Fish Pathology

THIRD EI]ITI()N

Erlircl h1'

Ronald J. Roberts
B V M S , P h l ) ( ( i l . r s g o $ 1 .I ' R C V S . F l { C l ' l t h , F l B r o l F l t ' S E ( l o m r r r . u r J c r o f t h c M o i t N o L ' l c ( ) r ' c l c ro f r h c C r o N r r ( T h ' r i l : r L t r l ) , L I n ' : c I t i f l tL ) i n l . , J t t i l t u lI l i i i t ( P r , t i \ ! 1 L r ' r i r ^ i r ) ' r ' / 1 ' l ' r / r oL : ' S ' . 1 l'tr lrt rin DintrLttt Ltndrarti Lr d St'tr tt 7t, Irr.rirr ., l ] n t \ n ' t l ' r r i r t r : ' r r 1, ' / S r r r / r r r yS r , ' l , r r r r l

\ ( / .B . S A U N DE R S
F d j n l . L r r s h t o l r t l 0 r Il \ r . r ' \ o r l i . . ()rr,IJ. 2 t ' r i l . r J , l t l r . r. \ 1 I o L r i \. S r r l r r e v ' T r , n , n r o { 1 0 1

2
The Anatotny and Physiology ofTeleosts

INTRODUCTION
Thc purpose of rhc prcscnt chaptcr is lo give a bricf out linc of the aspcctsof the anatony and physiology of teleost iish which are ncccssaryfor xn apprccialion of the pathological ch:nges which can occur in lish and lor thc uncler standing of the mechanisms underlying clinical discasc nlanifcsrations. Many basic functions ;rtc sinrilar to drose of othcr verte and knowledge of mamnals is nol ntirely ilrelcvinl brares, However. thc tclcosrsmust not be regarded:rspriDritiveforebcars of rhe mamnals; they are advanced,evolurionarily reccnt :tnd expanding into a frultitude of nichcs.There are morc spcciesof telcost than any othcr ciassof vcrtebrate are (Bone st d/. 1995),so that many generalizations obviouslv of dubious value.but thc prirrcipal exploited specicsrle of a restriccedrangc so that useful strtementscan still be madc. The fnain cmphasis in this account rvill be on spccific from the more familiat mammalian anatornyrnd differences physioJogy. A major considcrationis the rquatic envimnment and thc the constnints it imposeson fish.Ir is against physical:rndbiological degradativcinfluencesof this nedium that the rriliel interierrof the fish must ]:c mainrainedand rvith which nec of essary cxch:rnges materialsmust lakc place.An overriding factor is the high specific hcat of warer, which imposcs,on i.e. most fish,ectothcnny (poikilothcrn-ry), thc body rempcreNo simple turc conforrns to thc enviromnentalternperarute. physiologicalconstantvaluescan be given, e.g.lor hc:rt rate. rate of digcstion or rate of growrh; all of theseare subjcct to tcnlpclature and dris must alwaysbe borne in mind whcn lt The san, animal is often unrecogllizable e studying teleosrs. different tcnperatures, c.g. rvhether adrcnalineincrcesesor hcart rale rn rainbow trout depcndson the tcmper decreases aturc (Randall 1970).

THE INTEGUMENTARY

SYSTEM

The skin is chc primary barrier against rhe environmcnt, allowing norrnal internal physiologicalfunction. so irs condiproccsscs. The layers of tion is iDrport,rnt in nrany disease tclcosr skin, conprising cuticle, cpidcrmis, basementmembrane,dermis and hypodcrmis.are indicateddiagrammarically i n F i g .2 . 1 .

Cuticle
w.rsfirst described the Thc cxrernallayer, cuticlc or glycocalyx. in tlerail by Whitcar (197i]) as a mainly rmrcopolysaccharide laycr approximarelyI pm thick. k is normally formcd largely

Fig.2.1.

Schemaricdiasram ofnornul teleostskin layers (Frc

Bu ock & Roberts 1975.)

Ihe Anatony Physiology feleosts and of

a ,J

Fie.2.2.

Tail skin of\\,hiting, showing cuticle.epiderms rnd dernis. PAS x 142.

v -.v '_

rtom epithelial surface cells r.thcr

than by secrction fron of cell protoplasrn,

--,{4sq.

soblet rrucous cclls and is a conplex

doushed cells and any goblet cell mucus that has been secreted onto the sulface (Fig. 2.2). The physical consistency of thc cutlcle varics considerlbly benveen spccics, bcing especially developed in rock pool rlrd benrhic specic\.The cuticul:r hvcr contains specific irununoglobulins and l,vsozyrne(elthough rhe bctween species). and arnount of the latter varies very nuch

free fatq' acids.Thcsc arc believed to have anti parhogcn activ ity, as pafl ofde rn conjruction nucosal dcfincc system ofthe skin, working with cellular proliferation kinctics k) continu ftom the surface (Spearc &

ously remove nlicrcorqlnisms

Mirasalimi 1992)- NotmJl1,, however, small lumbers of bac tcda will still occur on such sulfaccs and there rrc obvious limits u'ith regard to thc elliciency of slrch rystems whcD pathoecn loading ofthe enviromr, cnt is high.

