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Diet quality and season affect physiology and energetic priorities of captive Steller sea lions during.. Jeanniard Du Dot, Tiphaine 2007

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ima oIG1 we associawi ces ioimesm i seaso  guces i coiowe ei emeasumes oose caci  coma mass ss wasseasoy e wi sea o sayi coma gw ( s ssiwieisumme Summeaea   a mo icu seasoseao  coom mi iosessese ysiogicais ca use oeec mo ee coeaooSesea ooiibl nttAac ^ iie ocoes  ^ iiisoes ^ sogus ^ Ackwemes ^ iCht  Intrtn ^ Sesea o a e iosess esis ^  1Eec ioes ^ Eoc guo ^ Coma gw ^ 5Oecs ^ es ^ Cht 2: Stl ln dtdt  bn dnrn rd r rlfr  l nt ^ 8Summa ^ 15Ico ^ 1Maaa meo ^ 1Eemeasig ^ 1 acquisio ^  19Sscaasis ^ sus ^ oiake ^ Ces i mass ^ Ces i comsio ^ Momeics ^ 3scussio ^ 31 mass a comsioces  e iosico ^ 31Coma gw ^ 3Cosio ^ 39es ^ Cht  A dt  ln fh prtStl lnfr jtthrphld bt lndpd nrn r ^ Summa ^ 51Ico ^ 5Maaa meo ^ 53Eemeasig ^ 53iiint  io ^ 5Qua eec esmas ^ Sscaases ^ sus ^ oeec io ^ Qua eec esmas ^ scussio ^ 7Eec ioes  summe ^ 73Eec ioes  wie ^ 7e oeec sagies ^ Cosio ^ es ^ Cht 4: Ch bld t d hn dn rlt nrn  Stl lndd t pd nrn r ^ Summa ^ 91Ico ^ 9Maaa meo ^ 9Eemeasig ^ 9 acquisio ^ 95Sscaases ^ 95sus ^ 9 mass comsioa mec s ^ 9oomes ^ 99Gcocoicoi  ^  1yios ^ 1Iuke gw c1 ^  1scussio ^ 1Eoc ces  mi iosess ^  1Iicaoomec ces eeeemeagu ^ 11Eoc ces  e coe-e ^  11Cosio ^  11es ^  11Cht  Cn ^ 6Summa ois ^ 1 ses ^  1Se a weaksses ^  13Imicao ^  131es ^  13i  ble 1oma comsiooe suse e summea wieeemes ^ e  Age i gss eysomass oia waiakes e summea wieeemes  ^ e 3Mass a  oma Sesea o  e summea wieeemes ^ 3e 3Gss ey iake masscomsioa ey coe measu e summea wieeemes ^ e 3 coooocoey a maiee ey  ecomaooe ey ci e sicoa e ey iake iase e coe-e isummea wie  e 1 mass coio gss ey iake a somass iake oeig Sesea o  e summea wieeemes  ^ 97e  Sem coeo oee  cao a 13-A oSesea o  e summea wieeemese 3Sem coeo oa ee coiso IG1 a GoSeseao  e summea wieeemes  ^ 1  rgu 1MeaSE ces i mass as a e oe iae-eemea mass measume eig Sesea o  e summea wieeemes gu  MeaSE ss o a a mass as a e oe  mass s e summea wieeemes ^  gu 3MeaSE ces i coiocom e se measumes e summea wieeemes ^ gu 3Scmac a simieio oe gss ey iake iaama ^ 5gu 3MeaSE ces ii 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ocoa maiee ocessesie gss ey iakeom wce ey sugcesua awas mo Iwas cauaccoi  e owi equao = GEI (E + U + E= GEI ges Eiciey (U + E(oiake (+ 5 kg  -1a ey iake (k -1oe ama wecoei (e 3a we use cau e gw s a gss eyiakes (GEIoSesea o oa weeks  e sesicoaco ases (e 3 ey we ao use esma ces iey iakeoccui eeese eaes (cioiase UE a E we cy measu isesmas om Sesea o a oei weuseia ges eicieies we coi  95% i a 93%55ock (seis a U ii was esma  7% oegese ey (g ea 19 wceses aage oe ocies (o 1977g ea 19 E is icu  se eemeayom e aime oe ( (g ea 19 Iese cauowas useas aamaooEseo om seea (1997 eyswas coi  15% oGEI  e se a e coe-eases e sico was assume  1 oGEI Gua15% Gu iSesea o is cyceoguoa isy sas ssia(seis En  prtn  (  ocoey is  oeiooe ey aoca oc o sucas sucgwsge o ey sees ooc o (egyco eIiscauas = G + S + E (3Ces i comsiowe use cau ces i (E ioueeme  mass oeacama(1 kgwas coei a comsiowas measuae e oe se a e o weeks  e e oe coe-e  e summe a uie e oe am ase ie wie ieyis was  usi e uum omeo(see Ceicouc e sea omeaoe y (-1 oig someasumes oe  comsiowe missi o amaiGue summeiaCes i comme ( us assueswe cauom acommoe i (se e sicooa e oe sicoe coe-e oEy coe ooi a  aseosicowe cauaccoi  sa ocmicaesmas (1kg   a39 kKi1975 Wiases i oassues occue aiocos