I . 11.

111. IV. V.

VI. V11.

VIII.

SOME ASPECTS OF NEOPLASIA IN MARINE ANIMALS

LIONE EL 14. MAWIMS~,LY-TEIOMAS l k x t w , IIuntingdon Research Centre, Huntingdon, England

E’cllow, Wolfson College, Cumbridge, England

I~itnit l i~ct i~~n . . . . .. . . Nooplasie in Maritin Invnrtobmtns . .

Taxtiriomic. I)ixtLrihut,ion of Nooplesia in Noolilaxia in Some Marine Mammals. .

A. Simnio . . . . . . .. 33. E’innipotlia . . 1 . .. <:. Cnt,eeea . . . . . . . .

Summrwy and Conolrinioti . . . . Acknowlocignmnnta . . . . .. Bibliography . . . . . . ..

Ncroplmia in %h . . . . . .

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15 1 152 162 183 196 196 196 207 219 220 22 1

I. INTRODUCTION In any p q w r pcrliaps the most difficult tusk for the author is to

write tho simplc, short, unambiguous English sentence. This is particu- larly true of any techiiical or specirtlist topic in which many words may be used in their true or original context as opposed to their current usage. It? is the problem of the pathologist who is essentially one who studies diseuses but who usually knows or understands less of the biology or ecology of the subject. This is all the more true when he in writing for biologists for whom disease processes may be of but passing interest. Some explanation of the title of this chapter might therefore be helpful. In iiiany of the infra-human species tumours have not been either seen or looked for : while many species will be omitted from lack of knowledge, this is not to say that tumours cannot occur in them but that, as yet, they have not been reported. The word “ neoplasia ” is defined by tlie Shorter Oxford English Dictionary as a combination of Greek and Freiirh words first used in I864 to mean “ 8 new formation of tissue in some part of the body ; a tumour ”. In the vernacular i t is the study of c ~ i c e r , or disease processes associated with it. Cancer may be defined 8s a “ malignant growth or tumour, that tends to spread and reproduca itself, and also to return after removal ”. A tumour may also be 8 new growth which docs not spread and is then known as a benign

t Deceaml. 151

1 52 I JON E r, p:. MAW 11 EYLE Y -THOMAS

tumour. Both ty1)c.s of trimour arc considered under the general heading of neoplasiii. Neopliksia was initially thought to be a disease of man, but the view is being increasingly held that, in all probability, i t is a property of life as much as rt%spirtition or reproduction. Neoplasia is nsually of unknown aetiology but, as research has progressed, vttrioua aetiologies have been found to bc reqonsible for neoplastic lesions. As more netiologies arc discovered, the diagnosis of neoplasia may disappear, since it may represent but a common response of the host to a variety of established stimuli. Some of these established aetiologies will be discussed later. In many instances, in the study of tumours in the lower phyla, one can only remember the advice of the great British neurologist, J . Hughlings Jackson ( 1 835-1911) : “ The study of the causes of things must be preceded by the study of things caused.” Within the classes of animals discussed in this chapter i t is the discerning observittions and the iLccuritt>e documentation of tumours which have been attempted. In the light of today’s sophisticated research techni- ques this approach ma,y seem mundane. The great observations of the past have often been tlcscribed in a detail which surprises the readers of today. With advnnced technology the simple observation is, all too often, never made. An interest in the past is often taken as a sign of senility, hut the reader of any text, on pathology since Virchow (1821- 1902) establiAhed ccllrilar pathology can appreciate that “ He who is not disposed to read history should repeat it.”

IT. NEoPrAsrA IN MARINE INVERTEBRATES

For maliy years it was tholight that neoplasicl was the exclusive property of thc vertebrate (Teiitschlaender, 1920) and that neoplauia was restric$ed in the phylogenetic tree to Class Pisces and above. Several authors have written at some length as to the reamns why invertebrate tissue cannot undcrgo neoplastic transformation but this view has subsequently proved to he unfounded. Of the tumourfi of Invertebrata only a few earlier references cnn be found (see Codremu, 1939; Finkelstcin, 1944: Wautier tind Wautier, 1953) and it was not until the 1950s that the subject of comparative pathology, in relation to animds in general, experienced a renaissance. Scharrer and Lochhead (1960) wrote a useful review with over 160 references to tumours in invertebrates. The only verificbd neoplasms in marine invertebrates have occurred in oysters and clams, and possibly one in a coral (Squires, 1965).

As early t i s 1887 Ryder desc+ribcvl it menenchymal t umour of the perirurdiitni i r t aii oystt.r (hzs.so,Pfr.m viwyinica (C>melin, 1791). A similar finding in the siirtw spcries was noted by Smith (1934). ‘L’hecre ticinours were po1ypoid:rl and coiisisted of t>ypical mcsenchymal c t 4 ~ eoverecl hy

21 singlv 1 ; y . c ~ of <boilttntiitr riliatcd rl)ithc~liuni. Lipid granules were also noted. Sprks r& a1 (1964) recorded i i mtsenchymal tumour in the Jap- i~ii~st? oystm- (Irawostreo g i p s l'liinnberg. The tumour was dorsal to the rwt,urn and had u yedicle. The tumour cvneisted of Leydig cells covered by t t luycr of t d l colurnnur ciliatcd epithelium. The presence of blood vcmels was n o t d and aii incrcased number of collagenous fibres were found just beneath the epithelium. The cellular picture wus similar to that described by Smith a d Ryctcr. There was also some evidence of lymphocytiv infiltration. These tumours, all of which presented similar histologicol cliitraoteristics, still present H. diagnostic problem. They iilso servc to illustrst,e o w lack of knowledge of marine invertebrates in their rrsponsse, if at, all tliffercnt, from that of vertebrates, tjo trauma, infection or nwplasitt. Taylor and Smith ( 1 966) noted polypoid end papillary lesions on the foot of the Wwhington clam Tresus (9hizo- thnerus) rr.utc'aZi ((loiirad, 1837).

0 1 1 microscopical exurnination 21 simplo papillary tumour was seen assooitited with a markcd infl;rmniatory response. An inflammatory respotis(', ;is poiiitct? out by L'aul(*y ( I 969), can indicate a reparative procc*ss i w woll it8 an iriflammatory one. In all probability, both occur (wncurren tly ILS often seen in the higher vertebrates. In this instance there was a suggestion of bacterial involvement although the evidence was far from i~ondusive as to whether it wus due to pathogens or not. T'irnley and Sayoe ( 1 867), in an interesting paper on some non-neoplastic (widitions in oysters, mentioned the presence of a tumour mass in an abnormal adductor muscle. The fumour was growing from the abnormal anterior p r t of the muscle and was thought' to be the first tumour reportled mising from rtn adductor muscle. Katkansky ( 1968) described t i t i intestitid triinour it1 the European flat oyster, O&ea eduZi.9, but the photogrtkphs S~IOH' I~ do not indjritte miything more than some epithelial hyprrplnsiti, possiil)ly a poI.yp, and should not be considered to represent true ticoplusm utihbss furthcbr evidenw crtn be produced (Fig. 1 ). Pauley trnd S;~yt.c (1968) notled ii benigrl fibrous tumour. the aetiology of which WIM unrertaiii. ;ind suggestodeither i k myog:cnic or gonadal aetiology, thc lilt ter bring thtb more likely of the two. 'I'here is such a paucity of in- formntiori on tho rcsponsr of invortehratcs to noxious agents that it is virtually irnj)ossiblc to make a definite diagnosis. Sparks el d. (I!jti!j) dcsvrihetl ttnot hcr mesenchymal tumour. but i t s histologid features W(VY extt.(~ttlcd!~ cwriftlsilig and, whilc on(' ('iLri ibgrr'c' that it W R ' ~ a tu rnour , t l i c w is litt 11. o t i ( 1 (*:tii ; ~ t l t l to t h c ;dwti(iy vorlf'lisirtg hi~t(rlogi~i~I patterns piwetit ivi.

It wiis t to t u t i t i l I!)(i8 thiit i k l i x ;Lttihi))i)t IVZIS m;itIc. t o t1.v t o c*o l l t . c* t tlicw isoli1tc.d ('~LSPS togptliw tm(l to ( * o t i v r i i ( ~ ii n1(,rtirig to clis(*lrxs the

FIG. 1 . K ~ m r rp~tl~rlial hyperplasia stXm In the put of a Europan flat oyster. (H. and E. %fag. Y 400.)

111aill.y 111a~jot' ~~t.ol)lc+tl~s of it~~(:rt.(h\)ri~t,t: pt~thology. Much praise must bc give11 fo I ) i~wc. i t t l t l H I L I W ~ I ~ ) ~ L ~ ~ ! ~ (I!fti!)) f i~ r t i t t e n ~ ~ ) t i n ~ this: in t'heir monogrup h ~ C I ( ~ V I L I I ~ , (il~t,t~. M ~ C ~ C ~css(!~~I b l ~ ~ l , arid ;innot.:mt~rd bibliog,rr:rphies revealed 1niLtl.v urt.ruspctc*t8r:cl sottrc!cs of illformution. Hershburger, the JXrector of t,llcr Iicgistry of ' l '~~mors of Lower Anirnals, has collect~ed nll~cli 1n1~tc~rii1'1 i m l while soma of t.hc cil,ses in his collectiorl may have ii,lrcv~cj.v I)(YI\ ~nc:~~t~ionc~tI ii, is M.OI+JI sttrnttti~rizi~~g thwn iigili11 ('Fable I ) .

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The f'ibct t h t the princ.ipul t111noul.s of tniwirlc invc~rtchriitc-x (.art 1istt.d i l l sucbt~ it short table intli~i~tcs eithcsr tllclir oxtrcrnc. rarity cjr poor obsrrvntio~i by the human eye, trainc~tl or othcrwisc. A filu. ~ ' ( ~ a r s ago t u ~ n o i ~ r s of thts ( '~n t ra l IlCrvolIs systt~ln of 1aboratory ar~irnalh ware thought to 1)c. vir*tually nor)-existant, and it was not until the brains of' thousands of iirli~uals had been examined routinely (Muwdedcly-Thomas c~nd Newr~lun, 1973) that this notion wits tlixpelled. Perhaps this will he

tlic. (.as(, wit 1 1 i ~ ~ \ l c ' t ' t ( ' l ) l . i ~ t ( ~ i I I I ~ I O I I I . ? '1'111~ ~l(~s(*ri])tion of' tlw ~norpliolo- gi(:i~l fin(lingh is oI)viol~hl,v i l l 111111~~1~stoo(l ; I I I ( I t / l t~~'(> is 111111'11 S(~OI)C lirre fbr c~xt)c~rir~~c~r~t;~tior, in ;r t , ; l t f t , t ~ 1 ) 1 to sc>pi~t';~t c3 ~ I ' ~ L I I I I I ~ L t'L'o111 i~if'(~tiolls. rept~ritt ivc. o l ~~c~opI:~st~i(a l (~s io~~s. kli~rsli1)itrg~r rt (81 . ( 1 !)7 1 ) 11oted the efkcts of' " Nitroso " c~ol t~pou~~ds o t ~ thc cn~ytisli Proc.rottbar~~s clnrkii (Girard, 1852) whicsh, i ~ t c.oncc~~trations of I00 and 200 p.p.tn., cailsed cleg(a11erz~tion of' the i ~ t i t ~ l ~ t ~ i ~ l glt~rld i~tid hyperplasia of the tubular cells of the h ~ p i ~ t o - \ ) i ~ ~ ~ c ' ~ ' ( \ i ~ s . One is tttrnp ted to speculate whet her the hyl)eq~li~sit~ of tllc I t ( ' l ) i ~ t o - l ) : ~ ~ ~ ~ ' r ( a i ~ ~ ttliglit h i ~ v ~ hc~c~ornr neoplastic

(Fig. 2) i f ' t l l c . c*o~tc*c.~~tr-i~tio~~s of' tllct c.ort~l~or~ntl Iiacl 1)c.c:n krigh(.r or the period of' thcb c>xl)c~rilnc.r~t inc.r.c.ssc~cl. I ' ~ L I ~ I c Y ( I!ftil)) has tvrittc.n ii uscf'ul criticit1 rrvic~w on llcO[)l2trtii~ i r ~ rrlollrrnc~n, with some 7 3 rr~fi,rc.rtc.c.s. Farlcy (I!)(i!)) i l l ;I f i ~ ~ ~ t i l ~ t i ~ ' f'e:it of' looki~ig at i~p~)r(~xirr~:it(.ly YO 000 oystcrs ovcr sc.vc11 Y(>:II-S, ~ O L I I I ~ six (li~('ilne(1 oystcr~ i n wt1ic.h thcrc. WiLS

mirroscol~ict~l invirsion hy Q,pical irnrnat ure harn~atoc.ytc~s (f"ig. 9 ) . These Iiut~n~afoc.ytcs wcw cxtrrmchly active allti rnitotic. figures were cot111ti011. Th(. aetiologj of this ro l~t l i t io~~ is unicnowrrl. A ~ i ~ . i i l artiology nll l~t t t Iwi~s I)fb ~011sid('rf~d w I I ( A ~ ( ' the possible [ ) r e ~ ( ~ ~ ~ c . e of' a c.urc.inogcn in (.st 11itriue witttlrs, uc.c.ori1pt~11ic.d 1)y biological c.onccsntrtltion within the ttni~nitl, t1111st L)c. c~ontc~1nplirt~t1 (( ' i i l i t i t t~i t t i t l ittlff K U ~ ~ ~ ~ S I I I I < J . 1957).

