ID~.rn.~ional COUDoil torthe Bxplorat1oD of ~h. Se.

0.1I.1987/Ha25,Pelagic p;8h·co..1tt••

. KOTSl L.I.

,ulantllIRO

1taliningrad

USsR

IftRUOPtlLATION GDETIC-BIOCHBIlICAt

DD'I'BRBlITUTIOli .llO) STUDYDfG O:r

POPULATION S'lROOTURB IN lIlRIlIE AJID

OO&lHIO :rIBH SPEClES

- 2 -', . (' ~~.'

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

:Pro~:the ana1Y81sot the relationahip ot geneUo-b1oohell1oa1

4it!~rentia~i~D~d's~tial ~t~eture ot'd1etr1butlon area~ and~ ~. '. : .'" .

, sea8o~1' ~io{~<. oi: th:r"e.' .pee1e. (SprattU:. sprattue 'L., Sthenohu-.' . ~.~ ." . .

thls 'pteropus ,St.. ~~hUrus s:rmnietrlou. murph;y1 (lUohols) a non,.;.

,'randoai ,diBtrtbution';t 8p~o1m~ 1dth different genot1P88 rlthin

the l~1tS' ot;the 'ci1~trlbtit10n'';ea18 8hown. TMs re8ults from a

eompl1eat e4 ,~~rrelii.ti'~~'b~t~.n the bloch,mieal' pol1morPhi8111 and •

euoh phenot7PeDan1~.8tat1oD8of ph781olog10a1 differentiation as: ,,' .;;.~

sex, growth and maturat1on~,Henee it can be 1nferred that tor '.tu41-~ - .. . . I

. ing the population etruetÜr.'o! marine and oee&n.1o apeoiee trom, .

tunet1oDatl7 .1m1lar parts of d18tribution areas al1 pos81ble popu-

lat10n parameters8hould be'obta1ned.

.. 3 ..

INTRODUCTIOX'

In the reoent 7ears. a question of adaPtive'Valueot'bioohe~, "

miosl pol~orph1sm is being wide17 discussed'(K1rpitchnikov. 1987) •.', , '

,In experimental,works on genet~and seleotion:ot,the .fi~h. difte-

·renoes between speoimens with various genot1Pes gOTern1ng the~

rate. development and survival (Shoherbenok. 1975; Iljin et al••

~ 1985; Danzman et al•• 1986) are noted. The studies of the popula~

~tlons'in nature have also revesled s'oorrelation between the gene­

tio struoture and intrapopulation difterentiati~n in sex. age and'

growth rate (lltukhoT. 1974; .lltukhoT et al •• 1983; KirplohnikoT.

1987. eto").

The aOkno.ledg~ment of adaptivlt7 of bl~ohemioal pO!7morphism

issue8 from the faots ,of its relation to suoh phenotypio mani­

testations of rh7siologioal intrapopulatlon difterentistionas

growth ~ate. sex and age. It, impl1es that the us~ of bioohemioal

pol~orphism in population studie. based onl7 on random.samples

from populations has no prospeots~

The majority of nekton1c pelagio speoles are characterized

by differential distribution ot the spe01me~ Tar,ying in sex. l~

and age during the spa~ .(Dawson. 1~86). S1m1lar difterenUa- .

Uon 18 evident17 typical ,ot the diadl-omous and semi-diadromous

_nsh during the spawning season(lltukhoT. 1974; Chikhatohev et

al•• 1979).

Therefore. the interpretation of the genet,io-biochemioal

differentiation' i8 possible only 1t the tunotlonal structure of '

the distribution area of the population (.llekseev. 1984) and.the

population l~self are regarded as'a tunct~o~l unlt1 (SchWarts et

al~~ 1972) •.

- 4 '-;:.,"

The papers; in ~h10~ the genetlo-bloohemioal dltferentiation.,; .. \ .. '~':. I •. .