osiowe cauusi 3kcessa kooi sia 17 kko (s You 19 ese es a cse  eoca5es (37 kk 1u eemeay i a gs (aWe1977En  tht  (ME  Maiee ey (MEcomisese ey aoca ysiogicao cessa  e  oe ama (aa  oco sucas e ey ee samaiee ossues a ocmicaocesseseoguooa ac as swME = E + AcE + E (ME caao  cauie   comsioces a i mec (  accoi ME =  (5is a measume oe ey eei oe ama o Icomises samesm (E ac (AcEa eoguo(Eeiesas we as ey sas a( a e ey u sioiomass (was measue o weeks coue wi  comsiomeasumes  esi e sea o ia e mec cmcoai a smaoa wi eugom e ama o i i u - u a s mec me (see wwe measu ociuismey we use o gooaiciuoie cma eouoe ama was coeusi a o cameA eomemeasueioom a a wasysm was ie say mis(wqui iecm e e m om iasi o muccom ouiAeaaqua equiaomeaiwas awom e mec cmaa co (usi a Sae Sysm Ieaoow ki mo5ow 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-e o sugges asoois owas a io isummeAc acse  oy ae e oe mo eaee amaiGuwesaye ce ieoguocaci com e77sicooe seoe agaiswi a ck oause  w eeccoioEnt prrdntEy aocaoa e ussiooe ey cioe ysiogicao oSesea o aeae  wiea e summeAma om  gu (a  s comae ey ci esico asi sey (ssue casm e oiooiscoo e ciwas we wie Uike isumme a sigca  oe ey ciwas ao coma a case imaiee ey (i amec essiosui om a case ieoguo ac a samesm  egu sweis ey-s case imaieeeywcseeme s eaieGuwas o gaaaoGuoe mo e sicoom e oeec cauo iis aa ae ama iGuasea sigca weamou osey  wieaama iGu(e 3eoug gu we si  mass ae same is ee sueom e ey oGu  aosesi oeamass ssues a  oeisees  coma e ey ci(e 3 w ama iGussigca mo massAs oiis sseec a ama iicamass sses asea wequai oeyom e es osea o iGuaGu e eec sagy aeGuseems   emo eicie aaoGu Ige e caysiogicase omosama  ioiois   emio ss a oy as a sso oe assues sees (Casi a199use acaa mo eme se om Guseems maeom aeec i oew e aoe amaeoguocaci was a io  e wie eemeagu osea oi  amicaa ai mecsms aaow em mimi e amou oey 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oeguoe  79Iis ies aGuac e ey sicoioosi ways iwiea summe Sea o iis o ey a omass iakesayea gi se ie summea a mec essioie wie eama ea a gomass oswi a w-ey iake aeeiac isummea aeioy sg  e wie is sugges aequaomass iakea seasoa i c ueyi e mecse osea o  iosessIoussimi ocaseey iake ekee e sagies sea o ea e e ie seasowe oey cicaao aece  oysiogicaaco Is eeas iSesea o iuces aessioisa mec sw a 57 case ioiake (weigiuces a gi se (seis  Coequeiis ima coie quai a e qua oseaas we as e oey ci e iosessa e seaso wcioccu  ue aecysiogicasese cauo case iaocao maiee ey (MEseems su mo om a case iac  aeogu cossamesm is e soaca aus case ME (g ea 19oo cque199 seis 1999 wcwas assessese iousO oe mosima o osamesm s  maiee oeoio oic ssues a co sysisea ga qui moey maiee (co o ae ss mecay ac ssues(e 19 Wee a i(199 sweau % oe s mecces ima was e  oio Ioueemee ama ock su % oeimass ss as amass ae e oe sicoCoequea ss o1kg omecay ac ssue cou a a case ieo ey use maiaie ieg ossuesa  a gaee GuCes isamesm meioa a scuWe measumesm 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se (weeka aweek a oe sico( a oe coe-e (Ce assks iicasigca wiiguees com sc  measumed hnIsumme  ces i o3  e sicoa ecoe-e we simi ea age gu (a  p >   e agu Wa ama we o  o3 we sigca wease  om e e oe sico(-35%uie e oe eeme (a  p Iwiewe e(agesigca aecces i3  e sico(gu 3  oGucase e sicocom e sc se e (-5 ae e oe sico aiasesg  e coe-eaugey maiwease  (-35% ae e oe coe-ea  p <   3  oGuwe gey se com e se (a  p >   ee am iase (7ie mi oe sico (p >   w3 we gGuaGu (p  iee age ieeces ia   esicoa coe-e  a seaso(a  p >  e Wa11 ^1-1 -1 1- - - -  ^  ^  ^ C ^ CEemeaasesgu 3MeaSE ces io o (3/  ( ieig ma Sesea