Laura Hinchliff: "The effect of infection with the swim bladder parasite, Anguillicola crassus, on the lipid content of the European eel, Anguilla anguilla"

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Ogpgj ambtebt mi t`e Eurmpenb eeo (Nbhugoon

nbhugoon) w`eb gbieatej wgt` t`e swgk donjjer

bekntmjeNbhugoogamomgjes arnssus

Onurn @gba`ogii

(^ueebs Ubgversgty Deoinst)

Qupervgsmrs

Jr. J. Evnbs, Jr. J. Tmderts

N prmfeat suppmrtej dy t`e Eurmpenb Ubgmb's GBZETTEH GYN \rmhrnkke knbnhej dy t`e Qpeagno EU \rmhrnkkes Dmjy

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Qtntekebt mi Ampyrgh`t

Z`e ambtebt mi t`e GDGQ wedsgte*(gbaoujgbh, dut bmt ogkgtej tm noo text nbj nrtwmrc t`eregb)

gs prmteatej dy ampyrgh`t. Z`e ampyrgh`t gs mwbej dy GDGQ mr gs mt`erwgse ogaebsej irmk n

t`grj pnrty imr use dy t`e pnrtbers.

Wmu kny vgew mr jmwbomnj nby ambtebt mi t`e Qgte imr prgvnte purpmses, dut ymu nre bmtperkgttej, wgt`mut mur perkgssgmb, tm6

stmre t`e Qgte, mr nby ambtebt mi t`e Qgte, imr nby mt`er purpmse3

prgbt ampges mi t`e Qgte, mr nby ambtebt mi t`e Qgte, imr nby mt`er purpmse3 reprmjuae, ampy mr trnbskgt t`e Qgte, mr nby ambtebt mi t`e Qgte, gb nby wny, imr

nby mt`er purpmse mr gb nby mt`er kejguk.

Noo mt`er rgh`ts w`ga` nre bmt speagiganooy hrnbtej nre reservej.

GDGQ knces nvngondoe mb gts sgtes kntergnos heberntej imr GDGQ dy nanjekga wmrc nt t`eUbgversgty mi Honshmw nbj nt ^ueeb's Ubgversgty Deoinst, gbter nogn.

Extrnats kust bmt de qumtej wgt`mut expogagt perkgssgmb. \erkgssgmb kny de smuh`t dy

usgbh t`e ambtnat imrk www.omuh`s-nhebay.mrh/gdgs/ambtnat/

*Qgte - www.omuh`s-nhebay.mrh/GDGQ

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OG\GJ AMBZEBZ NBNOWQGQ MI Z@E EUTM\ENB EEO (NBHUGOON NBHUGOON)

X@EB GBIEAZEJ XGZ@ Z@E QXGK DONJJET BEKNZMJE

NBHUGOOGAMOMGJEQ ATNQQUQ

Zm gbimrk knbnhekebt strntehges imr t`e Eurmpenb eeo

DNACHTMUBJ

Z`e reprmjuatgve kghrntgmb mi Eurmpenb eeos gs onrheoy jepebjebt mb njequnte eberhy reserves stmrej

ns ogpgj. Inatmrs t`nt kny gbiouebae t`e ogpgj ambtebt nre t`ereimre aruagno tm ubjerstnbj gb mrjer tm

nssgst t`e jeveompkebt mi iuture knbnhekebt strntehges.Nbhugoogamomgjes arnssus, nb gbvnsgve pnrnsgte

bntgve tm t`e Fnpnbese eeo, `ns deamke prmdoekntga gb Bmrt`erb Greonbj wnters sgbae gts igrst jgsamvery

gb Omuh` Erbe gb >770. Z`e strngb mi gbieatgmb wgt`N. arnssuskny resuot gb n behntgve eiieat mb t`e

ogpgj ambtebt mi eeos. Z`gs repmrt ngkej tm nsaertngb t`e extebt mi w`ga`N. arnssus kny `gbjer t`e

reprmjuatgve kghrntgmb mi t`e Eurmpenb eeo dy nssessgbh amrreontgmbs detweeb ogpgj ambtebt nbj

pnrnsgtga gbieatgmb detweeb twm eeo igs`ery sgtes gb Bmrt`erb Greonbj. Z`gs gbimrkntgmb kny de usej tm

gjebtgiy cey nrens imr t`e jeveompkebt mi knbnhekebt strntehges imr t`e Eurmpenb eeo.

KNGB IGBJGBHQ

Z`e ogpgj ambtebt mi eeos wns pmsgtgveoy gbiouebaej dy oebht` nbj knss, gbjgantgbh t`nt onrher,

`envger eeos `nj n `gh`er ogpgj ambtebt t`nt sknooer eeos.

Eeos wgt` n `gh` gbieatgmb gbtebsgty miN. arnssus `nj n sghbgiganbtoy omwer ogpgj ambtebt gb

Omuh` Benh` dut bmt gb Omuh` Erbe.

