaminant ecosystems experience mentioningthat,asarule, aquatic isworth It oftheirhabitatisalsoevident. preservation ofaquaticecosystems andthe the recovery and itsspeciesdiversity, of theimportance oftheEarth’swater for thesurvival population offresh viewofthecardinal importance In et al., et al. , 2007]. During thelatest , 2007].During 2007]. 1940; Moiseenko 1940; Moiseenko Semenovich, authors inthisregion [Krokhin, on more than30-year investigations ofthe of therelevant publishedresults and also This paperisbasedonananalyticalreview toxicafter disturbance. ofecosystemsestimate to theability recover andto according to theecologytheory ofthese changes to explainthetrajectory recovery; –through degradationcharacteristics –to and theirreduction:from background underanthropogenic lakes load northern of successionswater ecosystems of to reveal themainconsistent patterns the ecosystems; of space analysisofdominantcharacteristics reference conditionsonthebaseofatime- of ecosystem elementsandestimate their aretrospectiveto make analysisofconditions Objectives: of fauna. ofwater chemistry, aswellcharacteristics as forperiod; thisreason theyhave common ecosystem formation thepostglacier during Russia, are byonegenesisof characterized three situated lakes, of intheNorth-West (Fig. ofImandra lake the subarctic 1). These Onegaand Ladoga,Lake Lake Russian lakes: pollutionisthelarge multi-contaminating The representative exampleoflong-term water withtoxic agentsandnutrients. basin thatsuffered from thepollutionof and theBol’shayaBay ofOnegaLake Imandra for Volkhov Bay ofLadogaLake, differentecosystem changesduring periods benthos, andfishconditionsthatreflect indicators ofphytoplankton, zooplankton, andkey main parameters ofwater chemistry thisreview,In attention isfocused onthe paper isbasedondiscontinuousinformation. and, ofthelakes therefore,monitoring this there hasbeennocontinuouslong-term Although muchinformation isavailable, 2002].2002; Antopogenic Modification, Yakovlev, 1990;Moiseenko, Kudrjavtzeva, et al. , 1996;Moiseenko, 221.03.2012 10:05:35 1 . 0 3 . 2 0 1 2

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6 9

Fig. 1. Map of Imandra, Onega, and Ladoga lakes and locations of the main industries on their shores 221.03.2012 10:05:35 1 . 0 3 . 2 0 1 2

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7 0 anthropogenic stress [Falk to manage whentrying sites) isimportant found orminimallydisturbed atundisturbed reference conditions(ecological conditions is above thePolar of Knowledge cycle. lake ofRussia;Imandra the European part situated in theNorth-Taiga ecoregion in are ofLadoga,Onega,andImandra Lakes several millionpeoplelivingin258300km the Ladogaarea affects thelife standard of in theworld. The state oftheenvironment in is oneofthe15largest freshwater reservoirs ofEurope,Ladoga, whichisthelargest lake andecosystemwater recovery. quality provide forindustrialization abenchmark to prior background conditionsofthelake an area of813km Peninsula,in theKola Russia. has The lake issituated withintheArcticCircleImandra watershed area. Ladoga oftheLake equaling to aquarter about 56300km 120 m,respectively. watershed covers Its average andmaximumdepthsof30 temperatures, thinlayer ofsoil, slow inputs,influence ofatmospheric low (high oftheNorth The climaticfactors and 13m,respectively. with maximumandaverage depthsof 67 bynarrowmain basinsconnected passages, complex shoreline andconsistsofthree and thevolume of11km km 9800 km Ladoga. Lake after is The area ofthelake inEuropeOnega isthesecondlargest lake maximum depthis230m. intoof theNevaRiver theGulfofFinland. Its and Vuoksa, andthelake’s outletisbyway The mainfeeder are rivers the Volkhov, , of 17700km Finland.eastern Ladogacovers anarea Lake and oftheRussiannorth-west a great part watershed area,of thelake whichincludes CONDITIONS OF THE LAKES ANDCHARACTERISTICS REFERENCE 2 )

with acatchment area of12300km 2 , andthevolume is262km 2 (withitsislands, 18135km 2 (including the lake itself), itself), (includingthelake 2 (with itsislands, 880 3 et al., . hasa The lake 2006]. The 3 , with 2 ). 2 2

Voronikhin (1935);Poretskij The tableiscompiledusingdatafrom: period, are shown in pre-industrial Table 2. the during natural ecosystems oftheselakes quantitative indexes, conditionof describing arethe three shownin lakes Table 1,basic speciesin modification..., 2002].Dominant Ladoga...,2002;Anthropogenic1999; ..., of deepoligotrophic[Lake lakes mainly similarto thecontent andstructure phytoplankton from theinvestigated is lakes cold-water species. Content andstructureof of thethreeareoligotrophic lakes typical in theNorthern Taiga. Water inhabitants to Ladoga,located Lake from arcticImandra andorganicnutrients substancesincrease processes general, byphosphorcontent. In N/Pratio(43–45)limitproductional High its bioavailable phosphates) wastoo low. microelements; phosphorcontent (especially and concentrations ofsuspendedmaterial saltcontents, low hydrocarbonate–calcium were oligothrophicthe lakes typically with of ionsis20–55mg/l).Prior to the1930s, form clearwaterselement cycling) (thesum processes,chemical weathering andslow diatoms, in particular diatoms, inparticular (1940s) themiddleoflastcentury In 2002. Ladoga... 2002;Anthropogenic modification... B.P.Iliyashuk, Onega...1999;Lake (2002),Lake (1982);Sabylina (1999); Yakovlev (1998); (2002); Anthropogenic eutrophication... (1972); Petrovskaya (1966);Nikolayev Vandish and (1987, 1971);Moiseenko Yakovlev (1990), Pravdin (1956);Petrova (1948);Sokolova andSemenovich(1940);Berg and Krokhin in zoobenthos ofthe lakes. Oligochaeta relicta, Pallasiola qudrispinosa spp.) (Chironomidae), bivalves ( larvae 1966; Nikolayev, 1972; Vandysh, 2002].Midge 1956;Petrovskaya, [Sokolova, and Imandra dominated inzooplankton ofLadoga,Onega, Crustacea Cladocera [Petrova, lakes oligotrophic 1987]. northern biomass were low, of whichistypical Ladoga, andOnega. Values ofphytoplankton predominate inphytoplankton ofImandra, andcrustacea(