o isumme (esquasa wie (osquas Measumes wekeae e oe se (weeka aweek a oe sico( a oe coe-e (C C Assks iica e wiigueescom e sc  measumese ama we o  oa we aec ey sicoo e sueque co ie summe(a  p >  a syese ae sees ( 1   a  5 5 Iwie  maiagaico (ama 7 11 coeo casesigca  e sico(ama % p   a a  p   a i  se   e coe-e (a  p > Ces i3/os we aec eoage ieieseaso(a  p5; gu 3 Wa ama we ogee e 3/o casesigca  e sicoisumme( 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mi  a  p < Iwie  s a sus caseei oIG1 ocom e se (a  p <  we ces iIG1 om seesoo(mi osico  p  e s ae sus(e oe sico  p <  gu 1 Is oao es swea wecoeooIG1 asus oy ie mi oe sico(s11  m1 -1sus111175  m1 -1 = ; oeasesp > 13G a oy aie om e wieeeme (e ieeage imc oGos (a  p >  1 G ce e sicoa e coe-e wa ama we ogee(a  p >IG1 was sigca   mass i summea wie(a  p 0  Ces i oIG1 we ao sigca  ces ia mass i seaso  (p < 1was   coio(ap >   Ces ica we sigca ga  ces iIG1   isumme (p <   a iwie (p <  1; gu y1IG1 was sigca ga  ee coisocoeoi seaso(  p <   a si  3 a isumme(  p <  a iwie (p <  5; gu 5  oGwe  a oeameio coio oome aes (a  p > 5 oyi somaoca sess os imamma a s wi ey iake (qua a quai oose a age as we as eeame sucas seasoa m (Eas 19Kiysky ea meOki AioeKmeMye ea  ese osgu a oimesm  oimi ess oe ama  suimacoio (Wa Aige 19 Ces imec ways may i ec  ocac mes ( a (3-A  aca oi Ousus iica aces igcocoicoi asomaoc os we simi ose oeeiesiamamma ey sico a a se oos we ioeiguoomass ce iwie Isummewe oy IG1 aea  ioeigu  massguooamass ces was so i IG1 i seaso  oe os omes measu ces icoiooe sea oyios we ao cy ioeioeeces i mass omesmEndrdld nrn rCes igcocoicoi ese a maeoc se  sessoi iosess imamma a s (Oiea 197ea ea199 ocoisousuay iase  o oey sico(Eeauso19Oiea aOiea  Oiea aGuiea  mo ss ey quimese o somaocosGa IG1a e maiuce ogw iesiamamma (ea a a mosy ioeioimesm  iasi oisysisa ci oigo(eie1999e i e ea1 G is cay ugu iosess imamma ima (is oami eaiequey oaksMosie19Yamyam ea 199 Oiea w IG1  case  eaecoeaooey uga case iaa  ssue sysis a  ce uooaie ies as e s meossis eassueoco(ae 19Smi ea 1995eie1999e ea A ea Oiea Ces igcocoicoi a IG1 iousea o we as eec(iaiase icoisoa ee coiso a a case oIG1  iosess G sw aiaseis cou  e  e a eosecoois o om e i g maski ces iscmeasumesIwie  oIG1coiso a ee coisowe a ces i mass  e sico iica a coo guoomeossis iis seaso(GooaK19Ase-Ecksoea 19guoo si ( aa massis ke   ima iwieweec a eogu quimes a g(Wiea  Isummewe gcocoicoi we sigca ioeiguoo masscesw IG1 wasis iicas ae guooamass  IG1 is aio isumme(see ao Ce a 3a ae ge oioeme ogcocoicoi ie o sess se cases we io  maiaimass eoguois wesucas isumme ces i mass we age- oseasoe  o gcocoicoi a somaoc os we e eeage gu asomemes seaso cases iIG1 we ga sea o comsus isumme e somaoc as is ke mo ac a si ces iey e iyoueama sie eigw  is g Gaeoc ses seem   cessa s  esee amass  e sameas sus  o oey sicoisumme sie ces imass a  comsiowe age ae ca oIG1 weaays gisummeawieao iicas asucgw may occueiay isumme oimaiocoioA gacase iIG11 ao iicas ais sysm is mo si  ey sicoisummeawiee same sui ces i mass i seasoCes iee coisowe simiees a sus isummeao  oee coisowe gs aa me (see ao i199 Gai 1997 is iicas ae iosicowas mosesss isummeayoueama qui gao osess os  o e same mec auses as oeama iisseaso Iwie  sea o essea mo iee sess se comoeama  e sicoauge e ses we associawisimi cesCoequeyoueama may qui gaces icoisoos aoeos iwieas  ose  sessAugaiase igcocoicoi is  oe see  g osess os ca imea icuy ea cog cacies oyou amaKiysky ea (sweaccks oma s eose gcoicos  om iosess ieay sges oai a cog cacies a  wegi eicieiesIwi Sesea o eeee mi