Not; to be cited without prior reference to theauthor

•{J

INTERNATIONAL r.OUNCILFOR TUf, EXPT,ORATION OF THR SEA

C.M. 1978/H:I9Pelagic Fish Co~~ittee

Koval L.I. '

Feldman V.N.,AtlantNIROKaliningradUSSR

,\

1

.'

GENETIC-BIOCHEMICAL· AND MORPHO~mTRICAL

CHARACTERISTICS OF, THE NORTH SEA' SPRAT,

SPRATTUS SPRATTUS L.

2

Abstract

, ,

',' , }I'he variability of plastic charaeters and the nurnber of'kiel seales in two~and three-year-old sprat areeonsidered. The,stat~stieal 'analysis of the ,data showed the absene~ of sexualdimorphisffiand age variability by the majority of eharae,ters.The maximum depth' of body varies depending on the eonditionfaetor and fat content during thc spawning period. The sprat eof the coastal 'Danisll, waters ean be disti,nguished from the fishof other areas of the s~a (Farn Deeps, Dogger,: Southern) bythe lengths of snout and lowerjaw.

. There exists a good agreement between the results of; ,. . .

morphometrical investigations and genetie s~udies of polYmorph-ism'of muscle esterases. The'comparison of the allele frequencesof three polymorphie' loeuses oi, esterases and of ,one lOcus otmalatdehydrogenase of thesprat from the same areas demonstrated

, that the heterogeneity increases \Vi'th the, expansion of thes~pling a.J;'ea to the east, and that the sprat cf the Danish

, ,

watersdiffers from the same speeies of other North Sea areasin frequency of alleles of a number of loeuses. 4t

t

1

,

"

II', ,I

desprat de 1a mer'du Nord.. ~ ~ .-~ ." ", ': '. . - _.' .;

~ • <', •

II.,

~"'. . ,,

"I

", ,I

On' examina: la' va:riab'iiit6 des cara6tere~ plastique~~·t.....:-.

la quantit~'des' ,~caill'es de qUil-iechez 'le sprat 'd~ 'd~ux','e~'<"-'. ." " ' ..., ", , , . '.', " ,,',' , ,:';: :': . ;: ".. , ...

de, trois, ans~ L'analys~statistiqu~desdonnees a'·montre ... . ' .. .

d' apr~s la:':pl~partdes 'earacteres I' absenc'e' de dim'orphisme·.

sexu~i ~'~ .'de ·.l.~ 'V~~i~1Jili te. d ;'a~e. La plus g~ande haut~ur du, ....'. ,~ ": .

'. . .eorps varie en'fonetion du coefficient de,la condition et de ';\

: ,

, la. tene'ur en, graisse ,du po1sson' au 'cours .. da la saison. de :ponte~

Le 'sprat' des eaux cati~res de 'Da~ema:~k" s~ disti~gU~'~~elemen~','. : '. ... . . ..".. :, . .... ~: ~ ,,'. _~. ". . .." ,:', " . ,', ~; \

de' ce1u1' des· autres regions de la. mer' '(Faril~Deeps",Dogger, .:, ". '• ", ,'. • <

South) parIa ,longuetir', du mu~e~u,'et pa.r ia' mandibule. . : "., ..•:~ ',': . -; .:k ....~. .' . .' .' " ';' .~' .' ";:. . _::. -;'~ .' .... .; . .:.: J'". ' •• ' '~'. j ':. .' :. .. '". ~

Les etudes,morphometriques s'aeeordent bien'avec les r~-,':,.' ~'. . . ." . t... ~~'".. ." . .~, ~ .' \. ~ . .;, " .' ~- . . ' >. ".',... .

" sultats"de. eell:ese-ei' g~n~tiques du polyraorphisme' des esterases . "• ~ ... -:.... '. '.. ~"\'"," ~:, ," ~ .' , •• '. "'.'~ .' • ..~". '.~ .. :.. • ~ 0' " _ .tl'· ... :....'•• ~. '"

musculaires qui montrent' qU'en,comparant.les trequenees cles." , . :. , " ' ..", .:. ... '. .' ,'. .' ,,: .'. '... ' " .' .... .

. allele's de' J locuspolymorphiques des estflrases et un locus. ',. ". " :-: , • ~ ".' 1 '.' .. '1". • ',.\ " .. . • ... • • •

: de' -malatd~hydrogen~s'e du sprat de I' origine, des m~mes rflg1~ns' ..

~ ~h~t~r?ge~6'~~~. s'a~~ro1t'a~e'e 'I t ~i~~gl'SSe~ei?-~',de' ia' g~ographie;de' pGche. 'a,' l:~:est et' l~ :sprat', da la region danoise.::se distingue

par i:~ fr~q~enee"~~~' alleie's .~i'e la serie des loöus du 'sprat", "

.. ' ...des autres'rflgions de 1a mer du Nord.

. 3

. I,

~'I~lI

-l()<'-J~.

I

l·"

\

,pecuJ.'i,aritieso:f the ;spI'at from tbc ltorth Sea, Scagerrak :and

)Kattegat were studied, by Rodeko. 1910; 'Bjorkari.1929; Uo1ander,

•1940;' Poulsen,195b; Donnavig, 1951 i Jonsen, 196C? Theso authors, .

mainly -deal withthe -variability of tho meristic cho.ro.o'ters.

, Jensen'':(1960) presonted ,'thc mn'.iorioJ. on the morphometry of the, . /

. ,~

sprat .f~om' tlie: open North Sea' aroo.s. ' In, most, recent paper on 'I . ~.. . . . .

!,the sprat 'of the' Sweden ,\Vaters" Lindquist, (1968) milde a conc1us-

ion'that none of the considered meristio chnructors may be succ­

i essfully- used to single 'out the populations.