N. arnssuskny resuot gb n jearense gb ogpgj ambtebt mi eeos jue tmN. nbhugoon`nvgbh tm exert

kmre eberhy tm iubatgmb bmrknooy mr N. arnssusutgogzes ogpgj irmk N. nbhugoonimr t`egr mwb

hngb.

N. arnssushebernooy jm bmt cgoo t`egr `mst ns gt gs kmre debeigagno imr t`e `mst tm rekngb nogve.

BMB-ZEA@BGANO QUKKNTW

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NDQZTNAZ

Z`e swgk donjjer bekntmje Nbhugoogamomgjes arnssus, nb gbvnsgve pnrnsgte ebjekga tm t`e

Fnpnbese eeo (Nbhugoon fnpmbgan) `ns deamke n sergmus prmdoek gbN. nbhugoonigs`erges sgbae gts igrstnrrgvno tm Eurmpe gb Herknby nbj Gtnoy gb >704. N. arnssus anuses n kuotgtuje mi prmdoeks tm N.

nbhugoonsua` ns gbtestgbno oesgmbs, noterntgmb tm t`e swgk donjjer wnoo nbj jgsruptgbh mt`er mrhnbs gb

t`e prmaess. Z`e strngb mi `mstgbh n snbhugvmrmus pnrnsgte kny resuot gb jepoetgmb mi eberhy reserves

sua` ns houamse nbj ogpgjs.

Ogpgj ambtebt nbnoysgs wns perimrkej mb ieknoe sgover eeos irmk twm sgtes gb Bmrt`erb Greonbj

(Omuh` Benh` nbj Omuh` Erbe) nbj amkpnrej nhngbst stnbjnrj kmrp`mketrgas tm nsaertngb t`e eiieats

N. arnssus kny gbjuae mb N. nbhugoon prgmr tm kghrntgmb. Z`e ogpgj ambtebt mi eeos wns pmsgtgveoy

gbiouebaej dy dmjy oebht` nbj knss ns onrher, `envger eeos `nj n `gh`er ogpgj ambtebt t`nb sknooer eeos.Tesuots s`mwej t`nt t`e jgiierebaes gb ogpgj ambtebt gb Omuh` Benh` nbj Omuh` Erbe amuoj de

nttrgdutej tm jgiierebaes gb oebht` detweeb t`e twm sgtes. Z`gs gbjgantes t`nt dmjy oebht` kny de n cey

inatmr gb gbiouebagbh t`e eiieat t`nt pnrnsgtes `nve mb ogpgj. Eeos wgt` n `gh`er gbieatgmb gbtebsgty miN.

arnssuswere imubj tm `nve n omwer ogpgj ambtebt3 `mwever t`gs resuot wns mboy sghbgiganbt gb Omuh`

Benh`. Z`e jgiierebaes seeb detweeb t`e twm sgtes kny de expongbej dy jgiierebaes gb igs`

amkkubgtges nbj jgstrgdutgmb mi t`e gbterkejgnte nbj pnrntebga `msts gb t`e twm sgtes. Z`muh` resuots

nre gbjgantgve mi N. arnssus `nvgbh nb eiieat mb t`e ogpgj ambtebt mi N. nbhugoon, omw snkpoe sgzes

kenbs gt rekngbs ubaertngb.

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ZNDOE MI AMBZEBZQ

> GBZTMJUAZGMB """"""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""" #>.> UC eeo igs`erges """"""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""" #

>.4 Bmrt`erb Greonbj eeo igs`erges """"""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""" #

>.4.> Omuh` Benh` eeo igs`ery$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$ %

>.4.4 Omuh` Erbe eeo igs`ery$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$ &

>.1 Qtntus mi t`e Eurmpenb eeo """""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""" $

>.1.> Mverigs`gbh$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$ '

>.1.4 Aogknte a`nbhe$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$ '

>.1.1 \nrnsgtga gbieatgmbs nbj jgsense$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$ (

>.: Nbhugoogamomgjes arnssus """"""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""" %

>.5 Nbhugoon nbhugoonogie `gstmry nbj spnwbgbh kghrntgmbs""""""""""""""""""""""""""""""""""""""""" &

>.5.> Qpnwbgbh kghrntgmb$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$ )

>.5.4 Eberhetga reserves imr kghrntgmb$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$ *

4 NGKQ NBJ MDFEAZGYEQ""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""" '

1 KNZETGNOQ NBJ KEZ@MJQ""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""" (

1.> A`mgae mi stujy eeos""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""" (

1.4 \rmaessgbh mi eeos """""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""" )

1.1 Ogpgj ambtebt nbnoysgs"""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""" )

1.:

Qtntgstgano nbnoysgs""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""" #*

: TEQUOZQ """""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""" #*

:.> Qtntus mi eeos"""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""" #*

:.4 Amkpnrgsmb detweeb ogpgj nbj stnbjnrj kmrp`mketrgas6 jgiierebaes mi oebht`,

knss nbj ambjgtgmb detweeb sgtes""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""" ##

