--- This pc.per not to be cite<lwithout prior reference t~ the n.uthor

(1)

r - - -

-International Council for the Exploration of the Sea

.'

This pc.per not to be cite<Lwithout prior reference t~ the n.uthor

CollI o 1969 Specic.l Meeting on

''The Biochemcal und Serological Identification of Fish Stocks"

•

No 11

BIOC~jICAL AliD SEROLOGICAL STUDIES ON ATIJu"ffIC SALMON

by

(SALMO SALAR L0 )

-•

No P0 Wilkil1s

Marine Laboratory, Ab erde en0

(2)

. .

~l~CHEH~~AL'

'.

'~~D'

.

SEROL~~

lCAL '.'

S~JD'i:ES .'~

air' .

~

:

~ ,-~."

I : . . . : .: . '

:.~'

' . • ATiJillTlC 'sALMon (sAiJ;m SAtAR'L:).'

:::::':::::~'':',,;,,., ,.,~:.

.

-

~.: ...-:.:~' . ~. -::".:..:.. ~. .: ::..

", .'"~ ~. , •. .!' . . • .' .. , . . . .

• • . . '1

..".;

by

: ·I'··~"· ' . " " .::.'"'~..~'t ...; " ! , ' .

, .. :. .; l'f~.~ ~.,td..lkins·. . ., : , ' ..

Uc:.rine Lo.boro.tory,' AberdoCll;. .

. '. . ' Scotlo.nd~·' .' ::;: ..:.:-,>'- ;

I .. " : . ... '!.. ..J ;-', •:r.::: .. ;' .- ' ,

.

..

' ..

:~

.'

.

...""

•

, '. n~RODUCTICN

... '. . ':- .• " ; .. :'.' ... ! •.:" ;c-,.. ... " ...' . ' l . .,.

. . . ' ... .. " ... ; . ' . . . r " • . • - . . . . : , . .. ..."1 . - ." • ' . . .

Since1960....0. 'fishery for·Atlo.·ntic so.lIJon (s·o.1oo salo.r'L.) ho.s dovelopcd '.-off thc west eoo.st of Greehlnnd from 0. toto.l·o.nmmr-eo.tchof' 127 'motrlo tons ..

e: :.

in 1961 to ovor 1,500 ootrie tcns in 1967, o.nd, tloreover, ho.s e:~nndod f'rom an entirely coo.sto.l, fixed, gill-not f'ishery to include a large off'shoredXift not cOI:lponent. The o.nnuo.leo.tch in the 9f'fshorc fishery has grown froolcss tho.n 100motriotons to ovar 500 oetric tons in.tho·yeo.rs 1965tci 1968.' . : . : .

:",,".~. " ~.".-:: ,,' . : ' : .~ " , ~ [' '~ . . , ' : " , ~ • ',~'...: ,~l· :J, ' .. :';.' ,

Only one river in \7est Grcenlo.nd is knov.n .to ,support a spnwning:populo.tion .. ' , of' Atlantie .salmon, . und itsccntribution.to'the fishcry, 'while not acctirci.tely·~ , known, co.nnot ccnstitute o6re~thil.nn"minute fruction of'~the total·catch.·' Uore'·· signif'icantly, thc rceapture';' in' thc S00.' oft"GroeI1l.o.nd,·, of' salm:on' taggcd '0.5 ' ., . smolts in thoir rivers of origin, indicates ocnclusively tho.t s~on'originating

in both.NorthArncrica .(USA,and Co.no.da) ,o.nd, Europo. (Icelo.nd,. Sweden.o.nd the British'Isles) eonstitutc:the

Dajor

elencnts ,in this' fiSheI'jr.· .: .Tng6ing in ·the i1est Gre·cnland·arco.'husj .furthcmcre', "estnblished tho.t 'sone'ofthe so.loon ~ :.'., present in thc:.t areo.· return to the spo.Vming'pcpulations in Horth ..Aoc:dean and' ,": Europcan rivers •.. (lCES ·1969) .' ' . " . ., .': .. ; ,:.. . ...:.. ... ;',:.',,- ,',: .

.

.' .': '

..

" ' , " ' : ' , : :

-/::

,....

:.'

" .~'.

--

.. '.:-'.':

.~".::

:.:

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

...

The identif'ico.tionof saloon cf dif'fering geogro.phie origin, and 'the:', . estWo.tion of thoir. relative o.bUnd.."l.nce~-in .thc exploi ted' st'ock off West·: .': .';," " Greenland o.nd in other, S&"l. fisheries \7hich hnve 'rccently developed in the' north- . eo.st Atlantie has prcsEmted'

0:

problem of oajor ioporto.n.ce"in thc ass'essoent ·"of :. the effeets of this fishery o'n the saloon stocks in homo'water::i; " : ...

