r

International Councll

tor theErplorntlon ot th

1990 IJ t, 10

alt1cP1sh Committee

•

A r::.odel of cmbryon1c pcriod ofdevelopment in Balt1c8ca cod. . .. .

'Baltic Pisher1co' Reaearch. Inatitut~ (DaltNURB)

6 DuUga~gr1VWJstr•• l11ga 22'6049~ LatT1a,UsSR.

·Abstract. A' s:mthemat1cal ::odo1 pcrmitt1ng tOßDDcoi"codeSa:s.~

vivel on the bauls of' egga 01:0 e~oa1tlmi,and ~010ß1Ca1-Con';' ,

cU.~onD during tho öPaTm1ng'~~r1od.·13'ropi-csontCd' 1nthe "~per~ :

~emathod. or e~gB 'iiu:rT1v8le.aDoDsm~t1a :cioaCr1b~"acco~: to.' " ~ .' .; ", ' ~ .~'.:-'. : .. :' .•••, \..~.!.t. ~ ;.'-:.•. ' ,,' ,"' .

2ml1n1t7. temporntura end CX%7SCJ1ccntent in uater:and also Beoor- .

~' toegga.Dae~ fhO,' d~P'~deD~O of :ha~tan~oUamortaut7. at '

dif:terentetagcs ot' eggDdo~o1~~t'WB~O'v1thth'8~~Pci"\.' ... ' .. " :.. .'. . . . .... -

tho. special· Oo."'.l1putcr P1"O~(I. ''.. ~ " , . ' ". .', ' , . :,',', " ,',. : '

nesm.u~. Dans co rapport· on proP050 1m .=odcle nmthcmat1que pemet-• ...... ,. • ". '>. • ,- .' '. ... ,.:, •• :••. ~. ":,. : •• ..:.,.• .-.; ~~' ", '" ". ,.

, tant d' evo.1uer 18 ourv1c dca oouto 4c 1& mor\1C 80 baaant aur la .. ,,' .:.'\.....,. . ." .:.

compon1Uon' d1mDnD1onnel1o' dea oeut'o' et, 10scondJ.t10n~log1qu•." ' '. , ~. ..' '. ,..,.:.. .' .', ... , ...

P~lmt 10. trtü. Lo procode d'e.cc.roi.eGe:1entde la,'curv1e dee ooUZs.. ,. '. . '. -:.". " ". .. ", . '

, eot decr1:t C11 foncticm 48 1& cal1n:1to,' 40 10 temporaturo et du': . . , ...' , .' ,'.. .' . '..'. .. .; ' .. ,.. ' ..... ' . '.,

tau:: 4 t c:qgono danoltcau, ninel qua clOG ~tJ.UJ1onado8 'oeufs•.. . ;. '.' -~ " '. : •. .' '",' -' • '"•.. ". .'. ..' .... : tr' " . ;", :

A,l'c.1do d ~un programs upca.1n1m:umt ",.~c:.:cur W1 ol"d1J:latour on- ' ......." • ..•. ",'; .;. ,c":" ~'_.- .~... ~'.: ~";' :. • ~.:;.• ,'", . ~_. ,.' ..•. ~:.: '. ." .' .'. "'.". .'

Q eonstru1t 18 relation du coet.t1c1ent da l:1ortalite 1notantanee. .:. ';,." .. . '. . . ..' ,.'.:' l:. ..' :. ., " ",~. .. . '. : '\ ~.~ .~:.

C11X. ~orcnto ota.doo de l'cvoluticn deo oouto' dcpo1Boons... ~.. ~.:: :.::'~'. ,,='.. ' :..". . . ..

Long-tem inTco'tigatlons ot Balt1c eod reprOduot1cn (1954­

.1989) showed.: that' the embr;.on1c pcr10d ü'-tho dec1s1To momeht '1n1ta ~el~DeB ourv1va1. when t1ortaUt7 tluetUate4 trca78

to 99'% accord:lrig t'o' ,.~tiru(1-2. 4). Eggo surn~ :1a the most 'i >< .... ... , • , .• ' ~.

- ;-......,.

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:. relevant crtt;erlcm' of q,\~titat1yo'asaoomnent of 1 te year-olassee1,' • ..,.".. 0 < ' •

. ;' 8t~n&th.(5)::In-t&C,; ~cr n&lt1o c~·&dopted. to·the.r~prOduat1cm~ ..< .: :':'J" ~-'':'"'' .":.~. : .., ~ '.. "'. .. \' .' ~ :.,. ,--:~. t .. ~, '~ ~; _~.: .0·._ '. ;. .. ".:: '~ _,.• , .. ' '' •.j in: tbe' cendi'tions :1Jith lower. aal1n1~ ot _ter in thc 1".lesr bottcm·

·:··l~eri;or:·:.~e:.do·~PB~·:ltti:·.~w,irJ~~~Q!1tr~ll1nP;"f~oto~ 1~' :.a~er;; de~i:~~:·'!r~~·~i~~t1cm··.~:,c~: e~ .':ln..~':Pei~1ai':~"-d~termi~ed~~- ..... : ' ......' ~ ... .,~ .' ••••• ~. ' ...... ;' .. ' ....:. 0 •. 0 '< ••~ ~'.,' •. '.; : ....::.'" • ••• ...... '. ~ ;: .. " .'. ~ • • .... .'.-" ;

;,:·their·.'dens1t,,·:d1ameter and:h7dro!ogical' cond1tione. ·-re~tiCö&1. 410- .

, :<:·'b-1~ti'~,:~~"..'~~o' i&.:·~~~.U!C~~· ~th' the1r .~~e~er~ (3)'~ Large eg~: .". '" .-: .. ' :':"~' .. -.:.. .'.' ..... - '. h~'.'·· ...... -;'.•... ' ..• ' ..: .. ':" 'c",' ' ...... ': 4 :.'.. . .'

...:" ;lOC"t.ted·:1n·~the ..'upper:·lqtU:'8 .. sum'Yed:,better:thari"tnl:l:Ül enes,' ...hieb ..-: , :.~ :.''''l ._ .. , ..•... ~ ~ ..t ~.... • _.t •...(· -:.,'" ~":: .. ~ ~\. .;, ..'~~.;:~.,.. >•••• ~ ~;•••• ,\.: : öt-"-' . .; .-:,':<0-

.~:,:; dÖl'el.op: ~re :cl ose.:to ~~e ..·bottOm;:~~ ~O%yge~"cont~t' io "lmr~" ". .