I'lo. 3. Muuaivn irifilt,rution inCm;umo~tr~~rr ~ ~ i ~ u i ~ h i i . ~ with cnllu of the irnmaturo hec!matocytr: t,ylw. 1 \ 1 1 t,i~uutbs uh(~\v ovi(l~ticc 0 1 ' i11filt.l.tl.t ion (H(:o I L ~ I . I ~ W H ) . Helativaly r~~wrnal pit iu st.cvi t,owtbr.tls tho t,ott,om right 1)1'th1- ~~hotogruph. ( I 1 ant1 E. Mag. / 75.)

'I1ili~rc iL1.t' Inany c.xan~pltw of~~col)l;tsia ill fish found in the polluted rivers of' t hc ( ' i ~ ~ t coi~st of Arncl-ica. ( 'oi1(:11 ( I !t(j!f) dr~cribed ;L Icsion of thch 111i~ntle ill iln At)arric:tn oyster ('msso.vlr~c~ virqinicn which c~onuistatl of' " blsstoid " cclls which hut1 a high mitotic* i~rdex (Fig. 4) : t h ~ artiology is still ankno\vll. ll'olf' ( l!)fi!f) dcscribc~tl two turnourn in the Australian Roc-k 0ystt.r C'rrrs~oslren conztn~rr ia l i~ wliich were t~entatively clisgnosed ss " ~) i~pi l l t~r j . c.l)itl~c~lion~~it;t ". but t ~ n f ~ ~ t l i t l i i t ~ l ~ no search fbr metas- t i I S l l i . liOti1 t 111110111~8 \\('I.(' fi)l111(i 011 tilt' i l l l l th l ' llltblltlt' 01' tile ov~t(~1.. \\'oil' (1!131) fi)llo\rc.ci u p his I ! t ( i ! t fi~ltlitlg 1 ) ~ ~*t~l)ot.titlg a 1i~rgc~ ~~c~ol) l i~st t l i i r i111otllc.r Atlstl.;tlii~n ltocbk 0ystc.r \rllic*h Ili~d a volt~lnc~ rqual t o A th(3 six(. ot' tlic, oyster (Vigs 5 7). Lt ~)rc~sc~lltc(l as a pal)illif'erous

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t i imoor . 'l'lit> Ic\sion wiis i i ivirsivc. iLIrtt Irsd iilreidy pt.~~tbtr~~t~td tdirough t,lle niitiitle niiisclc into t,he o v t q . 'I'liis tuniour, like the previous two ciescrikwtl, rtwmbletl mc ire tl I P Iid(.rioc;ircin(,rna-like lesion of the verte- brate. '1'0 dihte twelve such t umours Ititve 1)een fourid and Wolf makes the following siiggestionu as to possiblc aetiologics :

(1) a genetic! mutation, whirli had pro'luced a susceptible strain of oystror ;

(2) infcctious agent, i s . virus, in oyster bed ; (3) infectious agent, non-pthogeiiic urit il oysters have been made

suuceptible by genetic mutation.

Dinammi anti Wolf ( I !)74, i n press) have reported multiple tumours in the pericardial cavity of an Aiistralian Rock Oyster Crmsostrea commer- cialis, t~nd cwnsidcred the differential diagnosis in the following order : (i) slow growing multicentrio benign neoplasm of Leydig cell origin ; (ii) congenitid anomaly ; (iii) hyperplasia secondary to infection or injury. Theso tiirnours were similar to those described earlier by Ryder and Smith. Brown ~t al. (1!)73) reported an incidence of tumours in 'L 121 fi.eshwiLt,er fish cxamincd from the Fox River watershed (4.38%) t ~ s conipurwt with 4ti3!j fish examined in Canada (1.03%). The FOX River watershtd arises from underground streams, presumably from Lake Superior, t ~ n d ww found to he i~ highly polluted system. The effects of pollution runnot bv disregarded, as will be discussed later in relation to vertebrate fish,

Pindly, t here is thc> txxtrornely interesting case of the coral in which a markd loc4ized lesion W ~ L S seen (Fig. 8). 1)oes this represent a neoplastic! rcvponsch or is i t merely u parasitic infection by Porites compw.unci. anti f'. Iobata (Squires, 1965) ! It will be obvious from the fore- going tltut only i n the Australian Rock Oyster doeu one see a histological pattern similar to thiit recorded in vertebrates. The mesenchymal tuinours, if indeed thc1.V are turnours a t all, tLre interesting, since the me.senchyma1 tnmour of vertebrate fish predominates over all other types. This is in complete contradistinction to the higher mammals, in which the epithcliul tamour is by far the iiiost common.

[rr . N E O P L A S ~ A IN F I S H

WIicn reviewing the* progrcw of fish p~~thology i r r this ocritury Snieszlm ( I !)72) statotl that '' AIIYC)II(? (:xpecting to firid thiit there is u wealth of piiblications on fix11 pat,hology inore than I00 years old will 1)e rliuu1)poiiitcd ", and with the cxcq~tion of u few caxw t h i H is t rw. This gives sonit' indication t lrat although I'irdiow wiLx t h c a J)Lt Irw of CthllllllLr I"tt I~ol~ggy, t htn tlin~;ircv I)rowns i l s dwwi t)(*(I i t i l t i ~ g:l't*iht hook

“ Ccllulai- Patliology ” puldishd in 1858 was considered to be almost cxcl usively the property of human pathology. Comparative pathology has laggecl behind general medical pathology ever since the middle of the ninetcwith century, with OIN: or two major exceptions. The great ,John Hutit,cr (1728 1793) c*oiilcl tw (IcHwibed i ~ s the English fihtticr of‘

FIG. 8. A pimo of coral &owing a lesion. Whether this Ionion is mopleetic or not is uncertain for the present.

comparative pathology and maiiy of his thoughts and views antedated those of Darwin and other great sciCiitistH by m;my yearii.

It, is difficult to iLttempt to uiiderstancl the rthnaisoance of com para- tive pathology in khgland. J . It. ill. Itincs, before the Second World War, a t Cambridge wa,s actively pioneering a new comparut,ivo ap- proach, arid in sucdi centres ~ L B thc Universities of Cambridge ;md Briatol

(lor~iII”l.iLt,iV(’ pithlogy is rcnliwtcd a s , and of, its own right. Relevant work i i i thc United States of America was catulyzed initially by Schlum- berger, whose untimcly death delayed it by several years. Fortunately, under the wspices of thci Nut ional C”uncer Institute and the Smithsonian Institute, the Registry of Tumours of Lower Animals has been estab- lished i n the work of its Director, I h tJ. C. Harshbargcr, backed up by Dr C. Ilawc, uiid him made great) stride8 over the past 4 to 6 years. The early history of fish disease has already been documented (Wellings, 1969; Mawdvsley-Thomas, 1970; 1971a, 1972) and it may be concluded that historicid evidence of fish turnours is non-cxistant with the excep- tion of somv borir turnours. The reason wus that until some form of

fixative wus svailablc, wine or spirit in the first instancc, only “ dry ” preparations could be prepared and examined in detail (Figs. 9 to 10).

The prewnt situation is extremely encouraging, as the field of patho- logy is no longer limited to thc veterinary and medical professions, and many others are now receiving fundamental training in basic diseases of fish. While Americu, leads the field for the present, it is hoped that fish diseases ibnd t,heir pathology might find a place in the curricula of some British rinivc.rsity departments of applied biology in the not too fur distant fiit tire.

The study of‘ pafliologirid processes is ofteri facilitated by the use of the poikilotlierttiic vwtc4~rate because its Irwtiholic rate i n slower than tjliutj of its Iioiiieotlic\l.i~iic couiittq)iwts so that onc is itble to ~tucly histogenesis niid pat8hogencsis iii “ d o w motion ”. This allows the pathologist to study more “ time franiw ” of any givcn promss wrl, in

FIQ. 10. Plaster cwt of male Pugrue unicolor. Thorn i R a tumour in the maxillary area, probably of bony origin. No original t i n ~ i i e available:.

so doing, Ijc given tlit1 opportunity f iw maximizing observation. Several inLtjor mctlicd diwovtlriw have I ~ c c ~ i i t t i i d e rising fish. Rant iiig and Best in 1!)22 cxprl.itiiciitect with fish to appreciate untl isolate t,he endocrine and exocrinca functions of the pancrea8, and even earlier in the 1920s work on fiRh thyroid, while lettding to some initial confusion, made tt

166 LIOB EL E. MAWDESLEY-THOMAS

significant contribution to medical science. As already mentioned, the term “neoplasm” is really a synonym for an unknown or as yet unappreciated aetiology, while various aetiologies have been associated with neoplastic transformation. In the fullness of time all aetiologies may be appreciated and the term “ neoplastic ” will pass out of our language. The study of fish tumours can often shed much information by using relatively unsophisticated epidemiological methods to point to a common aetiology, and there are already examples of this which will be considered lator. Factors of human ttppliciition and significance may bc established by the study of fish tumours in relation to their geographical distribution, their various phylogenetic, including immunological and enzymatic, entities, together with the part played by pollution, trauma and oncogenic viruses. The liver cell of some members of the Ralmonidae is the most sensitive histological indication that exists for the bioassay of aflatoxin B;. Stanton (1965) stated that “ The high sensitivity of small fish to a known carcinogen, their poikilothermic physiology and the ease with which their aquatic environment can be controlled are particular advantages in studies on carcinogenesis.” The sooner the cancer research worker realizes that there are animals other than rats and mice, the better!

Harshbarger (1972) was the first to record an olfactory neuroepithe- lioma in Coregonus hoyi (Gill, 1862), a freshwat er fish, while the documen- tation of a neural tumour in ih marine fish is still awaited. This remains the only mention of a CNS tumoiir in Pisces, excluding the neurilem- momata. The central nervous system of the poikilothermic vertebrate has still retained its fundarnen tal ability to undergo reparative processw, which in relation to man represents a peculiar. phylogenic phenomenon. The process in fish is at prcverit being studied (Mawdedey-Thomm el a!., in preparation 1974).

Epidemiological studies in animals and man have shown that there is often a geographical distribution of certain diseases, and appreciation of this fact may help to establish the aetiological agent. Examples in man include the high incidence of liver cell tumours in the Bantu, and Burkett’s lymphoma in certain parts of Central Africa. One cannot help but wonder if the lympho-sarcomata in fish, E.s+ox lucius (Linnaeus 1758) reported by Mulcahy (1970), Marek’s disease of poultry and Burkett’s lymphoma in man might not have a common aetiology, but present with different signs according to the species in which they are found. Other examples of pathological geography in Pisces are well documented. The papillomata of the Pleuronectidne off the American West Coast (Dawe and Harshbarger, 1969) and the stomato-papillomatosis men in the Anguillidae in the North and Baltic: Soils (I)cys, 1969) baing but two.

SOME ASPECTS OW NJNIPLASIA IN MARINE ANIMALS 167

Any netiology is usiiully mllltifi\ctorid but if one pitrt a t least of the Iwoblem (-mi tw solvcd t h i ~ (WI oftmi Iwd to more fruitful lines of investigut8ion.

It is oiily in the past 80 ycws that the niedical profession has seriously considered the problems associated with the genetic basis of pathological procesaes. In this instcince, the human geneticist has lagged behind the plant and aninial geneticist. Tumours of pigment cells, particulurly the melwomata and melanosarcomata, have proved to be useful animal models in which to study the genetics of neoplastic transformation. ‘I’hc works of Siciliano et nl. (1971) and Vielkind et al. (197 I ) with sniiiller tropical freshwater tish, usually of the Xiphphorin spp., are of grcut interest, although strictly out,side the remit of this article.

Of particular ourrent interest is the breeding of disease-resistant fish, or hybrids, which would offer great protein productivity or greater game fishing potentiul. Although genetic and chromosomal manipula- tion or mutation are possible, it must be remembered that several members of tnhc Sparidae are gcbnetically prone to tumour formations. The geographical distribution of the papillomata in thc Pleuronectidae on the West Coast of America could mean that there is a genetic pool, which could represent a possiblc aetiology.

The intimutt. relutionship betwcwn the fish and its environment makes water pollirt,ioii, ewpccially of coastal and estuarine waters, a particiilur hi~zurd. Major oil slicks in the ocean (lo not present too much of a h a m d cind w i i be simply uvoidcd by rnariy marine species. Of far greater Iiuzard is the cxplonitiori of tho HW bed. A t the present time the North Sea is heing SIU ssfully t a p i d as (L source of natural pis and, although geological surveys haw brcn tnntle, no attempt hw ycat been made to produccx an interface bet>wctw the physiral and biological sciences. The risks of‘ siting an oil rig in the centre of a breeding area of an important economic species could ultimately destroy major fishing grounds. This seems to be an inttmational problem which has not been appreciated by N.A.T.O. or the I3.Fl.C’. The sinking of oil wells in the middle of spawning grounds ot‘ fish of economic value could spell disaster. There is also some anxiety concerning the sludge being pumped out of these wells, for in association with the tides and winds potentially toxic chemicals could reach coastal water#. These Rubstances, as yet undefined, could well be potentially carcinogenic.. Brown et al. ( 1 $73) gave Table 11 in relation to freshwater fish and, while the hazards to marine fish are possibly iiot so grtsat, some attention should be paid to them.