In tlie p~~~ent p~p~r an att~~pt; is~o.deto analy'se 'tho'.. ~ ~ "'. ',,,. ,;. ." •• .' .' -'.., -", ~ • - .> " • '. •• • " •

polymo~phism of m~sc1e esterases and'malatdehYdrogenaze for 4t,,'_:._:~ ."~'.~ :' .. ~ ~. ~.!., ", - . t"· ' ,', . ',_ .' _ • \ '--~ . "

determination of the degreeof the sprat genetic homogenQi~'for.', . :\", -., ..' .t" . '0-., • ,'. .' • • .. , ." .. ,', _. ~

four 'clrsas of the Uorth"Sea, and SJ.so :for investigation of the.' ",'

varj:abiiity ~:f pi~sti6':ch~acters;of th~ sprät from these araas,

Although the spru~, Sprattus spruttus L., is one of the

'~ost abundant and widely distributed spocies in tho soas of the

.NortheastAtlantic, very 11tt10 in lcno~~ about gonetic-biochem­

',ical char~cteristics oi tbc aprat.The polymorphism of haemo-

;globin"t~'~~ferrins :Md. -se'r~Jm ~steras~s in 'the spratof the ,- "'.Norw~gian waters was iri~oo-tigat'e'd by '~eavciB.1· (1968, 1969)~The'

'data on the sprat of tho opon North Sea., arena aro. fragmentary .. . . '

: (Howlettand Jamieson. 1971). ~ho intraspecific morphological

", .,' ~ .'. :"

as·· anindex·'chnrri.6terJ.zing the degree of isolation arid 10001-," . "

, i zation.:' of th~' 'population. <•

.~ ;... ,........

4

1.tatorial und IIlothod

, '

and 'by a sampling unit from tbe Danish region :( tothe eas'ti6:f,' ,", . ' '. .

4t'6°E).

were analysed in' 1973. In 1974 tbe available material was' suppl-,:::," . l' ,.: '.. ' . ."' . ','", ,'. 0' . ',.' I. .," • , .

emented by'new sampIes fromthe.three areasinvestigated before,.. ....' -, .' ' " ;.. '~"

5 .. " . -....

, . . .usos were used', as genetie eharacteristics.' ' .'.• "".' . .. l "., , • .. '

prescription' (MaUrer; '1971), the lower electrode buffer was\ ... ".;" - .

, The fraetioning of water soluble proteins' ;wa~ ,made' in' a' :,' , .''\. .. '-. ' , . .. .' ,.' " .,

vertical block of 8.2% polyacrilD.mid,gel~ The 'upper electrode,

: and seI b~fers,werep~epared accor~ng,t~"the J~leyts et' al'.

, . . ..'

In ·1976' the museIes of the fish' sUbjeC'!i"to nlOx-phologic8J.' ,.-:,

analysis were,'used :for tbe genetic-biochemic'8J. inve'st:1.go:tions. ,

The ,'aJ.lel'e 'frequences' in the malatdehydrogenaz~,(iidh) ,polymorph­

ie ,loeus 'and in t~ee'ester-aze (Es3; Es6 ~;' Bs8 ) 'polymorphie loc-

• <:. .'-t· • •

prepared according to. Rinsfeld et alt '(MD.t:tr~r" .1971). To reveal,\{ ..

the fe;,mentative activitY o~ the blockn the substrat'e' ;Uxtures,". . . '!', . : h ~,' i .' ,A

j •. l. ..,.' ' • .~, • "

The material forgenetic~biochcniculinvestigutions TIas,,.' \

obtaincd in 1973, 1974' and 1976 in 'thc sun~ areas" whcre t~e ,:

snmplin(5, had ,boen made' 1'01' morphoI:1ctriclu 'studies: A total 'o:f .. ',, - . " ". " . - "", ",."".

2539 fish-specimens'was'collected. The,samp11ng units :from'3 '

arens:. Farn' D~'eps, D~gge;'andSouth~rn ,(te>' th~' south of 5~ON)"Y,:", ,," ... . ~

, The calculation ofthe ~lele frequences \vas~ade accorcli.n.i 1;0.. . . ... .. .. . ~..

the equat~o~ of Hardy-Weinbe~g~ the hC?mogenel1iY cri-terion-:, .

waro uscd. prepared ac c 01'ding .to the prescriptions of Shaw and• ~ , . ' - , ~ '•.-: ., • ,"'.. 11' ' .- • ' :" , • • ' • '. •

. Pra.s·~d (1970) .. ~e sche~es" of el~'ct~o~h'oregrainB' ~e, ~own iD, '. fig~ 1; ~et~ied des;riPti~n w~s give~before Ö(OV~,1976)•.

" ." . , .. "

was uscdto establish thc confidence,of differences betweon, -

the f'roQuences (Nill andShell, ,1958).. -. . ' ,

The' morphometrical data. were obtained in the Dogger and'

:'Souther~ 'areas (266 sp.) in April 1971,: in the Dogger Bank·

(172 ap.), Fa:i'nDe~ps: (1,9 sp.) ·'and Southern ,(1,4. sp.), ~eas

. in ~Jarch '1976;, :~d in:the Daltlsh area (110' sp~'j 'i~~ugust 1976.; , .' ~ ~ , . ~ . . . ,.' " . " . : .' '.: .' : .' ~:' ,.' ' : .

i ,'A 'scheme of Be~g: for .clupeids', was used in the measurements (Prav-'I.. , " "', :,",:'din, 1966).: All ~eaSurements were made' on the defrosted1'ish.

, Tbc ~atc,ria.1 was treated statistically (Plokhinsky" 1961 y, and.. ". . , " ' . ',.','

'. ,

the cri~erion 01' the ~elected diffcrence confidence (Student's 4t, criterion, t, named ,,,'j,1 difi~ in biological ,papers) estimated~

in order,togivethe' assessment ofthe difference degree be~veen i.- ~ -~ '. '

s3lIlpling, unit s •

The . fat- content 01' the sprat was. determined intheI

Sockslet's apparatus using the' standard method described in

11terature' (Laz~evsky', 1955) .. ' . ,", /' ' . " .,, .. I . .

. ,,' , I, .' ' '., /

Resultsand Discussion

" Genetic-biochemical characteristics'

I ..• ~' i:zofer~entn1 spectrum 01' m8J.atdehydrogenaze i8 represented,I. :' :' : '.' • " '_ ,',,', '. " , , ~, " '.", , " .

by three phenotypes (fig. 1)" which are,genetically controlled;; " " " .. \. ,- '.'.

,_. - ~. . . . .' - . . , ': . '.~ .'

.by~tVlo~alleles- 01' the mO,limorphal.locus. ,IIi. heterozygotes a

, 11Ybrid'f6;in(AAi)' 'ia 'obs~~ed,:'\~hiCh'i~'indi~ative.oibidi~-., '. ' . ~. '. . . ',": . . .', '. \ .. ,.~' .