:.1 Amkpnrgsmb detweeb ogpgj ambtebt nbjN. arnssus6 pnrnsgtga eiieat mb ogpgj

ambtebt detweeb sgtes"""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""" ##

5 JGQAUQQGMB""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""" #%

5.> \nrnsgtes nbj igs`erges""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""" #%

5.4 Qtnbjnrj kmrp`mketrgas6 reontgmbs`gp wgt` ogpgj ambtebt nbj eiieat wgt` N.arnssus gbieatgmb""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""" #%

5.1 Gbieatgmb wgt` N. arnssus nbj eiieat mb ogpgj ambtebt"""""""""""""""""""""""""""""""""""""""""""" #&

5.: Gbieatgmb wgt` N. arnssus detweeb stujy sgtes""""""""""""""""""""""""""""""""""""""""""""""""""""""" #'

5.5 Ambaousgmbs"""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""" #(

9 TEIETEBAEQ """""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""" #)

8 AM\WTGH@Z"""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""""" $&

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OGQZ MI IGHUTEQ NBJ ZNDOEQ

\nhe bukder

Igh. >. Ogie ayaoe miN. nbhugoon

Igh. 4. Qgtes mi stujy eeos

Zndoe >. Z`e jgstrgdutgmb miN. arnssusgbieatgmbs gb Omuh` Benh` nbj

Omuh` Erbe nbj ogpgj ambtebt miN. nbhugoon

Igh. 1n. \omt mi t`e ogpgj ambtebt (ns n % mi dmjy knss) nhngbst dmjy

oebht` (omh)

Igh. 1d. \omt mi t`e ogpgj ambtebt (ns n % mi dmjy knss) nhngbst dmjy

knss (omh)

Igh. 1a. \omt mi t`e ogpgj ambtebt (ns n % mi dmjy knss) nhngbst ambjgtgmb

mi eeo

Igh. :n. Bukder mi gbjgvgjunoN. arnssusper eeo gb Omuh` Benh` nbjOmuh` Erbe

Igh. :d. Ogpgj ambtebt (ns n % mi dmjy knss) gb Omuh` Benh` nbj

Omuh` Erbe

Igh. 5n. Ogpgj ambtebt (ns n % mi dmjy knss) nbj t`e bukder mi

N. arnssus

Igh. 5d. Ogpgj ambtebt (ns n % mi dmjy knss) nbj t`e bukder mi

N. arnssusns gbieatgmb gbtebsgty

9

7

>4

>4

>4

>1

>1

>:

>:

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%

> GBZTMJUAZGMB

>.> UC eeo igs`erges

Z`e Eurmpenb eeo (Nbhugoon nbhugoon O.) maaurs wgjeoy t`rmuh`mut rgvers, onces nbj estunrges

gb t`e UC. Qeverno jgiierebt types mi igs`ery exgst imr t`gs speages ns vnrgmus ogie stnhes nre expomgtej6

(>) t`e eover mr honss eeo igs`ery, w`ga` anptures ymubh eeos ns t`ey returb tm rgvers. Z`e Qeverb

estunry detweeb Ebhonbj nbj Xnoes dmnsts t`gs amkkeragno igs`ery, `mwever honss eeo igs`gbh gs

gooehno gb Bmrt`erb Greonbj3 (4) yeoomw eeo igs`erges exgst narmss t`e UC, imr exnkpoe t`e Z`nkes3 (1)

sgover eeo igs`erges, imr exnkpoe Omuh` Benh` gb Bmrt`erb Greonbj (Ebvgrmbkebt Nhebay 4).

>.4 Bmrt`erb Greonbj eeo igs`erges

>.4.> Omuh` Benh` eeo igs`ery

Omuh` Benh` sustngbs t`e onrhest amkkeragno wgoj eeo igs`ery rekngbgbh gb Eurmpe w`ga` gs

knbnhej dy t`e Omuh` Benh` Igs`erkebs Am-mperntgve Qmagety (OBIAQ), tm w`mk noo igs`gbh rgh`ts

mi eeos gb Omuh` Benh` deombh (Xnocer et no4>). Z`e OBIAQ `ns deeb gb ambtrmo mi t`e eeo igs`ery

sgbae t`e >79788). Z`e eover arns` gb t`e >70

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&

Imr dmt` igs`erges, restrgatgmbs put gb ponae sua` ns anta` qumtns, dmnt sgze restrgatgmbs, ogaebsgbh

sa`ekes, kgbgkuk eeo onbjgbh sgzes nbj natgve igs`gbh sensmbs perkgt sustngbndoe knbnhekebt

(JEITN 4.4.4 Omuh` Erbe eeo igs`ery

Z`e Erbe Eeo eb`nbaekebt prmhrnkke wns n knfmr Eurmpenb iubjej armss-dmrjer

prmhrnkke, set up gb >778 dy t`e Bmrt`erb Tehgmbno Igs`erges Dmnrj gb nb nttekpt tm eb`nbae

rearugtkebt mi eeo tm t`e pmpuontgmb, noomwgbh iurt`er expomgtntgmb nbj tm jeveomp n knbnhekebt ponb

imr t`e iuture mi t`e igs`ery (Knhugre nbj Hgdsmb 475777).