As the x t t i d y of cwnparativc putho1og.y embrtwex as rrii~ny phyla,

168 r , i o N EL E. MAW DESLEY-THOMAS

clrtsseR and sub-clwses as possible, an up-dating of the initial work of Schlumbergcr and Luck6 ( 1948) has been undertaken, and in this article an up-to-drbte t;ixononiic distribution of tumour bearirig fish has been included. While there are several omissions, it must be confessed that sometimes the taxonomy has become tangled. There would appear to be an increase in t8umours in the teleosteans aa opposed to the chondrich- thyes. The exact incidence of tumours can never be accurately assessed as the population from which they have been obtained can never be known with any degree of certainty. What can be said is that while tumours are seen in many fish species, the principal groups affected in the marine kingdom are the Salmonidae, Gadidae, Bothidae and Pleuroneotidue. This distribution may again be more apparent than

TAULIG TI. HOME ORGANIC COMPOTJNDS FOUND IN WATER (Modified aRor Brown et d., 1973)

Chemical - --

Toluetw Crude oil Ethyl etlinr Gaaolino Chlorinated Itydr( tcwl)ot IH

OrganoptioRpl iiitw

Ntlptittialrrio

Persistency

1-2 years 1-2 years 1-2 years 6 months 2-6 yearn 7- 84 day8 1-2 yearn

real as these four groups contain some of the more common food species and population studies should be included to ascertain the meaningfulness or otherwise of the preceding statement. It should also be remembered thltt 50% of the world's fish catch is caught from approximately 1% of the ocean surface. The incidence of tumours in freshwater fish would appear to be greater than in marine fish. The reasons are presumably legion, and will not be discussed in this chapter.

Few classifications can either be absolute or complete. It is essential for the sake of clarity to try to formulate some simple rules which will cover the majority of tumours. The most useful current classification is that based on the anatomical or histological type of tissue from which the tumours arise.

It must be stressed that a tumour is designated by the predominate tissue in it. All turnours, even epithelial ones, contain other ebmontrc, including connective tissue or blood vessels, but thew nhould not be

SOME ASPECTS OF NEOPLASTA IN MARINE ANIMALS 169

considered Its tnajbr entities when making a diagnosis of a specific tunlour. This c4ossifictttion ('hble III), while it includes systems in which mttniinolion tuniourH are found, has ctn additional group to include the varioucl pigment tiimours which are specific to hleostean species.

A malignant tumour can only be so designated if it can be shown to bc locally or distantly invosive. The only absolute criteria for malignancy arc it8 invaaiveness or the presence of metastases. The terms " benign " and " malignant " were initially used by clinicians to indicate a prognosis but, unfortunately, they have gathered pathological

TABLE 111. CLASSIFICATION OF TuMouRs (Modified after Willie, 1967)

Consist of papillomata. adenomata or carcinomata.

Consist of fibromata, lipomata, chondro- mata, osteomata, angiomata, leiomyo- mata, rhabdomyomata, etc. Malignant tmnours are described a~ sarcomata prdixed by the name of the principal type of t iww from which they have arisen.

Consist of turnours of lymplintic or other licmnopoiotic tissues.

CorisiRt of turnours arising from the central I I ~ ~ V O U R systern or peripheral nervouR syatem, viz. ganglioneuromata.

Consist of lesions arising from pigment cells, inalrrding melanomata, erythroplro- romata, guanopt rorotnata, etc. tc:ratomatrt or cxnbryonic ttimours.

overtones which are erroneous m d confusing to the zoologist (Mawdes- ley-Thomas, 1970). The only advice that can be given is that if a term is to be used then it must be so defined that even if other workers disagree they will at least know what is meant. The literature is cluttered with papers by cancer workers who consider that any lump " must, of necessity, be malignant. One of the problems in comparative pathology aa compared with human pathology is that so little is known concerning the general biology or behaviour of animal tumours that it is extremely unwise to asaume that they are anything more than analogous to tumours seen in man. AR all experimentaliRts are well

170 LIONEL E. MAWDESLEY-THOMAS

aware, " Only man is like man ". Wellinga ( 1 969) in his excellent review on neoplasiu in fish considered that fish tutnours were both analogous and homologous to those in inan. A t this time it is felt that there is no sufficient justification for this conclusion to be drawn.

While there are many reviews of bibliographies on the subject of fish tuniours (Wellinga, 1969 ; Mawdesley-Thomas, 1969 ; 1971a, 1972), some additional tumours seen at the Huntirigdon Research Centre will be included in this chapter for the sake of completeness.

The increase of polychlorinated biphenyls (PCB) in such seas as the Baltic and the North Sca is cause of concern to all conservationists. The relationship between these compounds end neoplasia presents a difficult problem and one which will not be readily resolved without the establishmelit of a multi-national interdisciplinary group of scientists. The problems of biological concentration have still to be evaluated, and this is particularly true of the clams and oysters, and many species of fish which are at predator pinnacles (Mawdesley-Thomas and Leahy, 1967). Migratory species, such a: many of the Salmonidae, which build up large reserves of fat, can soon raise their serum level of some of these chemicals, many of which are lipid soluble (e.g. the organo- chlorines), to toxic levels. Among the birds the same can be said of the arctic tern (Sterna macrura Naum), which has the longest migratory passage of any known bird. * The largest subdivision of the Vcrtebratu includes the fishes and i t is not surprising that there are biochemical and enzymatic differences betwcen species and sub-specirs. The presence or absence of EL particular enzyme system may detcrmiiic it specific metabolic pathway in relation to exogenous compountls. It is important to establish chemicaIIy whether a proximate carcinogen is found or not. It has already been shown that enzymatic differences occur betwcen sub-species of the Salmonidue (Mawdesloy-Thomas und Harry, 1970). I t is k~y defining such cherniral details that the difficulties associatcvl with species differences will ultimately be solved. The morphologist mus t become more chemically and biochemically orientated if he is to become fully integrated into the present-day problems associated with neoplasia. This type of lateral thinking could ultimately explain the differences in the reaction to aflatoxiti B,, which causes 100% tumours in rainbow trout when fed a diet containing one part per lo8, whereas brown trout are virtually resistant to it. The presence or absence of certain enzymes or iso-enzyme systems reflects, or may reflect, the genetic make-up.

The itnportantI part played by oncogenio viruses in neoplastic disease in animals is being increasingly realized, (Wolf, 1!)72 ; Mawdesley- Thomas, 1 !)7% 1973 ; hl;~wdexlcy-'i'horn;~s and Bucke, I 97:3), itnd great

SOM I< ASPEW" 01' NEt IPLASlA IN MARTN E AKIMALS 171

strides arc being made in thv ideiitificntion of the viruses known to cause fish diseases and fish neoplwsia. The possible palat played by oncogenic viruses has dreiidy been mentioned in fish-pox, an extreme epithelial hyperplasiii seeii in some specim, also the stotnato-papillomatosis of eels could well have it viral aetiology, as has the possible connection betwcen the lymphosarcomsta seen in the Irish Pike E'sox ZuciuS L., Marek's disease in aviaii species and Burkett's lymphoma in man. Until more laboratories hold sevwul fish cell lines and have expertise in electron microscopy progress will not be spectacular.

Fish which live at t h r bottom of tho oc*ean inay expose themselves to local tntuma by scraping thcrnnelves against a " rough " sea bed. Skin damtigo is oftcn the end result and, although often minor in itself, can be R potential site of entry of oricogeiiic viruses. The incidence of xkin papillomata in the Bothidae and Yleuronectidae could well attest to such an aetiology. Another predisposing factor is that, if there is carcinogenic mriterial anywhere in the water, it is more likely to accumirlatc on the sea or estuarine floor.

The relationship between phylogeny and oncogeny was initially thought to be absolute and it was believed that neoplastic transforma- tion depended on the presence or absence of a lymphoid system, upon which the iinmuw response was dependent,. The exact immunological response of fish to tumours requires further work and discussion. Some tumours have been successfully trctnsplanted. This work was performed in the early 1!)40s (Luck6 and Hchlumbergcr, 1941) in which auto- and homo-tr~~ns~~li~nts showed contiiiried growth (Good and Papermaster, 1964). I'hc ahtrophy of the lymphoid systclm with age, as seen in many of the higher animals, could well bc associated with the high incidence of neoplmstic. trimsformation seen in middle to old age, according to the species studied. Sonic strains of rats develop nearly 100% mammary tumours by 2 years of age. Sintilarly, some female strains of rats can develop u p to 80% pituitary tumours. Some work has been undertaken on the immune response of several fish species, including some Salmoni- dae, in relation to various infectious agents (Klontz, 1972; Dorson, 1972). Antigenic variations in 76 virus isolates of Egtved disease have been examined wing fluorescent antibody techniques (Vestergard Jnrrgensen, 1972).

Some tiimours of marine fish which have been seen at the Hunting- don Research Centre are illustrated in the following pageg (Figs. 11-21).

I72

a 9

f

m 9 E

.I d 6

175 S

OM

E A

SP

EC

TS

O

F

NE

OP

LA

SIA

15

MA

RIN

E A

NIM

AL

S

e .- k (r

FIU. 16. Soction of a fitwornn in Mcrlungius msrlan.3rce.

178 IJ

OY

El,

11;. M

AW

I)XH

LE

Y-T

HO

MA

S

SO

ME

AS

PE

CT

S

OP

NE

OP

IAS

lA

IN M

AR

TN

E

AN

IMA

LS

179

FIG. I!). &xction of livcr of cod (Vnclu8 morhm) showing parasitic lesions, due to B ~ p h a l u 8 spp., which could be mistaken for neoplastic lesions. (H. and E. Mag. x 100.)

FIG. 20. Section of kiclnry of wh~t~ng (Alerlnngiiu, msrlangua) showing metastatic deposits, posaibly from the liver. (H and E. Mag. x 100.)

FIG. 21. Section of skin of plaioe (Pleumnectes ~ ~ ) , showing lymphocystis, characterized by greatly enlarged cells. This was originally thought to be a neoplastic lesion.

SOME ASPECTS OF NEOPLASIA I N MARINE ANIMALS 183

1V. TAXONOMIC DISTRIBUTION OF NEQPLASIA IN MARINE FISH Tumours included in this table are considered to be spontaneous:

induced tumours have not been mentioned.

Class CHONDRLCHT HYES-(:ctrtilu~inorts fishes Order SQUALIFORMES

Family CARCHARHINIDAE- -Requiem sharks Prionace glauca Liver, icdenoma Schroeders, 1908 (Linnaeus, 1758)

Carchrhinus milbcrti Reticulum octll mrcoma R.T.L.A., 1972 (Muller and Hetile, 1841)

Carchnrhinua lewm tc'ibroacmorri;~ H arshbarger , 1 972 (Muller and Henle, 1841)

Family 8CY LLORHrNID AE-cat 8 harks

Odontoma Ladreyt, 1929 Chondroma, right flank Thomas, 1933(a) Osteoma, trunk Thomas, 1933(a) Epidermoid carcinoma, Stolk, 1956

oral

Syliorhinua cnnicicla (Linnaeus, 1758)

Family S Q A L Z D A K - D O ~ ~ ~ B ~ sharks

Sqmlua fernnndbme Molina, 1782

h'quah acanthiae Linnaeus, 1758

Thyroid tumoiir

Chondroma, lumbar

Fibroepithelial polyps, vertt hrae

oral

Family RAJIDAE-skates

Baja clavata Linnaeus, 1758 [ M J z t tnata

Raja mucrorhyncha Fibrosarroma Rafinesque, 18 10

Raja maculata Montagu Haemangioma (= R. montagui Fowler,

' ' Malignant melanoma,

1910)

Cameron and Vincent, 1915

Takahashi, 1929

Wellings, 1969

JohnRtone, I 9 12

Haddow and Blake,

Drew, 1912

1933

Drew, 1912

184 LION EL E. MAWDESLEY-THOMAS

Raja batis Linnaeus,

Malignant, melanoma, Johnstone, 1912

Malignant nielanorna Johnstone, 1913 posterul fin

1758

Class OSTElCHTH YRS-Bony fishes

Order ELOPIFORMES

Family ELOPIDAE- -Tarpon Elops saurus Chondroma, Surbeck, 1921 Linnaeus, 1766 subrntaneous

Fam i 1 y GONG RIDA E-cOnge I‘ eels Haemangio-epithelioma Drew, 1912 Fibrosarcoma, Johnstone, 1920

Lymphosarcoma, kidney Williams, 1931 subcutaneous

Coates et al., 1938

i Conger conger (Linnaeus, 1 758)

Family ELECTROPHORI1)AE- Electric eels Electrophoms eleccticw Papilloma, trunk (Linnaeus, 1766)

Muraena helena Adenocnrcinoma, oral Ladreyt, 1935 Linnaeus, 1758

Order CLUPEIFORMES

Family CI~UPElT,AE---Ht’rrin~

Family MURA ENIDAE-MOWYS

Lymphosarcoma Johnstone, 1926 Rhit bdorn yoma Williams, 1931 Leiomyoma ( 1 ) Thoma&, I 933( h) i Neurilemmomata ( !)

Clupea harengus

Linnaeus, 1758 harengus

Fibroma Johnstone, 1912 Fibroma Johnstone, 1924 Fibroma Biavati and Mancini,

Rhabdom yoma Mawdesley -Thomas, 1967

1972

I Sardina pilchardus (Walbaum, 1792)

Clupea harengus Linnaens, 1758

Order SALMONIFORMES

Family SALhlONII)AE-sUlmOl1, Trout Oncorhynchus Fibroameloblastoma Schlumberger and

(Walbaum, 1792) tsibawytscha Katz, 1956

BOME ASPECTS OF NEOPLASIA I N MARINE ANIMALS 186

Oncorhynchw ketn (Walbaum, 1792)

'Fi brosmconi a, body cell

Fi brostmconia, body cell

1, y m p h osarco m tb

Oncorhynchw gorbuschu Fibrosiirco m a, (Walbaum, 1792) body cell

Fibroma, hack

L y mphosarco m a,

Osteogriiic saroom&

Stomach polypi

Oncorhynchw kisutch

stornech (Walbaum, 1792)

Oncnrh:wtchus rhodurw (Jordan and

McGregor, 1925) Oncorhynchus mmou

ishikawae (Brevoort) Oncorhynchue nerka (Walbaum, 1792)

Salmo snlar Linnaeus, 1758

J'almo spp. Linnaeus, 1758

Rpp. not

defined

Kazama, 1922

Takahashi, 1929

Honma and Hirosaki , 1 966

Takahashi, 1929

Milleman, 1969 Milleman, 1969 Ashley et al., 1969

Harshbarger , 1972

R.T.L.A., 1072 R.T.L.A., 1972

H epatoma R.T.L.A., 1972

Rhabdomyostlrcoma Harshbarger, 1972

Thyroid tumour (!) Gilruth, 1902 Thyroid tumour (!) Wilkie, 1891 TAymphoaarcoma, Haddow and Blake,

Sq. cell carcinoma, Roberta, 1972

Thyroid tumour ( 1 ) Plehn 1002

kidney 1933

oral

Adenocarcinoma,

Thyroid tumour ( 1 ) Adenocarcinoma ,

Thyroid tumour (? )

Thyroid tumour (!)