,e'n~ionai .' stru6tur~ ,.~f" the "sp;at m'OJ.atdehYdrogenaze •.: ."". ' . ""

. ,~~ . • I ~ l. " ~ •.

A~ ·.üidic·~ted befere (Heavdal, ~969), the sprat 1s. distin~, I • :' . ' ~

uishedfor:extreemely wide individual variability ofthe ester-• : ',., " , ,,~', • ~ ~ I' ~. ~ , .~. :"; '. ". ' . -. •• •

aze spec trwn ~ Tbe examined' material possessed some tens of .."

phenotypcs; those which urc'most comnon are 6ho~n in ~ig. 2. Thc

hypotcsis was sUGsestcd that thc obnerved phcnotypcs are gcnet­

'ico.lly controllcd by tlu;ec locuses '(P, .LI, S)'. In this conncctio'n,

the freQuences 01' separate ollclesin these locuscs ~crc calcul­

ated: ·the allele Es3 f'rcqucncy for locu~ F, the allele FsG frcq-",· ..

uEmcy for locus lJ and thc all~leFs8 fre~ue~cy for 10c'us.S.The '.all.ele'· frequences i~ malatdehydrogenase and esterasesarc ~iv,en ..'

~ .' .in table ..1.' .

• <l;.. ~•.

. 0

. ',As shownby theassessment of the homoge~eity degree by the . :j- '," . . " ,.' "';

,e?-lle~~ frequen~y in maiatd~hya.rOg~n~se esterase~ ii 't~es~rat.,from,··: .

.differen1i areas t the intraspecific 'v'ariability cf spratbythe.:"::·'A '. 6' '. . '" ' ' .':<';-

alleles Mdh . and Es frequency is lower than that by the ·alleles' '.. '.

~s3 ~d is8 -frequency. It iso ~nlY due to' the allele lvldnAfreq~enci,','.' . , .. '~'

. that the Dariish sprat can be distinsllished from that· of· other . "

areas '(x~ 3.8"table 2)., ,

Insignificant differences by all locuses have been recordcd ..

between' the sprat from the Danish, area' end 'Farn Deeps. The '., ,.. . . , . '. , A

differences have been also observed in the f'requences of I\:dh , .

• ES~.' Es8 between "the spra"t caugh"t in "the smne area bu"t in diff­

erent :years. Tbe fly.ctuations were .thehighast among the sprat

fr'om the ,Farn Deeps' are~, ' arid l~ess pronounccd Alno,ng the fioh

from the Dogger Bank area•

. .

Morphometricat characteristics and'

morpho~physio~ogicalpeculiarities

, A hypothesi~ ?f seXl4al ~morphism in·the, sprat~.was tested..on the material collected in 1971. The analysis of each.sampling

....

7

unit showed insigni:ricop,t diffcrences in thc corwidcred features

b.etween 'l:ihe, sprut' males nndfcmales. Just in a sinßle sumpling

, Uni1:i,'of tVIO-ycar-o'lds an insignificunt' dif'ference bc't'\'lOcn the'. " .

'depth 01' the head at theocciput (Hdiff': 2.6) was recorded ".

in the'fi~h of'diffe~ent sexes. The absence cf seA~D.ldimorphism

.. in the fish. made 1t :possiblc to cax:-ry out the :follo\~ling anälysis '., . . ., "

.. \vithou:c ~ the ,divisio~,by sex." ,<

;: " ;A':definite morphological ,organization is knovm'1:io corres~ond.' . ...... , ,

t~" ~'a6h' stage;~f>1:ihe fish"ontogenesis.' The' variability:Of' merist-I· • ,..... .' .'

1c andplastic"cliaracters' is con~idered by'age 'groups. Two~and "t • ".(' ...

" three ye~ oid sprats, that'is 'the mature fish, 'at the same

,"ontogb~~'sis s1:iag~ were analysed~'The difference .in meristic undI' . •

, iüasti~' char'a.cters' b~twecn these age' groups 'Vlas' not 'considerable •. '. ~ .'. .,1 ,"'", ~ •• _. '. .: , ~ '" .•..• . " . ._ '. ~ '. ~ " •• ,~From 'thematerial'of1971 the differences bet;wecn two~andthree--

'I ' '" ' ',. ' , 0'

year-olds we~e revealed in the' sampIe from tbe'Dogger Bank in

three,characters: vertical'diamoter,oi: the eye, t11e length ofI ',," ,

middle fa::fS oi: caudalfin andtbe length of, pectoral fine In the' 1

'1976 materi~ the differences revco.1ed wore i,n the maximum body ," ~.,•!. ' • - (.'

length. (sampling unit from 0.11 areas),and in _thc"lcngth 01' 'middle,.' "

, '

fGYs 'of,lcaudal fin (:b'arn Deeps, sampling·.~t) (table ·3).'Due .to- "j '. • •

the absence of significant differences"bctwecn two-and three-year-: ~. '. - '.. '

I . ,

olds (by the ma;Jority of charactors), und also to a limitedI ' , ,volume of sampling units .obtained in 1971 ,the 'material for the

; , • , • • '.' \"'" • • ,.', ;. ,\. • ," \, < • , •

given year was compared without the division of the fishby age.:.

, \ , ,

1 •~e proc'e,ssing and an~l~si~s~f t~e ,ma~eri~ ~o.r.1976 was

made separately by aga group, , although the differences by the "

,,

,;:.' 8 'I"

I ~,"

majority of charactcrs wcrc;not rcvcalcd. The comparisonof 19

" I?las,tic, aÄd :5 mcri'stic , charneters(nWnbcr of kiel,soules K1 ;' ~2""

, ~d vcrtebra 'count) of thCfish' :c;..~:o tvlO Uorth Se~ regions ,:,~.',. . . . . . ',,'. : "'. ".' '.....