, . '.

--..

-.... '.. .,

-

' , . .

•

AB a contribution to the study of' this problem;' investigo.ticns .'are'in : :-r

progress o.imcd .at cho.racterising, biochenico.lly and serologically, individuo.1s of those spo.vming.populations known-'-to'contributoto the West Greenland.fishery. This, report,reviews 'so!ae of,'the results obtained 'When'n~nuober of protein .. : '.,

systems in salt1onwero o.nlllysed for p'olyrnorphis::::l using~serologieal imdbiochcoicoJ, techniq)les~'·.. ~ ...:..~.: ~... :::-

":',::;2'

:~

,.. :

" 1 . .':'!.":. " : ' ' ' ' ' ' , .. : I . . .

_ . , .• - .... -\ <I ~. ~'... •. . . : , • i .a:: . '. ,~~ ..L..- . : :; "' . • .".,e ,'" r ' • . " ~,~ .., •

. i : : . " , .

:.:~·':.'~:.'~·::lMbRiAtS

·Ä!m·:UErHOOO·,'·. . . :. , ... : ...

-

... _. . . ' " .

--

.-... '_:';J . , ' ': ~ : '" "', .; "tor· .. . "," , •. -'.: .• ", ...:~'.: ~.~ ,~' • . _" ~:: ',;-" ~,.}j.~... ".

SaI:lples 0:'·blood and other tissues ware colleoted f'rom samon froo: Cano.dD.; •. off West Greenlllnd, Md fr60 vnrious Europenn' cOUntries~: .... ; ...

• . • - .. .;~ ' ...._! ". ~ ..•.~': " . . . ...~ . - . .. ,

.. :..'~~, :'., ..",.' ...

. Details·of.the clectrophoretic proccdura and tissue cxtraction have been deseribed else'Where.(~ilkiDs1968;:0.'and b).·· Precipitnting and·ugglutino.ting anti-sero. ·...ere produced in robbits'.by in,tro.-ouscula.'r o.nd introvenousinjecticns.

: .• :, "':"';0' " , , ' : ~... " " ' • . (" ,'~." "':,',: •.••~' " r ' .~' .. , , : , . . . ; . . . ,,:_4, __.

>

,~.' :,'.1 . Except where ·indicatcd otheoos'c, 'the'.tem 'adult ·sa.loon' ;i8· used:hero to' . indico.te, individuo.ls. of~cver.50'cm, i.e~ .both~'grils c.' (1+ :3eo. rr.i.nter' saloon) .. and 'saloon', (2+.seo. Vr.inter,so.lri<?~)•. ,

.;,1.. -, '... ;:,

'. :....

~~::.':.

: .. .

.

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

.:-, I _ ~ , . • •; • . " ""

," ,., .. ~, 4, . . . .~~-.I':... -: I: ~ ..:-: ...~ f::.:.,. a

;' ,.'m:SULTs

. , 4 . .

,

.

~,'~~- ,

Hnemoglobin

:.. ~ ~',) ...''''- , . '

.

:- Thc'ha.emoglObins~01'. OVer.SOO. indi~idUii.l So.ioon' from.:Cano.da, Scotlo.nd J

and West Greonlo.nd wore nnalysed by starch 'gel electrophorcsis.-.:: Considcro.ble ' ontogenetic .variation "ms·.obs erved 'and described (ifilldns 1968a)~ but· no '.:'. '.' ho.cooglobin, or c?nplex of)1.ucnoglobins ,', 'Wo.s Ob3 eryed to . cho.racteris e.· . .': ..", '

exclusively the individuols of' o.ny one'area~' . The po.tterns observed did n o t .

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dif'f'er f'rom those obtaincd f'or Scandinllvian sllloon by Koch ct

.!l:1..

(1967)

or f'o:" Illndlockcd populations of Nor'l7cgilln salDon .(Koch ~

!:1;;.1964)

Ilnd Canad.5.an salr-,on (Wilkins, unpublished). Throughout tho 6relltcr part of' .its r:mge, then, the sa.loen exhibits no va.riation in the nuobcr er eIectrcphoretic nobility of' i+.s ha.emoglobins.

.