sicooa coc sis aaeay agesui ig osess os egw cacigieicieya immu sysm cou  imi(SAueuKiysky ea eKme Coequesuiow cos cou caseicuy as iosess is mo ke  you ama w qui a gmass-scic ey iake aoeos (Wiea yios gu gw a e imamma (Cksi eaa ay aima  isamec  a mec eogesis(usewo 1991Sia 1995u ea 199Ey sicousuaysus ia case iyiacwcsees  case samesm  saey (A ea 1979Eas 19Key eea  Coequeyio  we eec case  e ey sicosamesm was cy measuiousso eec ocesiyio  we cy assessaewe ces ii1mec s ( a sa mec s (SM Awe measu(SM A oyiwie see Ce3 mo i Uike ioei a woocks(You 19Ase-Ecksoea 19i 199 oe yioswe coe e mec s measu iica aces iyios we maeug imcoumec  measumesis eswemeaaey eei a samec  case aaWama we ogee iooo io  (as oeeom3/was wguisumme ec a case ie acac oe yios o3 ao caseisummewaama we o(eouge case was weak ese o ces may ewee geey eei  o oey sico em iase i3/ iwie(esciay Gu is mosusiesciay sie e ama we sw case eiey aocamaiee o iis seaso(see Ce3 a is oy e  e miase i3 om Gu e sicoCes iyios we o ss oious a oueaaicia   age iousea o (e ea 1977Eeauso19Ase-Ecksoea 19 1991SAuea 199 uea 199wcsugges aey may  emi ie guooeyeei  mi ey sicowe we measu ee  (3e mosmecay ac o a e  (3wcays aima ie guooyio ac  iosess (Oaea 19Key  Ces iei cou sw a mo oue oyios iSesea o ey guoaousus iicaIndn  t b pt pQua oea sigca imcoe  ossue see usee ey sicoiSesea oGucois smo  (aesi  isumme a ss oiaGu(w sesi oiiwie as measu e uum omeo Iis us eecaeoeeees eemacac us (ooioo wou 11ecie oome igca Gu agciu  a A  Gu as we as e  ogcocoicoi a somaoc oss mass ss a  comsioces oe mesmeasuece ee iysiogicase eee egu o e eig ama oiicaacasm occue e sicoa case e coe-ewe iice ee iiei ocasm eeGua   o (3-A we ao useic mec ceswe we oemeasumes oao ciu  oose mes wcieg s osysisuo ssues (ia a cae om e Iese ocesses simuaous wi eacoe o  casy uay se sces iysiogicaoIwie e ca owcec a ce ioimesmiasesigca mo GuaGu e  a case icaeaaiase i Ca  a iee oousis aooiie ea oe musc mass ie ama (yaA19 A case ica  Gumay ece g ooiss (ee a 1% oe  mass ssom muscs iisseaso(see CeIseemswe ais mewas ae  coyicuy maces ioicasm a may  useimo sucasesIeous sessigca ces ioomes we u accu ece  ooia mass ces  a come swe ysiogicaces a asc (ow 19a ea 199 Aou eaMeisIo e ee ie ge oiosicoise moske eoe ck ose ioomes ioueemecom oeses a mo sicosui ia mass ss o115% o mass (wcmay  mo asc oSesea oiosiaoi111e wi ese oome a sei iica omec a iossSomaoc os acas a eseaoage o oi ooey sico(eie1999 Coequeiases ie acoosomaocos we eec  gaiama iGuw sa weamou oamass com Gu e waswe  ee eeces i oGoIG1 Gua Gu Wa e ama we ogee(a us e sscaweiaseIG1 was sigca coeces iamass  iwiea summe is iicas aIG1 gueamou oamass s(ogai  e e ioases iSeseaoe caIG1  we ao sigca  ca ocois aa case ois o sus ia goigoSess os a usuay coe ces i coio(e oie ima mammawi mecay ac coiso(ee coisomo ey ss ugsis oacys  iosess (Oiea a Coequecoisoa ee coisowe eec iase sg moiGu(wcsmo aGu(see Cewe sussweasess o  we  ces i coioiouamauike eous is o ea seac Aac seaa Sesea o (Oiea  Oiea aGuiea Kumagai  Oiea ( auea ocoisoese a ccasig io e amaoces i coio is may  e case  acuiosico (esciay s imieos sucas ioueemeGcocoicoi we us mo aec ey iake  acomsiooe ooces isea o comsio(Kiysky eaKumagai Augee coisowas  aoi coio iwas   i iwie sugges acoisowas somewioeie mesm iouama iis seaso(Oiea e so o iwieeeee coisoa a eeee coisoa mass ce i11wieiica a si a mesm we uea gecois seasoweogu ois mo esseia(a ea 1977seu1997yios a ao kw  ioeimesm iosess ima mamma (usewo 1991u ea 199a  gu mec eogesis  ac eogec mecsms sucase /K-Aase oCa   cyc imuscs (Sia 1995 e oy eeeguee iyio  