1$ oonsidered against the background of the eoologloal struoture

of ' th:~" ~~P~~liO~ ,:~t; ~ine 'and; ooeanle' spe'oles. are still rare. .'-.,. .. ." ~ .' , .' .'

(.uek~e~v' 01. :8.1; ~:: 19'19).: ';', ,. in'the."pr~8e~t~~~~ra relation of the genetto-bioohedeal

..' :. :-and phYsiologlcal intrapopulatlon differentlatlon i8 ana11sed, and

, the dat~'of;t~~ ~e~et1o-bioChemiealstudies of tha populatlon

struoture are interpreted proceedlDg from the ideas of funotlonal

struoture ot'd18trlb~ti6nareaaand s~asOnal 0701es, ...

IUTERllLS .1ND IolETHOnS

The ~aper ls based On the data' of lntrapopulation genetio- .

bioohemieal ditterentiation of, the sprat, in phenotypes of malate'. ". • 'b

deh7drogenase (KOv&l',1979, ~986), of ,the PeruTian horse macker.l

in looua Est-1 '(Koval, 1986) and of Stenoteuthis pteropus ~. in, ' ,

isotermentl0 speotra of esterasee (Koval, 1984). Pol7morphiem in th.

above-mentloned loouses ls known tor other speoles wlthin the,

respeotlve familles wh10h proves its tenetie or1gin (Kolpin. I&li-, '.:. , .. ' . .

nina, 1983; Jorstad. Baevdal, 1981; Alferova, NefedoT, 1973& XOTal,'t " •

1983).RESULTS AND DISCUSSION

TheNorth Sea sprat. Tha data obtained during the 1973 spawn-, '

Ing season'indlcated that the malea and femalea dlftered in tre-

quenor of phenotypes of malate'dehydrogenas8' iemaie. ha~ a htehe~frequaney of phenotype BB. and males - of heterozygoue phenotype'

AB (Koval, 1986). A slmilar relation pattern was oharacterietio of

the Black Sea aprat ( Kalnina, Kalnin~ '1986); Aocordlng to our data~. ,

the divergenoe trom the equl1ibrl~ phenotype ratlo'took plaae in

March and November. In both cases t~e exo8ss of heterozygotes was

reoorded agalnst'the background ot'difterences in slze (in November)

and physiological condltion (ln March)~

- 5 .'

!he main Ilpawn1J:lg grOundll of the Ilprat in a11 the distribution

'areu are 001l'Z1 100 be timed 100 openwaters (FoUer et al., '1985J' ,

reldm4n, 1986). !he open part of the North Sea, where we oarried

out our .tudi..,. i. the prineipal Ilpawn1J:lg e;roUnd of the '.prat ,

of thi. area.' . '

I

I

I

Pig. 1 depiet. the variation Of the frequene7 of heteroz7soull

phenot1Pe KDa-U and' of the sex ratio during th~ spawning. 'Tbe

~3pawn1ng eontinued from lebruar,y 100 June. Th1s oanbe' s~.ste~from the value. of sonosomatio indioes in females (fiS. 1a). Tbe'

faotthat at the beg1nn1ng of the Ilpawning heterozlgoll1t7 \a8

higher in'speeimens Bt stage IV than in specimens at at'age. 'II' an4

nI (KoVal, 1986) can exp~in ,a gradual 4eorease of'heterozygos1tl

in a IlpawU1.ng population fro~ Maroh 100 June (fig. 2b~. 11; Ilums' ,

like17 that the IlPswning eontingent 1s being recruited bl the

spe01mens with ~elll7ed maturation during the second halt of 10M '

~wn1J:Ig season. At thetinal phue of the spa1l'Z11ng, when the,

t1ah start 100 teed intensiV817 (Jul7-J.u.guat) , , the above-noted ten­

,dene,. 100 a ci:lange of hehros,.go.1t7 upaets. , In November. apparent17

betore w1ntering, the mature eontingent 1s being reoruited bl first• • ". • • 4

maturing s~ec1mena. fhe proportion of heterozlgo.1tl 18 being

lncreaJled among the lII&1e. andr~c~ t,s : ,,' (lt~V~l, 1986). '

e ',' '.rhu.e ~ prOc'eed1ng froll the 1d8& of the open' partof ths N~rth. . , . . . - ..