(\

•

Tho polypeptide chain structures of' tho oultiple hlleooglobins ha.vo nO\7 been ox.a.minod. Scottish, Cal1.adian and Ca.na.dian lnncllocked soJ.r:lon all exhibit identical hllemoglobin sub-unit structures' (involving'S'distinct polypeptide

chains), f'urthcr conf'iming the absence cf vllria.tion bet'l7een different popula.tions. Intorpopulation dii'ferences in the'ra.t~ of'·ontogenetic development of the hacooglobins are evident, but these are not sufficiently pronounced to give an easily usable. index. of"-popula.ticn miXirigin' the Uest Greenlo.nd area~' .' .' .,'

• : ,,;1t .i. .:.~:. ..~,-.. =.. •. c·'~ ~~: ....~. . ' "" .., ...-.... ... ::: ...6& .. _ . . . - ~ .. • • • .. •

R d C 11 Ant- . .-..~-' \ ..,"\~ .~~.. ....: ... : .. f" ,~ I '

e

e~~.,. ::;~lg,e~~~.~

..

:,-;~:.~;

.· ..

>':.I:.:)~.,.:': ·.:i.~ ~.·.'l",'~'::;"~_:"~~'

'.:

"'~ .~

.. :'.

PreliDinllry studies esta.blished that .the nomal. serum of CO\iS and· pigs ..~· .... :

Ilgglutina.ted the erythrocYtes 'of' 'o.l1'sali:ion' tö 0. titre of' 6L.-and 32'respectively~;':

Cells :of:Canlldia.n f'ish exhibited

0.

:wider, range of' reaction TIith these sero. thlln did Scottish 'fish'(Wilkins

'19670.),

'thus indicating Il':degrec of Ilntigenic

<. :

dif'ference betvreen the 'Scottishand Canlldian·pcpuIc.tiöns··tested.·· Cells :f'roo ' .. '~. sllloon to.k~no.t i/est,Greerullnd exhibited·o.··sioilar'range··of'rooction to the {:,-.:'

Cllnlldian'so.mples.'r-·.--~."; ' ..1. • . • : . , ' . : ~.: 1 " - : ' " . ~ ','::,'<1 . . ~

, . . ' v ' . • • . • . . ' . ' : : :, ~. • •1. ~,::. : .•. • • • • . __ • • • , . . . •

~ i:· .~ .. ' ~"," ::.~ "l , ' . : • • •' I . . . . , . . . " , ' ~ • 1 . . ;

,.II:lI:lune agglutinllting:cntisero to tho erYthrocytes :of o.dult:(2+.sea:TIinter) : Scottish saJiion 'were pröduced in rabbits; "TIith ·titres· vcrying··f'rom .. 32:.to'5111-;:' . according:t'o:',tho' injecticn' schedule:f.ol1m1cd.t ·"'Yihoo"juvenile:Scottish slllmon ' ."

wero Ilnlllysed~\1ith the'se' unabs'orbed'sera;~the -agglutination' titre' of·.thc. cells' .. of one yellr old pllrr "IlS significantly lor:er than that o'f' 't\10 yellr old soolts,' ;.': roared under,the sane,conditions (Table

1).

,The titres of c.dult slllmen,

however, were not' ·sigirl.f'icllntly. dif'ferent froo those' of' wo year old smolts. Abs orption' cf' the'_{. .} _{. , . . ' .}

iI::ourie

_~ _, sera~_~ \1ith oucus 'or s_. _. _{• . •} _{I '}

erun'

_. _,.,of purr res ulted' in a slight . :._~, _. _• lewering of'.titre,· indicating _that some," at least,' 'cf tho antigens. detected',: . r:.ay be coo.ted on to'"the cellduring. the firSt 'je'ar 'cf life. All subsequent :'. Ilnalyses were theref'crö "confined to 'Ildult 'fish cllptüred.o.t .\lost . Greenland or ... cn entering,their spawning rivers. . ,

'. r ... :.: .',~.. ' • ,':'. r·-· - I" • . - . t.! " , ..':: ;':: ..~ :.:~ " . ',,';' ' ' . . . : . - : . .', _.';.'/1."') :

... All·.the,individuals "test'ed 'rellcted pos'itively'l1ith .the· unabsorbed 'serll ..~. -.' Ilnd absorption nith. evcn .the ';leakest~.ce11 -rE'sulted, gerieraIly, in'the ': . . .... ' •. , eliminationof ·rellction.W1th"llll ether ceIls~ '. A nimber of'specif'ic ;reagcnto:· ...

wore, however, success:rUlly pröduced.·

by

'absorption'TIith 'the' cells of',saloon . .' : .. ta.ken at West Greenland. The proportion of individuals tllken in Scotlar..d and' ._"

in West Greenland '\lhich reactcd positively with theoe reo.gents are shovr.'l in • Table 2. Uany of' tho rellgents indica.to-sisnif'ioant differences between the

sllr:rples from the t'l70 Ilreas, but'; ·iu'the··absence of suf'f'icient dnta cn Canadilln so.1oon, Ilnd. 'on samon from other.-Europoan countries, it is not yet p05sible to indicate with Ilny degree ,of accürucy tho'.rela.tive.'proportionof'·saJ.mon of'··" Scottish origin in the West 'Gree'nlnrid f'ishery~"'.. -- .' , .';' ... . .... '.; .'.: :!.~