and Copepoda typically and Copepoda typically Monoporeia affinis, M. Aulacoseira islandica, ) dominated et al. Euglesa (1934); 221.03.2012 10:05:37 1 . 0 3 . 2 0

1 2

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7 1 Table 1. Dominating complexes of community structure of the Russian large lakes: Imandra, Onega, and Ladoga during the key periods of ecosystem modifi cation 221.03.2012 10:05:37 1 . 0 3 . 2 0 1 2

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7 2 Chl “a”,mg/m Si, mg/l Ntot/NO Ptot/PO Number, spec. 10 Number, cell10 Biomass, g/m Phytoplankton Toxic loads(ΣCi/MPCi)** Biomass, g/m Zooplankton H (Shannon'sindex),bit/spec. 1 – reference condition, 2 – intensive pollution and degradation, 3 – decreasing pollution and recovery and pollution 3–decreasing degradation, and pollution 2–intensive condition, 1 –reference Table 2. The main indicators* of water quality and community conditions of Russian large lakes during during lakes large Russian of conditions community and quality water of indicators* main The Table 2. 4 3 , μg/l ,μg/l aibePeriods Variable 3 3 3 6 /l 3 /m 3 the key periods of ecosystem changes: changes: ecosystem of keyperiods the .2010.2 0.5 0.1 1.2 0.42 1.1 3 2 1121 0 034 143 0.9 80 2.8 110 0.6 1.1 107 2.9 271 0.1 1.2 3 1.7 2 0.3 115313 2.1 3 5.5 2 0.5 1 1.7 0.5 2.4 0.8 0.1 0.1 3.4 7.9 0.6 3.6 8.0 0.7 0.1 0.7 0.1 3 6.8 1.0 2 8.4 3.2 1 0.7 0.1 3.6 3 3.8 2 0.3 1 3 2 1 . . 3.4 12.3 0.4 2.5 3.6 0.1 3.2 3.8 0.1 3 2 1 . . 3.6 3.1 3.4 3.8 3.3 3.7 3.1 2.5 3.2 3 2 1 9/9688 890/230 920/240 450/130 648/85 750/120 350/110 395/19 436/102 260/17 3 2 1 622/ 34/9 10/3 178/100 24/5 54/30 8/1 26/2 26/21 6/1 3 2 1 mnr ng Ladoga Onega Imandra Lakes 221.03.2012 10:05:37 1 . 0 3 . 2 0 1 2

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feeder; feeder; Coregonus albula withthepresencelakes oftrout andloach: 1968]. were whitefishThese lakes typical 1940; Gerd, 1949;Sokolov, 1956;Alexandrov, andSemenovich, Naididae families[Krokhin were represented bytheLumbriculidae catchments. thebeginning ofthelast In inthelakelarge enterprises industrial the 1930sresulted inthebuildingof expansioninConsiderable industrial 1966]. [Galkin, lake ( arcticchar in LadogaandOnegalakes; salmon ( benthophage. Amongcarnivorous fishlake AND ECOSYSTEMS DISTURBANCE ANTHROPOGENIC LOADS 7 Salvelinus alpinus Salvelinus 3 Biomass, g/m Macrozoobenthos Sannsidx,btse.13526– 2.6 B.P.Iliyashuk, Ladoga…2002;Anthropogenic Onega...1999;Lake modifi (2002),Lake cation…2002. 3.5 (1972); (1966); Nikolayev Vandish (2002);Anthropogenic eutrophication… (1982);Sabylina(1999); Yakovlev (1998); Berg andPravdin (1956);Petrova (1948);Sokolova and (1987,1971);Moiseenko Yakovlev (1990),Petrovskaya The tableiscompiledusingdatafrom: Voronikhin (1935);PoretskijandSemenovich(1940); etal. (1934);Krokhin for aquaticlife concentrations(MPC) to maximumpermissible inRussia[ListofFisheryStandards..., 1999]. ** Toxic loads:ΣCi/MPCi Cu, –thetotal concentrationofpollutants(Ni, Pb, phenolandlignosulphonate) normalized, valuesofindicators from are* Numerical theliterature taken cited below; 1 H (Shannon'sindex),bit/spec. Number, spec. 10 H (Shannon'sindex),bit/spec. Coregonus lavaretus Salmo trutta Salmo aibePeriods Variable 2 3 /m (L)isthemainplankton (L)) dominates in Imandra (L))dominates inImandra 2 (L)dominates (L) isthemain . . 2.7 – – 2.5 1.7 2.3 2.5 1.9 2.8 3 2 1 . . – – 3.5 1.4 1.1 2.0 0.8 7.0 12.6 1.1 4.8 2.4 1.6 1.6 0.2 12.0 6.2 3 3.2 43 2 0.6 0.5 13 3 35 2 0.6 1 3 2 1