Epidermis
As in all vertcbratespecies, lindanrenral unit ofthe epi thc derrnjsofthe teleostfish is the fibror.rs malpighiance11.This is, however.thc only consistent feature,asthcrc is grea! diversiry in all of che other ccll types occuiring there (Bullock & Robcfts 1975)-ln adultsthe epiderr,ris a stradfiecl is squanous epithelium covering thc body sufacc and investing rhc rail and fins. Unlikc its mammrlian counrerpartjit is living and capable mitotic division ar all levcls, of cven at lhe outci-most. squamous layer.Thcsurface ofthc outermostlaycr is arrangecl in a rvhorling patteln of micro ridgcs (Fig-2.3). The rhickncss ofrhe epidernris variesrvith spccies. age,site and oftcn, staeeofthe reproductivccycle.lt is usuallvthicker in rhosespccicswith negligible scrle covcr (e.g.rhe eel) and

Fig.2.3. A.Autorrdiog, )Fh ofsccrion olphice skn fren I young llsh inocubted rvith trrtirred thynndDc 12 houn prevrousb.The specicrl]' labeled nuclci ofdrvidnrs cells (arro$,cd)rre round ar a[ ievelsofthe epidcrDrs.H + E x500. B. Sc.rnmngeiectron nicrogrxph ofthe surfaceofthe epiderns ofguppy shorvinqthe chancteristic ar ngcnrnt ofmicro Dr D.K. cone.) dgcs.x2200. (ts,by couftcsy of

also over the fins, rvhere it A prrticr arly well endowed rvirh ncrue end organs and lrucous cells (Fig. 2.4). The uralpighian ce1ls are always prcscnt iD teleosr epidermis. They arc rounded cclls vcn' similar in structutc at rli levels exccpt rhc outermost. Hcre thev arc flaftened hodzonrally, nith a cytoplasn composed largely ot an lccumulation

PathalogY Fish ofputetively structures' and macrophages large clear q'st-like are especially prominent in the .at.,l". -lgir-t, which Gadidac.

Dermis
The upPer iayer'the of The dermis is con-rposed two iayers' ^loose network ofcollagen and reticulin stfihtnl spo11giosutll,is membmne li basement fib..., "orrtigoo.r. *'th rhe epidermal mast cclls and .ot,"io. ,nJ pigment cells (chromatophores)' Thc lower layer'the the scales cellsofthc scalebeds and also dense matrir which is stfttltm .otttpactum' the coliagenous strength of the skin The capaciryfor fr.tto* ,n" structural or due to sexual .olo,t. .h".g. to match the cnvirotunent ir, many tcleosts ".,i.,i y o, dir."r. is very highiy dcveloped of the intetplay of ,rri ir'ina.t."a Uy controlled modulation of.the chromatophores' absorptiveand reflective propctties cells'are asteroid the Melanophores, dark,pigmen!-containing bound electroncellscontaining largenum6s$ of nembrxne which c'rn be movcd clensegranules of melanin pigtrent cffect' cytoplasm of lhc cell to give the desircd *t*"',n" soiventorgnic Lipophores rr. .hto-"tophott' containing conarc subdividedinto erythrophores' solublepigmentsrnd containing ye)low A oftne gurnard Bod\tutl"cernrr'ou( Fie.2.4. the eptderrnr t^ining red pigrnents,and xanthopborcs' tn ro '.,0 '. ' a-o L1+r"p^'s 'rrrrrar rho'efound , -i. ,.,. n"0 pigrncna (Bultock & Robcrts 1974)' fin B.ThePectorrl PASx224' which cannot be sPecies gadoid The pigmena are mainly catotcnoids come of necessiry must de synthesized noro by the fish and for are responsible liom food Leucophoresand iridophores degeneratingmitochondria and somc vesicles, gu2rmnc' of elongated purines'usually white and silvet colours and contain the more qpical widely a.nr. U"".rai., of {ibres' instead of ofreflecting matcrial uP io 10 pm thick which exist asplates mitochondria around a distributed bundles of frbres and arrays'rather like a and arrangedwithin the cell in parallel generallY ovoid nucleus (Fig 2.5) ' venetianblind (Denton & Nicol 1966) the ePidermis of all Mu.ur-r".r.,ing cells are found in flexible plateswhich lic of The scales telcostsare calcified These species posteriorly' but numbersvary great\ with site rnd teieosts within shallow \cale PocLets" orientcd partly '.Iwo layersof the epi..11, usually originale in the middle are describedgobi.t n-taintypcs,differing in surfacesculpturc' a mucous cell may in postcrlor on iermis, altho,.rgh a very thin epidcrmis' bear stiff spicularprccesscs lbeir Crenoid scales They its base on the basementmembrane Both rypes be seen ro have from cycloid scales externaledgewhich arc absent (mxinly glycoproterns) secretions elaborete in increase sizeand which' in many sPccres' have growth rings on their suriace' the asthey ePPrcach surface ege Ultmsffucturally' allowietetmination of the individual's found in the lower with a nratrix of Clrb ..11 ".. l".gt, tttually round' cells' consistof collagen{ibresi$tcnPe$ed scales teleostgroups' midclle layen of the epidermis of certain and depositcdhydro>'ryapatitc albuminoid matcrials in which are in rhc club cells ere ttre Shrc&stofzellenfound The classical crystals. which sccrete a potent alarm subepidermis of cyprinicLs, similar morphologically stanc.,b.rt many othtr sp'cics posscss cellsin their epidermis'which clo not Hypodetmis Iargeclearnon-mucoid appearto be relatedto such ftight teacuons' tissue'whicb is more The hypoderrnis is a looser' adiposc of teleost epic."rrol. ."11, are founcl in a widc variety ofthc dcrmis than tbe overlylng stratum compactun vascular ascribed to them' but as yet no function has been dermes ofinfectious prccesses antl a ftequent site ofdevelopment lymphocytes' cells found in the epidcrmis include Other