>.1 Qtntus mi t`e Eurmpenb eeo

Z`e stntus miN. nbhugoon`ns deeb ubjer gbtebse sarutgby jurgbh t`e onst 1< yenrs irmk dmt`

igs`erkeb nbj sagebtgsts (Kmrgnrty nbj Jeccer >778) jue tm jeaogbgbh igs`ery ygeojs nbj nwnrebess mi

pmmr `enot` stntus. Gt `ns deeb nppnrebt t`rmuh`mut Eurmpe sgbae t`e >70

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(Xgrt` nbj Derbnt`a`ez 4

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(

INM onbjgbhs nbj honss eeo anta`es dmt` s`mw gbternbbuno iouatuntgmbs ambsgstebt wgt` t`e

Bmrt` Ntonbtga Msagoontgmb (BNM) (Cettoe et no4

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+

>.: Nbhugoogamomgjes arnssus

N. arnssus gs n pnrnsgtga swgk donjjer bekntmje ebjekga tm t`e Fnpnbese eeo,N. fnpmbgan, gb

Enst Nsgn (Kmrnvea nbj Znrnsa`ewscg >700, Ceooy et no4

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)

>.5 Nbhugoon nbhugoonogie `gstmry nbj spnwbgbh kghrntgmbs

N. nbhugoon`nve nb gbtrghugbh ogie `gstmry gb t`nt t`ey `nve dmt` n ambtgbebtno nbj maenbga

p`nse (Igh. >), sudfeatgbh t`ek tm n vnrgety mi perturdntgmbs (Jurgiet no4

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*

ayaoe (\gerrmb et no4788).

>.5.4 Eberhetga reserves imr kghrntgmb

Z`e eiigagebay wgt` w`ga` eeos naaukuonte eberhy reserves imr kghrntgmb gs vgtno ns mut mi t`e

vnrgmus eeo speages,N. nbhugoontrnveos t`e inrt`est tm rena` spnwbgbh hrmubjs (5

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,

jeveompkebt mi iuture knbnhekebt strntehges. Z`e aurrebt onac mi jntn rehnrjgbh t`e gbiouebae miN.

arnssusmb t`e ogpgj ambtebt miN. nbhugoonoenves n hnp gb mur cbmwoejhe. Z`e extebt mi w`ga` t`ese

pnrnsgtes kny gkpnat upmb t`e strntehges w`ga N. nbhugoonreoges imr suaaessiuo ambtgbuntgmb mi t`egr

ogie ayaoe gs t`ereimre pmmroy ubjerstmmj.

Zm t`gs ngk, t`e presebt stujy exnkgbej t`e ogpgj ambtebt mi eeos gbieatej wgt N. arnssusnbj

amkpnre t`gs wgt` stnbjnrj kmrp`mketrgas sua` ns oebht` nbj knss tm nsaertngb t`e eiieatsN. arnssus

kny gbjuae mbN. nbhugoonprgmr tm kghrntgmb. Z`gs gbimrkntgmb wgoo `eop tm nsaertngb t`e extebt mi

w`ga`N. arnssuskny `gbjer t`e spnwbgbh miN. nbhugoon ns ogpgj ambtebt gs n vgtno amkpmbebt tm t`e

suaaessiuo amkpoetgmb mi t`egr reprmjuatgve kghrntgmb (\gerrmb et no4 ieknoe sgover eeos were amooeatej dy igs`erkeb irmk Omuh` Benh`, nbj 1 (Igh. 4). Z`ese sgtes were a`mseb ns Omuh`

Benh` gs t`e onrhest rekngbgbh amkkeragno wgoj eeo igs`ery gb Eurmpe, `ebae t`mrmuh` ubjerstnbjgbhmi pnrnsgte-`mst reontgmbs`gps gs gkpmrtnbt ns weoo ns cbmwoejhe ns tm t`e qunogty mi t`e eeos gb t`gs

igs`ery. Ns n prevgmusoy natgve eeo igs`ery, Omuh` Erbe wns a`mseb ns n amkpnrntgve sgte. Dmt` sgtes

were cbmwb tm ambtngbN. arnssus.Z`e amooeatgmb pergmj wns tgkej sm t`nt ieknoe eeos wmuoj de

anuh`t gb t`egr sgover stnhe deimre t`ey dehnb t`egr spnwbgbh kghrntgmb3 ogternture `ns s`mwb t`nt t`ere

nppenrs tm de bm sensmbnogty miN. arnssus gbN. nbhugoon (Z`mkns nbj Mooevger >774n, Ceooy et no