\:·..::;::·.;·>.::::·~O·.~~~~1·~~:~~~~.;~~t~~.·:.~~ .;~.~:~~a.;~o~·,.·e~ ~~~~v;'·..i-;' on ·'the :~1ö".ot;e8'ga;8izecomPoaitiön:,aUd.bittrolo~c&1.:cOnd1tl'w :.'. . •

':-...:.1n :tb~::a~~iJiji' ~':~le~:·~i"OP~ried~:;.~~~~U3~~~' ~t :.tn;~~:·~l~ ':..•·~·'··:·g1·~·::1'ae~o~:t.~·:~~~:·~t~·eggtJ·;~t::d;.tt"~;"~t·size~Was~~011~~~ .. ·"

·; ..~:.~.:~~814~i·~~·:,~t~:~"ai1rii~:"~~'·;t~~~~".b·~t2~h~"e":cn~"G~'.' .." .'. ~~.1m;O:r~~:t~:·~~~···:·~4~.:·dt:nB~·iy :dep~n4~' ~":t~~s~ .fac'~c~~ .:!h'a't~~:';:' :.-::.:.~~~~.:'. ~~::: ..~~~c1' ~~·n.~~c~ ·:~:.tho: d~t~~ .er ..·~n~~batl~ '~.; .."~.. ' ...', '. ~''':' .. ;' ':-.' ,',:',:"1':::_ ".; :':,::'-.: ~;':"',:,:"',. <,: :.~.,.:.: .•~ .. ~ • .'::~: .• >'. ~ '.\~' :.": _ "" .. ," .. "';. ~ ::~..

.::. r104•. O:7geD', cencetrattcm 1D.;.~: W.tel" :1&7Or,: "",AlBre. eggt! deft10p.....

<>~t~~~:L~:.;-rir.tti~··i·~f··~.zi,;~t~~~~··~ri~1~:··.~~6t~~~t.·Egp·.·':::

·.;,,:it~±i=:i7r ...~~;roua~?~~~ •.n:.,f.~7: o;t ••..•.•.•••.•••

>·:·;...~·i~~.~:.~d:i:~th~~.·:::1~'B~·:~ie~::~·~·;:~~i..~~·~o;·',~··i~~:':"~'.' .";':~~:.8pa""I1C_~' (O;t1~";d:~'1~6lÄt?&4;~9&T~t970 i ;t976-t?78~I.·'.•.: ,:,EornhOIm-'.195B.:1961-19W•.19&4.-1970· c.ud >.1976-1978)." Pour stageD .'

::.: ..·~~·~"~1·~~t .•~~:·t~·nu8 •. C1~~~t~:~tl~ C1946rnre 40- "

'''·.te~~.f~UT1ng··anc1'4ead ;e~s.· ·Essn.1n '.: '8~Pi~"wt:~ CCZ1yenUo-: :

:.,.~. ~~d'~'1nt~::4"1J1~e ~p~'~ ·d:~6~.:tic~.·1);:66:i4~70·CX .' 2>.' .:::1;~4~75~(X~:n;"76~4~eO CE ~··4)~:~·eU.~oter·18 ß1yen'ü m1~-""'" .'l"ClIl~ter md.is .( 1·unit· - 0.025 mm).·

, ··.:;~~itrd-~og1Ctll. data'; < nam~17:y~rt:1caI.· distribut10n of t~tUre; ". . .

.<aaun1 t,.·and .~gtm conaentrst10n.1D BornhOlm·~~._ter8 (~tat1on.~. . '. '.' ./' '. . . . . . .~.. ' -

.~: .,2) amcl.1n :·the 8outhern' part. of'· Ootl.and (station 4&) tor' 12 ,~ears ., . !. . . •

IlIl

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··b,. .~Vd&1.3CJ·(BalGmm)· wG~,u8ed •. :.''. ': .WO C&1au1ated S1s=a-t .(a~sume4 _t~, denettty)·:i.n 8tan~'hori";'

· ZOn,,' accord1ng tO·lf~ir.ZUbov"8·(6) ~o~~Wat~rde~lt7·end, tempora...:

~~. 'aa~1n1Q- ·and.0x7~ content betveen .thuo .11o·r1~ona 1I1!r·~·,de~<~·. '.. ' .'. ' . ., ., .' '

'·tomine4 lt,' ·11nearintorpOlatlcm.:... · ':." .,". ".:.... ,.:.;:

.' "3~CC~. to: th~ ~a~ .. ot: ~~i-t1ciü hau10 c4 egge' in· d11'1'o~t'lvJ~ '::

.~~;.: c~~ otit. ~~·.19b4.'·~d .1·19&9,·at·.»o~öbt: ana.·.Ootl~:d'~;~y.~lzi~·:·~\·:'. " ..' " '. . .' " . <: '. ,.. "', "', ' " .:.~. '. ,.. ',,, ". ,: ',,' ," ;' :~ ".' .. ","" :, \", _.,0. :", :::

'. gt'OWulä. egg iDean"cU.emeter· DU· aiuf-:-ter;denalt7 tor :eve17·.layer:' ':1:; :::.:, .

. ':'Were:'~et'driÜi1ed;' Shtc~ thO·,.a~~lt7·.'~~ ·e~·.~~t~~·~ ~t~~;,·~~··.:··,:·~·:>

[' "..... ~quä1' .to··~ter: d~~l~;'" ~,'d~:~b~:' ~.~~'~~': ~bt~~~:;q, i~o '. rihOB:~ri··:::.·:·

Ii . b'~t~~~d de;·~d~~~·~.~1:·.~~'::·d~~ty ·~;·~~"e~\u~e~~~:Dii'·':";.:···/..·, .:. arid '.;~~~~: ~ho ~~et~~~ot'·~~~·:d~P~~~c~·.:' .:.:':: .., ,:::~"'.'.":',~:'..<';,,::.. '':' .... <:";"

! .> .:4~Po; e'y~.~tiig~·.>· ~~·:·de.el~~~t ;:'~ :.·e·~~ri ~i~~e> ~~~:, or' '~ggS :t~'e'~~ .;>'.. ~~ ~~ d~et~·r:~~··Caic~at·~~·:·AC~~~:~to:·th~ :d'e~~c1eri~~"'riS1E (tl,;)·.. ·

: .e~<d~~l~~'~:~:l~~tGa;<~···thöi1:',ri··det~~~~·::·~~··d·e~t'b~ ·whcr~::;, .... .... ~'.' ~ ..:. . : '.. ".. '10': .~.,., '." ~ .' ,", .,l :C.' . 0', ,'~ :: ," ' ••• ,' " " ',' '. .'. '. ,

.: egge."ere ,tloat1nß.· \Vater ;tämp'eratun" and Ox7gen. ccmcientrattori OD):.' .'· .' .. : .'.' ':- . ,':." :.' '., :.\ " '" :. ' ' .,,:". ",~.,.:... . :;. ' "', .."'. '\.' . ~.':.', . '.:., ':"',:.: ':"':. .