Thyroid tumour ( 1 ) I Erythrophoroma, skin

thyroid

thyroid

Gilruth, 1902

Pick, 1906 Jaboulay, 1908

Marine and Lenhart, 1910

Gaylord and Marsh, I912(a, b)

Peyron and 'I'homcla, 1930

Thorn-, 1931

(a, b)

186 LIONEL E. MAWDESLEY-THOMAS

'

Salmo clarkii Rhubdomyoma, trunk Adami, 1908 Richardson, 1836

/Filmma, stomach Hunter, 1782 Fibroma, subcutaneous Bland-Sutton, 1885 Fibroma, stomach Bland-Sutton, 1886 Osteomu, maxilla Bland-Sutton, 1886 Osteoma, vertebra Bland-Sutton, 1885 SarcwmtL, swim bladder Bashford and

Huernangioma, pectoral Murray, 1908

Osteosarcoma, Murray, 1908

Fibroma, oesophagus Williamson, 191 1 Fi brosarcomil, Johnstone, 191 2

Murray, 1904

region

opercul u m

\ subcutaneous

Family PLECOaLOSSIDA&-JupallrSe fish Lipoma, subcutaneous Takahashi, 1929

Plecoglossw altivelie Myxoma Honma, 1966 Lymphosarcoma, ovary Honma, 1965 Adrenal tumour Honma, 1965 1 Haemangioendothelioma Honma, 1965

Temminck and Schlegel, 1846

F&mily OSMERIDAE-~tneltS Sg. cell ocircinoma, Breslauer, 19 15

Rhtibdomyoma, trunk Bergman, 1921

Oemerw eperlanw Linnaeus, 1758

Hyporn,asus nlioluv Sarcoma Gill, 1862

Hoshina, 1960

Family OALLLCHTHY tDA IC

Corydoras julii MeliLnornu, eye Cohen, 1965 Steindachner, 1906

Order LOPHIIFORMES

Family L0rHIII)AE-~oosefishes Lophius pdscxxtoriw Melanoma, skin Ingleby, 1929 LinnaeuR, 1768

Uadus m.orhua Linnneus, 1758

ROME ASPEOTR OF NROPIAARIA IN MARINE ANIMALS 187

(Osteoma, vertebra I Fibroma, orbit I Fibrusr~rcoma, trunk Dentigc?rous cyst, jaw Fibrosarcorna, oral Ii'ibron~a, stomach Haemangiomrt, swim

bladder (~mglioneuroma, trunk Fibrosarcomtt ,

subcutaneous Fibrosarcoma, ovary Osteotna, maxilla

Williamson, 191 1 *Johnstone, 1914(b) Johnstone, 1916 Williamson, 1919 Johnstone, 1920 Johnstone, 1924 Johnstone, 1924

Thomas, 1927(a) Thomas, 1927(b)

Thomas, 1927(b) Williams, 1929

Adenoma, parabranchial Peyron and Thomas, body 1929

Fibrosarcome, trunk Williams, 1931 Osteosurcoma, Thomas, 1932(b)

pectoral fin Fibroma, gut Thomas, 1933(a) I,yn~phosarcoma, orbit Wolke and Wyard,

1969 Melanosarcoma Mawdesley-Thomas,

\ 1971(a)

I Adenon~u, pectoral Takahashi, 1929

Gudus morhua gland

macrocephalus Adenoma, parabranchial Takahashi, 1929

(Tilesiu~, I 810) Adenoma, parabranchid Urellings, 1969 Neurofibroma, head Wellings, 1969 Neurofibroma Wellings, 1969

I Fi broma Lawrence, 1895 Fibroma, oral Fiebiger, 1909(a) Rhabdomyoma, gut Fiebiger, 1909(b) Fibrosarcoma, kin Fiebiger, 191 2 Fibrosarcoma, Johnstone, 1926

I ~ubcutancous Hwmangion~a, trunk Johnstone, 1926 Fibrosarcoma, mandible Johnstone, 1926 Osteosarc*omtl, Williams, 1929

vertebra Sq. cell carcinoma Williams, 1929 Chondrorna, head Thomu, 1932(c) Lipoma, livor Thomas, 1933(b)

188

Fibroma, subcutaneous Johnstone, 191 0 Fibrosarcoma, head Johnstone, 1922(b) Fibroma, subcutaneous Johnstone, 1924 Adamantinoma, maxilla Thomas, 1926 Fibroma, trunk Mawdesley-Thomas,

LION E L X. MAWDESLEY-THOMAS

Melanogramrnw

(Linnaeus, 1758) aeglejnua

Merlangiw m,erlangus (Linnaeua, 1758)

Merluccius merluccius (Linncteus, 1758) Molva holva (Linnaeus, I 758)

Therqra

( Pallas, I8 I I ) chalcograrwmut~

1 body Sq. cell carcinoma,

IFibromu, trunk Fibrosarcoma, trunk ’ Fi brosarcoma,

Hubcutaneous Osteoma, fin Melanoma, skin ,Sarcoma

Family ZOARCIDAE- IFelpou t H Zoarces viviparus Melanoma, rrkin (Linnaeus, 1768)

E’alnily BROTULIDAE

Parabassogigas crassus Sy . cell carcinoma (Vaillant) snout,

Family PoMAToMIDAE-B1Uefishes

Takahashi, 1929

Takahashi, 1929 Takahashi, 1929 Takahashi, 1929

Takahaahi, 1929 I’akaharrhi, 1 !b28 Nishikawa, 1964

Hergrnaii, I92 1

Mawdesley-Thomas, 197 1 (a)

Pornatomus saltatrix Lyinphangioepithelioma Harshbarger, 1972 (Linnaeus, 1758)

SOME ASPECTS OF NEOPLASIA I N MARINE ANIMALS 189

Family KYPHOsrDAE--Sea chubs Boops boops Thyroid tumour (?) Johnstone, 1924 (Linaeus, 1758)

Family CHARTODONTIDAE---BUtterfly fish Chaetodon8 sp. Osteomata Bell, 1793 Linnaeus, 1768

Angelichthys isabolita Adcmocarcinoma, Nigrelli, 1952 Jordan and Evormsnn, thyroid

1896

Family SCIA ENIDAE--~)rUMR Sciuena spp. Osteomata Linnaeus, 1766

Pogonias cromia (Linnaeus, 1766)

Sy. cell carcinoma,

Fibrosarcomtt , mouth

subcri taneous \OsteomiL, vertebra

Micropogon. undzclatita Odontotna, mouth (Linnaeus, 1766) { Sareornu., sulwutaneous

Micropogon opercularis OsteomaL (Quoy and Gaimwd,

1824) Yapillo I tia

Pal dlom LL

Papilloma

Qeny0nemu.u l inwtw (Ayres, 1855)

Chabanaud, 1926

Beatti, 1916

Beatti, 1916

Beatti, 1916

Roffo, 1925 Roffo, 1926

Bertullo and Traibel, 1958

Ruwell and Kotin, 1957

R.T.L.A., 1972 R.T.L.A., 1972

Fttmily EPH IPP 1 D A tc-spade fishes Chaetodipterus faher Osteoma, vertebra Schlumberger and (Broussonet, 1782) Luck6, 1948

Family STICHAEI i)m--Pric.klebacks

Lumpenw 8agittu Neurofibroma McArn and Wellings, Wilimovsky, 1966 1907

Family S C O R ~ ~ A E N I D A R - ~ C O ~ ~ O ~ ~ ~ ~ S ~ ~ ~

8corpen.a inermia Fibroma, eubciitaneous Takahashi, I929 Cuvier, 1829

Family TIZ~(*H I l r l tLI)A E ( ' ~ I ~ ~ ~ I L S S ~ ~ S I I I ~ S 1;rpidoprc.s sp1). Ostc*o~tl:t. f i l l (iervais, 1875 Oont~n Menidea Oer,tjlli~~/& hlrli~no~n:~, ski11 Nigrelli and Gordon, (Cope, i866) 1944

li:ilmily I'EIZC'IC*HTI~\.'I~,AE '1'(>111p(>ri~t,(\ I)~LSS Morone sn.tatill'cr N ( ~ ~ ) I \ I . O I ) ~ ~ I S ~ ~ > I I ~ ~ I Helnlboldt and (Wiblbaun~, 17!12) Wywd, 1971

14'aniily S X B ~ ~ A N I J ) A &---S~ii I)ILRHCX Serranus srribcc Tllyroici turnor~r (!) Marsh and (Linnaeus. 1768) Vonwiller, 1916

9armnus cabrilln 'I'livroid tumour ( ? ) Marsh and

(Linniieu~. 1758) Vonwiller, 191 6

t'ar'dental ttirnortr Roffo, 1925

tc'ibrosurcomn, l,a,/aobhrcr.r japonicu.9

Takahashi, 1929 ~t~bcutt~tieous

(Orlvier, 1 X28) ICiho~n~('OmiL, I ) ~ ~ L ~ Y I > X Takahashi, 1929

Itpincphelus ptrrzn Mt ' l i~1101 t ) i~ 1)ollf'~s at al., 193% (I,i~ini~ous. 1758)

Ll';tmily IAl1'I ' . lANII)A IC Sll;l])))(Ts

I , I L ~ ~ ~ T L I L , S grise t t s Nt~~lr.ofiI)ro~~lit Luck6, 1942 (l,innat~us, 175X) Nt.~~rofil)r-o~~lit R.T.L.A., 1972 Ltctinr~u.u rrpor1rt.s Nc~r~rofil)ror~lw J111ck6, 1942 (WnIl)tbt1111. 17!12) Lttlinn tts joc.11 N(~r1rotiI)r.o111i1 T~1ck6, 1942 ( Hloc'h slid Sclll l('itk'ls,

1801)

l~i~lnilp sr,raroa IC - Porgiclx

I'irgrus tt rr icwlor Ostconlittu Mawde~ley -Thomas, (Quoy and Cairnc~rtl. 1973

1824) /Osteomi~tt~ Kt~eama, 1924

C7hrysophryqs tnt! jor 'remniinck and t

Srhlegel, 1 843

Osteornil Stgawa, 1925 Adenoc.arcinomu, coelom Takahashi, 1929 Osteorna Takahashi, 1929 O~t~eoma Luck6 and

Schlum berger, \ 1942

A ehotvnr~jus U~~i~~io l )horornu , skin prohntocephnlus

( W a l b i ~ ~ ~ m . L7!12)

Family a0 l l l l l ) ~ ~ - - ( ~ O f ) i C ? ~

Bathygobius soporabl. (Cuvier and

Velenoienns~, 1 837)

AcanMogohiu.9 jlavintanus

(Temminck and Schlegel, 1845)

Sq. cell earoinoma

Papillomt~ ( ? ) Papillomt~, skin Sarcoma, undiff , Pal)illoma, skin I Papilloma, skin

I Papilloma, skin Papilloma, skin

Family TRJU Ll l )~~- -Sea rob ins

Lepido f rigln nbta Fibrost~rcomu, ( H o u t t ~ ~ y l ~ , 17X2) S I I ~ > C ~ I ~ ~ I , I ~ ( ~ O ~ I S

Chelidor~ichth!/s kum 11 Allol)horoni:i, lAesson nntl (:t~rnc*t, s ~ ~ b t ~ ~ ~ t ; ~ n e o u x

1873

Fnlnily I,A RRIDA R- \ l r r ~ ~ s ~ ~ - ~

I'latyglossus hivattnlu.~ T ' ~ ~ ~ ) i l l o ~ ~ l i ~ (Bloch, 1791 )

Labrzc~ mL.1 us Rtiat)tlon~yorn;t Linnaeuu, 1758

Tautoyolabrus Papilloma Giinther, 1 862

Family ECHB:NEIDIDAE--R~~~~~S

Echeneis naucrates Thyroid tumor~r ( 1 ) TAinnaeus, 1758

Thomas, 1932(b)

Mawdesley-Thomas, 1972

Oota, 1952 Nishikawa, 1954 Nishikawa, 1955 Oota, 1952 Imai and Fujiwara,

1959 Kimura et al, 1967(b) R.T.L.A., 1972

Takahashi, 1929

Luck6 and Schlum- berger, 1942

Order P L E ~ IWN RCTIFORRZ ES

Id'amily I3OTHl I ) A IC lA(lftlc>y(> iIo1111(1(~i-s

I ( )stJc*otnn, vcbt.t c.bri~

~S~ophth(ilm U S Myxott~i~. 11chitd nrauolicus Myxorrii~, tr~lttk

(Piklla~, I S 14) Pi ~ ~ O A ~ L ~ V O I U ~ L .

sub(~~~t,nnt~ous l?ibrosaruorita,

S U ~ V I I ~ , ~ L I I ( ~ O I I R

~Scophthal~~~us ttraritr~us Fibron~u, dorsi~l fin (Linnaeus. 1758) Rhabdonlyotricb. trunk

Pibrosurcotila, trunk Myxortiu, trunk

Lepidorhott/l)tca (:anglioneuromu. whiffingo n i s

(IValbaurn. 1 792)

Schroeders, 1908 Schroeders, 1908 Schroeders, 1908 Schroeders, 1 908

Johnstone, 1923

Johnstorle, 1924 Young, 1925 Johnstone, 1927 Williams, 1929 Haddow and Blake, 1933

Paralichthys olioncaus Lipomr~, trunk Kazama, 1924 Temrninak and Osteoma, vertebra

Schlegcl, I 846 Takahashi, 1929

Johnstone, 1912 Johnstone, 191 3 ,Johnstone, 1914(b) .Johnstone, 1915 .Johnstone, 1926

Panlily P I . I ~ : [ ~ I ~ O N TI I ) A tc -Righteyed flounders (I'i~pilloma, snout

Johnstone. 1926

Hippo!jlo~s rts

hippo!/loss (t* 1

(I,innne~ls, 1 75s)

Williams. 1929 'I'hornas, 1932(b)

Pi brort~ii, ooclonl tc'ibronlir, trl~nli Mclanorni~, skin I'ibrorni~,

si~l)c~itntttw~~s Id'i bros;~t*c.olt~:~.