and only to 8%ofthewholelake. be noted, thatthearea ofthisbay amounts should Ladoga...,2002].It total value)[Lake Bay’s was50–60%ofthe part (Volkhov phosphor loadwas6–7thousand ton/year hundred times:inthe1970s–1980s, increasedphosphor fluxinto thelake a town. According to scientists’ estimates ores) andwithwastewaters ofthe Volkhov apatite-nepheline phosphorus-containing raw bythealuminumplant: materials wastewater usingnew (especiallyafter with fluxoftoxic agentswithindustry 1960s. Phosphor loadwas associated reached itsmaximumbytheendof benzopyrene, andother toxic agents Plant pollutionwithphenols, lignosulfate, Bay). (Volkhov on theshore ofLadogaLake andother plantswere builtpaper factory century, thefirstaluminumplant,pulpand mnr ng Ladoga Onega Imandra Lakes Окончание табл. 2 221.03.2012 10:05:38 1 . 0 3 . 2 0 1 2

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7 4 to theR.A. Vollenweider classification, condition byphosphorcontent (according – Imandra White Bay) satisfyeutrophic in Bay; inOnega–Kondopoga Bay; Polluted bays (inLadoga– Volkhov (phosphorandnitrogen).of nutrients went withthebulkinputof thelakes ofthelakes. sections Thus, toxic pollution toxic into wastewater andsewagebled-off went withuncontrollable activity industrial thisperiod,of intense water pollution.In toxic stress for water dwellers intheperiod limited information isindicative ofhigh into theecosystems, buteven available to dimensionsoftoxic estimate exact flow isdifficultof substanceswasdramatic. It Pollution with toxic ofthelakes mixture area).all lake (i.e., Bol’shayathe lake (38%ofthe Imandra) of part pollution occurred inthenorthern nutrients, sulphates, andchlorides. The main metals,of contaminantsincludingheavy to wassubjected pollutionbyanumber lake effects ofpollutionwere mostevident. The Data are available for 1983–1992,whenthe 1940s andreached itspeakinthe1980s. began inthe pressure Lake ontheImandra beganinthe1930s. AnthropogenicImandra deposits inthecatchment area ofLake andiron apatite-nephelinite, nickel-rich, development ofcopper- andIndustrial severe pollutionthanmany Arctic lakes. to hasbeensubjected more Imandra Lake increased to 0.56g/m agents. Phosphor loadto theBay area compounds were in the1980s, however phosphorandnitrogen treatment system wasforced into application treatment for facility 30years. Wastewater The plantoperated withoutwaste-water area islessthan 3%ofthelake. nutrients. Its sewage, whichcontainstoxic pollutantand wastewaterof industrial anddomestic Bay receivesThe Kondopoga large amounts plant. pulp-and-paper largest inthecountry Onegaispolluted bywastewaterLake ofthe nitrogen loadreached 11.1g/m 0.1 g/m However total was phosphorloadto thelake 2 peryear [Sabylina,1999]. additionally appliedas 2 peryear andthe 2 peryear. sp water dwellers as time, percentage ofsuchspecificnorthern ofcontaminants. Atthesame the impact occurs becauseofrotifers’ highresistance to increased tostructure inImandra 60%,which Percentage ofrotifers inthezooplankton species. the dominanceofeurybiontic lakes, zooplankton structurechangedtowards ofintensive theperiod pollutionofthe In (see that period Table 1). green, algaedominated in andcryptophyte ofeutrophic waters:typical blue-green, promoted intensive development ofspecies, changes ofphytoplankton community they occurred to alesserdegree. Structural ofImandra inthearcticlake Onega Lakes; in Ladoga andblooms were observed cyanobacterial input.Intensive nutrients 20–30-fold becauseof large dimensionsof baysbiomass inpolluted increased lake thesummerperiod, phytoplanktonDuring of allecosystem units(see Table 2). occurred andinvolved changesofstructure –water toxicity –has and newproperty different from theirnative characteristics, organisms intheanalyzed bays became pollution habitatconditionsfor aquatic also occurred. Thus, of intense intheperiod havecontaminants withtoxic properties elements content hasincreased; changein level, sulphates, chlorides, andbiogenic hasdecreased; pH water clarity in type: ofpollution werein theperiod similar Water changesinallthree lakes chemistry mesotrophic condition. 1979) andadjacentlarge areas satisfy have whiletheirbiodiversity steeply risen zoobenthos inthepollution zones ofalllakes The total abundance andbiomassof decreased. crustaceanspecies northern of typical and Onegaincreased, whereas abundance decreased. Abundanceofrotifers inLadoga increased, whereas index speciesdiversity total biomassofzooplankton community According 2002], to thedatafrom [Vandish, ., Holopedium gibberum Collotheca sp ., decreased. Conochilus 221.03.2012 10:05:38 1 . 0 3 . 2 0 1 2