8ea aB of the ll&1n spa1l'Z11ng'ground of the NQrth Sea'.prat. the

lntrapopulatlon ~enet1e-bl00hem1Caldiff~r~t1atl~~'oanbe r8~ard8d, '

as a result of regrouping of speell11en8 of the two sens dUtering ,

',in growth &nd maturation rates.,. . . .

Sthenoteuthill pteropull of the ~8tern Trop1eal Atlantic.

, Genet1e':'bloehell1~;"l studies sbond thtit Sthenot~uth1l!1 ptero-

" - 6.' , 1"', • ~ i \.:.• ,. •.

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

,pUS of: the' Eastern Trop1oal Ulantio oan be Bubd1v1ded into three

;;ö~~~'~~th~p~~;'t:i.~~ii'oont'inuouB values of frequenoes of eshra-

~e'~h~~~ri,'~~;(KO:';~l;~ ,1984). Prem the material's for'1977-78, the•• • , .< , .••• •

;~tio:ot-:tr~q~~noy.;~f th~ ph~notype AA.BBo and' th. peroentage of;: • ~ • • ... '. ,.. • 'l' ". • •

large temales, in d1!fe~8nt areas of the Eastern Trop1cal AUantic~ • * • '.> . , .• ~'. .

'trom north'to"soutb.:'canb'e se,en,(fig. 2). The oorrelation of these

,ohAraoter1sUo~• ',18' obvious.', 'Evident17 the ßeneUo-bioohemlcal" -7' ' ..,:" ~... ? < ~ " • • •

di!!erenuat~on.is,s~~,t~ th!l d1tferentiaUon of the squids-' •

r.veale~ f~om,biolog1c~ c~aoter1st1os - into equatorial and'," •.... ;: f·.

per1pheral groups,dlftering in length of temales (Nes1s~ 1977).," ....- - . .

, i. ",' ~', • '. ",' .',.;

, Tabl.,1 gins a.ratio of the homozygous phenotype AAJm' and" '

'the het~~~lgowi phenoty.pe ,.u,']3B' in Bqu1d samp1ea trom the area'

":t the Gu~~a'H~llo~'~~ll~~te~~ :r~bruai7-AP~ll'1978dUl'ing the'

," p'e'r'i~d" ..he~ 8.1{'t~e~ "~OUPli '~f Bquidll' ~~re 'p~esent 'there. A gene­

;lI\l:,'r~~~i~;18 o~cteri8tiC of '~ll ~he three grouPSI a highe'r '",' ."',", ,', '. I>

h~teroz;ygoä1t3', in ~le~ and smaller temal~8, aDd a higher fr~queno;y

",ot'the:~be~~tri,e~Ium.: b luge temaleB~ An analou cu be dra~'. ,w1t~ t~~'~~ekeye h~~~;"1ri."~h1eh' small earl,. ~tUl'1Ds ~es hau a~, .)-:'... ;.:.,.:.... : .. ,..,','... ,"',.. \.' '. . ," . ,: ,- " "

bigher heterozygos1t;y'(Altukhoy et a1., 198). ~1 analoS1 with'

salmo~,"~~otlldeUiier(Nes~8, '1977), IIma1~ e~17' ~tUl'lJig male~keep to 'th~ rep~oductivepart'ofthe distribution ar.a wh1l11 large

iemales p~rt6rm length1 teed1ng'~~tiOns to the per1pher,y'of ~dll1tr1btlUon areas'.As '1s evident f~om table 1, in Februar;y-Ap~ll '

the proportion oi· males and email. fe~bs among the squid,s ot the, '