4

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Igh. 4 Qgtes mi stujy eeos3 Omuh` Benh` `gh`ogh`tej gb jns`ej ogbe, upper nbj omwer Omuh`

Erbe `gh`ogh`tej gb ambtgbumus ogbe. Tei6 Hmmhoe knps 44

1.4 \rmaessgbh mi eeos

Eeos were amooeatej nbj stmrej nt ->0mA. \rgmr tm prmaessgbh eeos were jeirmstej nt rmmktekpernture imr 4: `murs. Oebht` (kk), wegh`t (h) nbj sex were reamrjej. Nb gbagsgmb knje nombh

t`e ubjersgje tmwnrjs t`e peatmrno igb noomwej naaess tm t`e swgk donjjer, nt w`ga` pmgbt t`e bukder

miN. arnssuswere nssessej knarmsampganooy. N seatgmb mi kusaoe tgssue pmstergmr tm t`e nbus wns

rekmvej irmk t`e oeit sgje mi t`e eeo nbj stmrej gb 50mA.

1.1 Ogpgj ambtebt nbnoysgs

Kusaoe snkpoes were oeit tm jry nt 55MA imr >: jnys gb ondeooej honss petrg jgs`es (J6

7.:h mi

kusaoe wns imojej gbtm igoter pnper nbj stnpoej tmhet`er. \geaes mi ponstga were nosm stnpoej tm t`e

igoter pnper tm jgiierebtgnte detweeb snkpoes. Z`e wegh`t mi t`e snkpoe gb pnper wns reamrjej nbj

t`ree snkpoes were igttej gbtm mbe extrnatmr. Z`e ogpgj ambtebt nbnoysgs prmaejure wns njnptej irmk

ket`mjs ns jesargdej dy Imoa` et no(>758).

4 (v/v) a`omrmimrk-ket`nbmo imr `mkmhebgzgbh t`e tgssue wns kensurej gbtm

rmubj dencers nbj ponaej mbtm n `enter. Xnter rnb ambtgbumusoy t`rmuh` t`e extrnatmrs imr 9 `murs tm

sepnrnte t`e ogpgjs irmk bmb-ogpgj sudstnbaes. Qnkpoes were t`eb rekmvej nbj jrgej nt 55MA

mverbgh`t tm rekmve nby rekngbgbh kmgsture. Qnkpoes were t`eb ponaej gb n jessgantmr imr nb `mur tm

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:.4 Amkpnrgsmb detweeb ogpgj nbj stnbjnrj kmrp`mketrgas6 jgiierebaes mi oebht`,

knss nbj ambjgtgmb detweeb sgtes

Z`e kenb ogpgj ambtebt mi eeos ns n peraebtnhe mi dmjy knss wns `gh`er gb Omuh` Erbe (Zndoe

>). N pmsgtgve ogbenr reontgmbs`gp detweeb t`e ogpgj ambtebt nbj omh-trnbsimrkej jntn mi eeo dmjy

oebht` (Omuh` Benh` T4 ; >83 Omuh` Erbe T4 ;

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Igh 1 \omt mi t`e ogpgj ambtebt (ns n % mi

dmjy knss) nhngbst dmjy oebht` (omh) (Igh.

1n), dmjy knss (omh) (Igh. 1d) nbj ambjgtgmb

mi eeo (Igh. 1a) imr Omuh` Benh` (doue) nbj

Omuh` Erbe (hreeb). Tehressgmb ogbes s`mwt`e T4ogbenr vnoue imr dmt` sgtes.

Igh. 1n

Igh. 1d

Igh. 1a

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Igh. : Bukder mi gbjgvgjuno N. arnssus per eeo (Igh. :n) nbj ogpgj ambtebt (ns n % mi dmjy

knss) (Igh. :d) gb Omuh` Benh` nbj Omuh` Erbe. Mutogers nre gbjgantej dyMmr *

Igh. :n

Igh. :d

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Igh. 5 Ogpgj ambtebt (ns n % mi dmjy knss) nbj t`e reontgmbs`gp detweeb t`e bukder mi N.

arnssus (Igh.5n) nbj t`e bukder mi N. arnssus ns gbieatgmb gbtebsgty (Igh. 5d)

Igh. 5n

Igh. 5d

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5 JGQAUQQGMB

5.> \nrnsgtes nbj igs`erges

Z`e behntgve eiieats mi pnrnsgtgsk gb amkkeragnooy gkpmrtnbt igs`erges nre weoo jmaukebtej

t`rmuh`mut t`e ogternture, imr exnkpoe sen ogae (Oepemp`t`egrus snokmbgsnbj Anoghus aoekebsg!)

pnrnsgtgzgbh pgbc snokmb (Mbamr`yba`us hmrdusa`n) nbj a`uk snokmb (Mbamr`yba`us cetn), nbj

degbh anpndoe mi ogkgtgbh ambservntgmb eiimrts (Crcm"ec et no4

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detweeb ogpgj ambtebt nbj t`e bukder miN. arnssusgb Omuh` Benh` dut bmt gb Omuh` Erbe. Kenb

jntn s`mws t`nt gb Omuh` Benh`, mb nvernhe eeos were onrher, `envger nbj `nj `gh`er gbieatgmb oeveos

miN. arnssus,yet t`e kenb ogpgj ambtebt wns omwer.