~ ~ I I I I li

Bergman, 1 92 1 ,Johnstone, 1922(b) Johnstone, 1924 Johnstone, 1925 Johnstone, 1925 Johnstone, 1926 Sandeman: 1893

I I.ip011ii1. tr11111i Ith~b(tornyo~t~ii.

tl.ttnli (Myxorna, trunk

t'le trron~c.1~~ plntessr~ I,innateus, 1758 '

Fibrosarcoma, subcutaneous

Lipoma, trunk Fibroma. fin Melanoma, skin Papilloma, skin Haemangioma, trunk Fibroma, head

,Papilloma, skin

I'settichlhys nielnlrostir.l.us

Girard. 1866

Pseudop1~urorcrctc.s arnaric:cnius

(Wulbaum, 1792)

Papilloma. sltil~ Ly tnphosi~rcomr~, orbit

i Papillomt~, sltirl Papillomt~, skin

Pi~pillotna, skin Yapillorna, sltin Papilloma, skin pap ill om:^, skin

Pal)illorni~, skin

Wrc*llil~gs 0 1 ( 1 1 . . I!MR \Yc~lli~~gs I L I I C ~

(:htlinard, 1964 Chuinard et al.,

1964, 1966 Wellings et al., 1965,

1966, 1967

Sandeman, 1893 Johnstone, 1912

Wellings et al., 1966 McArn et al., 1968

Ketchen, 1953 Wellings et al., 1065 Nigrelli et al., 1965 McArn et al., 1968

Eryt 1lrol)horom:~ Smith, 1934

l'r~l)illon~a. skin l'aljillonlu, skill I'aj)illoma, skin

I Pal)illoma, skin Papilloma, skin Papilloma, skin Pa])illoma, skin Pii~)illoma. skin

I(irnura, st al., I967(a)

Young, 1964 Welling~, 1 !Hi!) li.rl'.lA.A., 1 !)72

i'acis, 1832 (:ootX, 1940 Wellings et al., 1 966 McArn et al., 1968 (looper and Keller.

I9fi9

Nippof]los~oit!cs WeUings i~tld

elttssodo?~ Clluinarci, 1964 (.Jordan tk11(1 (iill)(b~-t,, Miller and

1881) Wellings, 197 1

Hippoglossoi~lr~s t l / r b i ~ ~ s l ' t ~ l ~ i l l o ~ ~ ~ i ~ . s1ci11 I < i ~ i i \ ~ t - i ~ ~t / I / . ,

(Schmidt, I 904) l967(a)

Pibront;~, t rr~ltk Wellings, 1969 H i p p o g l o ~ s u ~ Fihronrt~, rittlrltliblo Wellings, 1969

stenolepis Fibro-lipotna, tail Wellings, 1969 Schmidt, 1004 Lipo-osteoma Wellings, 1969

Fibro-osteom a Wellings. 1969

Lepidopsettc~ hilin~crtn Nigrelli et al., 1965 (Ayres, 1854) Papilloma, skin Wellings, 1969

IJlyptocep11,alus Papillonla, skill Wellings et al., 1965 znch,irue

Lockingtoli, 1878-!)

G'l!/ptocephalz~s slrlleri Papilloma, skin Honma and Kon, Schmidt, I903 1968

Gl yptocephalus Papillolna, skin Hontna and Kon, cynogloseus 1988

(Linnaous, 1758)

Family ISTIOPHOR[I)AR

Maka.irn aurln.c k'ibromt~ Mawdesley-Thomas, ( Philippi) 1971

Family SOLEIDAB-- Soles

8olen aolen Myxom;~, cocllorn Johnstone, 1926 (Linnaeus, 1758) Papillolrla, skin Thomas, 1930

Family BALISTI I)A E- - Triggrsfishc~s and Filefishes

Alutara schoepji Osteomt~ Schlumberger and (IValbaum, 1702) Luck6, 1948

A. Sircv~ icl So t ;~r i ts (:all be i ~ ~ ( ' ( > r t i ~ i ~ ~ ~ ( l , no t~~rno i~ r s have been reported, as yet.

in the Sirenin.

I < . l'innipcdia The Pinriipedia include all seals, sea lions and walruses. They are

amphibious creatures, spending much time out of the water and belong to the family Otariidae, which includes many genera. To date only 7 neoplasms have been documentetl representing 7 species. From the examination of such a small number of tumours, i t is difficult to oom- ment, except to note that neoplasia of the blood and reprodrlctive systems account for all but one af them. Hepatic lipomatosis (Larsen, 1962), in all probability, does not represent a truly neoplastic condi- tion. Larsen said little about his two cases of neoplasia. one in a seal Phoca vitulina Linnaeus, 1758. This was a two year old female animal which presented with generalized lymphadenopathy, splenomegaly and hepstomc~galy. Hi~tologic;ll clia,gnosirs confirmed a lymphadenosis.

FIG. 22. Enlarged spleen from a Harbour Sonl suffering from lyrnphatlc leukaemia

SOME ASI'E('1'S Ob' N E0PI .ASIA [I\, MARINE ANIMALS

F~tr. S f . E~~l\tl.gotl lymph notlo from thv socond Herhour SoaI showlng ~ncrtba>ctl ret rculu- ontlt~t ht.lcnl syston.

The other ( ' ~ L Y C ~ . a four year old wa11'118 Odoh~?~zts roswur?rs (Linnaeus, 1758) also prcsc~lted with he1)rtto-syle~~o~nc~galy. Therc was evidence of i~ gttstritis, toget her with some ~l(~c1-atio11 anti haemorrhagr seen in the gastro-intestinal tract.

r l I hc tni~ligrtctnt letrliac~rnic~ Iy111l)lionictt2t described by Griner (1971)

wc.rcS SC(~I I it1 f wo tl~ikle sct~ls I'horrc vilz~lina gc~ronimensis Linnaeus, 1758 t~g(h(l I ) c ~ t I ~ ~ ~ c ~ ~ t I .I! R I I ~ 24 .ytba~s ('l'i~blt: IV) . Neither animal showed i~bnot.ttii~l sigtis : on(. i t1 facrt was fi)lllid dertd, and the other died after failing to ftbc.rl fbr two days (li'igs 22 to 24).

Two similar cases of generalized lymphatic leukaemia occurring in one areit, u.ci~ld suggest ~t probably common aetiology and in this case a virrtl agcbnt wi~s considered to be the most likely. Limited experiments wclre p~~*forntcd in which other seals wcm injected with 5 ml macerated rnaterit~l froni the spleen and lymph notles, but with negative results. An elephant seial Mirounqa Z~koninu (Linnaeus, 1758) also in the pool did not show tan-v signs of external disease.

TABI,E IV. (Modifiod aftor Grinor, 1971)

Ijlood I 'al~rex Diflerential Countx Seyvnented

Seal W . H.C.8 Cellx Lymphocytes Monocytes Eoainophils

' I ' h c b t t tcri~ic* trc~rnoluvs of tlic. gt.cby sc?ltl H'inlicltnerus grgpzcs (Fabricius, 17!)1) rc.t~)rtcbcl by Mawdcslc~~-Thottii~~ land. Bont~csr ( 1 970) were of interest as bllt*y tc.chl.cl Vtboni a 44-year-oltl i~rlirnal, the oldest living seal to have been ~.c~c+or,tlt*tl. 111 this animd, i l l c~ddition to an old pyometrtt which llucl bct,ornc~ ;I h~.tlrornetra aff(betitig thc left cornua, small pttpilliferoux tutuot~ra \rrcarcl ~loted which wtlre sl~own to be squttmous celled cctrcino- mtitn (Fig. 2'5). l'llis finding was of' interest as it represented a neoplastic c'hartgc ir i a ttiet-lplast~ic t issu~ (Fig. 26), the uterine mucosa usually 1)cing of tkc coltimnar type. This gynaecological curiosity also prc:- sent,cd sevcri~l cervical fibro-myomata (Figs. 27 to 28).

'rhr ovarii~n tumour in tt Southern Elephant seal Mirounga bonina (I,innueux, 1758) affechted the l ~ f t ovary. The entire repro(1uctive trar-t weighed 4-6 kg, the uterus and ovaries, including the mass, weighed 2.6 kg. l'he ovarian mass was lobnlated and covered by a thick capsule. Fibrous bands transvcrsetl the tumour, which was yellowish in colour, while ot1ic.r areas showed evidence of old haemorrhage and other dege~ic~rut~ive changes (Pig. 29). The cells which composed the tumour I L I I ( ~ irtvat(I~(1 th(> C L L ~ S I I I C wercB t?,l)ic.al gra~ittlosa cell types (Fig. 30).

FIG. 25. St.<: tion fro111 Grey Svnl Hrlli(*hOema grypzm) illustrating tli lutw 1 lcft cornua containing papilliferous turnours. The grt>.;.s tlistension is clearly shoxvn. (H. I I I I ~ I E;. Mag. x 0.5.)

Y I ~ X . 26. Trans\-em section in which ~vell differentiated squamouu cell caroinoma is seen infiltrating the surrounding musculature in a Grey Seal (Hnliehoerua qrypt tr ) . A marked clegrcur of i~lflnmmatory reaction is also seen. (H. and E. Mag. Y I%.)

FIG. 5. T\\.O cc.rvirn1 til~r~l-r~\yomtltn from n Grey Seal (Hnlirhoertr* qryp,rsL Typicill n-hitish \vhc,rlecl appearance. (H. and E. Mag. x 2.5.)

VIU. %. Tmnsveree section of the fibro-myomaka i n a Grcy Seal ( H a l k h r u s qrypue). The fumours consist entircly of connective tiw~le. (tT. and E. Mag. x 125.)

1'1(:. 20. 1.1%1~gt. t tlrnolir of thv left ovary of a S o t ~ t h r . r ~ ~ Elt.ptrfint Sf.81. T I J ~ I I I I I I ~ wr!iyherl C , \ . I~I . 2.0 kg, ~ I I S Iol)~~lttt.c?~l atl~t its ~.(lg'' 41o\v1~tl fitwotc- I ~ ~ r ~ c l s trav~,r-ilrg thr! .+trrzc- 1111.t. . 'I'II~. O \ V > I ~ ~ I I I I * < I ~ I J ~ ~ L ~ , ~ I I I I \v&s j,rllo1\i41 ~ I I I I I ~ I I I * Y I , \ V I ~ S ovi111.11r1: of w+rrIv olrl ~ ~ ~ I ~ ~ I I I ~ I I ~ I ~ I I I ~ ~ ~ s . l,c)\\or ~ I I ~ I S L I ~ L ~ I O I I 411.>\\ ~ I ~ I I ~ , * ~ I I < ' , .

E'I~:. ::I). 'Tht. rthil* fro111 s [ * t , c \ i ~ l ~ t ~ ~ ~ ~Ilustrutt.cl ilr Fig. L'!) ~,rt+sentetl as atypical g ~ ~ n ~ ~ u l o s d ~r l l* . 'I'llr cells \\.twL It~rgt. ancl c~~t~ri~inct l ~ I ~ I I I I I ~ I I ~ I ~ rytopla.%m. (H and E. 11%. - '300.)

I.'IG. :t I. Invlwioti of capsule of specimen ill~tstrated in Fig. 20. TIIP 1,n.clominance of small blood vessels IS Neon.

(H. nnct E. Mag. '< I".)

1 4 ' 1 ~ . 32. JIarlit.tl ckgcllrra~ti\.r c1lan~t:l. scc~l in hoth the int,ima and media of tphe blood vcuseb of t.hr qwcirnen in Fig. 29. the re.uw of \vhicll is tu~certain. It is prohahly age related. (H. and E. Mag. ,\ 20U. )

SOME AHPECTX O W NI~~Ol'lAASIA IN M A I t f N I A N I M I ~ I A S 207

The cells wero lttrgc! and c.ontuined obvious nuclei with abundant coninopl~ilic: cytoplas~n (Fig. :I 1 ). M11c.l1 arttolytic change had occsurred, :t11(1 hydinr drgctnert~t~ion W ~ L H H C O ~ i~r011nd mcmy of the nutrient arteries (Fig. 32).

T A I J I , ~ V. ' I ' U M O ~ J R S Ibic~~on~r~er~ IN THE P~NWIPEI>IA

Fox, 1923

Simpson and Ridgway, 1970

Simpson and Ridgway, 1970

Grirler, 197 1

It is sr~rprixi~~g that some of'thc c.ctaccaans, the largest animals on earth, have eve11 survived a t all ill their marine environment. This they have done only by caxtromc adaptattion of many of their systems ( Harrison, 1978).