"southern" group.1s greater th8.n that ot~large temales. Inaccord-. . .. :' ," '. '. ~ , .. .."."' .. :.. :, .. ' . ~., ' .. .. .

anoe with',the idea of a tuneUonal structUl'e of the dis~r1buUon

'area (Zuev et' al.~' 1985)il~ge !llmales oeeur iß'the peripher,. oi. . . , . .

the distribution ~~~ at 15-17°5 duriDg that pe~iod,the warmest• • \ • • ." ' 4. ~ '; ~

, ,J

- 7 -

in the.80uthern hemisphere (fig. 2). Large temales of the "north-

distribution ot.gentically different speo1mens in distribution

areaof squid groups. Obvioualy one oan speak'of a non-randoDl

. (-

ern" group·can.be observed in the periphery.of the distribution'.' .

area at 15-20oN in September-October. (fig•. ~).. .Thus, the genetic-biochemical differentiation of ~.pteropus

i8.cor~elated'with t~e' functional' stru~ture of the distribution

. .

studiea showed a ditferen-

[..•~r,eaSof.~aoh ~f the, thre~g~~ups.

; Peruvian horae maokerel •. Earlier'.

" tiation of the Peruvian horse mackrel into' the', "northern" and"

. , ."southern'~ popula.tions .w1th var;ylng. ratio of phenotypes of the- .: locus esterases Est-I. Compared to. the "southern" population. ,the

.. heterozygos1.ty in the "northe~".population i8 higher in a11 age'··,'. ,"

:~oups, beg1nning with one-year-olds•. Be~ides, the fOllowlng.pecu-

iiarities'of the intrapopulation differentiation in the loous '. , ,

Est~I can be notedl a faster'growth.of heterozygotes Be among one­

year-olds ~ompar~d.'.. to, ~mozygotes.M,. and disttirbanoe of.the'

• equllibriwnratio of phenotyPes dur1Dg the pre';spllWIdng season, .:

(Koval,1986a).

After,the example of .the horse mackerel of the "southern"·

population, aooessible to'the throug~o~t~the-yearobservation,' it

~iS sho~ that the seasonal differentiation 1sm~re olearly pronoun~

cedthan the differentiation by sex (table 2). The increase of the. ,haterozygosity during the pre-spa~in8 peri~d'~s asswned to be

related t~ the fact that the heterozygous specimensbegin to torm

pre-spawning aggregations earlier. The oomparison of the speoimens

at different matur1ty stages in the beginn1ng and·in the. end of

spawninc showed that in the taster maturine speoimens the·propo~ .

. '. '. ~ .. ' - 'S ,-: .'

' .....t10n of thephenet;yp8'BB ls'smS1ler 8nd the proportion cf hetero­

Zygote.' B~' ~&8te;'t~ in ~ho~. w1th delayed I118.turaUon (f18. Ja)." ' ". '. '. '. I . '

On the whole; iho "southern" population 18 ebaraoterized b7 tha~

at 'the first '~has~' ~f the spa~ ..aB'on (Ootobe~Nove~~er) in

ih~ nOrthern'~ .'o·l ;t~e 'di~t;ibut1on ~ea 6;-3605)~ whe're the ', . " , ~, : ' ,,-.. . .

spawniIlg b,g1ns, the raUo of, ph,neiype. RB/BO 18 .lo:wer than ~ater

in the' areu, where 1t eont1nue~ 1r1th' the eollÜJl8 cf, the southern .•' '. :> • ,

hem1sphere 8UIIIllle~ (f~.,3bh ~~ .change,:~f ,the ge~etie s,tnoture em&7 ind10ate _,change of the spawning.contiIlgent. The ra~ics cf

phenotypes BB/BO are the lOll'est ,in 'the north-western part of theo • , o. •

~1stribution ares ~ Ootober. lI'here the spawning ~f the large.t,

.peoimena of the population begins earlier than 1ri other parts of

the ares.