is a eiace i3 iwiewguooe  m is icuy ima ma mammaAcase i3 ama iGumay imy a gacase iooosees ao a case imec eogesisis s oe iuyeouemay aow casi mec eogesis (a us eyeeiwioucomomisi oeogu caciIcos yio  ama i (Gu may  iaseosyeco coee e mec eogesis cacicessa sie e ss o wasgaaGuiay sui iweueiuo(Ceaoyassessi aa ioss oma mamma aoa e see o  sui ieieies quis a maeae c ces i coio(i oiooamass ie wc oe measuos was ae  IG1 seems a ae maeoces ioimesm camaeoguoomesmemeeom ouguoocasm cou acay come om ecooooseosysms coec causi e oeeces eougiiaeec we sigcaia gea o oeos ke ioeicasmguo ey a o aagosc os   ae a ioss ugguoo (Weig ea 1997Gea  iis gaa geis si coe mass a  a kw  ioosico a osicoa io saao(Cummis Oui Iouse eemeae eicomsioas we as iomass113osi e same cac iakeGu(wcwas wequa sceiice as mucomass ooas Gu(see Cee iegooese e sig osaaoa mass ss may  sueigigeos Gua a gasigoooaGu esicois assumes aia geay e same  isaaoa mesm iSesea o as ey  imaa s wcmay  e case(a gy emiea Amou ea Oiea Iacaseassessi ces i coioiSesea o may qui iegseaioos  a oimesmEndrdthntrl rfde eoc ces aoccu e ey sicoo seueos ae gii oe -e aseese ae ke moe caci ooasms  coom iosess (Mosie19ema Wi1997mick ea  Iouse sea o sayea coma gwssicoiwieisumme(see Cee ocoocoma gw is we cumeeimesc ama a ma (Mosie19ema Wi1997mick ea a ioes coo ces mosy omsomaoc osao om gcocoicoi a yios (ema Wi1997Coma gw usuay ioes aiase ie moGceoesciay ie u-e (i a cessaooe Gsise see sico Sa i1999mick ea  is iuces aima iase iocooIG1 (esciay iG we ee sicoSimuaous g oGa IG1  e -e may se uooies ey mesm a iase o asmsucgwoey sge (Mosie19ae 19ema Wi1997Ious oIG1 maiw  e summe-e owe mass oe ama syese aa w  Oe oeiwie oIG1 iaseck was se es u-ewa11coma gw was oee is coeoeeces iIG1  ae occuee ocoma gw coos e esis asomaocos ay a ma ie caci  coaeiosess iSeseao (ick ea  e wiecoe-e o IG1 a somaoc os iasea  osess os case e coaoeeeiase iey iake a e sig om e eoc sysms  iasessue sysis okea s ogw ie sea o iwie ie same wayscesiamamma (ick ea  e caio sea o e eiss oassue mass  is seasowas  weeie ugces iese osWe mecsmsiseems eis a io Sesea o  gaieioimamass iwiecomsumme iay oc oeogu aso (Cea Ki1975ea 199Iis uaw IG1  iaseiwieisumme o a kw  seasoy (isseea 199 me Mye ea a ao coeo oIG1 we gisummeaiwie aeemeaasesis cou iica asucgw is seasoy aa gaisumme Iis casee seasoee icoma gw cousu om a io  sume sucgw ( e ime o eycoeisummea a io ey accumuoiwie(see Ce a3 gao coeo isummeus wiee same  cescou ao meaago  a quiisummee same ge ose  -ea ais seasoiic sise  somaoc oscou aas eiw e sea o we ue  s ei mass e summewe e somaoc as ie is aec seoemecosi gcocoicoi a yiosg ogcocoicoi eassociawi a sise  Gmec eec a smuooIG1seco(e i  eigeeec oe somaoc as115maiua(e ea  Isumme  oee coisomaiga  oyios maiw  e coe-e w eye eise  iwie e ga ooeeiouseeee coisoa IG1 sugges ae w gw se  e iase iies isummecou come om e oeeii eecogcocoicoi oIG1 ocoa ey sge ugiusise (Saky ea eKme yios ao smu e ocooGaei (ema Wi1997oea  so eiw  couco  e w sysis oIG1 a sueque ck omass gaiu-eIsumme e mass ss ia e sicoceaseeougeo  syese  e coe-e ae same  as aee oe sico is mea auee  oey iake (sicoa -ee same  oos iggee  ocacses a  mass ces e sico  oos sigeamass ssw e same os was  a cessaoomass ss  ecoe-ee w aie ies  e coe-eowe suicie  ecioces    e-eemeamass may  esie a ee issue sei  e osigge ae  mass ss (Wie Kiysky CnA e os scussea a ica  o ae ca acomeweagus ioa eec ss (oea  o o oy a c soe eec guo os ciia  comemae(Eas 19 eiaco a ao suec ces isei ae oa eieoa seasosis as  oe  cyc(me oas a secu coicase ococ eo  (SAueu Coequeiis eme icu  