Ihe Anatomy Physiology leleosts and af

15 rhe body lateral1y genemteproprlsive forcesby oscillarion !o (Gray 1968,Vidclcr 1993). ofbody and til

Axial skeleton
Thc layoucof thc skeleroncanbe seenin Figs2.6,2.7 ar'd2.8. The sku1lconsists ofa rigid cranium !o which are articulated the bones of the jrws and bnnchial and opercuhr rppaiatus. The structure is very complex and much of lhe skull moves dtring feeding;rnd bteathir4lrrovements. rhe components All arc intcrdcpendentand the structureofthe skull is bestundersroodin the contexr ofa descripiion ofbrcarhing movcmcn* such asthat ofBallintijn and Hughes (1965). lhe nrrnrber verre\merr nor.on\rrnr rr J rrven \pe(re\ ol and is allected by enviroDDrental condilions during larval developmentEach vertebralcentrum is a sirnplecylinder,the 'cross' secn on radiogmphs rcflccting thc conical rcccsscs
Fig. 2.5. Elccnon micrograpli ofthc dermis offie in hyersit1l = nelanin straturn spongiosum ofthc f= collagen Iibres platclcts.

cnclosingthc intcrvcrtcbral p:d.Thc cdgcsofadjaccnt ccntra are connected by ligaments And thc whole colulnn is hcld togcthcr by longitudinal elasticligameDtswhich run dorsal and ventral to the vertebrae. the verrebraehave a neural Al1 arch rnd a ncuLal spine, the caudal vertcbrae also having a vcntral haenal arch and haemalspine.In the thoracic region, insread of the haena) arch, there are pleural libs which support the lateral rvallsof the body caviry lD rrlany species. intcrmuscularboncs of various affangcncnls irlsondiatc out fiom the vertebral column in the septa between thc

plaice. bm = baseneDr tnedbmnei g[llules.AuoNs

indicate iridophorc

EM x2800.

THE MUSCULO SKELETAL SYSTEM

The ftisiforrn shapeof the typical fish is deternned by rhe rcquircmcnts of swilntning. Thc strcaniined extcrior minimizcs drxg and the main muscle blocks (myorneres)are arrangedon either side ofthe arial skeletonin order to bend

Dorsat I ?istal fin rays L J Proximal

I Centrum spine Vertebra / Neural J spine [/ / Haemal

Buccal cavity

Cranium

Swim-bladder

Dorsal

Hypurals
Mandible Premaxilla Pelvicfin and girdle Pectoralfin Ventralfins

/l

Branchiostegal rays
Fig. 2.6. Radiograph ofa rypical round fish,the saithe.

PathologY Fish

Operculum
Lateral line

Anleflor kdney Adipoee fln

Caudal peduncl

Mandible Sr,vim-bladdel Pyloric caec:r lntestine $omacn Spleen

6sh' Fig. 2.7. Diagnm ofthe basic anatomvofa "lmonid

duct lymph lnterspinal

Neural lymphaticduct

sinus lymPh longitudinal Dorsal

Mandible

cavity Buccal rays Branchiostegal fins Pectoral

Pelvicfins Kidney Cardinalvein

Cannula Hypurals lymphduct Interspinal

Bodycavity

'Ventrallin

sinus lymPh longitudinal Ventral

Fig'2.S.Radiographofatypicalflatfish,theplaice.Theinjectionofaradiopaquesubstanceviaacrnnu]ahasdelineated*renerrrrllymphanc duct and lhe rcnal portzl vein. (Bv courtesvofDr C R'Wardle )