• • • '.:- ~...: • [ ..-;'~"~: . :~:.: ~~ ~', ._~.; ~ , ' . I . . • .. • . • , . . , ••. .,

; :The serological:opecif'icities of' ,two of' the reasents listed'in Tllblo:2 Ilppollr to:be :relatod.·..L In both the .West .Greenland. und the Scottish material,'

tho factor C is present only

in

c·oobina.tion \7ith B,; o.lthough 'f'llctor'B can

,:<,

cccur ll.lone.:" Thus,:.the.llnti-B ,und anti-:G reagents r:.n.y represent sub-typing reagents .or, alternatively, ,-the. ant'igemic fo.ctors·B und·C.togo.ther .oay.foro Il distinct blood group BC.' Th'e 'occurrcrice of' such(pheriosroups~containinga ., nunbor cf' distinct antigenie f'actcrs y:ith overlappiIig ··spe·cif'icities could

Ilccount for both the roactivity of' 1111 individuo.ls \'Iith the unabsorbed sera, Ilnd tho diff'iculty ex.perienced in the pr,~duc~ionof absorbed rea.gents.

Seruo nroteins

'" i .,r. ..

In a. singlo test involving 22.Ca.na.dilln a!ld 12Sc,:,ttish

1l~'.J1.t (2~-'~~1i

..

~,

'--- .. .'

r.'inter) sllloon i t ~i:lS pcssi!>le to di5t~g~sh'indiyidua~s-fror.! the.two.arcas by the electrophorotic pattern of',their se~pr?teins; Ilnd' also by their ... reaction '\lith'imDune ~rocipitati?go:ntisern: in Oucht?rlony ~ouble·diffusion : _ plo.tes (Wilkins

19671» •.

:·-In t~~. CIl~lldilln'saopl,e, ,Il ~;inpI~ p'olynorphis~': " .

(4)

involving ~. codooin~nt.. nllelos'\1ns observEld e.m<?~ ~o?~ of, the iron-bincl:ine (transferr:Ln) prote:Ll1B., The frequency. of ,the c.llele governing. thi)·.fastel' : phenotype ,y;ns

q.29. ".

A' sinilnr .polymorpMso hiis not· been ·cloc.rly shown' in:. .' theScottish1:laterial• . " ':-. .... , ,,~. '. i , " ; .~; ••• , ; . : . ; : . ; ,••••. : • •

•~ . "... • ". . . " , • r 'O.~ _ ..

, ' . - ~"" "L ·• , . • • L •.-~· J : . " ; " , _ . \:t.~ ..

_ ; The

v.a#~~~ons

·.in

t.r~nsferrin

type

bec?~e

obscured in' sarnples l1hich have -' : been deep frozcn for ovcn n' short .period ef timc~":For this reason' it· has not .', been possible tö' ~ype.clec.rly sero. eoliocted at 'Yest Greonlarid and. trc.nsportcd ':' deep frozen to Aberdeeri;' .. " ... ,. " . . ... ;.:".. :" ;:. . '\" "~ .. ' ' ...

~'.: ~ .~',.. .' . ;• • ": ..- . ':' :~.. ".7 ~".' ~...'~ ~'...."" '~"' . ., :.-'...~... :,. ":: •• ,; .:'.:::". : ~ .• , '.

poiYmo'i:,phisn

ha.~

::be·en.

ob~ e~ea:

i'n' the'::s

ertb·

pre-albumins 'of Scöttis'h, ' , saloon smolts, but 0.1; such low' freqüency

(1"':2%)

üs 'not 'to'be of' practical'vnluo'; in racial investigations.

~issuo Enzymes

•

( i r Lactate Dchycl

ro

₆

_{enas e :(J.DH) ;"':':, ';.'}

..:~.

. .... ; ...

:.,~.

,:l!,' .' • •>.:' "'. . '" ... -,:. ,'.;; .. " t ;~ '. ,... :;. . , : ' .. ,." ;'. , I , , · , • • ...