Although apparently more inaccessible than the Pinnipedia, more C~LHCS of neoplttsit~ have been reported in the Cetacea than in the former group. Tho reported incidence of rleoplasia in both the Cetacea and, Yinnipedia can only represent but the tip of an iceberg, for unless a group of scientists is prepared to dissect large numbers of these animals when they arc slaughtered, which is often enough to deter the most enthusiastic, there is little chance of more than the grossest of lesions being recognized by the pathologist. Cockrill was fortunate in being appointed as Whale Fishery Inspector and Veterinary officer on the Sozethern Venturer in 1948149, and later on the Balaenu in 1961/52 and

muah useful i ~ ~ t b r ~ n a t i o ~ i wils gnthcred on these two voyages. Unfbr- tunetely t,lir zoonostts took priority ovcr the search for and clocumenttl- tion of ncoplasit~.

The first, threc tr~t-rlours rchl,ortcci in the Balaenoptera musculus (Li~lnttc:~~~, 1 758) wcrv it11 libro~tlyotllitti~ of the ut,c~rr~s; perhaps t h c ~ e records n1i1.y h i t~ t ' 1)(31111 ~ I I V t o tilth I \ I I I I ) \ ~ ~ H ~ S p \~( ' e ( l 011 the T(:(I\I~:H~H of the reproducat,ive j)hysi(~~ogi~ts who ot%(b~l itslcctl fi)r t,hc cntirth rc?procluc- tive tract to be removccl in toto tixctl i l l formalin i ~ n d sent bark to the laboratory. 'l'he fibronlyomik dcsc~ibetl by Stolk (1 950) was found in an cxtrcmely oltl itnirnnl which rnea~snrcct 26.8 n1. Thr tumour measured 45 cm by 50 ern. I t I)rcsc~it,c~tl no p~rticaular or clistjinctive histological findings. It WiLa ext1.cvnc4y similar to tliose t,uniours found in the human uterus. ltowell and Willis (I!)49) i~lso rec.ordct1, fiom an unidentified species of the Cletacea, ;ti1 ovoid well-cirrnmscribed mass of 4 cm in diametrcr attiiclird by a sllort ycdic~le to ;t more spongy piece of tissue. On hist,ologicttl examination, a typici~l structure of it benign fibromyoma was found, some areas of normal endometrium were also noted. Slijper ( 1 949) report(ct1 si milur findings, although fewer dotails were given. Uys and Best ( 1966) not,ed fibromyornata in Physeter catadon (Linnaeus, 1758). Thest: were scen in a 10-7 rn r~ninlal which was approximately 19 years of itge. Thtl uterus " mass " weighed 112 1 g and metlsured 13 x I2 x 8.5 om. 'Phis mass was found iu the right cornua, although several srnaller Inasses wchrc fi)uncl in the left, cornus. No degenerative changes wcrc wen in itny of the tutnours whicbh hi~t~ologically resembled the typicaal.fibrornyomi~.

Sevcbral O I L R ~ ~ S of O ~ i ~ l ' i i ~ 1 1 11(*0plit~ia Iiitve ~ C C I I docl~rnented. Stalk: (1950) rccordcd n citrcinoma of' tllr ovary in itn cxtremt:l,y old Balaenop- tera phy~nlus (I,i~ini~cus, 17.58) hut Sitilrtl to state in which ovary it had occurred. The tuti~oirr consistctl of a ~lociule 3 ern in which several stnaller iiotlules wvercJ noted. 7'11~ cells arc described as " b1addt:r- shaped ". 'J'hcre call be little doubt tlli~t these are tu~nour cells. There was evidencc. of son~e tlc~gcnp~-ativr change, together with areas of hya- line degeneri~tion. Thc t~~nioul- woulrl, in all probability, be a granulosa call t,urnollr i ~ s previo~~sly desc.rib(hd in Miroungn leonina. The occa- ~ i o n ~ l cystic spacc was also noted. Rewcll and Willis (1949) reportetl a muc!inous cystadenoma of the ovary in Halaenopterc~ musculus. It was a rnultiloculate cystic t,rlmour about 30 ctn in diameter. The superficial part of the mass consisted of layers of fibrous tissue, in the central part of the niass several cystic spaces were seen around a yellow core. There was no evidence of any haemorrhage having occurred. Histological examination showed normal ovarian tissue a t the periphery, sonle corpora lutea were seen but no developing ova. The cystic spaces were

SOMR AAI'ECTY OF NEOP1,ASIA TN MARINE ANIkIALS 209

l i~~e t l partly hy c.olurnnar mucr~s secreting epithelium and partly by irregularly strat ificd epithelium. 'I'hr c>ent,ral yellow core consisted of degenerate c*ell,.i. In the same paper, Kewell and Willis (1940) described grsnulosn (.ell trrmoura, two in Balasnoptera mwculus (Linnaeus, 1758) and one ill Hnlnenoptera physalw. These ovarian cysts were about 3- 3.5 cm in diameter, and all presented with partially cystic characteris- tics. The cystic spaces were lined by granulosa cells in various stages of degeneration. None of the cysts was large enough t o present the typical " rosette " appearance seen in smaller mammals. Some degenerative changes were aec.11, together with some dystrophic mineralization.

Although tutr~ours of the reyrodrlctive tracts of several Cetacea have been documented, other ocrasional neoplasms have been noted but as they are relativcly so fcw in number it is not possible to classify them into any particrilar grouping.

Rewell and Willis (1949) reported a papilloma of the tongue in Ralaenoptern, musculus. This was a pedunculated mass of 8 cm in diameter. It presented a coarsely papilliferous surface with some pigmentation of the deeper layers (Fig. 33). Histologically, i t consisted of a benign squamous celled papillotna. Pigmented dendritic cells were noted in the deeper layers of the epithelium. Stolk (1952) described a fibroma of the tongue in a pregnant Balaenopteraphysalw. It presented as a firm pedunculated tumour which measured 55 x 44 x 29 mm. The tumour consisted of dense connc.rtivt* tissue elements covered by squarnous t.pitheliun~. Stolk wcnt on to describe an adenoma of the liver in Tursiops lrzcncatus (Montagrrc, 1821) although the fact that these nodules w i ~ c multiple and the only microscopical picture shown is one of bile (1uc.t proliferation ~urroirndetl by dense connective tissue lind es no hepatic. parenchyma waa shown, this diagnosis is suspect. It is much more likely to represent a degenerat,ive change or a deposit from sornt. unexami~letl site. Often when examining the complex stomach of the dolphin particularly thc fundus and pyloric parts of the stomach rnliltiplc " grape-like " clusters are seen, often in great numberrl. On first macroscopicul examination each appears as a papilliferous tumour hut they are in fact parasites (Schryver et ul., 1967). They are strigeid trematodes known as stomach flukes (Brauninu cordiformis). It is a pity that although, in his day, Stolk must have been one of the most prolific writers on diseases of the fish and marine mammals, one is in- clined to be sornewllat suvpicious of some of his findings a8 he only used a small nun~ber of illustrrttions and later tended to draw what he saw. Stolk reported various other fibromata on the lower jaw, and subcut- uneous tissue but they do not warrant further comment. Uys and Best (1966) noted a dark nodule on the lip in Balaenoptera borealiv (Lesson.

FIG. 5-1. Group of nttrvo ceiis in a gangliolle~iroma In Bulaenoptcra ~nuacdu8. (H. and E. Mag. , 90.)

1828). This 11odr1lc Kiln ul )o~~t I .!i c:nl in tiismeter, and this Sei whale wiw t h o ~ ~ g h t to I)(. ;~ged ul,out 48. Histologically, this tumour had ~hects of c(4is hcuvily packed with melanin which obscured all else. Many srnall ;~rterioles were sc?en in the tumour. After bleaching the cells with hydrogcbn peroxicte the (~11s wen: seen to have vesicular nuclei, wit-h

WIU. 16. Fibro~nattl. of ~11out.a in Urcbenoptsm nhzcecul~~. The papillifernus natum of t h ~ t,rctnour.r, b clttarbly seon.

promiuent nucleolus, these cclls contained more cytoplasm than those cells which did not contain pigment. S o functional activity was noted at the epidermal-dermal junction; the overall comments by these uuthors was that the tumour showed evidence of hamartomatou~ proliferation involving melanocytes and arterial-type blood veesels.

E'inally, some tumours of the central and peripheral nervouu system urc noted (Fig. 34). Rewell and Willis ( 1 950) recorded a ganglioneuroma

FIG. 36. Transver* =tion of fibroma in Bnlnenopte-ra museul~ur, showing some cellular infiltration of the chronic inflammatory type. (H. and E. Mag. x 12.5.)

&>a. 37. Trenweme sect.ien of fibroma ahawing a far l a ceiluler picture than the previous sectioxr. (H. and E. Mag. x 126.)

F'IG. 38. W i o n of oeteom of t b glenoid fossei in Balaelaoprcm phywlw. (H. and E. Mag. X 145.)

Fro. 39. Testicular turnour in Tuc&pn truealw.

FIG. 40. Adrenal tumour in Tureiops IruncaCwa. s 4

'I'Alt1,IC VT. T ~ J M O I T R X ~~EJ'ORT~~:D I N THE CETACEA

Halaenoptera nbttnotlux (I.~IIII~WIIN. 17.58)

Not givc*ll

Ralaenoptern nrrtaotl~tx I1alaenol)tem /~l~y.qal?m HalaenCoptera nrrtnc~tl~rx Turaw~)a trtrncal?re

(Montagll, 1821 ) I'hyaeter catdon

(Lirinuntt~, 1758) Hahenoptera g~lryaaltu, Balaenoptera phhyaalw nfegaptera ~zo~iaenn,gllia

(Ron~wski , 1781) hfegaptera no~meam&&ae Iblaanoptera i ) l t y n a l ~ I'lbyaetcr cntodorr I 'lbyneter aatorkott I#alaenoptera ph,!jxnlt~x Hahenr~ptcrcc nburrrrtlun I'hpcter catn)do~t /{ahenoptera Ooraalin

( I r t ~ u ~ o r ~ , 1828) .4fylnptera r~)t~rtenngliao Olc>binephala ntelnenn

(Trnill, 1809) I{alaenoptern pttynalw

L'ibronlcb of lowor jaw

Sttbcutaneous fibroma Fibroma of tonguo Fibroma of tonguo

Fibroma of lowcjr jaw S ~ t h c ~ ~ t a r ~ o o u s fihrornlt S I I ~ V I I ~ ~ L ~ I O O I ~ R f i l ~ r o n l ~ H w ~ ~ ~ n t ~ g i o ~ n c t of livol- Hodgkins tliwon~c. Fihrolipollia of uton~tic!h F i b ~ v ~ ~ n y o ~ ~ t ~ t e of ~ttot*rln Botrigr~ ~ncal~ll~rttic! tit tnoltr

of tho lip 1.ip0111a of t11o I ~ r t t i ~ ~ stern Coitrc*

Hrwc?ll tuld Willis, 1!)50 Stalk, 1950 Stolk, 1950 Stolk, 1962

Stolk, 1952

Stolk, 1952 Stolk, 1952 Stolk, 1962

Stolk, 11153 Stolk, 1953 StoIk, 1963 Stolk, 1953 Stalk, IY5:4 Cockrill, 1960 Uye awl Rest, 1966 Uys 11.rlt1 Bout, 196fi

Mawdewluy -'l'llolnas, 1973

Mewde~lcy -'l'ho~nm, 1973

Mawdosloy -'l'hornt~, 1973

M a w d ~ ~ l o y -771orr1at~, 1973

Mawdexley -'l'l~ornns, 1973

SOME AAI'ECTS OF N E ~ P L A N I A I N MARINE ANIMALS 219

in the anter.ior rnediastin~~tn in Bcilnenopt~rrr 1r~uscu1u.s. Histological e~aminnt~ion xtiow(~(i H nli~t~rix of collagenol~s tissue in which large k)undlcs of non-rnc~dolli~tcd and ~nedullated nerves were found. In some xi*ctions woll tliffercr~tiatctl neurbons were idso ueen. A cerebral nenro- fibroma (T'illtlri. 1968) wihs re1)ortect i11 Ij(z2aenoptern ph? / sn l~ . TWO neurofibrornittfi locat8ed at the Ibase of thc ventral paraflocuhu of the cerebellum were ;~lxo re(-orded by this ibuthor. Both tumours contained a n~at~rix of collugolious fibres and rrlJcc1 to nome nervous elements. r 1 I t~ose werib remnsnt,~ of' atrophic niyclinated fibres, together with some ganglion cells. Pilleri ( 1 966) has previously described a lipoma in the bane of the brilin of n hlt~qaptem novomn!lliue (Borowdki, 1781), which WiLS fount1 i ~ t thc junctioli of the pons anti rnedrllla. The tumour was hon~ogcnous I L I I ~ contilincd angiomut~oes sl)aces within it. The adjacent cbhoroitf pltbxns sllowctl ovidcnce of xomc strand proliferation with i~trophy of t l ~ r 1)i~pillae and. h.yperplnsia of chorioidd proliferations.

Stolk ( I M t i ) described a case of (:ranulorna malignum (Hodgkins I )isease) in I3olnrr1,optera ph?jsalw. The paper is a disappointing one in that it spcilds too long discussing the disease in man, which in this perticulttr (*ontext is irrelevi~nt. Again, the illustrations are poor, and little cotnrneiit iasn be made. This paper, like so Inany of his fish papers, is bedcvillrtl by c*ibrneriL lucida drawings which are so stlbjective as to I ) t h alrnost vnl~iclt~ss.

Rcco~itly furt l r c ~ sprcin~onx 11i~ve 1)et.n cxamined in this laboratory, ~t~vcrctl of' whic*h canme from the K. R.S. l)iscovar!j expedition. They includc sevc-rtd i~~tcrcst~ing turno~irs (Pigs 36 to 38).