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1999; Lake Ladoga...,2002]. 1999; Lake had decreased Onega..., dramatically [Lake salmon abundanceinLadoga andOnega 2002].Lake andKudriavtseva, [Moiseenko in water anditsaccumulationinkidneys was closelyrelated concentration to nickel rate (%ofthosesurveyed) Case frequency lipoid liver, etc. cirrhosis, were recorded also. whitefish stones), asnephrocalsitosis (kidney abundance decreased. Suchdiseasesof commercial catches ofImandra. Whitefish These speciescompletely disappeared in of theirhighwater pollutionsensitivity. significantly decreasedcommunity because Abundance oftrout andloachinfish matter inthebottom silt. toxicity andhighaccumulationoforganic dominated inthezones withdecreased substances andnutrients. Oligochaetes onorganicwith naturalvaluesthriving 50-fold higherbiomasses incomparison chironomids large-size formed 10– Rather formed inconditionsofmoderate pollution. complexwerechironomid-oligochaete Communities ofsomeresistant speciesof 1992;Belkina 1999; Slepukhina, [Polyakova,communities wasobserved and thetotal annihilationofbottom production outlet from thepulp-and-paper offlood-release occurred intheperiod lakes Dramatic pollutionofLadogaandOnega and [Moiseenko ting thelake Yakovlev, 1990]. metalsandbiogenicelementspollu- heavy M. affinis has probably petered outofthefauna. before Lake seen inImandra (i.e., epibioticcrustaceans than 50%.Oneoftwo and bivalve mollusksdecreased bymore of pollution-sensitive Chironomidae larvae Atthesametime, Lake. theratio of Imandra to 200g/m less than1–2bit/spec. Oligochaetes (up index intheseverely polluted zones was in thecontaminationarea. Speciesdiversity tubifex, Limnodrilushoffmeisteri Chironomus, Procladius of biodiversity abundance andrestricted have decreased. Communities withhigh 50g/m 7 5 2 ) abundancedominated inbenthos appeared to bemore resistant to 2 ) andChironomidae (upto , Nematoda developed et al., M. relicta) M. relicta) , Tubifex 2003].

and kept decreasingand kept steadily, dueto more interest; itsconcentration didnotimprove ofsiliconisparticular structure. Dynamics utilization inthechangedtrophic ecosystem fold), and nitrates significantly decreased (6–20- Concentrations ofbioavailable phosphates decreased inall three onlyslightly. lakes level; concentrationofcommonnitrogen theyremained atthesamein Imandra, in LadogaandOnegadecreased; whereas Concentration ofcommonphosphorusforms Table 2). and phenolsinLadogaalsodecreased (see of Onega;concentrationslignosulphates pollution, decreasedBay intheKondopoga industry ofpulp-and-paper as markers concentration oflignosulphates andphenols, decreased fromImandra 150to 10μg/l; ofpollutionin asamajormarker nickel analyzed bays decreased: concentrationof Toxic matter concentrationinwater ofthe of Imandra. (Onega)Bays andreachesand Kondopoga anthropogenic loadto the Volkhov (Ladoga) occurred inresponse to decreasing andecosystemof water recovery quality decreased.lakes Tendency to improvement the1990s, theanthropogenicIn loadonthe ofintensive theperiod pollution. during state Imandra ofecosystem healthinLake fishdiseasesindicated thedramatic criteria, health [Adams 1994].Usingthese andRyon, are usedasintegral parameter ofecosystem forare assessingtoxic important effects and (by physiological indicators ofintoxication) to fishconditions determine high. Criteria lethal outcome for staying thefishafter was fish intheseareas wasdramatic, andthe and Yakovlev, 1990].Diseaseoccurrence of the effects oftoxic contaminants [Moiseenko to thesefood-rich areas, fishwere exposedto ased, migrating whitefish. whichattracted By developed andzoobenthos biomassincre- Productive areas ofbenthiccommunities TENDENCYRECOVERY TO which indicatestheirmore active 221.03.2012 10:05:38 1 . 0 3 . 2 0 1 2

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7 6 to 3.4g/m from varied 1.7 biomass valuesinthelakes remained thesame.in Imandra Average Bay ofOnega; phytoplankton abundance Volkhov Bay ofLadogaandtheKondopoga Phytoplankton abundancedecreased inthe diatoms. ofitbydeveloping absorption active the analyzedbays indexhadincreasedand thebiodiversity in abundancehaddecreasedcommunity of the2000s, theindexofzooplankton From theendof1990sto thebeginning withnaturalvalues. highincomparison very biomass andchlorophyll content remained of recovery, maximal andaverage indexes of the bays ofLadogaandOnegaintheperiod spite ofthedecreaseIn ofphosphorload in structure. algae wasstillhighinthephytoplankton bluegreen,of cryptomonades, andgreen Aulacoseira islandica Stephanodiscus genera Cryptomonas, still ahighabundanceofspeciesthe varied fromvaried 3.6to 7.9mg/m Chironomidae complexwith thedominance inLadogaandOnega.Oligochaete- rising decreased, biomass inImandra butitkept slowto recovery.Zoobenthos wasvery Its 2002]. Ladoga..., 2002,Anthropogenic modification, Onega..,1999,Lake [Lake Bay ofLadogalake dominated inzooplankton inthe Volkhov pen diculata) Eudiaptomus graciloides, Hetero cope ap- ( of themostpollution-sensitive crustaceans However, there wasstill only atraceamount before thepeakofpollution,recovered. sp., L.kindtii) Cladocerae ( pollutionindicators.typical Valuable food and depletionofratiosmallrotifers, ( ( increasing abundanceoflarger Cladocerae decreased, butnotsogreatly, becauseof (Table lakes Imandra Cyclops sp., Leptodora kindtii Bosmina