,',(Dogge~ imd'Southcrn, : '1971 )' did 'notreveal rclinbl'c differenc'es"':, ' . . ~'.' '" . ... . . '. ,

... '.. .' . '. . . '.' . ,,' .' .' .' , .. ~ "by thc'majority ,of characters., Thc oD.1y considcrablo differcncc >

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

,'- "

. rccorded waS thc: length of post-orbital 'distanc'eof thc hcc.d

(Mdiff ,4.8)~It'should be notea.. th~t thc 'measUreme~ts of ~h~' ,','"

, post~orb{tai :dis't~c~" of,:the hend. Gannot' be 'reiiable",enoU~~"~iriC:~'::• • ..... . . i,' i ", . ". "-.; . " f_ •• :' ",' :. 1

, membrana 'branchiostcga edging ,withtbc gi11 cover, posteriorly '1s .~',

• soft. 'Tbis Ileculiarity hadb~en:~sonot~dbY Lind'luist (1968).:." "Ge~g;aPhic~ vro;iab'ili-tY, ~f 'pias~i~ c~aracters can',be ~ se'en : ",; ::'

• ',> .. - . '. , ~,.' .. • - •

~ -ir~m', thc ,"materiUl,'for ''1976, coll~ctcd in thc:vast region (tab13 4-).:,,~'e coeifi9~icnts-o~ -variation ,(cv/'.~f:'the'~~jOrity: of plastic " ~.,.' ,

• , '1\' " " . '... -, • ,,' :',' .' ~. ' .•,:., : ',' '. - " ".'.> • , '. • ;:.

characters' do ,'not: exceed'10%,which is indicative of, homogeneit'y ,,:• ' • < • •

of s,~'pling units ',and insignificant measurememt errors. High -""

values of',thecoci'fici'ents ofvariation are'peculiar'to dei'initc

, characters' {n",thc saInpllng unit pi' ',two-year-oTds from·the South~. .. .' .' '- ~. ',-.

'ern a.rea~Certain,overcstimatio:o. may be, c2.'lsed'by a smallsample '., • ~J -. • • -. . ~ • '. ',' -.. ,

, • vo~ume'~n.,the:·onehrold',~d bythe proper' ~izc"of ~h~ charact.ef,,~

'on the other, whicn' is invcrsely dopcndenton the value, ofthe:. . '. . :,' '.' ...' , '.' .

v:ariation coefficiEmt.: S~ch a dC.PcI,ldcl1ce hadbecÄ ',note~ bY,Mair. ,',~ .' . t·

"(1971)~,':

-As is e~,iden-t :from 'the' Wl~~S !J:r thc !l1':l"terial'for ,'1976,

i;he ditf'~ence,sbe'~een;the:si>rat oi' 'the'op~ JloriJh Seaareas

, ('I-III)~e 5.~~ignifi6~t'. Co"nsid~anle' ~.ferenc~s "e~iSt h'",,'. p1o.stic :ch~act~rs'of' -t~: fish fron the'~eas'I~III':'~dthe,ai-ea~

" \ .... .,' .' ',.. .. . '. .

Iv. (~anish 'areal. Out of 1'7 eOinpared'piastic charac1:iers the' .

9

.1; ,

j ,

!

, '

Dnnish sprat cleal"'ly differs from the fioh of tho open soa

'e.' I ,

...., .

.: ',:~ .:r;egions in, 7 characters. The, sprats of the 'Danish waters have , I

" ,deeper body~ relatively larger. base of the dorsal fin, but, , .'

. '" r~la.t~~'elY~'niali.er 'len€;ths of' the' sno~t;,~ lower jaw, verti~al, 'r '.. • • •. • •, ,.' .;,' "diameter' of the eye, anteventral and: anteanSJ. distance compsx:ed; , ,' .. ' . , " ' , ,~, : ,: ' , :' " ' " ,".. : ' " .....'. ~to 'the.lndividuals from the' Farn· DAeps', Dogger, arid Southern

':1 :,~')~~~;. ',~e:~~Pth'o~"thO~: sp;at:bodj increas~s soh~heastw~d.';, .., ~'. • . • t .'. • • .....' ~ .'., ~ • . .'-. ." • '" • •

'.::::"Ariother importrint character",'the l~ngthof,the lower jo.w, varies"; t ...... .. . . I • : • • .' ~ , • \ ..... "e~: ." :~. ", • .... ". '. ", ' . • .',..' .'....., • .> : • • • : ' < • •

• t lnthE(same' direction. Other characters tendto be changed east-I' " " " '

I,' wa:rd·.~.~· '.'.

~e.depth of th~ bodi'is kno\'nLt~ be' important,"although, ' .

co~dition fa:ctor 'was shown in many-,pases (Milere~e, Shin.derite,

1976)":The s~e ;egularity of,'variation ispeculi'ar to .the'

. ' unrelia1?le, t~onomi~'~haracter. The rela:tions o'f this character,

!, t9 ~ther 'fea:tures in ,thc process'of its developm~nt, however~',areI, • .' , •• ~ • • • ',' . • • ~

, far; more 'importarit: than its invariability (Kirpichn'ikov, '1933).:t j '.' . f"

I The', existenc~ of a dir~~t d~pendence:of~he 'body depth on th~'; ,II

, I: ~,, !

, ,

" ' (table 7). From the, stro:t of., t?~sp,avming, wbi6h is of portional

., lorBcs-tdepth Qf, the sprat ,body, which i8, c,on.f:irmed b! hißb, ." . .

':"v~ues; o~' t~:e: c.orrelaiiion factor between ~liese, cha.racte~at, the e '0.01 significance level' (table 5). 'A dependence, of the variation

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

·of,the.Clark's condition factor and of the largest depth of the••• •• • '... • .' ,f. ' •

body'o~, the goIiad ma.tUt'ity'stage is undoubtful (table.6).' It:'·.. . . . . . '. '.

! can.. be se,en, 'that th~ conditio~ of .th~ sprat, 1s 1?he, lowest ,after.

Im~t~ati~~'of 't~e' first' portion o~,·'se~al pro~ucts '(maturityi ' ,',. " ",'. ",:. sto:ge' Ir,. II,I)~' thnt' is, immediately ,after overwill~ering, when~'I·." ' .'. " .' . .:' 'fat contentin muscle tissues of the sprat was·as lowas 5-78%,

•• I , ,.1 •

"!

'tIOII

..churo.cter and lasts. :fron Uorch to August, the sprat 'bcgins' to.

feed' and tho feoding intensity increascs ;;,ith tho advance;ot"·the..··. . .