Z`mkns nbj Mooevger (>774d) imubj n pmsgtgve amrreontgmb detweeb eeo dmjy oebht` nbj t`e

bukder miN. arnssuspresebt, sgkgonr tm mur igbjgbhs gb Omuh` Benh`. \mtebtgnooy, t`gs kny gkpnat mb

rearugtkebt ns resenra` `ns s`mwb t`ntN. nbhugoonrena` n argtgano oebht`, nt w`ga` pmgbt t`ey dehgb

t`egr reprmjuatgve kghrntgmb. Z`gs argtgano oebht` gs jgiierebt detweeb knoes nbj ieknoes, wgt` ieknoe

eeos hebernooy hrmwgbh inster t`nb knoes (\mmoe nbj Teybmojs >779). Dmjy oebht` `ns nosm deeb

s`mwb tm gbiouebae t`e up-strenk kghrntgmb mi yeoomw eeos. Onrher eeos were anpndoe mi kghrntgbh

t`rmuh`mut t`e sensmbs rehnrjoess mi tekpernture, dut t`e knfmrgty mi sknooer eeos were restrgatej tm n

s`mrter kghrntgmb pergmj (Kmrgnrty >709).

Ambverseoy, Cnbhur et no(4779) w`m imubj bm reontgmbs`gp detweeb gbieatgmb gbtebsgty nbj egt`er t`e oebht` mr

wegh`t mi eeos gb t`e Tgver T`gbe, Herknby. Dnru"nbj \rmces (>779) imubj t`ntN. arnssusgbieatgmb

gbsghbgiganbtoy niieatej t`e wegh`t miN. nbhugoon.

5.1 Gbieatgmb wgt` N. arnssus nbj eiieat mb ogpgj ambtebt

Z`e aurrebt stujy imubj nb gbarense gb t`e bukder mi pnrnsgtes presebt `nj n behntgve eiieat

mb ogpgj ambtebt (Igh. 5n). @mwever, gbieatgmb gbtebsgty sghbgiganbtoy niieatej t`e ogpgj ambtebt gb

Omuh` Benh` mboy3 t`gs gs jgsaussej gb hrenter jetngo gb seatgmb 5.:. Z`e behntgve eiieat miN. arnssus

mb ogpgj ambtebt kny de expongbej dyN. arnssusgkpngrgbh bmrkno swgk donjjer iubatgmbgbh gb eeos,

w`ga` `ns deeb s`mwb tm hrentoy rejuae t`egr swgkkgbh perimrknbae (\nostrnet no4

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Z`e ndmve rensmbgbh reioeats amkkmb t`muh`t t`ntN. arnssuskny resuot gb n jearense gb ogpgj

ambtebt jue tm inatmrs w`ga` resuots gbN. nbhugoon `nvgbh tm exert kmre eberhy tm iubatgmb bmrknooy

pre-kghrntgmb. @mwever, gt gs wgjesprenj cbmwoejhe t`nt pnrnsgtes nre anpndoe mi renoomantgbh

resmuraes, sua` ns ogpgjs, irmk t`e `mst imr t`egr mwb beejs (\ongstmwet no477>) ns gt gs kmre debeigagno imr t`egr `mst tm rekngb

nogve tm suppoyN. arnssuswgt` immj nbj `ndgtnt.

N. arnssusgs cbmwb tm gbieat knby ogie stnhes miN. nbhugoon3 Bgket` et no(4770 (Evnbs nbj Kntt`ews >777), dut t`ere

wns bm reamrj miN. arnssuspresebae gb Omuh` Benh` ubtgo 4

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sghbgiganbt eiieat mb t`e ogpgj ambtebt mi eeos gb Omuh` Benh` dut bmt gb Omuh` Erbe. Qgkgonroy,

Cnbhur et no(4774 >770, t`nb >777- 4

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(Hmoomac et no4), Z`e Eurmpenb eeo qunogty

jntndnse6 tmwnrjs n pnb-Eurmpenb kmbgtmrgbh mi eeo qunogty,Ebvgrmbkebtno Kmbgtmrgbh nbj

Nssesskebt,>016>-:, 481-40:

Devnaqun, J., Keog, \., Argveoog, N. F., Hnttm, K. nbj Je Oem, H. N. (4:7), Z`e jeaogbe mi t`e Eurmpenb eeoNbhugoon

nbhugoon6 qunbtgiygbh nbj knbnhgbh esanpekebt tm suppmrt ambservntgmb,Fmurbno mi Igs`

Dgmomhy,806>, 41-10

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Dmtgus, G. nbj Dmtgus, F. (>70, >-40