assess e migu sysms a e eoc ces  e ysiogicasesoee e sus oous sw a ee  oIG1 ioeia11mass guo e measuos a mes cao coiaeomae  assess e  coioa ioss oSesea o  ami iosess sucas iousese eec ioes eieecegu se  aeseoos (e ia geowe e  acomeiecooc yo e sime ao coeo oiiaos ie oo117fAl Mcie AWooo (Coma gw issa se gw essio sa ses 1-19Aou Gew  (s mesm iAac sea(Arh zl  Com ocmisy a ysiogy A1-1Aou YMois M w yI(aoiasma iise  s iAac sea  (Arh zl.  uaoCom ysiogy ocmicaSysmic a Eieaysiogy17-3Ase-EcksoSy E Wak (19Mec a ocoes omoi a geooge iAskaoa sesea  (v a  l.  Caauaoogy 1A  Maiwa  o (1979Eecowieseoiyias iAmecaack a Amecauaoysiogy 77-Eea Me M IasAoeMOis  (199 s as amec sess gm sec amies coisosec mass ays e me omamacmecoisocoeo ia meuaoCcaEocgy a Mesm 99ema  Wi (1997CaugwEoc ews 11i (199 Mec a oces  e mo ocai gy sea(lhr   Amecauaoysiogy 3911-yI(Eecscoequees essI eeA(e Mamammaogyaeo aac ackwe Sciees isOo ; Mae MA7-7eie (1999guoooia ey mesm  e somaoc asmesc AmaEocgy 17-11omo i CA (1991Iee oioiooiuke gwcIsomao a mec os iswiuaoAmaSciee 55-7eAA i (Coogw  e somaoc asGw oa e iuke gw c  a iee sAuaew oysiogy -1Cksi ke G Si CSy M ( oyios imaa  amaoc a c seaeaa oc cogy 79-1Cummis Oui(GasoiesguoooiakeuaoCcaIesgao1173-eMMicuCusee Iwi Issse  oue (Eoooome  weig ss ieemeaose ag gsuaoAmaysiogy a Amaio-171Eas  (19e Iee oios oyioiouseasAmecaogis51-3Eeauso (19Ai o ces i sea phnld  a gy sea lhr ,   e saoGea Com Eocgy 35GaiK  A(1997ea auaaoiasma coisocoeoamo wi a cai osea  (v.  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Ieg issisussismua eacoEoc ews 5-Sia  (1995yio cooeogesis a ey eyi5-Smi W UeooCmmo  (1995Eec ocac ooisicooiuke gw cI (IGIa IGi oi icaasuaoCcaEocgy a Mesm -9SAueu (Sess a ma mammaI euGua(eC ok oma mammameciC essca o w YoWasg3-9SAuway S Wes i (199 yia aeosCiu  iwi a semimesca rtrt,a iee ose agea seaso Ma MammaSciee 11-13S WA (19iosess a gw - seee iai seAmecauaoysicaAogy 3117-isseKege Ueoo (199 ioguooe iuke gw cEoc ews 151is AWey C(e c oSesea o  Ej  iAskaa ew oe iosess esisMammaew 33-ume S o Casi MA (ges coais oaaquaccaieec oe equey a ey comsioooseauaoCom ysiogy 173 -5ae  (19Caugwssie mecsmsAc EocgicaSumes 93-Wa Aige K(19Ciauayms oocoumio  massamesm iyeow iemaoCom ocmisy a ysiogyA-ysiogy -Weig e CooWESiSous M Kuie (1997Eecosea e sicooasma iuaoCcaEocgy a Mesm -51Wie Kiysky AS (Eoc ses  uce eieaesSess oai-sess os? Ieg a Com ogy-9WiA is AWseS (A oeec moesma e oquimes oSesea o  Ej  iAskaUSAMaEcogy-ogss Sees 991-31Yamyam ESice MA G(199 ossmec cesa s mec  iei ueoi coma gwuaoAmaSciee 757-You  (19Ieeo ee weigocoumioa asmayio coeocycs ie woock Mtn.Com ocmisy a ysiogy A-ysiogy 7733-531Cht  Cne goaoous was  i w seasoa qua oeimceysiogy a eec ioes ocai ma Sesea o  iosess a sueque o oco  sowe measuces oo 3-moeemes (summea wie i mass a  comsioas we as iey aoca e ysiogicao i maiee o(mesmeoguocacia ac  Eec oseasoa quaoeoe eoc sysm we ao iesga ue w ese meiecwi os   e oeeysiogicacisioe a oo a me coeo  iica  comsioces was aoiSr fdOous coieaseasoa qua oeiec ieee ysiogicases oSesea o aeeee iosess oey iake coee s omass sse qua oeieoiocasm o a oissues as we as e  ey sea oaoca ysiogicao sucas eoguoa acIouse ock e e sea o usi mo assues as ey soues com e g ouse osea o aa e isocac i eis eecwas seei seasoIse  ey sico cai sea o ea i ao ausio oeiey eiaccoae wi eieacoio ey woueeee ie wiw ama coumi ock  Mo scicayeama ea i iaseeiac  a caseeieoguocaci isummewwams a e keooi sie wia eec  e gs ey ao swea susicase iac w maiai a se eoguocaci iwiewgaiuowas ea we siooey ieieie mayweeigi eicieye esmaeoguocaci a ac1 osea o