. spuvlIling. The condition factor in 'also increased. Its hißbost".· :".'

values are eharaeteris'tie 01' ,post-spawners "at stage VI-II •..Dul"~, ",

'ing thfs period ·the ~at c'o~ten~ in thc muselcsis. hig~' (a~~~t.~>~~," - .~.. , .' .'....

18°') .:. ..,.... .," .:... ', ".jO ••.. '. :. .. .. : •• : ' : ••~ . . •.. ' .: ",.. ;': : ." . . < C.• :. ,:,"

; , .~ w" .. : ~ ." "" •• ' "." • '. .,.':; "':'. " ....... " .• ',:." <.. <,": : '~'.'~ '..~,:~ ..•~

: , ": :Similar changes' are .observed' in' the largest'depth':01' .~~~ .....:.:. '.:. ,.. ,: '.'," :. \, ~: ..~ "": ,:~;.'. ,~,~' : ...'"' . .- ...~.: ...:.'........ ... :, "~~" "',,.... :: . .. ~ .....~.~.'..-, ',: .. ,

body' during thespavm.1ng season 'dep;endin€5 on: conditi~n" :fat.::·,·::.,

.cOIlteIf~arid ~~t~i:tY::"~'~'ag~~:,Th~ :i~luen~·~.. ~f: p~s.iol~Gi~cÄ" ';~d~~'::.e ~rs on t~e de~t~ of:fihe\bo·~·c~·.:b~·~limin~~~·d..P~OYi~~d ..~h~~·>'::~.'~~~. . . .-. . ...,.' '. .'. ... . "' .. " ."., ...;' ""

exelusively large .sampl.esare a~~l~ble·•. Therefore,.th'~·useo~,··:,:·..

tlii~'cha:racter a~.~an i~dex'o~ th~' sprat' p~~ulat'i6ns 'i~~l~ti~~.:~. <~~d'l'~~~~z~jii~n·deßr.e~ 'i~ .·~ot;os~ible ~·o 'd'~te'~:' \ ... ' .. "'" .... ,~:':

" '

" 'From other'plastic,eharaete~s,.whichdistinguish.the,DaniSh'... ..: ' ~.. . : . "

sprat, thelength of the lower'. jaw. (lmd) andthe length.of .the, • r .... ' ~:. ... .. ~ , • < .. ' ~ .. ~. "J " " •., •• , •• ."" -," • • • • ,

snout(r) ean be taken, .since they have varied significantly::,.,:."',, •• ~, A. • _ "". .' " • ,-' ~. j, ' > . ,

. ~ , . . .in twosue,cessi:ve. ye~ elasses. The analysis .of correlatioribetw~

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

een these features and the lengths 01'· the head and body, accord.,..,

• ing to Smith ,(1); ma~eit POSS5.;l~ to estap11Sh their° rela~~v!,>

inciependence,,'(tabl~8). Therefore,.in a:series of.piastic,· eher;,:,;:. '., " .... ,.' ,., ~. ,'.... e. . '.. "'\.' '. ~ : .;:' <~). ;,'.'~ . ". .'- ,.:'..' .' ,,',';-:.' . ," ....•

.ac:fiers· which. may .be' used to distinguish the sprat of' the open', .. '

.sea 'regiOns frb~' that of th~ :'DaXusli~~t~~'~.~'iu;"'~h~'l~~~th~~ '~f '. i, . '." ,.-,,,... ," ; '\ • , • • .,' '.~ ,''0' . ,: •• ", ",

the snout andlower 'jaw ean be ~ealized.:· "... . ". ~

Conelusion .,

'. Th~', heterog'eneit:r of :1ihe Uorth'S~a sprat'r~veal~d :from t~e .

.genetic.,..bioghemi·ealeharaete~isties may 'be eaused by ~oth th~" " ..': .

II

'I

I

. :

I

I

·1I

I .!reproductivo isolation' und tho'diffcrences'in conditions that

;exert influence upon the yea:r class formation in: various regiö~s"I .

· I'·· .' . '. .' ..Proceeding from year-to-year differences, we 'Carl admit thattheso

. . " .,

conditions -are most variable" , . ., ' ... ~ . ...'

.! and D~nmark.·,·The offshore arens of. the central north 'Sea'(south":' .1 ' . ,'.- ~ ..... ~ ... : ...-.' .. ;

'!ward of:·Do·gger Bank), where thc spro.t· is charact'erized'by more':···.·J , .. ; " ; ",,""., . ,.~ .;', .',' ~ .' . ." '," ", ."-',' , ~. ',', ' ..'w,~, ..•• '.

:complex aga· structure, the. influence of pecul~arities,in form'7 .

.~: a~i~n :~:r "o~e o~,:'~ot~~~ year'class on ge~etic' '~~~u~t'W::e~ '~f"th~'.' . . ...... ~... '. ~:

.'

;)... Btable o.iffer~~c~~ 'm:;~~me :rela.tivelY··i:ndep~ndentplastic ':' e~ characters .of two' suoCi·essi.ve.yearclasses betw~en'the central' .

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

',i NorthSeaEiroas. andDan1sh waters .are' also· indioative of '0.' cert-.'

"!'dn 'i~olatiön of 'the s~rtit from' the"coastai Danish wD.ter~. '., '...t '. ' .1 ", .1

.; .~ .. The:'absence "of such' c1iffer~nce.s·in a number' of 'yeär ~las.ses; • • • ~ .. .. • .' • . • '. • f"'. " ....

, of' the fish' from other' areas" howevcr; "suggest that no isolated ~.',. ..,.. • .. • "'..' '. 0 " ....;

:' : groups caD.',oe :found i~ t~e open' arcas ofthe ccntra! North Bea•.~ ~ • ,0 • ~

, ; '" I ~ ':

"

.' :fu:or'der to' conf'irm· the 'de~ee of "i'solati~n and to' establish

'. lthe' locB.1itYof 'groups,' it' might beuseful, to consider some other

"lchar'~c't~r's'rel~te'd 1;0 '~·o\~h. ~d ~'~prociu~tiO~~' "\7e beli~~~ ,. how- e"1 t ~ .. ".,:.. .' • .:.,.' .. "" r~ .. ,.... ~, ..' < ':" ': '-' "':~"~ " •••• ' •• :, ,~\ .,:

· ever;that used cha.racters (inoluding ·the. maxim~ depth of the·". .~:.• '.' : • • •• .~,,'. " ,." " ,.·~ •.• ,.·I'"· .'.. ~",' .. '~'., ':.' ."":

body)a.re,reliable enoug!l.·'indices for differentiation 01" theI ;.' '.., .;:, ..'... "." '.;" ". ,;: .....'.' '. :',,:' .' '. .. <.',

.; groups by ·la.:r:ge, synchronous' sampling ~its', provided that .theI. '. .' ". ' .~" ." '. :. .\ '. :.... ~,' , I"·

, I mor:Phophysiologioa! peculiarities ond conditions' for the :forma- .·i'" ."'. '. .' . '., ..... .' .. :' '. ". . .' . .... .... .'.1 t~on of different year classes ~e taken into.consideration. Tae

I .