Dmtgus, G. nbj Dmtgus, F. (>705), Ogpgj nbj prmtegb ambtebt gbNbhugoon nbhugoonjurgbh hrmwt` nbj

stnrvntgmb,Jnbn, :, >->8

Dmb`mkkenu, Q., A`nssmt, E. \onbque, D., Tgvmt, E., Cbnp, N. @. nbj oe \npe, M. (477795), \rejntgmb, Dmjy sgze nbj Amkpmsgtgmb mi \onbctmb,Qagebae,

>577:n), Gs t`ere n

rmoe mi maenb ebvgrmbkebt gb Nkerganb nbj Eurmpenb eeo jeaogbe=,Igs`erges Maenbmhrnp`y,

161, >78-477:d), X`y Gs Tearugtkebt mi t`e Nkerganb Eeo,Nbhugoon rmstrntn, Jeaogbgbh gb t`e Qt.

Onwrebae Tgver nbj Huoi=, Anbnjgnb Fmurbno mi Igs`erges nbj Nquntga Qagebaes,5>64, :87-

:00

A`nrrger, I., Knzeoo, Y., Anrnhueoo, F-K., Ndjnoon`, W., oe Hurub, O. O., Oehnuot, N. nbj Oniingooe, \.

(44), Esanpekebt mi sgover-p`nse Eurmpenb eeos,Nbhugoon nbhugoon, jeterkgbej irmk

igs`gbh natgvgtges gb n Kejgterrnbenb onhmmb (Mr, Irnbae),GAEQ Fmurbno mi Knrgbe Qagebae,

976>, 1709), Eamomhgano inatmrs gbiouebagbh t`e amkpmsgtgmb mi t`epnrnsgte inubn mi t`e Eurmpenb eeo,Nbhugoon nbhugoon(O.), gb Greonbj, Fmurbno mi Igs`

Dgmomhy,4064, 47

Asndn, H. A., Onbh, K. nbj Qnoyg, H. (>771), Z`e bekntmjeNbhugoogamon-arnssus(Bekntmjn,

Nbhugoogamogjne), nbj gts rmoe gb t`e jent` mi eeos gb t`e Once Dnontmb jurgbh >77>,Knhynr

Noontmrvmsmc Onpfn,:06>, >>-4>

Je A`nroermy, J., Hrgsez, O., Z`mkns, C., Deopngre, A. nbj Mooevger, I. \. (>77

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&%

JEITN (4-:>

Jehnbg, H., @n`nku, @. nbj Oevnbmb, J. (>709), Z`e reontgmbs`gp mi eeoNbhugoonnbhugoon(O.) dmjy

sgze, ogpgj, prmtegb, houamse, ns`, kmgsture amkpmsgtgmb nbj ebzyke natgvgty (nojmonse),

Amkpnrntgve Dgma`ekgstry nbj \`ysgmomhy \nrt N6 \`ysgmomhy, 0:6:, 817-8:5

Jeccer, X., Ansseoknb, F. K., Angrbs, J. C., Znucnkmtm, C., Feooyknb, J. nbj Ogacers, @. (4

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&&

Jrnaubauomgjen) nbj sudsequebt kghrntgmb mi onrvne,Jgsenses mi Nquntga Mrhnbgsks, 8, 78-

>

@nebeb, M. O. K, Oe`knbb, F., Ebheoskn, K. W., Qtrebderh, I-F., Tmmzebdurh, G., Cercmii, Q.,

Dreteoer, FCoegb (45-4:

@mpcgrc, H., Xgoos, T. D. @. nbj Zmwbsebj, \. T. (>785), Qensmbno vnrgntgmbs gb ogpgj ambtebt mi eeos

(Nbhugoon nustrnogs),Nustrnognb Fmurbno mi Knrgbe nbj Ires`wnter Tesenra`,4964, 48>-481

GAEQ (4

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Cbgh`ts, D. (477>), Qwgkdonjjer bekntmjes (Nbhugoogamonspp.) nbj hgoo kmbmhebenbs

(\seujmjnatyomhyrusspp.) pnrnsgtga mb t`e Eurmpenb eeo (Nbhugoon nbhugoon),Fmurbno ju

Ambsego6 GAEQ Fmurbno mi Knrgbe Qagebae, :8, 17>-170

Cmmps, @. nbj @nrtknbb, I. (>707),Nbhugoogamon-gbiestntgmbs gb Herknby nbj gb Herknb eeo gkpmrts,

Fmurbno mi Nppogej Ga`t`ymomhy,56>, :>-:5

Cbmpi, C., Knjrgoes @eok, N., Ouagus, T., Doegss, X. nbj Znrnsa`ewscg, @. (4>->1>9

Cbmpi, C. nbj Kn`bce, K. (4-:79

Crcm"ec, K. C., Oewgs, K. N. nbj Ymope, F. \. (4777), @mw ebvgrmbkebtno stress niieats t`e gkpnats mi pnrnsgtes,