. .:: wen .·eat.tmät·ed tor: tliat ;depth~:Acoord1ng' to 'the .:i:lumber of ·llV1ng.<· .

. ..:,~;~~.,e~>~~~8~'~B;ed"'~~ ::~~'~tti~~{ ~f.e~~:,b8~tan~~d~· .. ::. ::'·· ..:·_~ ..·.~i-t~1~· ~:~: :(i<.·.· 1:i' ~~ ~·;:..:.jf:·(!rh~·::tormuiQo .. a~e\~i~en .~: ...:..

... ••;~e~tjc~Z:~:,~~ •••~~.the~'~ •..~+po~ts~cr.'doL:;:.·'.~w1th···the:.h~~~:·~f, 8pe.~.~·}n"O~~e .~~G·.fO~ :~.r~ ~.~' ..:: ... :.. :..,...../.<..~~.::

.'.5."0 ccmaldor,.thatoxygen concentrat1cn in l:'1tor detorminoo: l!i. .....: .

..~~e;.hich .18···c~~·~t 'd~1ng' 1-a~agO; ~:·dev~l~~e~t. we:c'ompa~::' ;'.'

·;~·".·~al~~: ~~h·~geri :ccmtont·. OxD-in :~~·l·aye~~·:.~e~:·e~'de- .,.: .: ~l~p; anci;~cul8ted tho '~rem~t~~'·~t· ~be:'de~eI1d~n~:eJ.t (OXD) ·at '~." ...

eaeh stage 'or' ~gga' de~ei·~ent •. These Ctll~ui~tl~':we~ ~~~eci ':.~''~t' a.Cco~~ toth~' 'data t~r &11 '7eara '(197~~'978'Gxcl~~)~.:. ::':'.'

6.!he data =ew' and .bydi-Ology ~orthree ~e~··1976,..197a we~.aa84'~~'control' ~t dep~DdenceD' J8.(O:X»)."-: '. .. ....

3.

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nABult~. Vertlc;L;. cUatrlbution cf """,t&r denoity md axygp,n,concentJ:Q­

,t~cm in tha epa-ming layar, (Pig.1,c.nd 2) emuddorabb Vß1'7 according

tri j~&.~.

~e ~Dntrol cf the. hypatheB1e CD egge vert1.c~ distribution 1n '. . .' .

. dOIumdcnco on eg21J d.1weter c.ocording to tho data or 19&4 anti 19&9. . . '. . .

,tor, ~th oP:\~11i: grounds 11&8, earr1ed. out. Egge Daltplezs, taken At,

,,'th&.~e t1to.tioncn·dj,tteren~hor1soimwere 'an».1~~. 'Egga 8W!tPlea, .

, at ,l..n atagea o~ development'; At·Ul-n etasea and at I-IV otagcS.'

, were a:u.1.I,sed ee~tel)r.·StUdent·s' criter1~n uhoWed~ that the 41ft!'" , (t

,'ferenoe of l:1eam e1~eter8 ,n ne1gbbour1Jlg l3ye~ 18 .rel1able to I

'0...9' or.O.95. )JeQD. d1ar,ete~ of eggGlltcUU'ermt e'i;uges o:f',dev~ ,

,1Cp::lcut (1-11 ' und nI-IV) are different', butthese ~U:rfeZ'enc~ß ar6.

not ~~at13t1c~17 trutb!Ul~and'ao~~cn,~cu1Qt1ng'~hedependence

, 'cf eese dO%1zd.tTon eggr;d1aTAster;' the of;~e8 ofdeyolopm~t JZliQ7not

, : :,btt,taken,intocona1derat1on.' "" "" . '.

, Bgzt'l "p4l'mod b7,' a' :tem::de·ana, fert111fOe4;' b&g1n to d~elop; 8nd.

'~'d~;the'tlrat')-4'houra,ere"bydrated; 1'or.i:l perlv1te;n:1.~e' Spl'lOe,

• ;a~becoms ,bi~r.'!fhe ~er1ment~ Dhond~,C1');that1n~t~rith ,

'~t"~1U11V1~~gg,d1=~terincrea8ed '~71~3$' th8t' ~~n~t areat'

'~C~,W1th'ems.~r1abl1lt7·'~,na~·cmid1t1ons~We,bad'1;o"" ...

, ,: '~le :.the eggo after ~tio~;vhen ,tho7t\1.reaeti",~ 8t the C~'

.tc1n :~or1'1cn. , , . ,"•.,..:The depandeneeQt ee;g dena1't1': attorJ:.T;rd.ro.tlon ml ita diameter ..

", 'mut ded~ced~:Let An . egg "!li, a tCca1e bad:' befcre by-ü.rß.t10n han the

',d1=eter dte.tter'b7~t10n-DY(7olkl',:and af'ter'gett1lig tuto

, w.~e%'.,- m!.'B~den81ty betore~t1cn1~ dec1gnated aa ('1; U~: qu:d dens1'Q";'l'\doorbed by the yon:, ~t the procee" cf h7draUen -e2,the de:Uslty cf theporlv1tel11ne ops.eo content -p'J. and e~ denal­

"ty ~ter gott1n.g "nt~ ,tho wator - DSTE. Tbc n1sht 01" an egg after

gett1ng 1nto 't,he water '18 tho Sw:l of the' welght or' YO~ proper be-

, ~ore bydr&non P 1.11/6$d';' the "C1ght or the adoorb9d b,. the ,"olk, ", " ... '

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., '.,

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llqtd.d e2*J1i~.' (nYJ '-. d'.' and. th:! W01,ght' o~ the, peI-1T1t~l11ne .. '

8paQ~ c~tentpJ~51i6* (Dl43 -DY».·.we cons1deranegg;tonn &e .

aban. D1:nding .thie wIue bythe·re~tedegg·~lume 71/6* "DM) .

