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International Council for the Exploration of the Sea

Ce

E.1971/F:26

Demersal Fish (Northern) Committee

DIFFEHElITIATIOlT BETW'EEH 11 11A.PJNUS n AND ^It 131ITELLA 11 TYPES OF :;::;EDFISE BY ELECT:ROPHORESIS OF HEHOGLOBIIJS

By

Gunnar Naevdal

Institute of l·.i:arine Research, Bergen

INTRODUCTIon

LlhLdbeck

(1940)

claimen that besidws the common redfish, Sebastes

marinus (L.), there excist a deep sea type in the Bear Island area and on the Iceland-Faeroe ~idge. Travin

(1951)

described the deep sea

redfish from the Bear Island area and the Barents sea as a new species, Sebastes mentella Travin. This species pref'ere deeper vlater and

somewhat': higher- temperature than S.marinus, although they also occur sympatrically. S. mentella is characterized by cl. beak on i t s lower jay{ ¹ a.."1.d this species possess a greater eye' diameter, grOirls slm'reri a..."'1.d reach a smaller total length than S. marinus. The colour of mentella is

bright red ~"'1.d of oarinus usually ora.."1.ge-rod

Andriiachev

(1954)

a..~d Templeman

(1959)

restricted Sebastos oentella

~ ~

to the status of a subspecies, S. marinus l!1entella,because the differences bet'ween marinus 8.:."'l.d mento11a ,'{ere not clear enough to

justify thorn as t1'lTO distinct species. A.o. Tompleman

(1959)

found that mentella was not restricted to the Northeast Atlantic, but ''las common also in the lTorthwest Atlantic, and in nost localities here occurred in greater quantities tl-l.an marinus.'

By meristiC, morpnometric or morpholgic metods clear:cut separation of redfishes liito marD"'1.US and mentel1a groups has been difficult

(Kotthaus

1961,

Templeoal1.

1959,

Kelly f Baker 8...."1.d Clarke

1961)

¹¹ A great proportion of the specimems posessed the characteristics of both groups in a varying manner. Eo"t'rever: genetical differences bet'\'reen the two

2

groups vrere suggested by TemplenaIl an.d SaJ.1.denan

(19.59)

who found great differences between mar~1.Us a11.d mentella types in relative occurrence of caudal melanophores of pre-extrusion larva~o

~ Ym1.ulov

(1962

a) found si~1.ificant differcnces between the two types in infestation rate of various parasites in parts of the North"\,lest Atlantic.

By imr::J.u...l.o'::'diffuison tech ... '1.iqucs, photron-reflecto:r.J.etric measurements a11.d two separate methods of chromatography 6 ^JRourke

(1961)

fOUl1.d evidence for biochem..caJ. specif'icity of the t,,·.ro forms to such a degree that they could be considered as two distinct species. Similary Schaeffer

(1961)

£ound that the two types differed significantly in content of certain free amL'1.o acids m1.d total nitrogen content in ouscle tissue.

In a series of papers (AI tu.1r ... hov et al.

1968,

Al tulc"hov and IT ef'yodov

196[,.

~ Al tulr,..hov, IT ef'yodov a..""ld Payus ova 1968, N efyodov 1969) dealing with

the redfish problem, "lOre described differel'":C1!0.§ in mus cle te;r'l"!lOS tabili ty, and also differences in frequencies of sone polymorphic serun proteins characteristics (haptoglobil1.s, albumins, aJ.1.d - g1.obulins) bet1wen sanples of mari..nus and r.aentella ty:pes froo t:;'le "I'I1"aters between Icela11.d and Greenland, ShoYling that the ful.alysed sanples were i1.ot drawn from one homogenous population.

In the present paper differences in electrophoretic nobility of

henoglobins of the two types of redfish are described, and the results are discussed in relation to their significance on the systematic and the management of the redfisho Besides 1'1eooglobins, total ^serm~ and mucle proteins, seru.. .. ^':1 and muscle esterase, lactate dehyd.J::'o:5'2;:-.l.B..se and aspartate ar.1inotra:."'1.sferase ,,'ere analysed by electrop~l.oL";)s:i.G~

The result of these analyses, ho'vever, ·were not vcsry eC:i:lcl-c.,s.ive, ruLd are omitted in the present report.

3

l·IA.T:;!;RIAL LIlD l::IETEODS

B.edfish for blood saopling was caught by. bcittOT~·.t t:-&.w.J." 'Blood was obtained by a syri..."-ge fror.l the heart region or the fish

... "as cut open and blood collected into snall glass tubes.