'. Up

;to 10:LDH' isoenzY6es,' cooposed of 4'distin:~t' sübüru..·ts·;~enccdcd. 'at

4: '.' .'

separate 'loei, "are 'observed~in'various .tissues ·of.~AtlnnticsalDon (Will::iiis ....~ ~

1968b). : . No' intraspecifiovariation in.the, tissuo-speoifio:isoenz;Yne patterns' . uere observed'uhon'over 60.individuaJ.s,~froo.Canada, Scotlö.nd,.West ·G~e'oiü.and·~' and Sweden - \7ere anaJ.ysed. Tho isoenzytle pattern cf the' retinä', . in' whieh'a .. thira.~'set' of 'iso:enzyoes has bcen shewn to oeeur. (Duke and Kelly .1968) has. not been investigo.ted;'· tho pc.ttern·observcd~in .tho eyelens,' however, -exhibited .

o~.;:;:. ~~e.

tetro..ocrs· kncun"froo ·the other

orgl.lns.:~

: . ... : .. ':

~

..

>'.:

~_;\~ ~

'.:,

.:",~::

':.: ..',~,

..

'. " ~ . ,.

.

•I'~ • . • • . .... . . • • . - . . . " . ~ , . . . : : . ; :., . , ' ; ." ' - . " : ' "... ••~ '.,~", • ~

, J"In white sooo.tic muscloj' ,all, the .isocnzymcs,are .obscrved, ,the most 'o.nodl.ll· . ('heart-type') tet·ra::J.ers' occurring only.in ·traoe quantity.- .In"tho 'white' . , Duscl'e 'cf juvc1llle' individuo.ls ofless than I.lpproximl.l~ely~'15.·cm;, tno ~~heJ.rt':'..:~ type' isoenzytlcs ooeur :at· rclntivolygreatcr intensity thaninadU1t's~.·andt . ' ameng the less anodD.l ('muscle-type') isoenzyoes, fracticri'5 i8 'relativoly :-. more ·ccncentratcd.'uhile fraction 1.occurs. only in trace. quantity., _

.' ,~"" ~.:.. :":".:./. '~"''':~''_'~':' .:...~; :: :'~''','':':::.<\_.. ~1..• ' : . ~~ , ~

'-(~~~";':

..

·g~l~te D.Chy~~o?~na''~~)::

,': .'".,.:.: .... : . "

.~

..:.,--: . ..' ..

~':/<'':

.

:. Fiftoen :1.soenzymcs cf mal;te:

dehycir~g~~ase

'"':".'

:i~~i;nS'· ~:.iniiu.rau.r;. :~f.:

6.--:' ..:

.', • . ... ,.; ,. , . 6. "., I"

,distinot subunits - are:observed'in~extrc.cts 'of salmon hcart ouscle·~· No _.: . intraspe'cific variations'have been observed 1n ..

·16

aa..U1t Scottish ·ol'.1Ocn '. ','"

s·o.far 'analys'cd; saloon'·from otherareas.havo·not.yct b'een:tested for 100 ._'... . polymorphiso.-' .:' .i "~:.: : . / : . ~~'.'~': :', ' : , .. : : : ' , , ' t . • . • . : ,

Oll' • • • • , •~ ,1 .~.':"'"~..: ..' • -- : . ~ f

. . .~.. . . -. .; .. - ' ; " . ... ":."i,;,. -.I . , .~, ~:. . . " . . \ :. . ' , ' . . .~, . ' . . . : ~: .. (iii).. Ncn-specific·Es'tcrnses·: of,:the Liver ' .. '.' .' .:.~.',: ',~ :::- :.~

•• .. ,.~:.. .... - ~.. , , ' J . ' ' : . • . . . • • • • • •~; •~. : .~ , : . . . : - ~. ' . . .. . : ' \ - ~' . ;'. - • : ,

. . Seven" distinct patterns 'of .non-specifio esternses have b'ecn cbse~ed:in.

nomogenates 'of-thci' livcrs' cf over 250 ·samon.: The patt'ems

-äre

:'illus.trntod ,

diagraIr.Ii~tically in Figura 1'~ .:.. ,.,' " ' . -,",.:"-: ".' :. ~','" . ' ,";.'

>:.... ':., .

. .; ',::.' - :: ' .~. .~:•.'.. ~-...~... t.''t_', ,: .. \ ..!._. ' , ' ' . a , ','" ... _'~~'•. ' A, ~ •

Each pattElrn"coopris os il nur.:.ber of, individual·ba.naB .cXh1biting' esterase': activity• . By uso of different substrates and inhibitors all'tha individual'"

components "ho.ve been' charocterised aS"ltl1iostcras es· (for terminology see ..

Augustinsson;.19~1)~·:-

:!',.:

~.;

. .,.: ..., : ..

t:~·;.·:-·i

::;-;':"

'.:.:~

• .