·we. hav~ 'got '. ;DST~ ~.. 'P1.;' ~)/DM)- p2*d"J/DU3+ f2.DY~/DltJ+:'('3:*('1 ~py3lP~j:: ~::. We :auume th4t : ..a)S1nce. egg d1m:1eter'~ whexi gottinginto the' "at·or•.:.~ .. Cbange' 1mi1gn1f'ieant~1n:Dalt1C cca :'C~Par1ng 1dth'oth~r nah' s~~~:·,·.~1ea;':~ c'onsldcr~tDr', Jm'. ~" ~ conat~.··" '; .: .. ';. ,".,.', ':"::.::;:' -'.'

.: b)Thed~1t7p1~p2··andf3d·onot.'depe~ on. mt~.:.. .' .": ;'.~. : :.,.. nua.. we ·l1äve tot f'rom (t ) :.. .' . . . . " '.:- ,'. ..: "

"DSTE • a)"cp 1:~ p~)/Dn~;...p 3(1:-·E3),.'·~.pz. &.3 ":'~Dm "~ .Af.IlInUf :.' Where.::A ·•.P3 :(1:- ·t3); +e~ :1t~~' a;ud.B'.~ '~3 ;'~p1 .. -e2):':· ...:.... :: .. :.~....'.;:::.'.;.APpaiUt17d:·c~eD less·.. tban ;DM;.,1~e;··the·dependenoe·can"betoUnd';

.h theequat1on:~D~:;'~'·A-:+B•.~DU~:(ri~O) :<: :(2).···,'··;· ...... ".'~""'"... ~'::",'::.:.'

'.. ' ,: ··l'or,·.th~"'~je~~nien~:~t"~q~tlon'param~t0r3'"(2·)".t~~·.~t~:on.Te~·:':. ti~"·~~~91J:' ~.·19G4:~d··'1969:"e;G·aJmi.:raec1··'l:· th~~·m~~'.d1ameto~·,,/~::·

• • c, ,'. .." '.::, '" ", ,".,' • ' •• :' .....;_ .• '. " •• ' • :.' '. ~~.' :'..... , •• ," ".,: ',. ,; ' •• ,",'" ~ .. ".' ," 0'":· of: egge 111: each1Sater layor. W:UJ'~determ1ned 'and was; compared vith . '.

'~'_ter 'a~1t7 at .~1i·e·:· d~~th'; 'wh1c~'18':'the' ~Wi~e~ot·W.a"liqer~,<.·~us~·::·.1f~01nt8:t~~·:Bomhö1D·~;11 ,~inta·ior:aotl~1rer~~~to~·,

..... ·Ddned. ~h~ 'dep~cle~ce~,obtd.n~d:'-1n 8~rch··~t·two·~~~e~·.(A··a~':;·.', B)·:.:~·thed ·~··~~·~.(c· D -,'~d "0 ;.":,-1'> 'lo~."l1Jr~ the'toll~i··.:.. tor B~mbo1m·-:.·DSTE.~: 2.0:'';''3.34. 106.. ~~j·.-· :-·"(3) :.,:'. : ":>... '~' ..

.. .'. .' " .'. .- ,.: .' .,' . ". .. . l' :., . ..:,. '. :. . .: . : :.'.:0.

• '; " .'.";,, 1)89,•. ';'11.61:+:.2055.6. :W-. " ":(3'),.: " " ~ :.' :.',: ;): .

, .. and tor Gotland . DS'l'B • 4.60·.+ 1.«*.'106* rot-:-' ,.... ,,'(4)·' , .::. . :"~.' .' . .... ." .. , .' '. . . ',' .' .' .... '. .' :'. . ' ......

'. D~B.••'.42.'" 885.95. m4 .. ,', ",:- (4') ... ' . '.', .' ..

·!rhedependencea' (3)' and'(3') p~tlcal1J' ~o1ncide'~·the·d1a~~""'''':the dependenccB (4) and.' (4') alao co1nc1de~ that:1s'~ tor.tw:th~,~:·

· ca1culatlons we'.ca;y tnk~' oither of them•. S1nce' .6.<0 haa.no ~'.':.. '. .' .'. ". . .

wo have chosen the f'o1-=t1ae tu and.(4)' g1.'9'en in F1gure·). ' .

:~~ dep~nt\enCeD f~r dit:rerent· areaa ~oD81derab17 Var7;" the d~·: .."'. .." ". . . . . .., ','

·pencltmce· f'or Gotland puses lowertlum f'ar Bornholm. It 18 .known·j. ' ... . ~ .· that '1n Gotlanc1 apavn1ng ground water ua11.n1:t7 and. water dens1ti·

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· are 1O\t'er·:(Pls.2)~·.that·:cr~~telJ ~rae' eoDditlo~'~~rcod egge ~de­

··.;"~lC1JfDent.:·:·.But':WejC~~d~~;;·;·~t·:tho.. pi.oce8BeB,:ot:.egg.d8Dll1~~~-·'·:.• .':, "". .:.', '... '. • " ...'.',.. I • .... • •. ' • • • '. • .' .' • ~...

·sule.t1an·dutl,to·the cbe.ugo;of.1ts·d1ameter.IIWIt·,be f.he:8&IU' at 411-'

,~~~t.·Dpal!lTlt~·g·\~e"::AP~t1:r.··th~re1."'8~'83'.teD*t10: emr '.'at ·dGte.~cm;Ot:Rter·;d~1t~·1n.~~~:·.(1~th70~~OD:~:'~eyi'V8re,~rrl.ed·;Cut.:.~~·b1::~~he::~ct.'&r~~:.~h~~·li;Vdro~o'S1cal.. ,', ~ .