After centrifugation the seruw was pipetted off lliLd the cells lysed by adding destilled w·ater.,

The hemoglobins were analysed by the agar gel electrophoresis at pH 7.2 described by Sick (196.5). Selected speciDens were also fuJ.alysed by the conbined starch and agar gel electro- phoresis at pI{ 9.0 described by Doller (1966) ^$ TI1.e henoglobins vlOre always al1.alysed. wi thLn. 24 hours, and usually wi thi...L 12 hours froE sanpling, and part of the naterial were reanalysed after one or t·wo days 9 Sone henoglobL"1. specinens vlere also

frozen and later reful.alysed at the Institute of' Harine Research in Bergen. The henoglobi...l.s were stained by Amidoblack 10 B.

All sa~ples from the Barent'Sea were collected ful.d analysed during a cruise by

Riv

"G.O.Sarstl in Novenber

1970

and

the saoples from Icelllil.dic waters lil:ewise in hUgust

1971.

Fishing localities, depth, date of saoplu1.g and nunber of specinens Ll. each saople are listed in Table 1, and fishing localities are also shown in Figs. 1 ID1.d 2.

All redfish specinen vlere separated norphological.ly into Sebasted viviparus (baclGlTard directed fu""lterior p:':<~~~o~orcular

spine (hudriiashev 19.54») ¹ nentel1;,§: type, E3E:].-~'.::·L::'::<." type

and intero.ediates (according to thG c."Gc:ri.bticH'"l by T::~:::'·Y-i.n(1951»;

and separation by :oorpb.ology was 0<)":::~J<~_:,>.,0r,0 tl."le resar..':::S froD. the hGI::lOg1obin a."t1alyses. Total 1.3LZt:.l/ 1)3ually cclso se;c and age: were recorded for the :fi.sl.:..es of' which blood was collected"

i=ffiSULTS

I-IEHOG-LCBIH T'£P~S OF ?-2DFISIi FROM TI-:I:i5 BJ.!.a~lTT SEt.!.

Anong the samples from the Barent Sea were found two nain henoglobin patterens, Fig. 3 • The first pattern occurred in specimens

,l-rhich according to Travin (1951) were of the I:1entella type. In agar-gel at pI-::: 7 .. 2 this' pattern shov;ed a strong but rather diffuse ,anodic moving component, fu~d one weru: component stayed near the

origin. By conbined starch and agar gel electrophoresis at pE 9.0 the strong conponent shm..;red high fu~odic mobility, two weak conponents also shovred anodic nobility llThile one very vreal: conponent noved

slightly towards the cathode~ This pattern was tentatively called,the nentella pattern.

The second pattern was foUnd in specinens morphologically deternined to be of the narinus type, and consequently was called the rlarinus pattern. At pE 7.2 all components noved tOvlards the cathode! tyro weak

components showed the highest nobili~Y1 and OITe strong componentr moved only slowlYb Individual variations were indicated in strength fu~d

occurrence of the tw'o weal;: conponents, but this variation was not clear enough to foro the basis of clear-cut classification of the specinens. At pE 9.0 the strong conponont of the narinus pattern noved slower towards the an.ode than thE:{ strong conponent of the

nentella pattern. Also in t~l.e narinus pattern was seen two vreal~ COI:1PO-

nents, one with internediate fu~odic nobility and one with slight cathodic nobility.

One Sebastes viviparus from the Barent Sea (sacple 1) s:::l<.nred the sar..1e henoglobin patterns as inviduals of t::'l.e t:1[.Lrinus type.

Sone specinens possessed norphological cL.u.r2:.cteristics both of i::l.arinus and nentella types. This l'ras expccially evidcnt at one iocality on the IJorwegim~ coast (sanple 8) ehere the greater part

of the specinens werc'recorded as such t1interoediates^lt (Snestad, unpublished). EowGver, vi th no exceptions all these specinens. show'ad

the narinus henoglobin patten~.

The ru~alysed specli~ens varied LL length fron about 15 to 40 cn,

m~d both henoglobin patterns were observed aoong the snaller as well as anong the greater specinens. Consequently, no indications

of ontogenetic variation in henoglobin patte~~s were fOCULd ll~ redfish.

The he~oglobin pattern could be recognized also after freezing and thawing of the henolysate, but the clearest patterns 1~S

obtained with fresh naterial. ".: post, Dorten variation, except that the total patter~-s became di-ffusEi' could be observed after prolonged storage in the refrigerator.

The distributions of the two different patterns ih the sanples fron the Barent Sea are shown in Table 1 (sanple 1 - 8)0 The nentella pattern w-as found nost frequently in the area betwe~n

Bear Island and Spi tsbergen and we're rare in the eas tern parts of the Barent Sea and near'the Norwegian coast. Both were found together in the sane hault, but nentella occurred more frequently in deeper water. This is in accordance with the general appearence of the nentella type of redf'ish for instance fron the des crib- tion of Travin (1951).