~. '.~.~::,~~.' ~

'.l.' ': '

... ./ I, ... .~. : ". ; .. ... .:1... \.!_ ;. _" ; .• _' :., ~ . . . .:!. :. j'/ _ . '. ..'. . . : . . . , • •

','" : ·.";The'estcrase zone. cf lcmest Modal oobility' -;labcllcdG .·in, Figure: 1 - ' . .i~·. coopos cd:ofa~ ri~obero~, ,\"'enkly~ rc:loti~o ~ cOI?pcnonts·w~e~·.

nro;

not. c~~~u;:l.y '; " separntcd undcr the electrophoret:Lc' condit:Lons. eoplcyod.:Ln those.s~~~es..~ ....

··l1.

higher rclo.tiv·e int<msity 'oi"' these cc:lponents· '7ns obsorved in'pnrrnnd" 'smolts :tli\li·in:iidU1t.fish.··~···':" ··.::i,··-n - ::.,,~ ' ':~'._..

.. '. : '. - • • • • r .' ". ," .• :.: ',: ( ; .~.: • ' . .~.

~ '. • .. • . . . ' . , • ~• .~ ( . : : : _•."~: ' . ' " . . . . .:" •• ' . ' J ' . " : ' ,.• r " " , ,..\

CotIponents E a.nd E₂ are a110lio,: o~e".or:bot~,.!,.ei~g p:resent in.' ~e!y ,fish analyscd. Becnuse these oooponcnts 'i1cre not I.lh;ays clearly separo.ted··

'frain .eo:ch~thcr

1. ₁

:i.-~ ·.no~ yet possiblo to dotcrmine the :frcqu~noy.o~,the alleles (E and E ) J.n dl.ffcront populc.tions •... The ccoponcnts Ai B,

C;

D. .;: ~ore'al1'elearly soparatcd,:'nnd thoirprcsonee or.absence foros thQ basis' • of the cl.nssif'ieation into

7

distinct po.-t::tcms. :. The froquenoy cif.' occurrence

cf the patterns is not associnted V1ith length in cithcr thö Scottish"or West' Greenland samples. Tl1e oode of ir.hcritc.nce of ccoponcnts A-D has not yct boen elucidated.

(5)

,

Tho froqucncy of 'tho ·7 patterns, in 'samplcs' fron a.

n~bcr: o~r" a.iff~r€nt·

'. :. ,: arms is presontcd'in Tablc

3: '.

Pattern 7has. not bocn cbservod'in the ' ' .. : European oo.terio.l, whcroas it

i3

thc cömonest' type in'tho Co.nidian oo.tcric.l',··· .: o.nd is also rolatively CO:JI:lon in the Uost Greenla.nd saople:" ·Individuo.ls '01' " ' this csterase type at, West Greenland can, .thcn, roosonably bc assuned to , rcpres ont· fish of, North .A6crlcan orißiIi~'.~ On ,this bazis, the r:iirlii:l.\in· contrlbu-tion cf North .Aoericanfis~to the 'West Grocnland.fishory·nus't 'eXcie'ed': ' ... '" approxino.tely

17%.

On the other hand, tho pattern chscrved

'in

the S\1cdish fish (pattern No. 2) has not bcen observed at West Groenland~''The "Swodisn fish analysed·may ·reprosent individuals of, a population vlhich feeds in tho Bo.ltic and does not ccntri'büto. to" the West' Grecnland'fisherY~'·:.- ,:,': ":.:., ::

- . ~.. , - '

..

)... " ; . ,~~,. .:~

..

:... ' " ,'~.-;..

:.

~., " ~ ;...":-\; .;. -

-

:.'.

':; .

DISCUSSION :~. ~.. .

•

,;.~

.

'

I ,

. '" " -

...

Fron the results prcsonted it is evident that North Anerican and Europoon populations of saloon. diffor in sorae, at lcast-; 'of the biochemfca1. und" - . s crological chäract'ors: investigat.od... In particular, liver esteras.es appcar to beospecially~useful:in .dis t inguis hing them,and:~tho~cste.ro.sepatterns·:~'... observod in s'amon' fron- the' T:Cst: coast of: Grecnland indicate: clElarij-' tho' " , . presonce'of individüals o'f different 'geogr:lphic origimin this

äreiJ..

~

.- .

. . ,

_-.,..

~ j.~;~,.J/,

.. ;

~.,..~.' ~

..

i ' : . ~ ":~-.I~.

\

,::.;. __ :..

.: -/",.