Xbus. the ~eaaonal genet1o-b1oohemioal d1fferentiation of the",' .

Penvian horse maokerei 'of the "southern~ population 1. the reali-.

zation of physiologioal differentiation ol genetioall7 Tar,11ng

speoimens under the eOnd1t1~~Of th~ pr~traoted spawn1ng season

and differential distribution of the speo1mens~in sex,

lengih and physiologieal oondition.

1'18. 3e shOws a ratio of phenotypes in the horn I118.clcerel of

iho area between' 20° and 40°5 dur~~the 1984 8pawniIlg saason. The

sam;>le. are presented ,bl the .speo2,Jnens with II1m1lar age OOmpO.sit1ce. ..._,'

How,ver in the area to th, n,orth of. 30°3 ihe, sP':wn1Ilg beslns earl~er'

than'in ihe area 80uthWard öf )0°3. Neverthele.s, the'differenoe.'

in the ratio of phenoi;ype8 between the northe~ and southern group•., '

.ere of the other 1c1Dd ihan the7 mtght hATe boen in tho group. of

one and ihe same' population. ! • • hown alter the ezample ol ihe, .

80uthern populat1on, the faster maturing specimens and groups

t."I

,

I'

Iwould hl:\ve had lower values of tho ratio of phenotypos 1m/Be

... '. . -.' .. '. . " ' . " . '. ~ I

'~f1g. Ja.b). Th1a oontrad1otion oan be attributedto the fact

, t~t the groups oono'e~dbelo~'to variOWI P~~~t1cn. dUfe;1ng

'in reproduotive 070les (.uekseeva. ~986) an4 ret~n1ng the1r gen~, • '-. "•., •.,j. . • ",

Uo spec1t1t7 du.ring the live ;rem of observations (JtoVal~'i1986a).

CONCLUSI01I3 '." : ': -,

, 1. !hlll anal;rs1s of the l1terary andour oYm data shotlS a

~OO~11oat~d correlation b~tween the b10ohemicalpOl1m~rp~man4. .' . .. '. . .

. th. 8ex, growth, a.nd mature.tlon. .~ . ,.~. ;, " ...'. • < .'\. '. , { .' • :' .• ;,~. .'

2. M' a result of this crorrelatioD. the sp.o1e~ with a pro- ", I." . . - . • ~ . .

trBotedspawn1ng season'trom a rUnoti~nal~,8truCtur1.ed'd1str1b~..:. • '.. • .'1. . ......' • " ',....,. '

t10n area manifest intrapopulat1on differentiation in trequences ' .of ~henot7Pe~.), ' ",', ,.", , ) ".., ,

;.. ',"

1" j.!lie .interpretation ol tha gl9net1o-bio~hellid.oai,d1Uue-nti..•.• , . • • _ ", ,." t .'"

l. tion is pOlll8ible onl;r if tbe distribution srea cl, the po~~U~n ". ~ .:. ': - . . , . '. .

, . 18 ,regarded as. a funotio~~, llItruoture.

......-------------------------~--- -~ - ----- ----- ~-

10 --, '

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1~ Alekseeva'E.I.;1986~ Coniparat1ve characteristio of maturing

and spawning of horse,ma.ckerels of, the g~nus Trachurus of the

: Atlantio Ooe~ ~nd the P~citlc. "Lite ,cycles.,distrlbution and

migrations otthe commerc1al tiSh~~ ot the Atlant10 Ocean and the

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•.

AtlantraRO, 5-19.

3. Alekseev F.E., E.I.Alekseeva, N.V.Tltova,~1979. Pol~or~

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the tishV L., Institute ot Cytology ot the Acad.' of S01. of the \'

USSR, p~ 58-63., ,

, 4; Alferova N.M~,. Netedov G.N.', 1973. Electrophoretic studles

of musoie esterases of some ~st Atlantlc species."Blochemical' "

genetios". Inst1t~te of, oytology ~'f ~c~d. ot So'..of th~ US~R.: L: ,':p. 195-200.