HEHOGLOBIN TYPES

2E

REDFISH fE..Q1:! ICELA.NDIC WATERS

Sinilar nentella and narinus henoglobin patterns occurred aoong redfish sanples fron Icelandic waters as in the samples froe the Barent Seao In sone specinens a nodified narinus pattern ^occ~red~

This pattern showed an extra conponent which in agar gel at pH

7.2

noved slightly towards the anode; and in conbined starch and agar gel electrophoresis at pH 9.0 noved towards the anode with a nobility greater th&L tho connon strong conponent of the mentella patterna

Totally 88 specinens of sample10 and 19 specinens of sanple 11 which were supposed to be ~. 'viviparus fron their baciDNard

directed-preopercular spines, all showed one of the two narinus patterns.

Few nentella patterns were found in Icelandic waters. West of northern Iceland (sanple 11) were found three specinens out of

159 which showed the nentella pattern, and the fishes which

possessed these patterns were easily distinguished norphologically.

West of Reykjanes (sanple 10) were found no nentella patterns, and on the Iceland ~ Faeroe Ridge (sa~ple 9) were fO~Ld 9 nentella patterns out of totally

38

specinenso In this sanple all but one of the snaller fishes (9:13 en) showed the oentella pattern;

6

and also tP....I ec fishes about( LW CD in length) showed this pattern.

Ho.weye.:r:, L"1. contrast to all other saoples 1 norphological differ ....

ensiation of the fishes silowing the different hemoglobin types was nearly iopossible$ even for the greater specinens.

Four specimens, two ll~ each of sanple 9 m~d

1o,

showed a heno- globin pattern .vith both the cOLlponent of nentella ond Lmril1.lls patternJ Fig. 2. Morphologically these four specinens could not be distinguished from the oarll2us type, but because the pattern indicates hybridization between individuals with

different henoglobin types, this pattern tentatively was called :the lIhybridl! pattern. The fishes wIrlch showed this pattern

ranged in length from 36 to 45 cmo

DISCUSSION

The compositions by anino acids of the"proteins polypeptide chains rukd consequently the electrophoretic mobility of the protein~

ar.e contrOlled by genetic factors (Mmkwell a..'"1.d Baker 1970) ⁰ The electrophoretic patterns of fish heooglobins usually are species specific: illustrated by a.o. Tsuyuki ~~.

(1968)

who described species specific electrophoretic patterns of henoglobins of 28 species Pacific Ocean Scorpaenidae. Intrnspecific variations of hemoglobL~ patterns, controlled by co~doninant alleles_l have been described for several-species (see de Ligny

1969

for references).

In other species: a,o. saloon (Kock, Bergstr00 a..'"1.d EvrulS 1966) pnd herri...'1.g (~filkins and Iles

1966)

intraspecific variations have been observed to be connected with ontogeny. Also variations

due to changes in heooglobin components after prolonged sto~ge

pf samples have been described for several species; but these usually variations are found :in the minor heooglobin conpone~ts

(Sick 1965, H0ller and Hoevdal 1969, Tsu-jUki.§.!~. 1968, I:Joevdal 1968)0

Neither ontogenetic variation nor post DorteD changes c~~ account for the observed patterns of redfish henoglob:L.'"1.s, because the two connon patterns were observed in fishes ranging fron less than 10 cn to Dore than

40

cn in length, and all specinens were given the sane - treatnent and all were analysed 1".rithin 24 hours f'roD

sanpling. Control specinens did not show ruky major change in patterns even after two or tp~ee days in ref'rigerator or after freezing

and 'thawing.

7

Segregation of two co~doninant all?les witllin one species could

produce tl~ee patte~~s i1ke the nain patten~s observed (the nentella and narinus patterns as h~mozygotes a..'J.d the flb.ybrid^tt pattern as

heterozygote). However, 'according to the Hardy'::"lleinberg law tt'le hypothetical heterozygote (here the Ithybrid^ll pattern) should then be expected to occur ouch more frequently than observed. I t therefore

seeos unli1r..ly that her:lOglobin variation within one species can account for these patterns. But the modified mar~'J.us pattern in redfish froo Icelandic vr.aters may be nomal intraspecific variation,

probably genetically controlled.