3 obtusirostris ; chlorophyll “a” concentrations , which used to affect the lake , whichusedto affect thelake . CladoceraandCopepoda Ноlopedium gibberum, Daphnia Ноlopedium gibberum,Daphnia Cyclops , of Polyphemus , Mesocyclops leuckarti , Mesocyclops . Relative abundance 2). Biomasshadalso Ladoga, Onega,and ) andCopepoda 3 . There was . There pediculus, и ) et al., 1999;Falktime [O’Nail, with development andstructure occurring investigations ofprocesses ofcommunity apply theoretical ofecologyto principles to processes, recovery itisnecessary why, inorder to understandecosystems do nothave analoguesinthepast;that is ofanthropogenicunder theimpact load changesofecosystems modern Often, numbers offishcaughtfrom thelake. aproperimpossible to take accountofreal of successionsfishcommunities, sinceitis complicated to factor definethedetermining is fishcommunitiesstructure. It could impact together withpollutionandeutrophication an increase ofillegalcatch volume, which noted, thisperiod, there thatduring was shouldbe data abouttheotherlakes.It [Moiseenko of 2003,fish’s physiological state improved fell. Lake Imandra According to thefindings decrease, theincidenceoffish diseases in increased. response to In thetoxic load andperchwhereas abundance pike didnotgrowCorigonidae infishfauna, Abundance ofvaluableSalmonidaeand 2002]. to 60%[Iliayashuk, abundance grew –from almosttwice 36% stretch ofBol’shaya Imandra). Their relative in profundal(the benthosofthearcticlake M. Affinis supply,good nutrients epibioticcrustaceans significant decrease oftoxic load andof conditionsofa In of theobservations.) average, compared to 1964(thebeginning its biomassincreased over twenty-fold, on of speciesrose more and thanforty-fold Bay improved;the Kondopoga thenumber polluted waters, theconditionsofbenthosin facilities thatpromoted dissipationof of operationthewastewater treatment not found there. a30-year During period butepibioticAmphipodawas Onega Lake, Bay of oftheKondopoga part northern of wormsstilldominated inbenthosofthe DEGRADATION AND RECOVERY SIMILARITIES OF LAKE ECOSYSTEMS 2007].Ecosystems are self-regulating dominated amonginvertebrates et al., 2006]. There are nosuch et al., 2006;Palmer 221.03.2012 10:05:38 1 . 0 3 . 2 0 1 2

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become reserves ofintensification ofthe become reserves at thisstageoftransformation andthey are notableto utilize bioavailable forms grow too;(orthophosphate) theecosystems of total phosphor, bioavailable forms Together withthegrowth of content of multi-pollution. underthe impact oftheirdisturbance period water ecosystems ofthethreein lakes us nowdiscussattributes, characterizing be considered as adevelopment stage. Let stable naturalstate to anewphasethatmay ecosystems have transformed from their pollution, itbecomesclearthatthelakes’ of allthree of intensive bays intheperiod intheecosystems indicators observed key unstable stressed ecosystems [1985]to the succession stagesofdevelopment andof Applying Odum’s oftheearly theory influx andstress oftoxic contamination. –energy factors of two inputnutrients lakes’ ecosystems changedundertheimpact factor, intensifying energy dissipation. The input, whereas atoxic contaminantisastress ecosystems represents anadditionalenergy influx oforganic into matter andnutrients Accordinglakes. 1985],alimited to [Odum, the ecosystems ofthesamezones ofthe release oftoxic elementsandsewageinto on theshoreshadresulted ofthelakes in For more activity thanhalfacentury, industrial North. ofcoldoligotrophictypical water inthe consisted ofpsychrophilic stenoecic species microelements. ichtyofauna Dominant of nutrients, suspendedmatter, and as oligotrophic withlowconcentrations decrease. Allthreewere lakes characterized processes theanthropogenic after stress degradation ofrecovery andsimilarities ofecosystemscommon characteristics ofRussiahasrevealed the European part of the bays intheNorth in thelarge lakes The investigation pollutionof oflong-term stateto 1999]. [O’Nail, thenearequilibrium natural processes thesystem back bring decrease orremoval ofanthropogenic stress, for self-repair 1985].Following [Odum, a systems thathave developed mechanisms 7 7 ( indicates thedominanceofsmallforms phyto- andzooplankton communities, of the conditions ofintegrated pollution.Decrease incomplex inthebenthoscommunity organisms of the chironomids-oligochaete process offormation ofthehighbiomass ispossibleto retracecommunities. the It Small size rotifers dominate inzooplankton dominance inallcommunitiesincreases. speciesto toxic Eurybiontic impact. water,dwellers ofthenorthern vulnerable with the typical competitive connections andlackofconcentrations ofnutrients benthos communitiesgrows owingto high speciesinzooplankton andeurybiontic species diversity. The abundanceof fells, thatresults indecrease ofthetotal in zooplankton andbenthoscommunities vulnerable to toxicants (see Tables 1and2), species, northern The abundanceoftypical their smallsize. inecosystemsbiomass turnover dueto algae,cryptophyte andtheyprovide rapid that mixotrophic isafeature nutrition of isknown, of pollutionresistant algae. It green, algae, andcryptophyte aswell as towards thedominanceofblue-green, of thephytoplankton biomasschanges (phytoplankton) occurs. The structure producersgrowth ofbiomassprimary Due to a highphosphorconcentration, processesproduction andbiomassgrowth. [1985]. in compliancewithOdum’s ecologytheory with amore stable(mature) modification pollu they conformtion andto what extent ecosystems have decrease after oftoxic Let us consider, ofconfiguration whatkind unstable stress state. oftheircorrespond to thecharacteristics in thezones ofintensive pollutionand state oftheecosystems ofthethree lakes featuresThe observed indicate thecritical species inzooplankton andfishdecreases. additionally. Percentage ofpredatory utilization ofenergy subsidies, received intheecosystembiomass turnover and r -strategists), providing more rapid nominal individual mass,for typical 221.03.2012 10:05:39 1 . 0 3 . 2 0 1 2