,. ~., .5. Altuchov Yu.P~,' 1974. Populatio,n genetlcs 'ct the t1sh.M.~•Pishch. prom., p. 1-'247.

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struoture and 1ts relation to 1ntrapopulation different1ation 1n', '.: ,. . \ .sex, .age and growth rate in the P~c1t1c aockeye salmon Oncorhyn-

~ nerka (Walb.). The genetics," vol'." x~x. No. 5,', p~: ~9~~807 ~.:7. Danzmann R.G., M~M.Ferguson, 'F.W.Allendorf. K.L~Kludsen;

, "'.' ,1986. He~rozygosity and development~l rate in a straln of rainbow'

It

~,t

III

~,Ir

t.III

II

", ,11 -"

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of the, !iah", ,L•• Ins t. ,o! Cytology of the 'Acad~ of Se. of the" ,ussa, :p.111-115.

,,, ,\

9. Dawson W.A., 1986. ChAnges in western mackerel (Scomber.. , ~ . - ,. .

~,sCOmb~S), s~awnin8 stock compositi~n during,the apawni~,aea~on~

J.l.lar. biol. ,Ass. U.K•• v. 66, 2, p. 367-383.' " , '

10. 'Feldman V.N., '1986~ Q1ielq~estraite particuÜers dela

repartition et,de la biologie du sprat de la merdu Nord en fone­

tion de la differentiation a l'interieur de la population. lCES.

C.M. 1986/U132, p. 15.

11. Iljin l.I.:,A.S.Golubtsov, 1985. Eleetrophoretic analysis

of isoferments of superoxiddismutage and octande dehydrogenase in,

interbreedings of "Amur sleeper" Perccotius glehni Dub. Communica-:­

tion II. 'Inheritance of electromophs and dependence of survival on, ,

'the genotypea ofspecimena. The Genetics, vol. 21~ No. 2. p. 1542-,,'

1547~

':12o'J9rstad K.E:. G.Ilaevdal, 1981;,Ensyme polymorphism of

sprat fr'om Norwegian Waters. "lIprel1minary ,results". ICES;' C.Il.

1981/HI,65.',

'13. Kalnina O.V.,',V.V.Xalnin,' 1986. Correlation between the,

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, I.lDG in the Black Sea sprat. Thes~', o~ Rep~' ~t the Jrd All-Un1ol'!

'Conf. on' g~netic~, selectio~ and hybridizationof' the' lish. M.;. " "'.

14.' Kirpltchnikov V.S., 1987.'Adaptlve value of blochemlcal

poiymorphi's'm. "Journal of General BioloQ''', XLVIII, llo. 1, p" )-14 •. ,'

12 - ." , ....'" ','

15. KovaJ. L.I.~ 1979.,Il1traspeo1tlo'variabll1t,. of,non-speol­

Uo e!'te~~~ and malah ~e~ge~~;~ th. North ses aDd ~ltio.. ,

sprat. "Ecological ph,-s101ogy end. b10cheminstr,- cf.th. ~~sh"; val.

,2. ~etrakhan. TsNIORXh. p. 215., '., ..

16~ 'Xoval L.I.. 1983. neotrophoreUo'analysis of p~te1n.a '~t .

squ1ds. of tami~ ·Ommastrephide.e•. 3ystemaUcsand eoolog,- of oephalo­

podell • ZIN of .lead. '~t So. of the USSR•..L.:.•.. p. 141.. . . .., . .