Howeyer, the two common patterns may be explained by assuoing that the oentella and riarinus types of redfosh really belong to

different species, each with their OWlL hemoglobin pattern. The tlhybrid^ll pattern nay be due to occasional hybridization between individuals of the two species. The good agreenent~ except ll'l

sample

9,

between the results :from the analyses of henoglobins and morphological differenciation strongly support the theory of two species •. Four hybrids out of' oore than 650 individuals do not

show that the two types are conspecific, because hybridization between related species is not uncomnon among fishes. According to a second theory the IIhybrid" pattern nay represent an intraspecific variant of the narinus type. This teory is supported by the fact that the four specimens with the "hybrid^ll hemoglobin pattern could not be distinguished morphologically fron the marinus type.

Although the mentella and narinus type of redfish seen to represent different species with only occasional interbreeding in the Barent Sea and Icelandic l'Taters, the possibility s t i l l excist that the

two types nay be connected tlLrough intermediate populations in other areas. A definite conclusion about the species of red~ish L~

the North Atlantic c':'W therefore not be drawn u..."'l.til samples have been collected f'roo the, total geographic range of the redf'ish.

However, the marinusandmentella types of redf:l.sh surely repressent different gene pools' with ,a"In:i.nioum exchange of genetic material, and they therefore should. be treated ~~ separate units in manageoent of the Northeast Atlantic f'i.sheries •. In addition both the ruentella and the narinus types oay be composed oi'smaller units (populations, stock units) as clniocd by 0..0. Sinderrna.:.-m (1961) and Yanulov (1962a, b) in their stUdies of parasites and meristic charateristics.

8

The fact that Sebastes viviparus shO'l"led the narinus hemoglobin pattern has no effect upon this conc"}.u.sion.Sebastes viviparus doubtless is a lfgood!! species (Andriiashev

1954,

Trout

1961),

and two related species may have similar henoglobin structure, while

•

others differ widely.

SUMI4A.RY .AND CONCLUSION

Heooglobins of 225 specimens of redfish froo the Barent Sea and

357

specimens from Icelandic waters were analysed by

electrophoresis to search for genetic differences be~.een the morphological marinus and mentella types. Two main hemoglobin patterns were conmonly found, one characteristic for redfish of the mentella type and one characteristic for the Oarll1.US type.

Specioens of Sebastes viviPA£qs showed the oarinus hemoglobi,n pattern. Horphological !!intermediates!! showed the narinus p'p.ttern in the Barent Sea, while some specimens which showed the mentellu hemoglobin pattern but could not be separated morphological.l.y

from the marinus type were recorded in the Iceland~Faerbe Ridge area.

Four specinens show'ed a hemoglobin pattern which indicated hybri- dization between individuals with differnt henoglobin pattern.

The results indicate that Sebastes'oentella Travin is a species distinot from"Sebastes Marinus (L~, possibly with occasional interbreeding. However, for final conclusion about the species of red:fish in the North A tla...""ltic samples have to be collect'ed from the total range of the redfishts distribution.

~

REFERENCES

Altukhov, Ju., Artemjeva, K.F., Borisova, I.V., and Nefyodov, GoN. 1968~

Immunological analysis of serum proteins of redfish in connection with maturation. Res.Bull.into~.Northw4Atlant.~. ,,.2:44-480

Altukhov, Ju., Nefyodov, G.N., and Payusova, A.No 1968. Thermostability of isolated muscle in determining the taxonomic relationship of the marinus~ and mentella-types of redfish (Sebastes). Res.

Bull"int .. 6omm.Northw.Atlant.Fisho, ,.2 : 130-136.

Altukhov, Ju. and Nefyodov, G.N. 1968" A study of blood serUm protein composition by agar-gel electrophoresis in types of redfish

(genus Sebastes). Res .Bull.int .Comm.Northw·.A tlant .Fish. ,

.2 :

86-90.

Andriiashev, A.P. 19540 i.Fishes of the northern seas of the USSR;.

Zool. Inst. Akad. Nauk SSSR,

...21 :

1-566.

Kelly, G.F., Barker, A.~io, and Clarke, G.M. 1961. Racial comparisons of redfish from the west;ern North Atlantic and the Barent Sea.

~. P.-.Y. R~uno Cons. perm. into Explor. Mer, 150 : 28-41.

Koch, H., Bergstr0m, E., and Evans, J"C. 1964. The microelectrophoretic separation on starch gel of the hemoglobins of Salillo salar L.

Meded. K. vlaam Acado, 26(9) : 1-32.

Kotthaus, A" 1961. Contributions to the race problem in redfish" ~o

P.-v .. Reun. Conso permo into Explor. Mer, 1.2.Q : 42 ... 46.

de Ligny, W. 1969. Serological and biochemical studies on fish populations Oceanogr. Mar. BioI .. Ann" Rev.,

1.:

411-513.