..d8~:~e~ .~~el~ri) .:~!~t ;'~1~.,.:··~·:..further~ ~~uiät1~ 41rf~rent .,..'.. .:.... .: ..' .... :. . .• ........:... ..;..~ .•....> .:. .... ,'..;. ~.. ..... , .' '. -:....... ... ·depen4cmce8··DSD,:(m).:1"or>'tho:'c1ata:.;0D~·each::BP8u1ng~g:ro1Ul! ,wen .,

'i""~~~ ,.e ,<:":;'.:.;'i:/:<::<:;·:i:.,; \,;.:, ..''[ ';"'.' :-;;...'...· .: .. :~.. ,: !rhe.:dependenC8r;.DSD,'.Cw).':·are'.Tert:Dteep ·.~t.~101f:.DJI.'I:U..,·:.the ..,.~

": Ca)~~atl~·.:.~~~d·;·:"~~·':·~~P~~~·~~.;:(3)· ..:·~U.;::'~~l/~~::·tmi~t "8lnk~: .';.:,

·....•.. ,;0 .•~~.·~~~i.~~~)+i{4 ~~~~~,+ )~9~~~?~~~, ..t.',~.~Jri;"...•.•..: -(Dm:~: ccmo·.' er!t1CA1;;.Ta1110·· ~,;*,~ diGetcrr·DSD, .:.ccml!lt~.;.::; .:.,>. :;.' ../:..•.,'; ...

:". ',:.,': ....••. ,~'" •• :.,:.t'.'.'. '.,,,'-':'::" :~'. "'~'::'-"~"~'" •..••• .': .. ,.~., ~.' .. ,~ " ,', >: , :.{ ......•. ;:: :.. ",j ',:"!.:..;',::~'·...,...:. ~~ :. por.::det'''minst1=,;·ot.:döp'cnd~~c·f):'':of':.iius~tm\eoiul:··mo~ t7.,. ·c~ :~:'.

~ " ""', ', ··':.,·r:. ~., ,,-,,'. ::' .···:.·.1 · .. ,.~",;"'; '::;,,: '·, l •.•.•:.,,; '" ..,,' •. ~.. ~, ••.• ; •••• \ .. :: •... ".: ~ •. " " ", •.:;. .• ~,,' .'

f ~erf1cd..ent~·eD.::C%7,;cn\etmtoD~·:w:·:~,tCk8.:·thD:~f~li~··.~thOeCta~,:}\ ..~<',': . '; ..... "' .... ': ('" ':&.: ': ...., .. ::'z·o:-: .. ..~ ....':.,., ....:..:: .-:.:, •.., ......'.:.. ,.. ;.~ :,.1' ...._'..._~ :.",::,.,,: ..' .,.:: ...." .~:.:•.. ' '. I

, ': 1·~.A.t~: 1-u:mse,:of..·eggrr\doyolopmeDt :;(1·· .',1;~~4):" the .;·o~ott1016nt·.cf.::;·. :.: .. ..,:. . . . '. :':.' ", -. ,I.:, ':~ ':. " '••:, .:..•,"'~'" .. ' :. '.~...... ~ .',': :.- r .: ,\••. ~. ".; • , :.: : ~~ :...~,<.... :.~. ",\ ". ,t ~•••••.~~., .. ~::. '•• ~ -' ':.' :•••• : :::"',:' .. ,,: ... ;" •• ':~ ',. : "••,::, ~ : •

·; ....~t~~e.ou':J:JD<)~t7:.&?~~fJ.·:.oo~~~ •.~.:~.1~~:,:.~1U~··18::.det~.· :::,.: : :,.'.

·:.·zdned;··bi{:~~,,"~~t~·t~.öXD··:·:of:>~~~·::AP~nt1t~~:.·1d'th'::th~ 'inc~~ •.:".,~ . " ~ ", \ ....d : : .....; '....... ;'" .~~': .,'::: .:.>: '. r......, •• :- :..:' •.::.. ~.'.. ;.~ .",~ .'. ~~ ., .... !.:. '::< .•. ',:1.'.:,... ~~\: ~:\ •.... :\1.. ·.. '",,''' ':'~:': ..•..~: .... '. ": ,'. ','.' . .: '., ','

:.'o~.:.0ID,:·.~11·d~.~et"";~:: .~·~b1~·:~~~~:c~Jic~zitr4L~~o~)uJPi~.·:;tcr·~,,:,:.·• .' •••••' •• :. "I •• ··:·:~·;~;.·: .~,:.\•••.• :,; .,••.•••:~' '.', • ":.',•. ,:", .,.;::,••••:,.~ •.• , ,.'",:,." ~', ", '':'::''''' •• ':' ,.:".' ••• , •• : ••••••

: ·,.,omo·:.ccnzst=t.'.'YfA1uo··.AJ4~·:s~.<n·:;.hävGjdete1'idn04·:".'tho·.'4e:peDdaDe.:·....·····.':...

·.'..•.21..~~~'~i#1b#.7'~.'~5).. ';;~ '.;;.)l,::· ;~:':p ~':\\;':' ·::/V·?}.':·:;'··.··f ..;..~. '.:;··.~~'f&.::~d·}l~~1'~:'F?<~:~~~~~.~~~r;:f~.~a,-;:~.:~~~.~o~nt.~ /:.: \' ~'.,'.: .•...:2.It·.;·1~ ~i:ot .):ZlCwn( 1zi>hO;;~z:niCh:::tim(" .:-I( doad ~Gßs·::lea ...ea.'& ;.'S1~cD :h~' ....;...

'. .: '•• ' t • '. ':" ~. • ", •••• • ••' .. ". '. • .' ••••, .':. '. ..'.' " , • • '. ,. ' •• , •• ' • • ..:. _ ••• '

\-riUon.· :\'Ie,ouppQGo.·.:th&1t:··th1.·.t1mo l.Can ba··.c~d·"1th ·tho: dun- .

· . ':t1~ ,~t: '.deyel~~~t, 01;Äse~~ ~~·~.:·d~ta~ri'~B ,':tho' a~: of.' Gmr'4~:·. '.'. ." '., .... . . . ' .'. :' . , . . :.. . ..... '.. .':",' . :.:. . .. ::., " . .'., . \:. . .. ~

. "Ielopment;"'1'lO'do :Z1ot.'Dpec117'··1i'1.t '1a ·at 'tho··be8:1nn!ng·· O%'·at· tbe: .. :'.' .'ond .or· t~~';:8~•.: ~~ 'oCma14~;: ~t '~'" öt;· .th~· .~. '1~: ih~ ;Jd'441~:' ... o!' adoyc!o))inent:' etage'. ~.: ~o·.· the' rat~~·,of IlnliB. egp' ~maberXA ·(~.·k) ~t'k-e1Se ..~p.at .1~tase·.ot .. 40"010~~t to th•. e.. er .

• '. .' '. ! '. . • • ~ - 1 .'