Ihe Anatomy Physiolagy and of Teleosts Fins

The pelvic girdle rn iorvcr tcleosts (e.9. salmonids) n embed dcd nr thc ventral boc\ nluscuhlurc. Irr rrrorc adv:rnccd typcs it is in a more anlerior posirioll rcsting lgAirlst thc pector:rl girdle- The pectoml girdlc rs suspcndcd inmcdiatcly thc opercular reqion ofthe skull. Thc mcdian dorsal and venllal 6ns ate articulated to thc pterygiopholal rnusclcs which contnrue the line ofrhe neurel bchrnd

and haemal spines.Thc caudal lin is rrticularcd on a series of flat plarcs, thc dorsal cpural and vcnrral hypulal bones. The fir'r r:rys can bc oi two types: spiny or soft. The struc ture ofthc firr rrys ofthc tclcosrs rvas used at one time to sep ar:r!e thcn inlo t\.vo major sroups: lhe Malacopterygii rs r)ughly (rvhich

cquivalcnt to the lsospondylii of ptesel! systcnur(roughly

ics) or soft-rayeC species, and the Acanthoptcigyii

equivalent to thc prcscDt day Pcrcifor rnc$. The spiny rrys are sirnplc sinqle bones as in the firsr dors3l fin ofPercilbrrnes.The caLrdalfin rayc in all teleosrs lc ofthc soft khd, as aLc also all Fig. 2.9. Secror dnollsh r cranirl crrtilagc ofr young L nborv nout at dre edee of I centre of ossificatioD. Thc darkcr arcars the calcifrcdcoDpoDent.H + E x-100

thc other fins tu lsospondl.lii (c.g. Salmonidae, Clupeidae). The sofr rays arc scgrrcntcd. oftcn branched and formed of two idenrical latcral components either side of lhe nidllncThe fir web of wild lish n cleaL and veLy 6ne but in lanned 6shcs it is often thicker.

of eels (lngrillifbrm

movement) in which thc rvwc is gcner'-

rtcd by sequentiai contraction [.onr hcad to tai] ofthc lmscle blocks or n1'omeres. In shorrcr-bodicd, morc wpical. frsh. the

Bone
Thc mcloscopic structural elements of frsh bones :rre slnilar to rhosc of othcr vertebrares and eenenliy two typcs ofbonc atc found, cellular and acellular. The former contarns osteocvter and is coDfincd to lorver orders. e.g. Clupeidac. bone is

rlechanism is thc s:rnc, but during swirnming the flexurc of the bodv shows less than a cornplete lvavc and only thc oscil lation of thc tail is rcally apparert (camrgifurm locomotion). Somc fish srvin by scullrrg or waving nTotions ofccrtain frns,

in u'hich casc rhc appropriate muscles are highlv clcvclopcd lld thc nain nyonrcrcs rnay be considcrably rcduced (e-g. se.r

Salmonidae and Cyprinidac

(Fig. 2.9). Accllular

uniquc in verlebutes: it contains no osreocvtes and is found in advanced teleosrs such as Pcrcidrc and Ccnrrarchidae, ofren having a solid Gatureles rnatrix (Moss 1965). Thc lack of ccl1shas bccn shorvn !o preclude resorption of calciulD from thc bones so thar acellular bones canno! function rs a calcium rcservc. I{cpair ofiacurcs under ;rcrlcaennc con Thc most obvious fe:rture of the rnuscle of a roLrnd lish is the folding and inrerlocking of thc rnvoncrcs (Fig. 2.1tt)

Externalll'. the body musclcs occupy the quaclanls of the body. separatcd from cach other by che median septurn rrrcl thc fiansverse horizontal scptum. The two blocks of rltsc]cs donal to thc horizontel septurr erc callcd qrarial nr-,r/tr, wlile thosc vcrtrrl arc ca11edlypariai rnrsrirs. Supert'icially, borh

diuons in advanced releostsis thercfore lindered (Moss 1965). 'marrcw' Despite the presencc ofvascular canals and spxces i n ' o r r c b o r e s o f b o r h n r . r r nt r p c ' . r o h r , n . p o i e r , r*ue '.

cxparial and hypaxial myomcrcs arc foldcd in the verrical planc. Holvever, dissection ofthc nyoncrcs shows that they

present in such spaccs. k is cvidcnr that in releosts there ate ma.jor clepartur'csliorn other verrebntes in bone structrirc irnd physiology.

are secondarily loldcd in dre horizontal plane so th:]r below erch flexure in the cpanal and hypaxial myolncrcs thc rnusclc crtcnds backwalds xnd towards the nlcdiaD scptum as posrcr

Muscles
Most fish $e'jm by passrnga rvavc ofincreasing anplitlrde pos teriorly along thc bod;,. This is most evident in lhe rnovernenl

ior cones. Ar thc holizoltal

septum the rnuscle prqects for-

*'.rds :urd inwards as a single anrcrior conc.Thus contr:rction wilhin an arnnsclrlcnt ofcpaxial and hypaxial musclcs causcs

the body io bcnd (Fis.2.l l).