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7 8 forms inthesezooplankton communities increase inthepredominance ofpredatory 1)an factors: which canbeexplainedbytwo time, the biomassofzooplankton decreased, numbers; introducents appeared. Atthesame innature,are greatly solitary increased in communities changed, e.g. species, which only inisolated cases;dominanceinthe did notrecover occurred ortherecovery species, for typical thenaturalconditions, differed from thenaturalstate: anumberof ofthecommunities The speciesstructure Ladoga...,2002]. [Lake biomass turnover have ahighP/Bratiowithrate of period. developed therecovery during They algaehavecryptophytes progressively Ladoga, 1987].In inSweden [Willen, of lakes the decrease ofphosphorloadinanumber phenomenon alsooccurred inresponse to period. therecovery the N/Pratioduring This the phosphorconcentrationandincrease of algaealongwiththedecreasecryptophyte of was replaced bythedominance of the algae The dominanceoftheblue-green 2000]. Ecosystem1994; Great report, Lake only inthesubsequentyears [Grey of thelake’s oligotrophication appeared beginning ofthe1980s, andthetendency chlorophyll “a” didnotchangebefore the the production of phytoplankton and they have decreased two-fold. However, decreasing inOntario, and, bythe1985, phosphor concentrationshadbeenslowly load. For example, from 1968to 1985, phytoplankton to thedecrease ofphosphor Great Lakes,i.e. thedelayed response of A similarphenomenonoccurred inthe ofintensiveperiod pollution. decrease are the almostashighduring ofpollution theperiod content during and average biomassandthechlorophyll the polluted bays), thevaluesofmaximal the ecosystems ofLadogaandOnega(in spite ofdecreaseIn ofphosphorfluxinto in anewtrophic structure oftheecosystems. decreases becauseoftheirrapidutilization concentration ofbioavailable phosphates of total phosphor inthethree lakes,the At thebackground ofhighconcentrations et al.,

amphipod ofinvasions oftheBaikal under theimpact transformationsconsiderable structural of LadogaandOnegahave alsoundergone and favorable feeding. Water communities in conditionsofthedecrease oftoxic load species hasanadvantageinitsdevelopment indicates formation ofanewstructure. This crustacean However, large growth ofprey species–relict wasstilllow. theirbiodiversity recovery; Benthos communitieswere in lessactive mass oftheorganisms increased respectively. its structurerose also;thenominalindividual forms (K-strategists) andprey organisms in communities grew, andthenumberoflarge ofzooplanktonThe speciesdiversity to reduced pressure onthepopulation. and 2)theincrease inanumberoffishdue upper levels ofthe ecosystem trophic introducents, increase oftherole ofthe species, appeareance ofnewnorthern recolonization with individual ofthelakes the decrease oftoxic pressure, specifically, The features oftheecosystem state after conditions? theycorrespond withmatureextent (climax) andtheirnewstructure? trajectory To what are: Which features ecosystems characterize 1999; Power, 1999]. questions The important stablecondition[Chapman, near equilibrium regulation andreorganization to the turn ecosystem, theprocessesdisturbed ofenergy thatinany isknown succession conditions. It ofareturn to theearly the complexity ofanewlyformed ecosystemstability and these mechanismsisthemaintenance of complicated ecological mechanisms. Oneof only onimprovement ofhabitats, butalsoon ofanecosystem dependsnot that recovery The information presented above indicates matter andenergy. ecosystems’ processes oftransformation of production, itrapidlygetsinvolved inthe 2003]. Dueto highnumbersandrates of 2002;Berezina[Ladoga Lake..., and Panov, CONCLUSION Gmelinoidesfasciatus M. affinis– in Imandra Lake Lake in Imandra (Stebbing) 221.03.2012 10:05:39 1 . 0 3 . 2 0 1 2