17. Koval L.I•• 1986. IntrapopulaUon d~ferent1at1onot the

North 'Sea eprat Sprattus sprattus L. in phenotypea of malah da-

hJ'dro~ewa. ICES. C~I(. 1986, 1l1n18;ymPO~1~'- N 2,' p. ~-18,~ . '

18. Koval L.I•• 1984. Genetio-b10ehem1ca1 differentiat10n of·

Stenoteuthis' Ptero~U8. "Jtntr8a~eC1f1o d1fferent1&t10~,ofmar~e .

oommereial t1sh~ inv~rtebrateS";'Eaibüngrad. Atl~tmo.I

••p. 77-82•. " .

. 19. Koval L.I •• 1986&•.!he analys1s of intrapopui~tion'var1a- "

blllt,. of.th. ratio of'~sterase Pheno~~es in Peruv1an hor~e'~eke~• ., I· •

10el Traehurus symmetrious ~hyj.. Thes. of liep.· at the 3rd J.ll- . ' .'

UnionCont. of Genet1~, 5eleoUo.n ~d Hybr1~i1zat1~n ot..-t~. 11~h.: :.':.'. ... ,

,',\ ..

20. Kolpin v.v.• O.V.Kal1.rl1D8., 19a2~ Po17ßI0rphlsmof malate.. . " '.debydrogenase' ofthe BiaekSea sprat· 8.Iid ~o/lle' genogn.ph1~ (;~~a';':':

..... ~. " . • >'" • " ': '.,~, •• . ,,",

riUes. ".B101ogy of shelt watere of the World Ocean". DVlro of the:

~cad of ~o. o~. ~h&' ussa~' nadivoatok. part 2,' p~ Ti-79~" .,<:':.. -:' e, . , . ~ - ...

21. Nss1s X.lI•• ·, 1~77 .'. Intraspeo1.fl0 groups 01 equide. Thes,.: .

of ~8p. a1; the All-Union Cont. on Utll1zation. oi Commerolal Inv~~'. .. ' : .tebrates tor :l'ood and lleal PrOdueUon &nd Techn1cal' Purp~8.8. 11•• ',

VNIRO. p. 53-54.

22. Pott~r I.~•• P.K. Clarldge; 1985. Seasonal oatehes, slBe

and nerlstl0.data tor 8prat SprattuB sprattuB in the Bevern

, -\ 13

••tWU7. J ....r. biol.,.1sa.U.JC., T. 65, J, p. 667-67S~

23. Shoherb.nok lU.I., 1973. Oorrelat10nof pol7mOrphOU8 878­

tem8 of .sterase8 end transt.rrlna with economioally important

. ohar~ot.ri8tios ot the carp.. MBioch.mioal g.netic. otth. f1Bh".

L.~ Inat. of Cytology of th. ~cad. of Se. of the USSR, p. 129-137., ',.

24. Sohwarte B.B., B.D.GurTltch, ..V.G.I8hohenko~ V.:r.So.in,

1972. lUnctional unit7 of the population. JQurn. of General'.. ~

Biology, vol. 33, No. ,1. p. )-14.

~. 25. Zuev V.G., Ch.K.N1gmatullin,'V.I.Nlkolsk7,:198S. N.etonie

oeeanie squids. M., Agropromizdat, p. 1-22).

14 -

Table

AABBIRatio ot esterase'phenotypes ~ (1) • and ratio

of specimens of different sexes and size (2) in three', ,

groups o! ä.pteropus in the equatorial zone (II-IV, 1978).

males 20 cm' 3.96 39%

temales 18 cm 3.71 20%

females 18 cm 5•.18 41%

. * -, mantle length

, Groups,sex,size·

"northern".2

~...

"equatorial" "southern". e" 1 2 2

0.29 40% 0.81 34%

0.20 46i'b 1.14 ' 38%

0.58 14% 1.26 28%

- 15 ;..