I8

Fish Patholagy The folding and interlocking of rnyor,reresproduces smoorh sequentialcontraction along the body,since the contnction of an individual rnyomere also influences several overlying and adjacentnyomeres.The mucle fibrcs in thc -un in rn ;nreropo.tcrior .up.rbci,l l.rycr'ot fhc InyomcrF direction, patallcl to the medirn seplum. Contraction thcrc fore occurs in the plane of thc a-tisof the fish. However, the deeper down within the myomcrcs the fibrcs are, the morc thcy arc anglcd to the axb of rhe fish. Thus the extcnt of shorlening in this plane will be reduced thc closcr the fibres for lie ro rhe median septum.This clear\ compensates the

@ x) 1'

--.' .:l:=----,/ a -

changes absolute in velocitieswith distanccftom the centreof the bend end preventsfolding ofthe skin. Histologicaland biochemicalexanination ofthc myomeres in h:rs revealcd rangcoffibrc typeswhich areorganized marry a as species distind zones-tn most tclcoststhere :re rlvo main consisting; thc of subdivisions: ttlusoia^ latuttlis thc supctjcralls, profuttdus, so-called red nruscle fibres, and thc nrlrdrldlir lateralis which consists whice 6brcs (Frg. 2.12). From mcchanicel, of electrophysiological and biochcmicrl diffcrencesit has been fibres, shown thar the red fibresare aerobic,slorv-conrracing, similar ro rheir counrerpartsin manuulian musclc,and that the whire are anaerobic,fasi contGcrine and fast-fatiguing fibres (ayne & Lauder 1993). ln the Salnonidae and lo an cxtcnt in Cyprinidee,sand$''iched befiveenrhe red and whitc are found the pinlk fibresrvhich appcarto be intermediatein Also,in salmonids. tunction betweenthe red and white fibres. the bulk ofthc myomcrc is rnadcup ofa mosaicof white and pink 6bresrathcr than whitc frbresonly @ilinski 1974). In gadoidsthe red fibresare about 50 prrr in dianetel and the white about 100 pm, brt this varics considerablywith dcvelopment, exercise and starvation (Greet-Walkcr 1970, have shown that thc red 1971). Elecrromyographicsuudies and thc white is muscle is active during low cruising speeds recruitcd as mosaic dudng strenuousswimming, particularly in sprintsor burstsofectiviry There are also diffcrcnccsin vascularization berween the fibre types, rhe red muscle being generouslysupplied with blood and providing a good sirc for rhc injection of drugs,

D
Fig.2.10. Pattemsoffolding ofmyomeres ofa typical tcleost. A. Lateralview with detailsofsuper6cial loldins of epaxlal(1) and (2) hypa-xial muscles. c, Secondary B. fblding ofthe myomeresftom o, trunk rnd caudalpeduncle rcspectivcly. Planevielv ofsection to ofthe through epa-xial muscles shon dre arciforn appearance myomercsabour the medlan septum.(Redmwn fron Greer-Walker 1e70.)

Fig. 2.11. Action offorces appliedby a myomere to bend the the body. The line AB reprcsents median septum and axial skeleton. the anterior and posterior The paraleloeramsdb',/lnd .; !,r represent conesof a singlemyomere in the rcla-xed condition.The broken lines indicate the posinoa and shapes ofthese componenrsafter conffaction ofthe nryonere. The diagonallines ivithin the anterior and posterior co4esindicate the direction ofthe resuhantforces acting wrthin during inirial conraction and tend to bend the median septum and a"xial skeletonthrough the arcsQ. (Alier Nur$[

anticoagulantsand anaesthetics, sincc it only lakes a few seconds material to reach the central nervoussyslcm and for other parB of the body. The innervation ofthe typesofmyofrbrils is alsoquite difirt fcrcnr. Lhcrcd m.r'clethe nerveendrng:occurrrrgrnpp, ln thc middlc of thc nusclc fibres,while in white musclc the nerve endingsarc enplatte and are terninal where the while however, fibres arie from the nyoscpta. [n higher teleosts, white muscle has n-rultiple,et1gnppe inerv^tion The pink

19s6.)

Ihe Anatomy Physiology Teleosts and of

19

- 130/"

making its structurea ma_jor considcrationin the homeostasis of the milieu interieur the fish. The epirhelium is thin to of allow gas exchange and rhis aL\o renders it particularly vul nerablc co invasionby parhogens. well as having a respir As atory function, the gills are rcsponsiblefor regulating rhc cxchangeofsalt and water and play a major role in rhe excre tion of niirogenous waste products. Even slight structural

0.93 body lengthvs 2.01bodylengths/s

o;^ oo \-o /

dar,rage thus rcndcr a fish vcry vulnerableto osmoregulacan tory as weu as respiratory dificulties (Hughes & Morgan 1973).