" .".

. . material' .for the genetic-:-biochemicaJ. studies . should' alsC? s'atisfy.' , '

, .I

.!I

':12,

rI

Table 1

Allele frequences by 4 locuses in the sprat, ,

of different North Sea areas,

NorthSea areasYear Alleles, I , 11 IIr IV

Dogger ' Farn Deeps Southern Danish "

MdhA' 0.315 0.293 0.314:-, '

0

E~3 0.284 . 0.:;49 ' 0.:;63

1973 6 0.537 ' 0.516 0.551Es . -Es8 ' 0.574 . O~528 0.5~0

n 324 252 ' 526'

MdhA '0.254 0.233 0.282 0.364

Es3 0.326 0.245 0.?J'+7 0.402 "'

.1974 Es6 . 0.-530 0.515 0.500 0.527

Es8 0.469' 0.399 0.484 0.674

n' '181 163 124 169 '

MdhA

Es3 0.290 0.235 0.200 0.335 .

19?6 Es6 ". 0.490 0·545 0.485 0.475Es8 ' 0 •.515 0.655 ' 0.550 , 0~540

n 200 200 200 200

I3

H i Table'2~

,Homogeneity criteria '<:Z2) of' the aliele .frequences in, ~,

"

,,4 locuses er t:he Spr~t from different N'orth Sea areas

..Alleles Yearoa 01 1-11 11-111 .,I-III lI-IV '. III-IV I-IVobservation'.'

10/7} o "'(i. 1.8 0.47 -.. t::./

MdhA 1974 0.41 1.9 0.61 . 1}.2} 4.59 10.011976

1973 4.65 0·58 ) 11.04 - - -Es;

. ,.

1974- 5.12 7.17 0·77 21.;6 0.94- 4.071976 2.57 1.49 8.76 9.80 18.9 1.86

.'1973. 0.08 0.56 0.92 -:- -

Es6 1974 0.00 0.40 0.;2 0.24 1.12 '0.01

1976 1.60 2.88 0.76 3.94 0.08 0.02

197; 2 ..15 ' ;.1; 8.11 - ~

Es8 1974 3.26 ;.88 ·0.24 50.65 21.00 28.86

1976 5.·.38 10.25 0.2} 10.04 0.08 ' 1.02

,S & 1223 • •.. ~-- ,)

e eTable :;

, ,

Age variability of some plastic characters of ,the sprat'in

, sampling units trom thedifferent North Sea ,areas, .

•••• M diff.• n' ·• ·

: ,s Char- :: Two-year-olds: Three-year-olds <. ,.~,Are... *: aC""'ers . ::,-.---::.=.:.;+~.=.::::::.....;;=;;;;....-----------,-=;;;;;;..;:;...;;...o~:;;;;;;;....;;..=.::::=. _

". M -m': G ': n: M:;: m " :" Gi

1971 'Per, centof·head length

, ,

:Fer cent of bodY J,ength,

0.56 6'+ 5.94+0~05. - .'0;' • ,-

O~41 64 7~68±0.06. '.'

0 28.42+0.21-Dogger

10 5.26;;0.0'7

IV 8.08+0.05. -

1.7'+ 64' 27.81+0.17 '- ,1·35

'.'.

63

63 '

63

Southern' HFarn Deeps '10

18.81±O.11

.8.71±0.07, 0.790.61'

19765276

19•33±°.1'08.29±0.08· .

76Lj.9

3.5,4.12

;" ,.

" f'

." : .. "

. .' : .

-- i·'

i

Table.4'

", !.

Comparative c~aracteristic of major.plastic ~haracters'of the sprat from

the four different No~th Sea' areas

: II-;.: ','" ' ..... ,.' Farn. 'Deeps ..,nos·::: ·139'

. :.:.(? : CV. . . .M±m

•· '.

, .•·CV :.•

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

." . Two-yea.r-oldB' ;'.,.

,of head·lengthPer cent

"

,. 33.5lJ+O~17 1.69' 5.03 53.06±0.40 3.91 11.82" ~.

47.87:tO,,16 3~11 e47.90+0.16 1.58 ,3.29 1.49, ... - .28.52±0.13 '.'1.24 4.34 ,29.22±0.12 1.06 3.62

, .(

per cent o'f bodY length.18.8Q±0,,18 . . 0.79 4.20 18.26+0.15 1.32 7.22. . -46.15±0.13 1.22 2.64 46.28+0.20 1.77 3.82-.63.72±0.18 1.70 2.66 63.25.i0.32 2.89 4.56.

. 12.03±0.0'7 . 0.73 6.06 11.7.6±0.0? 0.60 5.•10

Three-yea.r-oldsper cent of head'lengt;h'

'35.44+0.18 ' 1.56 ,4.66 , 32.28:tO.18 1.32 '3.96. . -,"

47.62+0.18 1.60 3.35 47 ~58*Ö.~6 1.20 2.52 e. -28.45:t0 •14 1.21 4.25 ' 28.93+0.16 1.18 4.07, . .._' .... l

per cent of body length", ,

" 19.12+0.24 2.16: 11.29 18.5Q±0.12 0.89 . 4~81'I

..,'" 45.9lJ+O.12 . ~ 1 ~05 ,2.28 45.50+0.25 .,1 .87 . 4.10.. .. . . -

:. 63·53±0.14 '1.20 . 1.88 63.45+0.22 .1.61 2.53, - , . - -". 11.92±0.O8 0.75' 6.29 11 ~58:!:O.10 ,0.75 ' 6.47

.

rImdo

. H

aVaA

. eD i"

. rImd'

'. °

.', H'

'aVaA

ID

,.'j

....