iGumai co ugu eemesee iwieweiac  case e o weeks oeysico e caama ea w-ey s a wecaci  auseiey e ces i oey iake may see ooeisise  mi iosessIousiwas ua wee ysiogicaces we gugcocoicoisomaoca yios oee coisoa IG1we   mass oe sea o iwie oy IG1 was sigcaioeimass guoisumme iica aguoo mass ugamass may  a io iis seaso OIG1 guces iamassoeeie sea o  oe os measuwe a gooiicaoces i coio(oioo us amass ie oeamayio sus we associawi a oeeces imesma oomes (eseey acike es a ooua ogeca oao accu ec oooicasm oe sea oCoequeeougoimesm caassesseom  oIG1a oe measuoome i accuecysiogicass oSesea o awe mi ioy sesseSesea o sayeseasoy e ses  iosessIaeaeasieem  sisiosess a  coom iiwieaisumme wcwas co  eco om es oeec mo(Wiea  Scicaye sea o smass ae same  iwieas isumme e sicoeouge ey iake ci(com sewas ga Sea o auseiey e so ae  gaiosess ey iueiwiea we imccom summeis aea   eaceugouasci a ces iacSea o i eemeagu ao sayecoma gw iwieweiac  maiw aee sicowe ue  so isumme1Sesea o caaeeiou i oe seasoa eieee eec oe eieacoioese auses we mosie  e o oey sicoa we susi i e quaoeiesea a case iey iakeOusus sweaacase isiake was eiaio e case iey iake  iggeoua ysiogicaauses eoug mass sses we simiCoeque osaaoa ke   ima me ysiogicaa ouauses  ces ieIis ies   aauses ma  e sea o  e sicoa coe-e o we ao sie  e e-sicoases isummea wie Ey iakes  se ases we i eouiecoeeama a ai (i wee ama sweaiesio a sueigase iakes iwieaisummeascommoy oeeiiwe s omass gaiwe simi ese i seasois imies ae aioey ies e wiese was aoca o oeagwsucas eoguo(eougueiuois oug   ae e seaso oac ysiogicaa ouasci may us ee seasoees ieieacoio a ey quimes oui oo oiosesssus om Ce  3a sweasummeis a mo sei seasoioy sesseSesea o ais wie is i  co eecoOu o ee asea o caoy wis iosess eiwie ey caao coom suimacoio mo easi wieas weIaio ous coieaa w-ey eces ecaci  ouy a ysiogicay aus eieacoioiay e  e iic coc sig eeey iake a saaosui om w ey-i eycom e oeeces icomsioa oguoiicas agw o suc was aio isummea aiasi ey sge was a io iwiewaioeos we ma  esee ss a eoguocacieseseasoees i o comme sysicou eiw sea o1sayea coma gw iwieisumme sie iis sesss a i co sucstComa gw me s cause oe imcis oeaa  me oamaSes ocoma gw omamamma a cki a may co  euei e oes oSesea o a oesciesaso  e seasoee icaci oSesea o  coom iosess maiuaa su  assea cei ses aioo seasoas a co-c    coio owi amaicou ao  ies  assess e imcoa w-qua seoe caci osea o  comaComa gw usuayquis a oogey iuwcmay  asie iama oy access  w-ey sa  miges cacioome sucas oomes oos  ewi useassess ysiogicass oama  a s(Cos Oi19Aou eaMeisIoGuiea  weese me seem  ss accu  mi sico sucas iousIwou us  usea  a ae oomeacou  easi coecom seao  assess eiysiogicas a o  coio Iis ao ima keeimi aces ieoc ways gu ysiogy a ouoccuae  ie  ugo coeo ie oooai a gosa ceo ai a aiCoequea aem  ue e eoc guooey e a  coiooSesea o su ioo measumes oceo a i oiAwie oos (e i ge a oe os (sucasee  a e  ie case oyiossu ao  iisese ee ioicoe eew- a gey e seose eie ous eec oa ock eoma mamma a1sea swe e  osaaoag a w-ey so simiey iakes cou  e key me eiees ioua ysiogicaauses iSesea o seasu iesga eeec oe  osaaooeoc sig a e ioss oSesea o  eue w w-ey e imcess oese amaStrh d nSes iesga e imcoces ieoe ysiogy a ess oama a seewe amais coiii eie scaacwe aemeos a wi sco ouei oe o se oiiasea o ae ysiogica oeeceoc aou Ieg e auses ma aeacoese  aows a moaccu a ougasis ow sea o ac eieaces a eiacoequees is oeisuia oc caciCai ses o a uque oou  acqui  aa icu oimssie  coecie wiey ao o a coesei  ase a eeec oica ce simeyeasc eemeasigoousece esis asea o om e c wes o may mi sicoa coc sis w coumi emi w-ey scom ose ie