. red libres o while fibres

The structure of the gills

Thc gills of a typical teieostcomprise two sctsoffour holo branchs,forDring rhe sidesof the pharynx (Fig. 2.13). Each holobranch consists rwo hemi branchsprojecting ftom che of posterior edgeofthe bmnchial arch or gill arch in such a way that the free edgesdiverge and touch those of rhe adjacent holobranchs.Close cxamination of the hemi-branchs of a ftesh gill shows that they consistof a row of long thin fila-

starved
Fig. 2.12. Transverse sections ofsaithe rt point 0.34 ofthe body length frol1l the tail.The open circl.s rcpresenr L\e larger diamerer, fast-contracting white nluscleflbres, while the closedcirc]esrepresent the do\ red 6bres.Th. percentaee changein diameterofred and white muscle frbrcs follorving exercise at two d:fferent swinning speeds and stanation,comparcdwith the conrol, are indicated lvirhin eachDluscleDa$.Thc percentage decrease bodyweighr in (indicatedaboveeach scction)r represented a decrease total as in cros-secrional area.(Redrawn fioD Creer w.lker 1971.)

menls,the primary lamel1ae, which project ftom thc arch like the teeth ofa comb.The suface areaofeach primary lamella is increased futher by the formation ofregular semilunarfolds across dorsal and ventral surface-the secoDdary its lamellae. The dorsal and ven[ral rcws of seconderylamcllae on each primary are sraggered that they complemenr the spaces so in the rows of lamellae of idjacent fihments (Fig. 2.14). This arrlngement of archesand larnellaefbrms thc sides of the pharynx into two sctsofcorrugated sieves thrcugh which the

frbres have an innervation

intermediate betwccn these two

extremes.Although thc transmittcr in all cascsis acetylcholine, therc appear !o be differcnces in the rcccptor sitcs, as shown by the varied responses to drugs like dcxamethonium, which blocks red receptors prefcrcntially. The motor neurones of white

units tend to be of large diameter and myelinated,

whilc thc neurones of red muscles are slightly or non myelinated and thus seem thrnnerThe form ofaction potentials in

thcse neurones also differs in time coune rnd enplitude.This disdnction between red, pink and whitc 6bres is also found in the respiratory and frn nusclcs according to the nature oftheir mechanical function.

THE RESPIRATORY

SYSTEM
Fig. 2.13. The opercularcrviry ofa rainbow tlout showins the gll arches, with rakers, and prinury lamellae.

The areaof epithelium ofthe gills is comparableto the tolal area of *re skin and in many speciesis considerably larger,

20

Fish Pathalogy which is downstre:n opercularedgeofthe primary lamcllae. of the water flow. Blood enter the blood spaccsof the secondarylamellae by short afferent lamellar arteries.This blood flows in the opposite dirc.tion to water deoxygenated The resulting countercurrent pumpcd through the gill sicve. ofthe orygen in watcr being trans leadsto 60-80% exchange blood lcavesthc secondary fbrred to the blood. Oxrygenated larnellaeby efferentlamellar artcries to fecd the dorsal aorta by way of efferent frlament and efferent branchial arterics Wwithin ihe primary lamellae orllgenated blood is shunted through a nutritive circuit to the activetissueofthe gills. arch is coveredby typical teleostePidcrmalthsue The gi11 but at the origin of thc primary lamellac the epidcrnis is much rhicker and usually extremcly rvell endowed wilh Bclow this epidcrmisthere is usuallyan xrray of mucous cells. lymphoid tissuc, comprising lymphocytes xnd, rn Dlrny specics,large cells containing eosinophilic grantlcs These lactcr rre also frequenrly found along the lcngth of thc primary lamellaand may occul in largemrnbers at its tip Thc primary lamellais covercdby a mucoid epidermiswhich rrray lhe havc within it, in euryhalinespecies, pale-stainingsaline, cells and, beneath lhcsc. lymphocylcs' and or salt secrecing, eosinophilic gtanule cclts, and phagocytic cells, varying in nunber between specics.

by Fig. 2.14. The interdigrtanngproEle prcsentd the arravsof adjacentsccondarylamelae to the direction ofwater flow (After Hughes 1961.)

Thegill arch
structurefrom which The teleosrgill arch is a cutved osseous ndiate the bony supports(thc gill rayt ofthe primary lamellae.The angle ofthese lamcllar rayscan be altcrcdby a set of the adductor musclesto ad-just :mount of ventilation of thc arc larncllae(Fig.2.15).Alsocontainedwithin thcsearches lhe afferent branchial atteries fmm the ventral aorra and the efferent branchial rrterics scrving the dorsal aorta.The ventral up aorta divides into numerousfine branchesasit Passes the holobranchs.The aferent flament arteries run along the