16 i'

..Table 4 (continued)

·••..·· III : rr.

Southern . Danishnos: 134 :. nos ~ 110

:--;1r:-l-:±~m~-~:~6~~L:~C;;-;V":"'-;'-:---'l:-:-'l-:+--=m~-:.=-:~,:"::Ö=-=",--,---:,-:-,-:-.,"'i'iCmV.......-

Characters'

..- ',,"

',,'

-' "'. ','"'1" ; ....

\ ',' ..'".', ,

,.4:19

14·971.88

21.35

.. T\vo-year-()J.ds, ..

'per cent of head J-ength,'. . . .'

. per c;entofbody length' ",." _.. , ~""~ . .. :. ,~',. . .. ~

20.53+0~09'·O~74' ' - . 3.60'45.17;0,18,' 1.52 3~36'6i.45±0,18 1.52', " D.i?12~22±0.11. '0.92 .~ 7.52:

0.79.,,6,80

1.21,2.56

18.81+0.11 .. . .... .45~42±0.98'

6L~.17±0.18

11;99+0.37.. .....

'.1. .

:' .." -'

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

'.: ,3~~11±0.21

'48.88±0.22 .. · .

29.11+0.17, .-

" ~ < : / >

'_ '~. • M

" '",

' .....

"32.72+0.15- ,

,48.91±0.15

28.62±O.16

per cent of body lengtho '

19:3~±0,18", 0.87'. 46.25±0.14 ,1.28

6l1-.17±0.14 ' ,1.2311 ~84+0~07 '0.64-. --,

, ..4.50·2.'7~ ,

, 1 ~94

5~40

20.58±0.1'7 ,1.06'45.51:t0.24 : '1 .'45­

~2.96±O~33·, 2.07:12.1-2±0.16 0.98, '

"5.15'3.18 '

'., · 3.28'8.08 "... ,.

17

lI-IV·

Table 4 (continued)

.. • t 'C r.1 diff.)Charactcrs:-,-....:... ..;.... ----:.......:--~~-....;::.~===~----~-- -----

:' .I-lI I-III II-III I-IV III':"rv

. .'. ,~. ",'

.' .Two-year";'olds

per cent ofheadlength .. -... ,'". . . ... , ... ,

, .. ,,.. ..• ~ ._ .' , " • • • f

-', ' .. "r: ' 1.<?~,: .1".55 ....

lmd" < 0.1;, :', '~.56,; '"" ," , ,"'.' - . " ..

, 0" .:', '~~9~ :. ,'. 2.79 '.'

,.'

":':':2,,5~.'·

.. ,' .,0.55 .',: 0.'1:;~:

':" ',2.74 .'

6.55 ,: 4 ~ O? :,' " .2~17: .7.05, .', 8.95,:', ',.6.,9", .1.74" . 4~26" 5.86..

-.','. \.

"H

aVaA

.. ID

'. ;," .,.

... '.

0~11

:;.66·O.5?

per' cent. of boay l'ength

O.w: ; .,2;96 ' 4.470~74 .' 0.86' 4.47

. ,

·1,82· " 2.58 ~ 5~06

0.11' . 0.62 . 1.44

12.00

0.256.8:;

" ,~', 2.60

5.:;44.182.12,.63

!

r '0.64 ... . .

, lmd ' .' 0.05 '

O' '; :;~~9:

Three-year-ol9-s 'per.centof headlength

3.00 . 2.:;0 . ,'4.67 2.46 4.15,5.62 .6.13 .4.58 9,,89 2.33,0.92 1.:;4

'.

0.40 0.33 . 1.59

per' cent. or bodj' length! ..

H , .. 2.28 0.80 5.49· . 4.16 ' ·6.25 9.92., ','

aV 1.59 1.66 2.59 1.62~ .2.67 0~05 e' .aA ';0.15 :;.4:; '2.76 :1~95 3.,34 1.24 ..'·ID. .i,; 2.70 .O.7? 2.15 ',1.16 1.65 ' ,

i 2.95" , '.

'. , . ,.; ,

Notes: r :.. .':·l~ngth ,of'~hesno~t; Imd - length of' lower, jaw;O - vert-1c~'diameter of'the eye; 'u - the' lat'gest body 'depth; aV ~

. anteventraldistance; aA - antearial distance; lD - length of'

. dorsal fin.base;lc - length of.m±ddlerays of caudal fin;IV.-;length of'ventral'fin; C - length ofthehead.,

18

Table 5

A dependence be~veen the bo~ depthand the condition factor of "the spr~t,

, Condition,

• · Maximum · " • Correlation· , . • •Age • • body. · factor (Q), · f'aci;or,• A:rea • • •

of the • · ' depth, • · "betweeri '• e ,e •fish,' e • ,(H),% :' Clark's:Fulton's · Qkand H •• · '

•

I 18.80 ',0.67 ' 0.?6 ":0~66

2 years II 18.26 Cl 0.65 0.?7 0.81old'III 18.81 0.69 ' 0.80 " 0 .. 75' 'IIV' 20.53 0.86', 0.96' 0.87· Ie I 19.12 ' 0.?3 0.81 0.74,' I

3,years old II 1.8.,50 0.67 ,0.76 0.81III 19.33 0.72 0.83 0.69 '

IV , 20.,58 0.90 0.98 0.73,

.e

I 19

Table 6

Thc'var'iation of thel Clark's condition,factor'" - .

and the maximurri." depth "of the body depending on'the gonad maturity stage

. .Gonad, : Areas I, 11, 111, (March): Are~ IV (August) ;Nos.

maturity';: :-----------~., stage," : Clark' s Maximum :' Clark' s M~"'Cimum:'" "~ .condition bodydepth, : condition body depth; :

-: factor ' mm, : factor mm:. . . .•• 'cI''/:

11,' ­

111r/',

\ IV-V!V;,

VI,VI~II,

'0.680.690.720~73,_ 0:

o' •• '

0.66,,0.720.730.750.770.76

'21.4621.5422.0123.00

2 year' old fish

20.4521.84 -22.24 ',- -22.88 - ' ~

23.6322.84" 0~90 0.92 22.05 24.07

3 year· old fish: 11 0.63 0.67 21 ~79 22.07 - 53

111 '0.72 0.73 ' 23.21 21.93 - 28IV 0.77· 0~78' 24.40' 22.98 - 90

J:(~V' , 0.77 0~77 23.90 25.11 - - 29V ..... ' 0.79 - 25.19 - - -: -: 16.,

23~87 'S-' VI:, 'VI.....II: 0.81 0.76 25.45 2}.48 0.92 '0.88 24,.35-, . I

• .-:'_1•.2 e:! IIJ oe

20

SE

Table 7

3

The dynamcs ot· the tat coniients in the spraii muscle$ in 1976 (%)

1\1 0 n ii h sI - 11 111 IV V VI- VII VIII IX - ·X XI XII..