iasi eas oaa coumi iesCoeque a -3 case oey iake wama a swiom gw-ey ock  a -y ois a asoe amaoowisoug   occueie wi eaacu eemeasicoAiic coequee osimieses ia coeeie is ae ama a suec e mui-cacoio aeiwi coueaseoue Wi ama a ke  coume mo aa si scies osas iousaugsea o om e wes o eima (95%coumi w-ey s(Siieis ea As suc usia si scies ow-ey ey oa miu ow-ey scies su yiesimisus13gi a ac eees a uuesoy wesea o icaia cou a sigca ees ioeec cisio ma  amaIs ooeec cauoa momome we seoassumioa yieeesmas oey a maiee ey awe wiie e owwe u ioeses (see Ce3 Osus oee eemes eseequa ces sea o  e sicoa sueque -eey sweasea o (eougcaisigcaaeei oac (iase ocase i oe seaso  oimiey eeis gi io e o oi sueecoio ey wou  seasoy eoueie wiGiaysiogicaaco su  simiamo Sesea o gass oweey a caiowi amaae cosio ca awaue ysiogicaeec oseaso qua oea w ey iec su ie ysiogicaa ouces oeeSesea oSame si was  a se a a weakss oousOo eeig ma Sesea o aok  ioueemes eseea e same a cai s oe ma mammaOe oea samesi oeig caseem w om a sscacis was iay oome  a sime s siga emoyi ameasus oeacama iasee sscawe e sigca ees u s e w same si aus ke   ascOous oae ioao ue ioysiogy oSesea o uee ea seasocoioaugima caa su  ke imi we ousus  wi oIlnOusus  imicao mageme owi Sesea oiosess s ecommoy assume  mo seous  wieasumme(Meick ugi1997Wiea  we ousus sweoyasea o wis o omi iosess mo easi iwieeyao a ae  como easi om icom e summe131Sesea o aea ea e iase iey iake isummecoom a ss o massoeise ey may maiia suimas ie summeawi uima case eiessIasi ey iake ssicoa a gic se wou qui a sea o iase i gieowcmea ayou a e ama mig eose emsees  a gask oeoisummecom wie Suce iases iey iakemay ao  imssie  aai ama a a migescaciicuy ieie e gic se is maiy comseow-ey i s Coequesummesu  coias a mo ccaooe yeaSesea o awie esciay suiosea ima w-ey s Mageme cisio su ao coie gaeosea o  suiay iase eiey iake  coaea miey sicoisummea e sui iagacoc amig occuwi sesoiooie  ( ao cae coio s  euseas aie assess aoi (Cos ea 19Gas u199a ea a we ousus sweaw sea o ea ock wesi mass ae same  as ama ea iei coio syeseosg imoe  a emi ss oassuesesciay iwieSusioiss ca mo ous oa coc sis ass ossuea yeiwas ce  aomeasumes o coio e o as aieoe  coiooama may us  e siicaoeiaaasSesea o a a emi eow-eys Ioaooe  coiooSesea o io wiemi w-ey se may us  esea oeia asu  keio accou i e sesesciay e wes oOusus ee uei oe coequees aiosessmay  oe ess oSesea o a os ies w sus oeseasoeecaa eow-ey sca osea oey sw aa eoock may  su eec yo e  omass ss ae sea o omoimay aus eiysiogies  coma suimaeiea13coio a  iosessCes osuimay  weamacewi coc sage ow-ey ey com ama ea geys Uimaouis su    oeec mo e assiswi e mageme a e coeaooSesea o133fAou Gew  (s mesm iAac sea(Arh zl  Com ocmisy a ysiogy A1-1Cos C ck C(19gi eecs oAac sea io ces iey aiEcogy 759Cos OiC(19oocmisy meossis  oes ie ea sea Amecauaoysiogy 91-595Gas  u(199 Assessme o coioo seaogy13-3GuiCSee  MaiSGeoes ciA (Ce iasma coisoa mes  e aeae oas is csuac seaCom ocmisy a ysiogy A-MocuIeg ysiogy 137-531MeisIoS(oomes as iica oie uoisc ociseaes eooie sees ico Caauaoogy 79-311Meick ugi (1997gi ooa ma a you-oyeaSesea o iAskawaCaauaoogy 7577a seis AW (Uoosey sees  ses  age a seasoiSesea oCaauaoogy 9SiiE e (Seasoa saees ieie wessck oSesea o  (Ej  uaoMammagy 73-99is AWMieA MasceGAeMAgS CaeCaiACoy KO o Eey GgE GscCE S  Gcke  KacKim  CMaua MaCMaswski WMeeso  ioOkkoS Oa e-MasceKyeSci Wa  WiA(om-ui a e13c oSesea o  (Ej iAskaassessi e oceacma esisses Oceagy 1-WiA is AWseS (A oeec moesma e oquimes oSesea o  Ej  iAskaUSAMaEcogy-ogss Sees 991-31135

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