Lundbeck, J o 1940. Rotbarsch - Goldbarsch - Tiefenbarsch. Deutsche Fischerei-Rundschau, 63(2) : 32-33.

Nanwell, C. and Baker, C.H oA" 1 970. Ivlolecular Biology and the 0 rigin of Species: Heterosis, Protein Polymorphism and Animal Breeding.

Sidgwick and Jackson, London 1970, 394 pp.

M011er, D. 1966 .. Polymorphism of serum transferrin in cod. FiskDir.

Sy~. Sera HavUnderso, 14 : 51-6J.

-- -- -

M0ller, Do and Nrevdal, G. 1969. Studies on hemoglobins of some Gadoid fishes. FiskDir. ~. Sero HavUnders., 1.2 : 91-97.

Nefyodov, G.N. 1969. Serum haptoglobins in the "marinus" and ^f1mentella^tl types of North Atlantic redfishe Into Coun. Explor. Sea,

Special Meet. ~ "The Bioche~ical ~ Serological Identification of Fish Stocks", Dublin..l.222 (29) : 1-6.

Nrevdal, G. 1968. Studies on hemoglobins and serum proteins in sprat

from Norwegian waters .. FiskDiro Skr. Ser" HavUnderso,~:160-182c

10

O'Rourke, F.J.

19610

An immunological and chromatographic study of Sebastes marinus (Lo) and Sebastes mentella Travin~ ~.

P.-Yo Reun. Conso perm. into Explor. ~, ..!.2Q :

100':"1030

Schaeffer, Ho 1961~ A biochemical contribution to the redfish problem~

~. P.-Yo Reuno Conse perm. into Exploro 1vler, .l.iQ :

104-110 0

Sick, Ko

19650

Haemoglobin. polymorphism of cod in the Baltic and the Danish Belt Sea. Hereditas, ~ :

19-48.

Sindermann, C.J.

1961.

Parasitological tags for redfish of the western North Atlantic. ~~ P.-Yo Reun. Cons. perm. into Explor.

Mer, .l.iQ : 111-117~

Templeman;

W. 1959.

Redfish distribution in the North Atlantic. Bull ⁰ Fish. Reso Bd. Can~,

120 : 1-173.

Templeman, Wo and Sandeman, E.J.

19590

Variations in caudal pigmentation in late-stage pre-extrusion larvae from marinus- and mentella- type female redfish from the Newfoundland area. 30 Fish. Reso Bd. Can.,

12 : ^763-7890

Travin, V.Io

1951.

En ny art av uer i Barentshavet (Sebast~s mentella Travin sp. nov.) Doklady~. Nauk.§.§.§g,

.11(4) : 741-7440

L Norwegian translation by K.F. Wiborgo'i

Trout, GoCo

19610

Sebastes viviparus (Kr0yer) and the redfish problem.

~. Po-ye Reun. ~, perm" .i£!o Exploro J!.fer, ..!.2.Q :

94-99.

Tsuyuki, Ho, Roberts, Eo, Lowes, RoHe, Hadaway, Wo, and Westrheim, S.30

1968.

Contribution of protein electrophoresis to rockfish (Scorpaenidae) systematics. J o Fish" Reso Bdo ~O!

25

~

2477- 25010

Wilkins, N.P. and Iles, T.D.

1966.

Haemoglobin polymorphism:and its ontogeny in herring (Clupea harengus) and sprat (Sprattus sprattus). Comp. Biochem. Physiol.,

12 :

1141-1158~

Yanulov, K.P.

1962

ao Parasites as indicators of local rosefish stocks"

Po

266-276

in Marti (edit,,): Soviet Fisheries Investigations

in the Northwest Atlantic" "English translation by Bo Hershkovitz,

- - -

^,

Israel Program for Scientific Translations, 3erusalem

1963:'\

1962 b.

On the groups of rosefish (Sebastes mentella Travin) in the Labrador-Newfoundlru~d area o Po

277-289

in Marti (edit.):

Soviet Fisheries Investigations in ~ Northwest Atlantic.

:~nglish translation by B. Hershkovitz, Israel Program for Scientific Translations, 3 erusalem 1963.J

Tahl.e l . Dis tri'butions of t_8Lloglooin pa tter:;.i.s L1. reclfish frorcl t:::_~e

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Fig. III. Outline of hernoglobin patterns in redfish obtained by agar gel electrophoresis at pH 7.2 and combined starch and agar gel electrophoresis at plI 9.0. Legend: Filled in bars: Strong bands.

Hatcliedbars: Hoderately strong bands. Single lines: Faint bands.

Arrows indicate the points of application,