.~ 11V1ng. anddead egßs.ID. the 'eamplo "clu&raotenaria egp ""1"1&1 .',·:ttor·theper1od ~ER' U,)/2;" vhe~ PER" (1) ~. tho durat10n o!.1....taa8. . . ~~, .

6

. ~

j

01 dnelo]mlent~· and Y.D (1;'3) - the .n~ber of ~ead ee.gs in & zsaeple•..

.help cf the rCgN,seion Xi ."ertnl8 ·1/CXD tor .~e .t1nt threa 'staeee" ~.. " .,

..'

.', '.

. .

So,.egga 'BUrY1Y81ct k..-else grouV. at ..1-8~eot deYelopnent 18 :

S ·U:~k) ..- (Ut1;};) I. CU -( .1',k)+' %D (1~)}')2 U;)';. '.

and the1r 8UrT1"a1 dur1Dg the Who1.e.1ncubat1on per10d l' in :

SURV(k1 ',. n1:. ,: S ,(1-_k) , . ' .. (7). . . . .

. . ' Uean em,: 8ur.n1va1 .. tor ~the. ,.ear1s •. calC11l&ted .aa .meai.·· .e1~·ed "... of ~a1ues''mmv;'(1)·.acco~·:~.~~ 'nUillb'~~ .1n' 'H'cb ·'-81:0.~p'~f" .. '. ." , . " .' ,. , .. ;, ". . .. '; .'

.. a.aample·.. ". .: .. ..' ,,:, . '.' .:., ,,' ;.... '.,~ , ' ": -' '.':.

. ' In -.that .. CA8e ·the 1D8tantGneoua: mortal1t,' coettic1·e!lt. Bi" cl"'" ....· .. 1-8tag~'~:r de~e,lopment;i,j: o&1cUi~te4·.· tor cach ·kb;.·.·~~ tO~ll/ .

,',B (~.':k) :•.-. C1iP1m (1).~·;8 ü.~ ~k) .":'.-.>"cEn .. :.:, '.:. . , ..~3~!rhe 'd'~pen~'eD~e '6t .. e~··1nc~~at1~n·perlOd:. ! .~.:tempe~ture of th~', _tu iqe~~ whe~e.e~··de~~~p; ~:1•.:d~B:~1beei::~.the·tlqW\t~·~.or-.:. t~red:~i·'Z·.~~~~:·:~d·:I.H~~·:(7{:~~~\~tl~t1c··'~::.::.:':".>'. .. . ','

.' . !f;'; 1;~,~.3~ezpC,;o.21~#o\~" C9~~ ,','):> •...... : '~enn caruS1der~.thatdumt1CD.: c>t. eaCh.lltage: of dGTelolZent

..~ i..~ ~·~f.~· ..··n~t· ci~~~~ng·.~.t;P8rit.t~, ,<.~ER·.,u3:~ ~(l).T." "

':; :.s1n~e;.there.ä~·:~o·'·~ta~Eai~iO·.cod·.·~eD~~~~'the.dtiratlori .':, ot'deT~lopm~t:-a·t8ae·~~···'" :tälCe;ih~'jalUGS .~t.':co~t.~1ci~t~·: g{~')'~ . ....

,:O~t~8d':1D' :the"~~ent~",w1~h':~~' e~"ot' ~te"S~':a~ At' the···

.. tempe~ttire' 6~O·at··I~xV.,~tag88 .~t.~·eT81~pzent. (8) :..... ".. <, . '.'.g.(1r.~ 0.16;,. g~2).:·~ O~~2.~··~·tn~:~ ~~~I :-~(~)':;. 0.21' "i"" .:< ..

flle pam:ametera of the· dependenoe ,(5): 1'r82;'e. obt&1ned ,d:th tha. " . ,

I'

IriI

I

I

II

I,

'"ri

o~ de..,.elo~ent. Por thatOurth u·tage. mortaUt718 pract1ca1~ sero. '.

Per eac~ 7e&r 'anti ~ch ·"tase ~.d.,yel~:tt "f: .baye got .~ot lm~e' ~ .....

t~ 4 po1nt~ '(ao 8;" 11'8 ~cÖn.dder thec1a~~D 4 ~e' groups' ~t eE8~)·~.~ far~ T8&ra' ~ leea;becaWte .the pointe 11'1tu eggs' n~b~"'" ,.,lesD' tban 10' ~rc dtc;:mizure'd, ·or W1th ~ggtI ~orresPond1ng'to elS'Ei

. . . .

graupe wh1ch can not tloat in the spaw.!ne '1a78r c!uC!J to not p1"Cpe~

wter dena1t7 thie' 7oar.

------ ------------

~'.

.... .......

I

. ." .

... ',

,

L __

~he:re8ult8 of ealculo.tioM aocording to the date ~orthe .. I . .

. .first per10d or ,.aars are giTtm :1n !t'able 1 and P1guree4. a.nd~5.'

'. '. ~he ·P~~tfl., corro~ond1ng' to the aeocmd perlOd of 7can are .plot..,; .• '

.ted' b ·tbc tield otthe obta1ned curYea. AB we. can 800, tbc points

;o:ttl:e ·.ti~t ~d tbe .cecon6.pcr1od.. or~Mrs are .D~ttercd.a.pProx1­

.;mste1y~dent1eo.l.1.J't&md.the acatter1ng .181arge. '1'his 18 causedb7

': the ::f~ct~··1.) ·that esssdie not· only dueto: Q:ygcn d:~f'1c~enC,... . "" . ." . . .

.:2l·~I.argeseattcn.ngifJ alDo cauncd.by the ~act that bydrologlcal

·,ccmditiona;·1nelu~.c:ygen:conoentr~t1on;'aro' eotimatod at Btan~

. :-·dard atat1OM;' ~d 1neCQe,y~~ xnay', not co1nc~d.e \rlth real ccm..," . '.

··:.d1t.1.oJUJ'ot egge development on. tbo apnl:n1ng·ground.· .

:"·3j.:~o:tb1:rd.·::and·poa·s1b17 thQ:~reasön is tho ~~~t~ that. the e·~.::~::~~~t~.ot4CPen(loncO·Of ogg dendt7 ~n'ito' diameter trere'not ::: ...