The secondaty lamellae Gaseous cxchanse takes placc across the surface of thc thcsc consistof an sccondarylamellac (Fig. 2.16). Esscntially envelopeof epithelialcells,usuallyone layer thick, supported by and separated pillar cells,which are arlanged in rows 910 Fm apart.Where the pillar cellsilnpinge on thc basenent membrane of rhe epithelial envelope thcy spread to form

adduc'ior secondary lamellae

branchial arcn branchial anerles

waler tlow

with thoseofneighbouring pillar cclls which coalesce flanges which to complete rhe lining of the lamellarblood channels connect the allerent and efferent lamellar arteries (Hughes 197s). The pillar cells havc been shown !o contain colunlns of also contractileprotein similar to that found in amoebae.Thcy conncctive tissuewithin lhcir invest columns of extracellultr cell memb$ne. Since the blood entering the lamellar blood comesdirectly from lhe ventrxl aorta at high pressure, spaces the prcsenceof fibrous and contmctile clcments in the supwill serveto rcsisttheir distcnsionunder porrs ofthese spaces (Frg. 2.17) (ci normal circumstances lamcllar relangiectasis) thar piilar cells are used to it has bccn suggested Although control lamellar perfusion,there is, es yet, no experimental evidence to support this hyporhesis.[t has bcen shown,

oporcurar cavity
\water flow

Fig.2.15. Diagramulic rePrcsenutionof segnentsoftwo showing the flow patternsofwater and blood adjacentgill arches, In the upper segmemthe conEgura$onofthe branchialarch' the gil ray and the adductormusclefor a singleprimary lamella* indicated.(Afler Hughes 1961 )

The Analony PhysiologyTeieosts and of

2I

p
t?

Normalgill secondary lamella

f$'

f f6

Epithelialcell

TELANGIECTASIS secondarylamellaof gill of

& r'
ga
f

"8

Fis.2.16.

pillarcells Ruptured
Sectlon th,ough dre gill lrmell.re of ,r r,,1 bo\ trout Fig. 2.17. Diagrammati. repres"-Dtation sectioDtliLougli of se.o,)drr,v h.ie r of sil sholing locrtion ofpilhr cclls rnd ctTcctof ' r 1 1. . 1 1 p t r . ' - l ' . r 1 , , " U , t i ' r ' a , ' t ' . . r

rlLo\.]]rgthe lrinury h,uelh (P) sid us rfrrys ot delicatese.o.dJy L,rnlellle(rr1o\!ed).H + I x100.

hog,'cver. that ftom srmd:rrd to lrraximurn aerobic rnctabolic r,rtc thc tunction.rl arca rncrcascs bY a l.rctor of 6, marnly by rccruitncnt lnto irctivity of lamclhc from thc b:rsc of thc blootl florv conrrol has

2. Rcduction 3. Ilcduction though

in c)<posulc of body fluids to ionic cxchangc.

of rhe slze of the presure los ar 2,rragc thc gills, posibly by shunting blood dircctly hom

prnnarv hmellac distally Regulation ofbranchial is not iu11yundcrstood. lm'olvement ofauronomic

allerent to efferent lanre11ararteries, rhus enhancins the eficiency of the svsrenric circularion. The surface ofthe l.rrnellar epithe)iurl rs irreeular. althorqh not. as rvith the epiderrnal slrface, throrvn into such dirtinctive nricrovilli- These irregularities serve to aicl attachmen! of the cuticular nllrcus, which, jn addition to its role in reLlucin!! infection and rbrasion, has r siglificant role irr rcgulaling the cxch:rrgc of g:Ls, w:rtcr rnd ions. The cornbrned thickDcss of cuticlc, rcspiratory epithelium and flangcs of thc pillar cc1ls ranges from 0.5 to '1 pm, and rcprcscirts thc total diffusion diamtcr dist;rncc for of thc rcspir)tor,\7 blood

been shorvn thraugh the xction of neuro-transmitters and !'r)docrines on branchlal resistirnce to blood florv Adl.enaline rnd rrolldreurline both reduce gill resist)Dce, :rcring through ;rcts ;rDt;rgoiistic;llly ro incrcasc

p rcccptors. Acctylcholinc

bloocl pr'cssur'eby rvry of muscarinic rcccptors- The sites of thcsc rcccptors hn'c not bccn idcntificd but thc alhrcnt and t l e r e r r tl . , r l e l l r r . u r . r i . , l r d r h e - n e L r o r c r o u s c o n n . . r i o n ' L o rhc brauchial nuffitivc circuit havc bccn suggcsrcd as likclv

locations (Perry ct al. 1992). Restriction rrain eflects: 1. Reduction in rate ofgaseous exchange. ofblood florv through the lamellae has three

cxchange, sincc thc

lamcllar

chrnDcl is virruelly thc srll1c es thc diarlctcr erythrocvle.