~requeno7 of h~c~-oz7gotOU8 phenotype"BC

of Bat-1 looua in Peruvian horae maokerel

of the "aouthern" population by month

Monthl Males t8t· P'emales Mean tst

-VI-IX 0.515 :.!: 0.012 1.3 0.492 :.!: 0.013 0.505 :!:0.009

0.468 : 0.006 0.465 :!: 0.006+ . 4.0·

X-lI 0.4 0.466 - 0.004

0.462 :.!: 0.016 . 0.460 :.!: 0.023 0.462 :.!: 0.0130.3

III-IV 0.1

Nean 0.477 :.!: 0.005 1'.6 0.469 :.!: 0.005

• - p 0.95

" tat - Student' B oriterion

, J'1g. 2.

- 16 - ",\." ~ .'

J'IGURR LEGENDS

, l1g. 1.' GOnOs~maÜ~ 1%id1ces of f~male8x (a), frequeno1 of hetere­

"~igotu~ge~o~1P~MDC.1B (b),' per cent ol _leB (0) 1n

:samples of the North Sea sprat (lI-XII, 197) •

.x -'all niaturit;r, stages' except tlW&e with the lQ-dratation, .,features.

:Fx"equeno;r of Phenoty.p8,' of esterases üUt (a) and .per oent..

of large females (b) of A.pterqpus in different areaa of 'Ir, '" " '~. .

··the eastern trop1oal Atlantio (JZarch 1977-Aprll 1976).

Area " 15-20oN (September-october)

Area 2, north-westward of the Guinea hollow (September­

March)

Area ),' equator1al area (March, ~ebruar;r)

Aree. 4. 80uth-eaatward of tbe Guinea hollo" (lebru&r1-" 'April, June)

Area 5, 15-17°5 (Maroh)

1!g. 3. Ratio of ma1n phenot1pes of esterases in PeruT1an horse' ,

mackerel

a ~ in speoimens d1ffering 1n maturit1 'stages in the

b~g1nn1ng (1, 2) and in.the end () of spawning.

1-16°S, 2-)8°S,' 3-450 S. Shaded are .ltforestallinglt '.

maturit1 .t~ge.s. ,(IV in thlT beg~_ ot the ~P8.wn1Jig eand VI-II 1n the end of the spawning)

. . '

b-during the spawning season in the Bouthern population

(southward of. 32°S)

1 - 1n the beg1Dn1ng (bc~ober-N~Tember, shaded) in.the

,m1~dle (Deoember-Janue.I7), and in the end (Pebrt1ar7~'

April) of the spawning

I

~

17-" I

2 - in lat1tudal sone•• 32-360 3 (ehaded),38-400S

&DA .1-46°8

, - 1D Ootober-Nov••ber inthe areaof 32-)6°3. at

1)00. (_bad.d), 114-95°W,'86-800W. -.-",

o - dur1Dg the .pawn1ng .euon of 1984. northward of, ' ,

)0°3 in Ootober, and .outhward of joog in November-

Deoe.ber, 1.e. dur1D&, the !l1III1lar phase. of the

spawniDg .....on.

- 18 -

,.7.0 ,....:. a5.0

3.0

1.0 ~

" AB . f ß50

45' . f J t40 J . t f J.

"l! tc

f t I50

f I4D

3J

20 J

n· m (,)' ,)' Yl .YIl ,)J XI

- 19 -

- AAlB1.0 !

a

t I0.6

•0.2 t

I 2 3 4 5

" Cl>1

I· .8

60 I40 I

aJ t •2 3 4 5

P1g.2

- 20 - .

2' 3

8

2. 3

• Cl

••

•• • .a•• • ••• •• •28° ~ 36° ~.•.

..1"1g.)

!tBe

1.0.

0.8

0.6

t

BaBe

0.8

0.6

t

SBBe

0.9 • •• ••

0.7

aT' 24°

. UCl,IlIDtCaHO Jt nSllaTl'l 06.07.87

06~eM1.25 n.~. ~OpNBT 60x84 1/163Üaa 29011­TKpaa 215

POTanplllHT lIll1POI07140, MOCRB8. BepXBJUI KpaCBOCen.CKaJt, 17