·:·.c1e1:am1ned·· ,correct~. . .....>4)..ne data' on 'cU'fdseß1'U~'ootoggo 1» thc .~e:e>endenoesU(OXD),.~'ccribined~;"::but in f~t; for mtsl1·egga.highcr mortalltY' 1s '

. . ..:·~ater18tfc·.th8.n·f'orb1g onoa.,:··'

....~;.::\'; After "ccmb:Sn1ng·thc da~ ~or -both pcr.1"X1uthccorrccted dopen­

:: dences J4 (OXD) ',~~rO o~Otllated (Taol.. 2· tmdPlg.6) .'At· 'e~r11er 8~-,·geset·doytilo~e.nteu;s mortc.llty :t.s b1ghör. O.%Ysenco.D.Centmt1cm .

, : "',. ." . • ~. > • • -

.~18:the.:11m1t1ng fact'or .tor formation of thc yc~cl6DS atOXD

..le86 ~tb:n 2 'ml/l.

'ConolWflona;'.. '.. "

<·;t·.!fhe.dcpendonccs otwummod egg dena1ty onlts diameter "are c6l- .

<~·"cu1a.tcd.

'2~The depc,11dcnocs or egge 1natu.ntoneou:; mortalii;y on oxygenconoen~. r~

·tro.t1on fo:: each etago of deTelopmcnt u.nd caeh spemüngground

·'.Ware obta1ned.

)~Uea~ dlsmete~scf' esg8 in each 81:e group DM(k) tor tho'respec­

.t1ve·perlode Q'Cre detcr:rlncd. : Por Bomhom opt1mllng sroundDM(1) •.

• :64.25C6 .1.27); DM(2) 111 68.27 (0 lEl 0.85); D"J (3) .. 72.28 ·(6.t.1)'

8,

I

I\.

i

'DM (4), .,76.70 ((j'. 1~42). Andt'ar'Gotlo.nd op:mdng sround,mr(1),'~', 64.27 C6. 1.1);,Im'{2). 6S23,(D.t.4)';mrc:n ,.72.3S'C6. 0.(2)'• .-

DH(4) -7b.8 C6. 1.27";: ( cf ~ 8tand~ deT1~Uon er mean, d1e:meter)~,' .

c' G) 8~tter1ng '10 'Do'negllgible;·that'~th~re'18nO.ne~d"t~Ca1cu~.:::':',1&te :'m! ever.r7w.,assam1ng,1t--to be ,oS .:eh~above~

' ..4.!ho· obta1J1od ab,ove d.ependcmCOD·mn ,pm:mt.'1:1 .fumre :to changG·:"

the ~ethcda Cf' 7ear-cln~'atrength7prognoa1Dg.·It.1D':im~,;,tha~·.:>:, -ratio ct.big and' f:n:~11:egga'in ~'ef"004' i~e8;:~1','d1ffe;et··: /.

. ,aga ~.var1~.jt'!B-J1scesca17 ;t~, ~voat1SQte ~tiie; cU~trt~t1on<. ',,><" Oi,~tcd,'eggun~cr-~~·,tO',d1C171~ter':1ri'-fi~"ot:,eU:rt~i·,>,'

':, 8ßO.''-AP~~, • .-t~'.eU.a~~t1~:~O~b~':~~~''~cc'~~,:;tO:'',:··.

~ .fOr. ·thc.tJCO-::oge;:;~ '~.:~~~ ,·,~~'~t:';'~~:~~.',:c~\::::':·:·" ': B1t1onot'::the·.otOc.kthe,:ratli) :Of',b1ß'and.:cmn1~"eggBt·.aPevm.ed.'b7< :" ':,'" thewhole.popuiat1~:1r111'~tl;,~•• :kt'.~';tako'll(a);.~. f~~~':,':"':,mmber.A'(al"-o.bBOlute·1iul1-v1dlm1,t-ecimuv"at,:~' ~a~:~':,iuul'·;n'(~.', Jd~

. . . . ,..... ".' '.,. '. ... .. '. '... . . - '"

. ".part'Of',~ggs·ot'.k-6g0<,grc)~ i1:1,~a·:age t'eica.le:.s•. 'S1Dce·We :'" ~ ,<". . . .. .. . ....

, ·:'know "erttca1':dlstr:lbUttcm cr''tm::!pGmtllro.·;S811ntv, ana'CX7«d foio,

"e., gj:rcm")Fea%", '~8·tJa7'. ~u1Ato'~ '~oem~~t'CJ',,~C~g:'~, Otlr,';,>., ,ma~oda."· and, then' detem1nO-eggn curr1w1 fm-c each '81s~" group .ot': ':"

, egg~' Sk.' So; :tho.number,'orWrv1TOd:'cgp,in,t.\ g1~en7ear,:(~8'',:tnda%)·;·18'ca1cuJAted,b,-·~·~Ollow111g:;·:r~',:·.', " " '.";::'..,;,.';,.:

L"~ :La;S (a:)~ (a):.~· (a~':k) 8,'0:)., .' ... ', .,' ,.... ',~'• e bopst ,that theoo, calcu1nt1cmtJ 'l:J.rq change co::p11cated ;rear17 ,'.

:.'8amp}ha,'~ :~oce~dng"o1- .cod,·' Cg-gD aDdw11l':~ :.th,~: bCa1lJot'7~,: . '

,claas :otreDgth"prognoBinB with'cdvanoe o:t"'3'7oars~ ,. . ' .... ..

9

.... Referenoes .1

.'.. ;.,'''- c' .'.

:·,,:,b-udy·naltNDiO•..IsB;7~·.. p"10)..

'·::2~·G:rtlumsn·G;B.:·:1967.~~hctora~detemin'ng.&.1tic·cod :·abundance;. '"-

•. ~ ;:~·b~·nauc~o-teem.,:·intcmriatati.,.vmO~·~SS~2e· ..

.,:'3~Graumari·'·G~.;i·,1969~·:~.ne·~~Ort~C(f:·~f'J3aitic: cOa.·e~'·81se:f'or ':"'. . "," .:: .' ..,: : .. ::'. ":,." " ' ' ,:-:. . ' : ' .

.';':'':<'k emb1708:,.surn~.: :-'~'~~rud3" ·~AtlantWmQ·;·:':I8s~ :'XX:r; ::1l.86"95~ "

;'..·~~~~~DU1n,:.a·.1l}·~~9~;"':.~·~~~~ '::~Us~~ ·:ri.~ttiatlon~· 'b;'~ ~od :~dspi-a~'

':;~;;+~;~~;9~j~:~~~~!~·~~:§::~:;t;c:c~;.: .~.····>';:.·FGar-Clus>:ft1tb :ad~anoe.cf>3',earsii,:""...;Sbom1k. n6Uchrqkh .trudo... :'·· .