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about climate warming, inconclusion,we isanxious Since thescientificcommunity natural state. identified withthenotionofreversion to the the term “recovery ofecosystems” can’t be presented inFig. 2. Therefore inthiscase, oftheecosystem modification is trajectory to thenaturalstate.revert The theoretical ecosystems, theirdisturbance, donot after showed thatLadoga, andOnegalakes) ecosystems (polluted bays ofImandra, water modifications ofthenorthern The analyzed exampleoftheanthropogenic Odum’s [1985]. theory mature (climax)modificationaccording to state through development to amore stable of ecosystem successions:from anatural corresponds to themechanisms recovery from anaturalstate through to disturbance natural one. oftransformation This trajectory stable modification,whichdiffers from the indicate formation ofamature andamore the toxic pressure, discussedinthispaper, of theecosystem state decrease after of the share ofK-strategists –allthesefeatures forms ofbiogenicelements, andincrease of structure, successfulutilizationofmineral 7 9 Fig. 2. of A trajectory an theoretical ecosyste the attributes of the disturbance unstable stage and of the new stable stage, stage, stable new the of and stage unstable disturbance the of attributes the after aafter decrease in toxic impacts m under modification toxic and nutrients impacts: of amphipods in warmer temperaturesof amphipodsinwarmer will changes. For example, higherbioproductivity may benefitfrom advantageousecosystem as whitefish, perch, minnow, andsmelt char andtrout, althoughotherspeciessuch to favoris unlikely fishspeciessuchasarctic pollution-resistant species. Climate warming it willprovide for increasing productivity to increase temperature withrising and elements from thecatchment areas islikely species. eurybiontic The influxofbiogenic towards thepredominant development of conditions communitieswillmove warmer utilizedactively introphic chains, becausein 2005]. Accumulated willbemore nutrients trappingandstorage [Cairns,(5) nutrient (4) rate of suspended–solidtrapping, and processingrate ofdetritus andstorage, increase/decrease,rate of nutrient (3) fixation,(2) (1)rate ofcarbon functions: influences thefollowing ecosystem impossible [Harris areturn to referencewill make conditions temperature asaresult ofglobalwarming isprobable thatincreasingIt water climate warming. ecosystems dueto place inthenorthern will note obviousphenomenathatmay take et al., 2006]. Temperature 221.03.2012 10:05:39 1 . 0 3 . 2 0 1 2

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8 0 of anthropogenic pollutionandimpossible conditions water ecosystems undervarying changing effects ofman’s onnorthern impact Studies reviewed inthispapershowthe conditions ofbiogenicpollution. inpresentin someareas, whichisobserved to change from mesotrophic to eutrophic increase intheirnumbers. islikely The lake and growth ofwhitefish andwillleadto an create more favorable conditionsfor feeding 10. Depledge M.H. (1999) Recovery ofecosystems andtheircomponents following exposure M.H.(1999)Recovery Depledge 10. 9. samplingdesignsChapman M.G.(1999)Improving for restoration measuring inaquatic 8. J.Jr.Cairns (2005)Restoration EcologyandEcotoxicology. D.J. In: Hoffman,B.A. Rattner, 7. Berg L.S.and Pravdin I.F. (1948)Fishes Peninsula. ofKola Lenigrad, News VNIORKH, 26: 6. Berezina N.F. andPanov amphipodGmelinoidesfasciatus B.E. ofBaikal (2003)Invasion 5. N.A.,Polyakova Belkina T.N., Timakova N.M.(2003) T.M.,The state andKalinkina of ecosystem (ed. Imandra Moiseenko 4. Anthropogenic oftheLake Modifications T.I.). (2002) 3. Anthropogenic Eutrophycation (ed. Petrova ofLadogaLake N.A.).(1982)Leningrad, Aleksandrov B.M. ofthebottom (1968)Aboutstudyingstructure In: faunaofOnegaLake. 2. ofhealthassessmentapproaches M.G.A.(1994)Comparison Adam for S.M.,andRyon eval- 1. REFERENCES to pollution.J. 6:199–206. Aquat. Ecosyst.Stress andRecovery habitats. J. 6:235–251. Aquat. Ecosyst.Strees anRecovery 1029. andJ.Jr.G.A. Burton, (ed.), Cairns HandbookofEcotoxicology. Lewis Publishers, N-Y, 1015– 37–45 (inRussian). (Amphipoda, Zoological Crustacea)onOnegaLake. 6:731–734. journal 138: 277–283. LadogaSymposium2002.Publications Institute Lake ofKarelian the Fourth International sediments asaconsequenceofanthropogenicOnega.Proceeding influenceonLake of (inRussian). Moscow Nauka, (inRussian). Nauka. 37–39(inRussian). vodsk, Preliminary Resultsof Works ofComplex ExpeditiononResearch Petroza- ofOnegaLake. Ecosystem Health,3:15–25. uating theeffects ofcontaminant-related stress onfishpopulations. ofAquatic Journal Government ofRussia(№11G34.31.0036). no 10-05-00854)andbyagrant from the Foundation for BasicResearch (Projects bytheRussian wassupported This work new parameters attainstability. becauseaquaticecosystems with nutrients), anthropogenicof heavy stress (toxins and return to aperiod naturalconditionsafter ACKNOWLEDGMENTS 221.03.2012 10:05:39 1 . 0 3  . 2