V I . SUMMARY A N I ) (:~K(:I~LTSION

All the a~iirnnls c.onsitlc~red ill this t~rticlr h a v ~ one thing in common in that tl1c.y live either totally or partially ill a watery medium, and it is ciiffioult to dismiss the eflect of the environment on them. There are lnnny questions which hnve been raised, and only a few have been answered. The reuson for this is simple: there is much ignorance surrounding the i~~iinlalv examined. Yet one cannot escape the conclu- sioli that some of these animals if, and when. suitably investigated, would possibly show an aetiological agent, either local or general, which clould be responsible for some of the neoplasias described.

I t is difficult to appreciate why oysters and clam8 do not have more tumours, particularly when one considers their biology, ecology, their mode of feeding, and the fact that they are " bottom " filter feeders. It is an irlteretlting fact that over 10 p.p.m. of arsenic are permitted in shellfish as opposed tJo land animals. Arsenic is one of the

few, if not tjhe only, tc111)stancc wl~iul~ cituaBs tunlours in tnan but not in animals.

The usc of oli~tns iud oysters as itssessors of biological risk should not be beyond tjhc ~oopo of' the! interested parties. Since they are biological concentrtztors of ull types of niatericbls, their study should be extretnely rewarding scientifically aR well as being highly profitable. The choice of site for such an investigation could be of the utmost importance as Psuley (1969) examined 30 000 oysters and found only six instances of disease. Of d l the animals examined, the oysters and clams offer the best aetiological monitor for the environment in that they are essentially static animals. Marine fish could be useful in the study of the genetics of fish neoplasia, the environmental hazards essentially being of less importance, apart from such instances aa fish inhabiting brackish water c~rld the Salmonidae, for their avoidance reaction. The only problem which could be relevant is the effect of pollution on essential food chains.

The Cetacea and Pinnipedia are of especial interest, since many of the Cetacea have been " over-caught " for many years, and now are on the list of endangered species. The industrial value of whale products is also an important factor. The possibility of sharing cancer research in the species is remote. The dolphins or porpoise again, while contributing little to neoplasia in general, have proved to be of great interest and value in experimental physiology.

Thitl review has been essentially an exercise in bringing together in one place t~nd at one time the majority of current data. Some of the uses that some of these species could I)e put to hsve been briefly discussed, in the hope that ultimately the tu~derstmding of neoplasia as a whole may be inlprovcd.

VII. A(~KNOWLE~~UEMENTS The author would like to thai~k Dr J. @. Harshbarger for permission

to publish Fig. 2, the editor of h'cience and Dr D. F. Squires for Fig. 8, Dr L. A. Griner and the editor of the American Journal of Veteriw~ry Research for Figs 22 to 24 and Drs Rewell and Willis for Figs 33 to 44, Figures 25 to 28 were previously published by the author in the Journal of Pathology and Bacteriology, and Figs 29 to 32 in Pathologia Veterina- ria;, and thanks are expressed to the respective editors for permission to use these illustrations.

Thanks are also expressed to I)r A. N. Worden for his continued help in the author's pursuit of comparative pathology ; Miss Margaret Shafto for her unstinting help with the bibliography; Mrs Sue Oxb~~ry

SOME ASPECTS ON' NF.llPI,ASIA I N MARINE ANIMALS 22 1

fix nlr~cll help with M~ct i~rticltb; and to Mrs Pauline Heppolette for typing the ~naniiscript. Witllor~t the support and assistance of these colleagues t h i ~ article wor~ld never have seen the light of day.

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Kimnra, I., Kugiyt~mn, T. and Tto, Y. (1!)67t)). I'npillo~nrltons g rowt l~ in sole from W a k m i ~ Bay I C ~ ~ I L . I'roceeriing~ of the Society of Experimental Biology and Mediaitte. 125, 175 177.

IClontx, Q. \\'. (1!)74). Htrumnto1oplc:tll t(.cllnic~l~cs ant1 the itnrnc~rlr* response in rainbow trotrt. 111 " Dtuc\usc*u of I+'txl~ " (Ed. Mowdosley-l'l~omas, L. E.). Sy~npo~viv~nc of the Zooloyicul Rocirty of /,omion. iVo. 30. pp. 8!# U!). Academic Prnns, lAottdor~ t ~ n d Now York.

LnbbP, A. (1!)30). 1Jno trtmoltr corrlplex c . l ~ r b ~ rln rrrt~rlnn. IJtilletin rle I'Asmciation F r a n p i ~ de 1'Et~rde t l z r Cancer, 19, I 38 1 58.

l,ctdreyt, P. (1929). Sur ull odor,to~nr e~lt~ttl(h c1lt.z r l r l Srylliuva cattcltco. Bulletin de l ' l~utittrt orc'anographiqtle, Afonaco, 539, 1 4. (011 11, cut~tnrolln oclo~~torrtc~ ill a Scyllium cntulue.)

Lt~drtayt, P. (1930). Sur r l r1 rhahdon~younrcotne c11c.z lrr~ Labmu mi3-~TM. Hullelin de l'lnatitut ocdanographiql~e, Monaco, 550, 1 4. (011 a rtlahdr>niyo~arc:orna in a Inbrus mixtua.)

Ilrtdroyt, F. (1935). Hur un epitheliotnn d~ la nltlclrleusc~ palatine chee nrtc. mrlrerrrs (Muraena helena). Bulletin de l'lnstittct ocfanographique, Monaco, 677, 1- 16. (On a n cpit,l~olioma of the palatlne mucous membrane in a marine eel.)

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I~uck6, 33. (1 938). Studies on tnmt)l.s In cold-bloodad vertcbrater. -4nnua2 Hepurt of the Twlugaa Laboratory, 11137-1 938, 92- 94. \Vash i~~gto~l D.C'. : C'arncpio Inst.

226 LLON'EL 12. MAWDEHLEY-THOMAS

Lrtck6, U. (1942). 'I'tuno~trs of tlitb IIt3rvct sheaths itt finh of tlie snapper fa~nily (Lutianidae). Archioea of Ihthology. 34, 133 -150.

l,uck6, B. ant1 Scl~l~~rnb~\rger, H. 0. ( 1941). Tra~iqlantable ep~theliorria of the lip attd rno~ttti of catfish. I . Patl~ology. 'Cransplantation to antertor chamber of eye and into cortlett. Jorlrnal of h'xperi?nerstal Medicine, 74, 397- 408.

Llok6, B. a ~ l d Schliin~bf~t-gar, H. a. (1942). Common neoplasms iu fish, atnphl- biann and reptiles. Jurcrnal of the I'echnical Methods and Bulletin of the International Aaaoriation of Medical Muae?tme, 22, 4-9.

Marine, D. allti Le~lhart, C. H. (1910). On tho occurrence of goitre (active ti~yroid tiyporplania) in fis11. Bulletin of the Johna Hopkine Hoapitul, 21, 95-98.

M a ~ s t ~ , M. C. and Vonw~llor, 1'. (1916). Tllyroid turnour in the sea bass (Serranzu). Jortrvaal of Gavwer Research, 1, 183 -1 96.

Mawdesley-Thomas, L. E. (1969). Fish farming and tho veterinary profession. Veterinary Record, 85, 134-138.

Mawdedey-Tliolnaa, L. P2. (1970). Significance of Liver T~~rnour Irlduct~ort in Animalu. 111 " Metabolic Aspects of Food Safety " (Ed. Roe, F. C. J.), Rlmkwell, Oxford, p. 481 531.

Mawdesley-Tl~omas, L. E. (1971a). Neoplasitt in fish: A roview. In " Current Topics in Comparative Pathobiology ", Vol. 1 (Ed. Cheng, T. C.), pp. 88- 170. New York and London Academic Press.

Mawdesley-Tliomaa, L. E. (1971b). An ovarian tumour in a Souther11 elephant ma1 (Mirourga leunim). Veterinary I'athology, 8 , 4-15.

Mawdesley-Thomas, L. E. (1972). Some turnours of fish. In " Diseasee of Flsh " (Ed. Mawdesley-Thomas, L. E.) Symposium of t h Zoological t society of London, No. 30, pp. 191 -284. Academic Press, London and New York.

Mawdesley-'l'l~onms, L. E:. ( 1 973). Present and Potent la1 use of Tissue Culture in Toxicology. In " Prog~~t~sn wltho~it Pa~ti.'' Murray, Edwards & Co. London.

Mawdonley -1'llomct~. 1,. E. atld Harrq , I). H. ( 1970). Aeld and alkaline phosphatase activity i r t tho Ilvor of br.o\cn NII(~ ~ i l ~ b o w trout. Nature, Lo?ldon, 227 (5259) 738-739.

Mawdesley-Thornp~, L. IC. arltl I<ori~tt.r, U'. N. (1!170). Uterine turnours in a grey soul (Halict~oer~is (lrypue). Jour?tccl of I'atholoyy, 103, 205 208.

Mawde~ley-'rhornfbu, L. E. arlcl U~rckc., 1). (1973). Some di~eases of coarso fish. Proceedings of the lith 13ritish C'oawe F i ~ h Covtference, lJniversit.y of Liver11ool.

M~wdesley-TIio11i~s I,. E:. a11d I.rahy, d . S. (1907). Organocl~lor~ne pc.sticide ronidue~ l i t piko. I'rogre~siz~e Ir'iah Culture, 29(1). 64.

Mewdonley-'l'ltotna~, L. E. acid Nowrnan, A . J. (1973). Spontanc:oun turnours of tile C.N.S. in the rat. Society of Toxtcology, 12th Annual Mor-t~i~g, NPW York, Mt~rct~ 1!)73. Abstract p. 115.

McArn, C. atid Wellinge, S. K. (1967). Ncurofibr-o~na in a tc.lnont fish, Luvrq,r,nw aarlitta, Journal of the Fiahery Hesearch Board of C u v ~ d a , 24, 2007 2009.

MoArn, C., Chinard, R. O . , Miller, B. S., Brooku, R. E. and Welllr~g~, S. K. (19fjB). Patliology of ukin turnow found on English sole and starry flounder frarr~ Yuget Sound, Wasllington. Joztrnal of the National Cancer Institute, 41, 229-242.

McIntoslr, W. C. (1908). On a tumour in a plaice. Annals and iMagazine of Natural HLtory (Ser. 8), 1, 373-387.

Millernan, R. E. (1969). C~ted by Wdlings 1969. Miller, B. S. and Wellinlp, S. R. (1971). Epizoot~ology of turt~ours on flathead

Sole (Hippylwoides e l m s d o ~ t ) in East Sound, O r c a ~ Island, ITashlngton. Tranuaction of t k American Fihheriea ~Society, LOO (2), 247- 266.

SOME: ASPECTS OF NEOPLASIA I N MARINE ANIMALS 227

Mnlclthg, M. (1!)70). '1'110 t l~ylnus gl~lltl I I I I ~ ly~npl~onarcoma in tho pike, Euoz 1ucilc.u I,. 111 Ir18lt111cl. I'roreedinp qf tltc lnt~rt~dionnrl Sympoeium ojCompra- tizw 14etrkraenitcc Ilexc,rrrch, i5ik)ltllcn. 1.11o1nat ., 4, 600 -60!).

Murray, J. A. (1908). 'rllo zoologicel diet l'il)utiol~ of cancer. Scientific Heport and Inz?e.utig&iona of the Imperial Cancer Reeeard Fund, 3, 52-60.

Niy~utll~, R. F. (1952). Spontunsot~s ~~uoplwms in fishes. VI. Thyroid tumours in marirln 6st1c.u. Zoologica, New I'ork. 37, 185 189.

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Nigrelli, 1%. B., KoLc11ol1, I<. S. ancl It~lggieri. O . I). (1965). Studies on virus disonsc~ of fisltoe. Epizootiology of epithelial t ~ r ~ n o r s in the skin of flatfishes of tlie I'acitic c~>ast, with specla1 rofolu~nm to the sand sole (p8ettichthy8 melanoatdoua) from Northern Hecatcb Strctit. British Columbia, Canada. Zoologicn, Nett, Tork, 50, 115-122.

Ninhikawa, S. (1954). 011 the tu~nu,ro~~s growtl~ observed in two Ashes. Collecting and Breedilcg, Z'okyo, 16, 236.

Ntuhikawa, H. ( 1955). (A caw of spindle cell sarcomu, developed in the Japanese common goby ( AcanlhogobiusJEavimantm). ) Journal of the Shimone& College of f i h i n g , 5, 171 174. (In Japanese.)

Oot,a, K. (1952). An opidemic occurrence of tumor-like hyperplasia of epidermis in a specias of fish, Acanthogobi~mfivimanua. Qann, 43, 284-266.

Pacis, M. It. (1932). An opitholial tumor of Parophrye vetulua. Master's disserta- tion, Ulrivor~ltg of Wasl~~ngton. (Cited by Wellings, S. R. 1969.)

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Pa~~ley , G. 13. ancl Sayco, (:. S. (1907). Description of sornt, abnormal oysters (Craaeo~trea !~ipt) from Willapa Ray, Wwhington. North Weut Science, 41. 155 169.

I'n~rloy, G. R. ttrld Saycc*, (!. 8. (1968). An intmnal fibroun tumor in a Pacific ny~ter , (?rmxoutrea gigaa. Jou~ttal ~f lauerter',rde I'tathology, 10, 1- 8.

1't*rri11. W. 1'. (l!)(i!)). Tllo bnr~liurle, Cr,n*.lrockrma auritttm, on a porpoise (Stenella graflmani). Jottrnal o j Alamnmlogy, 50, 149 151.

l'ttyron, A. nnd Tllolntm, L. (1929). Contl.ibl~tion a l'otude dos tr~mcure du revete- rneut brericl~it~l cl~oz len poissorm. H~tlletin de Z'Amociation franpiee pour I'Pttde dti can(-er, 18, 825-837.