:"<"~'\~~~..~~~ ..i/{ ':';i.,"i.'~!'·'; .}:';,"':., .•.....<'. •.•.' •...:.;~'~i1boy~n.lJ~'·1.957.~·.Oceano~rh1eal..':tBb1es.,:-,JJ;yd.roJüt1saat~"'406·;p.· '. :.

"'.. :::. "~ .. '" .' "'.. "., " ~ . ','.. :. \. ; .' , .

',: .;:.7.m8~~~·~~~·~:"B~~:;;j:.~.~~74~~~J.eiu1.'~~·:'·~~Ography ~:..:;. ..H1~~t1.i; . :;.

)·::;cI&t·':*~f~~·.~~ ..··••··'····.··;:.';:··;,.:.'C',.;,~ ..,•.•·...,.:'"".•. >:·, ;::' ...... '•.•.•..:.•..",; e.l4akhOtin :7V.v.·;;·~~ ~c~ o.~..:fiJle1lco :)r•••~>S01r1"~S.G~·::19B4.'.~S0m8'.' .;....

:,;~~~~~~~#~~~~:B:l~~;~~:~:~~: ..:<" • ' •.~# : ....

•

10. I

I

--, ---'-- ---

,!fable 1.',

'Parameters qf equat10n (5), and 'correlatlon coeft1c1eut3,~t­

'ween ll, and ,1/OXD ,acccrd1ngto data ~or 1'.aY1958,'196O-1904

and 1967-1910.

0.27'.

',' 0.. 69"

. 0.&1

, 0;'70

o.~ ,_e.52

.. 0.63

, 'Co!TelationcCie:r.t'ic1e~t ', ' R

,.....

, ,

"

0.3&

0~29 '

'.. ' .

·0.37'

0.12

·",2.11' "

",2.60

. "'0.)8

, '0;'226

.0.98 '

"O~14:

, ''. . .

,:0.05

;-0.02

, '

'0.34'

0.32

',0.06 '

.:' -0.005.

. ,#'

,'1' .

, ' ~.

'1 '

:'2, '

>,'.3 '

',' 2.

','1

2

"

'. J,

,.~ Stage cf Parat:2oto:n, 'o:t· eguat10n ,(5) ,,development ,'::.A.U " ',',,: ..': ,lm

.. '

" ":

Bornholm

:Bomholm'- "

" , ..

I

I

~.

rII

III "'::', 1'able:,2. ' .' ,

: ":ParameterS of ,equat,ion ','('5) auci ..corretstlon coet1"lc1~t~'I

.',' ..I '. "":" betWeen 1{ Eind 1/OXD;: ca1culo.tCd nfJ1ng'addlti~: dataror :

I ..' .•..• ';I7~'97e (for ~~e :rearo thl>r~ an. da~f~r l&>7 ~~unej·.

'Area' ", stage of" ',,:" Para.met'cm, ot, öguat10!1" (5)" ,acrrelation ....I ,.!developm~~ : ~Dl",.: '. ':', .':SB. , " ",:: _ . ,coe~ic1entI

1 ' 0.80. '0.49

",) Gotland . 2 .,0~11 0.13

') -'..0.01 0.17

11

Pigureo .;

,.Pig.1."Verticaldiatr1butlon 'of' aaaur::edwater density (A)DENSW .

'. ard. o%yscn concentrat1on OD CB) 1n Dornholmarea (sta.-

'. :··<t:1on.~2) for.any4··Yoaro •

..:Pig.2.·Vcrt1cal distribut10n or aosumed.watcr density (A)DENSW

and ~gcn concentrat1on OXD (B) 1n Gotland area (station•

. . . .

.. _" P1g~3., Dependence of· egg dena1t,- on :1ts diameter sepnrately accor-'. , \. .'

···'ding to' tbe dato. tro!l1 'Bornholm .Ccurve 3,· elrclee) ,and Got-

. '... ·· .. :.:lancl Spawn1ng'grounds (eurve 4.' oqunrea).

·'tfig.4~:,DependeneoD·cf egg ;!natailtaneouB mortal1tY" coetnolcnt' on .

-. ·.-::.,~gen:eonccnt:rattonat .Bornholm apawn1118 ground; ·calcuia-· ", . ".' .,' '. ." ",' .'.

::.,~ed·:accord1~-tc the dct".:r~ 1958. 19&1-19b4. 1967-1970:

;",a)a~;"thc'f'irnt.otage: cf devt'lo~ant. . '

:'\::~,:~:b,:;at.~'tlie:'DQCCndat:tge· of devolop:1ant.·

,',,' :' :c);,at'tho·'th1rd 'stage ot':devolopment.·. '. . .

':"::'::9~\cimfidence ,l1m'tn aro'·ehade'd. crasees cerroopond to .the"',

: ·, •.'i ;:,,:::'·data.~fer '.;976-1978•." ' .

.., ,P1g~5~;·Depan.de~ceG' o~. ~ggS natural mortiil1ty on o:z.,.genconcentm-:,>.." , ,.,,:.: ' ' . . .,". . .'. ' .

·~::::::t1on'atCotlr.md· opavming' groundlJ'calculated accord.1nß'to .

. , .. ~,:::.th~da~ 'tor '1900-1964~1967~1970. .e;Des1grw.ticns are'.tbc< same nt11nPig.4•.

. Pig.6.·...Dependoncce.JJ/0D)·i"or dU'ferent stages cf. developmcnt~ cal.

,- ..culatcdinthe same ocale and corrected ac:c:ording to the '

:d~ta: fer1976-1978 I,' a) for B~rnholm opawning ground.

b) for Gotland epamdng ground.

/

, 12

•

13

4

.,...

t3D.110 t2.D

H D 1'190 .200

fig-. 2.8a

O ............l...Ji-l.....l...J-l.-.l-l-:....L..,.J....1-.L...l-:...L.-l...l-J.-i-...l-.:.....J.:::L.!-J......\-LJ

itJ

4

2.

e ~ .. BCI)

Z~ 6

II •

Fig. 3

· . . . 411.Flg•.

.... -"". '

15

" ..

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