",<' .~.~..~,;.~ ,"~,;,:~~ "i~' , . The relo.tivo"proportions"at:\7ost Grcenland,' of individuals, origino.tirig' from 'different geogro.phic': areas' caImot', bo accurc.tcly estiraatöd fram thc dato. ~,

so far availablo. Iro inforno.tion .is 'yet:available on the ,'serological und' " biocheoical oharacteristics of salmon from the USA or froo IcelaInj" furlnor:J.ore,

only a' soall; nuinbcr ofind.i.viduals of Canadian origin, most .of i7hich originated fron.a· single 'rivcr, huvo ·been analysed.'" The: esteruse results als ö'indico.to I that', '~ithin.the British Islcs,'" tho proportions· of .. the. dif'foren( ph?notypös :'may vary betw.een 'populati'oris sp:lrmingin different \latersheds.: .'C . ~. : . ~"," .

....;

''''.~ ·-.~7 I: :: ....,.~ ..:.: • . . . . c· ::.: (':,~,,:;-,"':1:' ,; i :.~::, .:..~ • . . .~::_.

In ouny of·the chnructcrs 'analysed·consJ.dcrablo:oor:lploXl.ty.~,i.Il~oh:ing " ontogenetic variation und the occurrenco of nultiple r:lolocular foms of various proteins - was observcd. Considerntion of th~ sub-unit structu~e.?f,many/of

these proteins.indicatcs that a greo.ter nu~ber of loci'encode tho30-moleculos in Samo ·salo.r: thtin in~möst :fish nnd other .vertobrato: spooios. . Thus,-, thora

app~o"b"öätleo.st:S·Hb; '1001; '4' tnoroal'::LDH loci~and:6·heart-~.mH,löci:" .

expressed,in' the so.lcion.' Uodern:Atlantic '-samen are:considerod~"on·.öytölogical eVidcnce, 'ta ho.vo·cvolvcd'through·tctraploidis:ltion, although uhotherby autö-~

or allo-tetrnploidisnticn cannot be ccnclusivOly detemined.· Tho ovidenco~for multiple loci prosented hore, and diSCUSSCd, in grcatcr deto.il olser.hero·(Wilkins,

in preparntion) confims the hypothesis:of_eyC?lu~i~nbytetrnploidy: in m0d:0rn SalDo so.lar.. In addition, it indicates that many of the individual proteins

cbsorved on electroph'oresis are .thyb'rid noleculos',.:1.0. moleculos .composod cf tho polypeptide "prodiicts of the tno cistrons resulting.fr~ia t'etruploidisation~ In the analysis and interpretation of protein polynorphisms,. thon" (arid, ,... ," indced in 0.11 aspects of tho bioche~stry, physiology arid ecölogy cf the'spccics) the hybrid status 'of thc··saloonl_{is -a 'factcr of.. oajor importanco •..}

... . ....~,l:..~ .;-; ,...~. ",".: '. \"~:.: J"l' .~,:' : •..••. : •. ,. ... ' . ' : . " • • •, . ~". . , : '; :~ ~( " . , .

The" occurronce ·'of ontogenetic .variations in.nost, of ,tho charactcristics studied is also of najor significanco, especially in intcr';'poPulation: :, '. '. '.-coopo.risons. .In.anadronous populations, cnly tho juvenilo and the largcrj . sexuo.lly.i:ui.turing individuuls are:nomally availo.ble in, and. in the vicinity cf, thc- river. of. 'orlgin;" wher"en.s only the pre-mture und recove'nng spent indiyiduals are takön in' the,'open" s·ea.· ~ :\iith:the' cxc€ption of. the h~eI:loglobins·, hovlever/ the biochelnical':lnd s(n~ological''chnracteristics of xaigrating soÖltin:re' similur to those of the adults of the same locality und valid coopnrisons can be' drnwn betwecn smolts in their rivcr of origin, 1+ soa .,inter sa.lmcu·in·tho· opcn seä, o.nd· iidult"saJiaon' on thcir, spamung xaigraticn•.. :.~ ...." ~.:: " . ' ' "

.. • ~ ~... ... ~. . ; ....\ . t • • "'r "'. .". .. ... " " ' . ~

.

. .. . ..'. ..

-

." ... .... ....

..

~.~.,..

,Fino.llYi' thä' :occurrenco"-of cntogenetio ·.variation is cf considerable interost in view of the tetrc.pl·oid nature ,of'.. tho samon, suggestinG .tho.t at different·

stag~s cf the life cycletho 'charactcri:stics cf thq':priid.t~ve dipl~id .ance~to.rs

,'. . J. ' . \ . . . . ' . ...I- '

nay bc~)~r~f~~:ntiall~ ~~:css~~;:':'..,' .:: '; ,: \. ,.~-' .':: .. , _'. .H· •

"

(6)

Augustinsson, K.-B.

1961

DW{e, E. J. und Kelly,

M. 1968

I. C.E.S.

1969

Koch,

S.

J. A., Bergstrao, 1964

E. and Evans, J. C.

REFERENCES

'Multiple forus of esterllse in vertebrate

blood plasma' •

Ann.