14-14- .1!?-12 8-5 7-5 6-5 6-5 6-14 6-18 14-18 14-18 14-15 15-12

NN

, Jable '8

Correlation factors betweensome i>iast1c CharacterS.\'. .\

............ l'

I1!.....•

c

......•.... I

IV••.... c

Two year old fishr 0.20 - 0.27 - 0.12 - 0.07 - 0.26 - 0.35 - 0.02 - 0.012

Imd 0.06 - 0.01 0010 0.03 .- ·0.22 - 0.29 . - 0.17 - 0.22

Three year old fish,.

r - 0.04- - 0.01 - 0.25 - 0·31 - 0.04- - 0.03 - 0.40 - 0·37Imd -.0.25, - 0.34- - 0.03 - 0.04- - 0.02 - 0~11 0.0:; - 0.12.

2 r

e.

References

,. BJERKAU P., 1929. On thc validityof thc "RIlCC charactcrs"

ofthe ,food fishes.Rapp. et ~roc.vcrb.,

54:73-77·" .::' .'.

"

z. DANNEVIG G., 1951. Sprat from Uorvegianwaters. An analysis

of veruebrae c~unus.Fiskeridirect, s~~.

'S:er. 'Havunders 9(2) ,'22 p.o

. "

3. HOWLETT G., J~lIESON A., 1971.'A system of muscle esterase

,:.va.rlants in the sp~at.(Sprattus Sprattus L.)'

Rapp. et Proc .~verb. , vol.161 :45-'-l·7 • .. ,

~. JENSEN A.J.C., 1960. Danish catches of sprat from theNorth

Sea, Skagerrak and Kattegat. Ann. BioI., 15

(1958) :189-191. '

5. KIRPICmUKOV' V~S., 1933. On the problem of race' formation in. ,

the fishes. "Biological Journal"', P.6.

, 'pp. 609-627 •.

6. KOVAL L.I.~ 1976. Electrophoretic versions of malatdehydrogen-

aze in 'the sprat (Sprattus Sprattus L.).

... ,Trudy Atl,antNIRO, vyp. XIX, pp. 19-23· . "

7•. LAZARl1VSKY A.A.; 1955. Technical' and. chemical ,contro'l in the

fish processing industry •. 111., "Pishcheprom-. '

. izdat", pp. ,519.

8. LINDQUIST A:, 1968. Meristic and morphometric c4aracters. . . . . .~

clas~es and ttraces" öf the spra~ (Sprattus

Sprattus) Institut of 1I1arin Research"

,!(yseckil, Sweden~Rep.',No. 17, Sero Biology,

L~d: 26 p. ,', '

23

9. MAIR E., 1971. Principles of zoological ta.xonOI:W. ,M., "Mir lt,

p. 454.

10.MAUREH G., ,1971. Disc-electrophoresi"s, M., "Mir", p. 2 ll-7.

11. MILERENEE.Yu., SHINDERITE. V.s~, 1976. Tho peculiarities o,f

meristic and'plastic features roldthe'charac~­

eristic of,cariotype~ The vimba (complex stud­

ies in different points of'the investigation-. . ."

area). Vilnus. Izd. tfMokslas" , pp. 11-25.

, 12~ 'MOLAIIDER A.R., 1940. A research upon the sprat of the west. ,

coast of Swe·den. Svenslc'e 'lIydr.Biol.Komm.Skr •

N.S.Bio~ogie, No. 2(4):63p. '

13. NEAVDAL G. ~ 1968'. Studie.s,' on hemoglobin and serum proteins in

sprat from Norwegian water,s. Fishdir.Skr.Ser., .Havunders, 14.:160-182.

'iih N:mAVDAL G:, ~969. Studies on serum esterase in 'herring and

aprat. Fisk.nlr.Skr.Ser.Navunders, 15:83-90.

"

15. NILL.G~, SlfELL W., 1958•. Th~ hcreditY of the human being_ Izd~

Inostr.lit., M., J? 388.

16. PLOIGfINSKY N.A.Biometry.M. Izd.1 MGU, p\ 367"..... ,17. POULSEN E.ll• ., 1950. La peche la poute et les races de sprats

-'. ,dans' les.eaux da.noises. Rap.· et Proc .Verb.,

126:92-95·

.e

;18.' PRAVDIl~ I •.F.,~ 1966'. The manualfor the studying of the fishes •..

M., tfPishchevaja. promyshlennost tl , p •. 376.

:19hREDEKE H~C., 1910. Uber' den Sprott und die Sprottenfischerei

in Holland~~ütt.des deutsch.Seefischerei-verein&'". ~ .

v. 26: 158-183.'

24

20. SHA'lJi G.R. t PRASAD R. t 1970. Starch Gel"'Eleciirophores:Ls of

Enzymes - A Compilation of Recipes. Biochem.

Genet. 4:297-320.

25

--••--AA---AA

Fig~ 1. Phenotypes of the sprat malatdehydrogenaze (A'A', .AA ­

homodimers,·AA·· ~ heterodimers).

- --:/

- - -- - -- --- - - -Alb. -

---8] S-7 .-6 ] M-s--It 1~3 F-2

Fig. 2.\. Phenotypes of 'the non-specific esterases of the sprat

Alb (albumine of the human blood sertllD:, control).

F - fast zone, M -moderate zone, S"- slow zone.

26

IIo,I(II. B ne'I: 26/JI 1978 r. 06'DeM 3,5' l1BiI•...l: 3a.Kas 1225 TPIpalit 220

WOOIT3MPX.· 101.925, MocKJ3a, 111. APXHllOBa, 4/2. . . ,