1'nytso~i, A. n~id T l ~ o ~ ~ i a s , L. (1 1130). I,om t111n~11rs tti~yrctidionnen des salmonides. Bttlletin rle /I'dx#(>~;dio?l jranpiee potir 1'Ptudc du rancer, 19, 795-819. (The tl~yroid tu~noum 111 e~ln~or~ids . )

Pick. L. ( I 905). 1)er Schilddriisenkrt.ny tlcr Sltlmonideri (Edelfischo). Berlin tierarztl. FVschr. 42, 1435, 1477, 1498, 1.532. (Cancer of the thyroid gland in snlmonids (rarefish).)

Pilleri, a. (1966). Hirrilipon beim Buckeral Megaptera novaeandiao. Pathologk C'eterilbaria, 3, 341- 349.

I'illcbri, G. (1988). C~t.rohrn10 Neurofibrolna boirn Finnural, Baloenoptera p h y d w . I'athologia Veterinuria, 5, 35-40.

YIHIIII, M. ( 1902). Bosartiger Kropft (Adenocarcinoma der Thyroidea) bei Salmoniden. .4llgemaine Piachereizeitung, 27, 117-1 18. (Malignant goitre (adenocarcinoma of the thyroids) in salmonids.)

1'1.irtc.cj. 11:. 11;. ( I H!)I ), Mt>It~tto~t~r(.ot~tit 111 (I Jtt~tldock. ((>it0(1 by Tho~nmi, L. 1931a.) LCo\r,~.ll. It. 16. tttt11 \Vtllis, It. A. (1!)4!)). Sor~ro trlnrol~rs found in whales. Journal

of J'trt/~olt~!l,~l a~ t r l Hnc~terioknlti. 61, 454 456. 1<1\~~11 . I t . K . tt1t11 l\'tllts, I < . A. (1050). Sottlv 1111nottlw in wtld t~ltilnals. .Jot~rnal

r,f I'crtho/o!l!j cr11ol llnrtario/o!]!/. 62, 450 452. ZCobcrlq, LC. . I . ( 1!)7Z). Orttl ctrrc.~ttctrtttttt~ t r l r t Hc~lrrlon, Saltno aalrtr (I..). I'etel-inary

Hecotrl. 91, l!)!l. ltoff~o, A. H. ( 1 !)25). S o h t ~ 1111 l I I I I ~ O ~ puntdt~rltar~o crr~ la corvina. Boletin del

I&vtitc~b tit] medici~arc r.cj>critnewtnl pnrn el estrdio y tratcamienlo de cdncer, ~3tiotlos Airos, 2, 28 42. ( 0 1 1 II I)(~tttgtt t l trno~~t' in tho sea-bass.)

ICoWo, A. H . ( I 92ci). 8.let-colnt~r t't~nocol~tlt~r d c . caorvir~a. Bolet4n del Inetituto de ~rrrtlicir~a e.r/~arirnrntul para el rs~rtdio y trata~niento rle ccincer, Buenos Awes, 3, 206 807. ( V I I H ~ ~ O ~ I I I nttr~.oitlt~s of tlln scqtr-hass.)

lCiiss(~Il. k'. It. ~itt(i Kotitt, 1'. (l!t57). S(~t~nrnotts p & p ~ l l o ~ r ~ a i r i tlte white croaker. ./otrr~tal rf the Nntiottol Ca~~c.er 1?1ntit11tc, 18, 857-861.

Itytlcsr, .I. A. (1HH7). 0 1 1 11 t l l t~tor in ~ 1 1 ~ 3 oyster. Proceedings od the Acdemy of Natttral iVr.ir,rtces I!/ /%,ldelphza. 44, 25 Z7.

Hagt+wti, 1':. (I!)25). ZI IP Ktrttltirtis (lcr ~ ~ ~ c h o l t g ~ s c ~ ~ w i ~ l s t ~ e . Canrb, 19, 14 15, ( 0 1 1 tlil. I<~IOWI<YI~(' of lint1 tlllnollrs.)

& L I I ( ~ P I I ~ I L ~ ~ , ( i . (18!)3). 011 tho ~nttlliplo t t ~ r n o u r ~ in plaice and flounders. Repart cf 1/14 fi7c'ix/rer?y ljoard of Scotland, 11, 391 3Ib2.

Schcwrt~t.. i 3 . I I I I ~ l,ot~hItot~d, Ma~rgar.~t H. ( 1!)50). Tumollrs in the invertebrates: A rc*vio~-. C!UIZCET tle.sr~arc.11, 10. 403- 419.

Bcltli~rnk)orl;ri~r, H . G. and Kutx, M . (1956). Odontogonic trimom of salmon. Ca?rc.er Hoviearrh, 16, RG!) 370.

S e l ~ l u r n l ) c ~ ~ ~ ~ n r . H . (1. nncl L11ck6, 13. ( 1948). Ttrl~rotlrr; of fishes, amphihiann and roptilox. C'ancer 1lc.venrc.h. 8, (157 754.

Sclt~~ocxj~~r:.l. V. 1). (1 !b08). ('l'ttmot.s of lishox.) Dissc>rtatiou ( I n Russian.) (Transla- t,iott I r i Ar111y Mtwl~cttl Lihrnry, Wanltirtgton, D.C.) St. Petersburg. (Cited I)y 8cltIritttl)orfic~t~. H . (:. crrld I ~ l c k b , 13. 1948.)

Scluryvcbr, 1-1. I?., Mc>tiwcry, Mr. l b r r t l W~lliuinx, J . 1'. (1!)67). Tlte stornach fluke, 13rac~nirto corvlijorniin r t l tltc. Atlnrtt~c f % o t t l t - ~ ~ o ~ ~ clolphin. Jolrrm$ of the Anccricf~tr I'eterinur?/ Aunoc-idiot/, 151, 884 888.

Sir~ilit~rro, M. .I .. I'cril~t~tt tcr~', A. t~ttd ('I1irI~, E:. (1971). Effoct of sex or1 tho dc4velop- rncnt of trlt~lnnolnn. 111 Iryl)r~tl finlt of tltc gtstrrls Xiphophorus. Cancer R w e a d , 31, 725 7!!!).

S i p s t J . . I I , . H ( 0 ) . Unpuhlishod dtabmostic data from tho Mttrlnt. I31onctertc~. Ffictlt t j , Point Mrigu, C'nlifornia, 1966 1970.

B1ijlx.1'. b:. t J . ( 1!)4!l). On some ~)ltt~ttornt,r~tt concerning pregnancy rtrlcl parturitilies of tlte C'cbitic.t.a. I?ijtlmyetc tot rle rlierktorde, 28.

i t I . . M . ( I ! ) .A t n t ~ ~ ~ t t c ~ l t ~ r n c ~ l t r ~ ~ n o r in nn oyster (0,vtrea krginica). -4111rricarc .lor~rttn/ of C'c~rrrer. 22, 838 841.

Btnitll, G . M. (1!)34). A C I I ~ ~ L I I ~ ~ ~ I I S red pigrnentod tumor (orythrophoroxna) with tnettwtancs. t r l H flr~tfislt (I'n~.,rtdopleuronectes arnericanus). American Journal of Catlcer. 21, 596 5!)9.

S~lies.r.lco. 8. I<'. ( 1072). Progrcass In fish pathology in tllis century. I n " Di~eaxee of F~nlt " (Ed. Mawtlnslcy-'l'homtts, L. E.) Symposium of the Zoological Rociet!~ of London, No. SO, pp. 1-16. Acadernic Press, London and N I : ~ Yoi-k.

SOME AHI'IC(:TS 0 1 ' NEOl9IAAHlA I N MARINE ANIMALS 229

Spttrkx, A. K., I'tr~~lc~y, C l . I(. I{tbltw, I<. I<. trt~tl Htryco, (:. S. (1964). A t~ic~xor~c.l~ymal turllor t t ~ IL I't~t'ific oyxti\t'. ( f ' r u ~ o ~ t r t ~ r g l y u ~ ) . . / o u ~ I of Irwect I'athvlrq,~/, 6, 448 452.

Sptrlcu, A. K., l't~c~lcy, U. H. tlt~tl C'l~cw, I(. K. (1969). A second mescncl~ymal t~lrnor fi*ort~ 11 I't~c~fii: oyxt(,~' (C'ra.wostr~n gigm). Proceedings of the National ~Shelljislt .4~*0r;rrtiott, 59, 3.5 39.

Squires, TI. 14'. (l!Mi5). N o o p l ~ ~ i ~ . in 11 ~ o ~ ~ t r l ? iScie?tce, New Ymk, 148, 503-505. Stanton, M . k'. (l!l6B). I)iot,l~ylr~itroaurnirtc~-~r~rlr~cnd hctpatic dogeneration and

nooplauta it1 tlro ctclittu'iutn fish, 13rachy~la~io rem'o. Jou rnd of the Ndionul Inutitulr, 34. 1 17 130.

Stolk, A. (1!).50). 'I'~~rnorlt~x i t1 wJ~ttl(-x. Arn&errlavn h'att~rdiet, 1, 28-33. Stolk, A. (1!152). Hot110 ttclno~~rs in whalox. l'rocaerlings, K. Nerlerbndee akodemie

van IVrtsna~ltopl~en. Moc:t9io1r C, 55, 21 6. Xti~lk, A. (1!)53). Sorutt t~itnour~x i t t wllslen 11. IJroceedings, K . Nederlandae

aknrbn,ie vczvr CVstenarhuppen. Sc,ctio~~ (', 56, 369. Stt~lk, A . (I!%!!). 'I'tcinoctrs In tvhalt\n 111. f'roceerlinqs, K. Nederlandae akademie

van IVete/~.vchu/)/)ev. H(.c:tiorl C, 65, 250 268. Stolk, A. ( 195ti). 'I'rrmo~rrw of fishos. IXA and IXB. Epitholioma of the oral

rnucostt, ill tllo sc-ylliid 8cylliorhinue oatulw. (L). Proceeding8, K. Nederland8e akadernie van Retenschappen. Section C, 59, 106-210.

Hurbeck, G. (1!)21). Citod by Thornas, 1936. l'okallnnhi, K . (1!)2!f). Studic, rlbvr Piscl~tiimorcn. Zeik8hrift fur Kreb8fwachung,

29, 1 7:). (All the. turnorll.s roported irt the Japaneue journals which follow arc corlxitlorc~tl in tlettail in t11t1 ttbovc mentioned reference. Bann, 19, 5-8 (192.5) ; Tranwactionx of the Japaneee Pathologird Society, 14, 274 -276 (1824) ; Tranuactiow of the Japaneae l'atathological ~!ociety, 15,294 (1926 ; Trafwdona of the ,lapaneas l'athological Society, 16, 212 (1926). (Study on fish turnours.)

Tavolga, W. N. (10.51). Rpidor.ma1 fin t111nom of tlte gobiid fish, Bathygobiua noporat*)r. Zoolo!/ica, New l'ork, 36, 273-278.

Taylor, It. 1,. ttlrct Sr11it11, A. (J. (1!)66). Polypoid and p~pi l la ry lssiol~s in the foot of the jittpc~ c+ltltn. (Z1rc?x~r& ntcttcrlli). Jvztrnd of Znrccrtebrate t'athology, 8, 264 2(11i.

l 'o t~ tsc l t l t~r~t~ i i (~ t~ , (). ( I !t2O). I4c.itrac~gc~ mrr v c ~ ~ ~ ~ l t ~ i c * l ~ i ~ ~ ~ ( I t ~ n On kologio rnit Rorclvck- sic11 tigr~ t ~g (1t.r Idcvrtit not3~frttgib. deitxrltrift fur Krehfm8hutyj, 17, 2 8 5 407.

'I'lrort~n~, L. ( 1'32(i). Epitllc.liornct o t l o r ~ t o l ~ l ~ t i c ~ r ~ c dew maxillairt*s ctrez une morue. Htcllstirt rle I',4uxoci1ztio)t frart(.aixe yottr I'gtvde du Cancer, 15, 464- 470. ( O c ~ o ~ ~ t o l ) l ~ ~ r t ~ c ~ opitl~i~lionrt~ i r ~ tllo rnaxlllrt of a cod.)

Tllolntw, Id. ( l927tr). Lcx mc~rco~ncm~fibrc+blwt~iquoe chez la rnoruo. Bulletin de l'Aaxorirrtio?t fra?t(.aixe poctr I'lfitrdr du Cancer, 16, 78- 89. (Fibroblastic snl*romrlm in t ltc. cod.)

'l'lromm, L. (L927l~). Stir 11n rnu dc ganglioner~n~rne abdominal ches la moruu. Iic4lletivr. r t ~ I'Axxoc'iation jrunwixe pour l'gtude du Cancer, 16, 282 286. (On tl. c.axa of nl)dorl~itl~l gnrlgliotle~rrorn~ in the cod.)

'~'II~~IIIILH, I r . (l!):IO). Cot~tribl~tion a l'htudc dos lesions pre-cancert!l~rcefi chez len p~i.wo~rs. L ~ H pnpillo~ncs cutallus de la sole. Bullelin de 1'Aeeocialion f r a n p a k

l 'fitudedu~ancer, 19,'31-97. (Contribution to the study of pm-crtnccro~~" - - lesioi~s in fix!^. (:c~tctt~oo~~x papillomas i r k the sole.)

'I'llomnn, I.. (1!)31u). Lou tctrnonrs d e ~ poissons (6tude arratomique et pathoghi- ~ I I U ) . ljtdletin rle L'Aaaocktion fran.pi8e pour l'#tude du Cancer, 20, 703-700.

l'l~c~nltn, L. (193Pn). Ttlrltbdomyt~me chez iln flet. Bulletin de l'd8aociation fmn- cake pour l'gttttule flu Cancer, 21, 225-233. (Rhabdomyoma in a flounder.)

230 L,TONEI. E. MAWDESLEY-THOMAS

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