N. Y. Acad. Sci.,21±-:

844-860.

tL~ctate

dehydrogenase isoenzyces in hybrids

of salmon and s ea-trout' •

Is oenzyme Bulle,

January 1968.

'Report of the ICES/ICNAF Joint Working

Party on North Atlantic Saloon, Hay 1968' •

Coop. Res. Rep. Cons. pero. int. Explor.

Mer, Sero A(12): 18 pp.

'The oicroelectrophoretic separation on

starch gel of the haemoglobins cf Salmo

salar L.t.

Meded. K. vlaam. Acad. 26(9):

1-32.

-•

• Koch, H. J. A., Wilkins,

N. P., Bergstr6m, E.

and Evans, J. C•

Wilkins, N. P.

VIiIkins , N. P.

Wilkins, N. P.

'Further studies on the multiple componenta

of the haemoglobins of Salmo salar L.t.

Meded. K. vlaam. Acad.

~(7):

1-16:-1967a

'Scottish Biochemcal Studies on Salm on ' •

Res. decum. int. Coumn. N. Atlanto Fish.,

(67/99), 2 pp.

1967b

'Iomunology, serology and blood group

res earch in fishes' •

In:

Pol:,"OorphiswGS

biochenique des Anirnaux.

Proc. 10th

Congr. E.S.A.B.R.

Paris July 5-8, 1966.

Inst. Natl. Rech. Agronom.

Paris, p. 355-359.

19680.

'Multiple haemoglobins of the Atlantic

salmon (Salmo salar L.)'.

J. Fish. Res.

Bd.

Can.~12):

2651-2663.

1968b

'Lactate dehydrogenase isoenzyrues in Atlantic

saJnon (Saloo salar L.)t.

Proc. 11th Ccngr.

E.S.A.B.R., Warsaw, July 2-8, 1968.

In Press •

(7)

TABLE 1

Agglutination of the erythrocytes of one year old salmon parr and two year old salmon smolts v:ith unabsorbed rabbit anti-salI:lon

inmune serum at a dilution of 1 in 8.

9

Number

Parr _I

_I

I

9

11 l ₄

-

-1

-15

I

II

3

10 1 I ₁ ₁

-

-14

-

2 -

-

2

15 1 2

2

-

₇

I

Smolts 16

₃

4

3

2

-

12

17 1

3

10

.

2 1 17

18 1

2

6

1

-

10

;

~---,---_._---;

I

L. Degree of Agglutination

I

(cm) (+)

+

I

++ +++

,...--"-7--.;...-.1--+---1

l~-....;---+---2---'

I

8

6

2

I

!

1

(8)

I"""

•

TABLE 2

Agglutination of the erythrocytes of salmon takon at West Greenland and in Scotland

with specific absorbed re['.gents. The Scottish salmon are all 2+ sea winter fish, taken in their spawning rivers. All the W. Greenland individuals had spent 1+ winters in the sea at the tioe of sanpling, Le. if allowed to return to rivers in Europe or North America they would represent 2+ s ea winter fish.

Reagent Proportion of Positive Reaotors

•

Scotland

w.

Greenland

%

Anti-A

76

65

Anti-B

75

58

Anti-C

25

34-Anti-H

65

46

Anti-J

87

48

Anti-K

83

62

Anti-L

48

55

Anti-M

74

43

(9)

,...---.

-•

TABLE

3

Numbers cf individuals cf the different liver esterase types observed in sampIes of saloon from different areas

44

17

7

51

No data available Smolts

1+ and 2+ sea winter salmon

I

2+ s ea wint er s aloen Ireland

Sweden England Soetland

I

r-

Esteras e Pa.ttorn

I

~

_ _A_re_a 1-,

lI_'at~_r:L_'_al

WI

3 i

41

5 i

~+-7--t_T_ot_a_l-i

Canada 2+ sea winter saloon

I

0

I

0

I

0

j

2!

3\

2 16 23

I •

I

I 1 0

I

10

I

7

26 0

2

0

291

8 2

3

0

o

0 1 14, 2 0 ;.

o

7

0 0 0 0 0

•

West Greenland 1+ sea winter saloon· 0 0 41

4 30

33

22 130

I---.!..---+-+--~_+-+-+_-+__l~--Totals

7

81

35

44 64 38

272

(10)

•

ESl'''i1\SE PAT'~, OBSERVB L. SALI>iON L1VER

.~-':I I'!{_!i,

'(ti ,;~

~~

-,~ ;;} ~'.

•

~f' .'" ,~~~ .~ "

-.M."! ~ ~~.t~

A

B

D

c:::J

I

f:::J r.=J

3

4

C:J ORIGIN