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8 2 41. Yakovlev V.A. (1998).Response ofzooplankton andzoobenthos communities onwater Willen E.(1987).Phytoplankton andreversed Malaren, eutrophication Central Swe- inLake 40. 39. Voronikhin, andNotozero N.N.(1935)Algaeandtheir grouping Imandra inlakes (Kola 38. Vollenweider R.A.(1979)Advances loading levels indefiningcritical for phosphorous in 37. Vandish O.I. (2002)Zooplankton. Moiseenko In: T.I. (ed.), Anthropogenic of Modifications 36. M.F.Sokolova (1956)Zooplankton Proceedings ofLadogaLake. of VNIORH. 38:53–65 I.I.(1956)ZoobentosSokolov oflittoral zone halfofLadogaLake. ofsouthern Works35. ofthe Т.D.34. Slepukhina (1992)Features ofdevelopment macrozoobentos indifferent zones. lake 33. Onega.EcologicalSabilina A.V. Problems: Lake oflake. water chemistry (1999)Modern 32. andneeds. ofknowledge Power inaquaticecosystem: anoverview M.(1999)Recovery Poretskij V.S., Zhuze A.P.31. andSheshukova V.S. Peninsula (1934)Diatoms ofKola inconnection 30. Poliakova T.N. (1999)Bottom cenosisinconditionsanthropogenic eutrophication. In: 29. Petrovskaya M.V. area ofzooplankton in lakes. ofMurmansk (1966).Characteristic 28. Petrova N.A.(1987) The phytoplankton anditsrecent ofLadogaandOnegalakes Palmer M.A.,Ambrose R.F.27. restoration andcommunity andPoff N.I.(2007)Ecologytheory 715–723. Imandra). (bytheexampleofLake Water lakes change ofsubarctic quality resources 6: den, 1965–1983.Br. Phycol. J. 22:193–208. 107–150 (inRussian). Peninsula). In: Works 2.Sporous ofBotanicalinstitute ANUSSR.Series plants. Мoscow, Idrobion, 33: 53–83. Ital eutrophication. Ins lake Met Moscow, Ecosystem. 162–199(inRussian). Nauka, Imandra the Lake (in Russian). Кarelian branchofASUSSR.Petrozavodsk, 5:76–87.(inRussian). St.Petersburg, ofecosystem condition.Nauka, –criteria Ladoga Lake 218p. (inRussian). Research Centre,Karelian Petrozavodsk 34:58–109.(inRussian). J. Aquatic 6:253–257. Ecosystem Stress andRecovery AN USSR8:96–210.(inRussian). with microscopic Diatomite. structure oftheKola Works oftheGeomorphological Institute 211–227. (inRussian). Filatov Onega:Ecological N.N.(ed.), Problems. Lake Research Centre Karelian Petrozavodsk 3: editor.G.G. Galkin, Fishes 84–107(inRussian). Area. ofMurmansk Murmansk, successional changes. Arch. Hydrobiol. Beih.Ergebn. Limnol. 25:11–18. ecology. Restoration Ecology5:291–300. 221.03.2012 10:05:39 1 . 0 3 . 2 0 1 2

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8 3 sustainability andenvironmentalsustainability policy, modelintegration andinteroperability. Alexey A. Voinov Alexandr D. Shalabodov Andrey N.Sharov Tatyana I.Moiseenko assessment, energy and natural resources, participatory modeling, assessment, energy andnaturalresources, participatory modeling ofaquaticandwatershed ecosystems, integrated environmental assessment,environmental assessment, impact models, systems analysisinecologyandeconomics, strategic particular, In hisresearch making, isinspatialdynamic policy and scienceinapplicationto decisionsupport and sustainability professional are insimulationmodelingofecosystems expertise fromEcosystem Modeling thesameuniversity. The areas of 1982,heobtainedhisPhD inBiophysics in1978.In and University received from State hisMScinAppliedMathematics Moscow (ITC)of attheUniversity Observation Twente since 2009.He Natural Resources, Faculty ScienceandEarth ofGeo-Information advisor ofeightcandidate dissertations. the authorofover 90 scientificpublications. Hewasscientific He is includes biological membranes andmembranetransport. Human Anatomy andPhysiology. The area ofhisresearch interests 1985. Atthepresent time, heisProfessor of oftheDepartment Functioning of Transport ATPases”. Hehasbeenwith TSU since in Erythrocytes ofMammal Skeleton RoleofMembrane “The degree from thesameInstitute. was The themeofhisdissertation ofRAS(Leningrad). 1997,heobtainedhisDSc In Biochemistry degree from ofEvolutional Physiology I.M.SechenovInstitute and the Faculty ofBiology 1985,heobtainedhisPhD TSU in1980.In of Innovations Hegraduated from Tyumen State (TSU). University Antropogenic Impacts (with Antropogenic Impacts T.I. Moiseenko). Aquatic under ofRussianLarge Ecosystem Modification Lakes (2009,withco-authors); recovery The Retrospective Analysisof ecosystem: referenceof arcticlake condition,degradation and in the Water (2008);Long-term ofLarge Lakes Quality modification Phytoplankton in Estimating Long-Term asanIndicator Changes Phytoplankton Peninsula from ofKola thelakes (2004); pollution, aquaticecosystem andbioindicators. publications: Main research interestsprimary are focused onenvironmental from ofLimnology(Saint-Petersburg) theInstitute in2001.His (Russia), Faculty ofBiology, andreceived hisPh. D. inecology than 250publications, including10books. levelsecosystems; critical andloads. ofmore Author andco-author toxicafter diagnostics’ impacts; inwater worrying ofearly criteria modificationinaquaticecosystems andrecovery structural-functional water quality, toxicand ecology; impacts,eutrophication, acidification; St.-Petersburg, 1993).Sphere ofscientificinterests: biogeochemistry 1984); DScinBiology(Specialized inecology, Research, ofLake Institute Biology (Specialized inIchthyology, ofFishery, Institute Leningrad, ofRAS.Ph. Din andAnalyticalChemistry ofGeochemistry Institute andEcology, ofBiochemistry Head oftheDepartment V.I. Vernadsky Corresponding oftheRussianAcademy Member ofSciences(1997), isanAssociated Professor of intheDepartment studied atthePetrozavodsk State University is Professor, andFirst Vice-President on –Professor ofEcology(1996), 221.03.2012 10:05:40 1 . 0 3 . 2 0 1 2

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