Ogkbmomhy nbj Maenbmhrnp`y, ::61, 745-75>

Onrssmb, \., @nkrgb, Q. nbj Mcon, O. (>77781), Ketndmoga eiieats mi stnrvntgmb gb t`e eeo,Nbhugoon nbhugoonO.,

Amkpnrntgve Dgma`ekgstry nbj \`ysgmomhy \nrt N6 \`ysgmomhy,::64, 198-18:

Oeiedvre, I., Ambtmurbet, \. nbj Argveoog, N. F. (4), Nb evnountgmb mi gbjgaes

mi hrmss pnt`momhy nssmagntej wgt` t`e bekntmjeNbhugoogamgjes arnssusgb eeos,Fmurbno mi

Igs` Jgsenses,1:6>, 1>-:5Oeiedvre, I., Kmubngx, D., \mgznt, H. nbj Argveoog, N. F. (464, 485-1

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&(

Kmobr, C., Qzceoy, As. nbj Dnscn, I. (>77>), Knss kmrtnogty mi eeo gb Once Dnontmb jue tm

Nbhugoogamon arnssus gbieatgmb, Duooetgb mi Eurmpenb Nssmagntgmb mi Igs` \nt`momhgsts, >>69,

4>>

Kmobr, C. (>771), Eiieat mi jearensej mxyheb ambtebt mb eeos (Nbhugoon nbhugoon) gbieatej dy

Nbhugoogamon arnssus(Bekntmjn6 Jrnaubauomgjen,Natn Yetergbnrgn @ubhnrgan,:>61-:, 1:7-

19700) Tevgsgmb mi t`e hebusNbhugoogamon Wnknhutg >715

(Bekntmjn6 Nbhugoogamogjne) mi t`e swgkdonjjer mi eeos, gbaoujgbh jesargptgmbs mi twm bew

speages,N. bmvnezeonbjgne sp. b. nbjN. pnperbng sp.b.,Imogn \nrnsgtmomhgan,15,>45->59

Kmrgnrty, A. (>709), Eiieats mi t`e swgkdonjjer pnrnsgteNbhugoogamon arnssusmb t`e kghrntgmb mi

Eurmpenb sgover eeosNbhugoon nbhugoongb t`e Dnotga Qen, Igs`erges Tesenra`, :6>, :1-50

Kmrgnrty, A. (>779), Z`e Eurmpenb eeo igs`ery gb >771 nbj >77:,Grgs` Igs`erges Duooetgb,>:, Knrgbe

Gbstgtute >779

Kmrgnrty, A. nbj Jeccer, X. (>778), Knbnhekebt mi t`e Eurmpenb Eeo, Grgs` Igs`erges Duooetgb3 >5,

Knrgbe Gbstgtute >778

Kmrgnrty, A. (>777), Qtrntehy imr t`e jeveompkebt mi t`e eeo igs`ery gb Greonbj,Grgs` Igs`erges

Duooetgb, >7, Knrgbe Gbstgtute >777

Kbjeroe, K., Qures, D. nbj Znrnsa`ewscg, @. (4-447

\ejerseb, K. G. nbj Jgepergbc, A. (4169, >47>->1709),Nbhugooamon, n pnrnsgtga bekntmje mi t`e swgk donjjer sprenjgbh

nkmbh eeo pmpuontgmbs gb Eurmpe,Jgsenses mi Nquntga Mrhnbgsks, >, 447-41

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\gerrmb, I., Dnujrgkmbt, K., Dmssy, N., Dmurjgbenuj, F-\., Drt`es, J., Eoge, \. nbj Knssndunu, F-A.

(4

\ongstmw, Q. F., Zrmussnrj, F-\. nbj Azgooy, I. (46:, 1:9-15>

\mmoe, X. T. nbj Teybmojs, F. J. (>779), Hrmwt` rnte nbj nhe nt kghrntgmb miNbhugoon nbhugoon,

Fmurbno mi Igs` Dgmomhy, :06:, 911-9:4

\muogb, T. (4761, 141-11778), Omw int ambtebts gb ieknoe sgover eeos6 gbjgantgmbs mi,

gbsuiigagebt eberhetga stmres imr kghrntgmb nbj hmbnjno jeveompkebt,Fmurbno mi Igs` Dgmomhy,

5788), Z`e Eeo. Dgmomhy nbj Knbnhekebt mi Nbhugoogj Eeos,Ombjmb6 A`npknb nbj

@noo Otj, >-:1:Z`mkns, D. nbj Mooevger, I.\. (>774n), \nrntebga `msts mi t`e swgkdonjjer bekntmjeNbhugoogamon

arnssus,Jgsenses mi Nquntga Mrhnbgsks, >1, >95->8:

Z`mkns, C. nbj Mooevger, I.\. (>774d), \mpuontgmb dgmomhy miNbhugoogamon arnssusgb t`e igbno `mst

Nbhugoon nbhugoon, Jgsenses mi Nquntga Mrhnbgsks, >:, >91->877778-

11