of thc tclc'ost

22 Ventilation and gas exchange

Fish Patholagy thc gill arches reversed. is Coughing ftequencyhasbccn found to b.^ related to thc lcvcl of irrirant pollution. There are and chemorcceptors the gi11s, in arrbient conditionsoflow on Po, and high Pcor, gill venrilation and hearl tate incrcasc. of \\.ith increases Ventilation shows the greaier resPonsc, thaD exposcdto severe hypoxia (less morc than ten fold in fish An important receptor site in this 20% air satumtioll levels). a contexl is the pseudobmnch, rudimentary gill locatedunder by It rhe operculum,dorsalto the main gill archcs. rs perfused oxygen:ted blood from thc t$t gill arch and probably moni pressure, to ton arterial Po2 but it is alsosensitive hydrostatic pH Na* ions, osmotic prcssure, and Pcoz.Thc pseudobmnch (IXth is innervated by a branch of the glossopharyngeal craniel) ner-veand has other non-sensoryfunctions. such as hypcroxygenation ofthe choroid ofthc cye. Centrally,the breathingr\thm is coordinatedby neurcnes Motor innervation to dlc in dispersed thc ncdulla oblongaca. "***' runs in thc Vth,vllth. and rXrh cranial :.;':j:::"

During breathing, watcr is passedin dtough the moudr, over rhe gil1sand out thrrcugh rhe opcrcula.The venrilatory flow is and contractionofthe buccaland dtiven by altcrnatccxpansion in such a way tha! a continuous operculat chal,bers,acting rl'ater florv is maintained over the gill-s.In comparison with airbrcathing animals, the encrgy cost of ventilation rs very high, especiallywhen thc orygen contenr of rhe water is low, in warm in in or polluted col dirions.Thisis bcst manifested aquaculture thc respimtory distrcsssyndrcme,which ariscswhere the energy the required for gil1 ventilation exceeals encryy rcleasedby lhe exlracled o,Jgen. Carbon dioxide is highly water soluble so drat there is little dificulty in its rclcaseftom t'\e gills. the pump is stoppedassoon as ln Dranyteleosts, respiratory to a suficient speedhasbecn achieved allow vcntilation ofthe gils simply by opening the mouth and lc$ing the current flow ovcr thcm- Known as ram jet vcntiiation, this systcmcnables considerablc energysavingand indecd fishessuch asthe larger tunascan only respireby this method. For the control of ventilation, proprioreceptors and mechanoreccptofi,which rcspond to changesin gil watcr flow, are prcsent.For example,if thc branchial water flow is artificially arrested,a reflcx cardiac inhibition occun. Also, mechanicaland chenical stimulation of the gils can trigger thc cough ieflex, by meansof which the wacerflow through

THE CIRCULATORY

SYSTEM

The gcncrallayout ofthe circulatorysystcmofa typical teleost is shown in Fig.2.18.A usefuldetailedaccountofcircr ation in 6shes given by Satchcll(1971)and Farrell&Jones (1992). are

common cardinal vatn branchial efferent arteri6s

supelror meenteric anery coeliac anery

renal

anery
gonadal arieries

branchial albrent arteries wntral aonl

ventricle hepatic portal sy$em intestinal venous sy$em

Fig 2 7

renal portal sy$em

caudal veln

ofthe circul:tion ofa typical teleostfish. ComPatelith Fig. 2.18. Schematicrepresentahon

The Anatamy Physialogy and of Ieleosfs The heart

Thc hearr in tclcosrs is situxtccl insidc the pericar.diuDr of its o:,1'gen sr.rpply ftom thc 'vcnous'blood

23
in thc lumen.

Fjshes with hiqh ircdvity levels. such rs thc rlrnrs. 1]]xy hivc coronary supply to the spongy h,\'cr rs *'ell (Tot:r 1989). lndividual cardiac muscle Ebrcs arc apprcxirutcly diameter, :rbout half rhar of namrnalian 6 p"rr in

.rnlerior to the nrain bocly cavity ;rnd usuaily venrral ro thc pharynr. lt has lour chambers rhrough which btood florvs in sitttplc succession (Fig. 2. 19). l)eo\l,genercd venor.x blood

rl-rusclc. ln other

cntcrs the sinus vcnosus from thc ductus cuvied and mein veins.Therc arc no inler vah'cs and rhe sinus is so snall rhat it can hardly be rccognized as a discrctc cardiac chambcr. The rvall is thin, colnposcd mainly of coll.rgenous connective tissue, althoush in sone species it is nluscular ancl conrracrile. k is in rhe rvJl ofthe sinus vcnosus that the pacetnekcr. u.hich initixtes thc cardiac contacrion, is locatect. Through two sino arrial mlves the blood passes inro rhc

rcspects lhe fibres :rre similu ro mammalian oncs. u'irh inter calated discs berq'een iDdividual cclls. Frcm dre ventriclc the bloocl is passed ir,ro thc bulbus iutcriosus through x pNir of vatves.Thc bulbus h.rs a rhick rvall corrposcd ofa nrixture of clirstic tissue and smooth rnuscic (Pricde 1976). It has a colnpler stmcturc but acts basically as :l pllssive elastic rcscrvoir which snoothes the pressruc pulsc lion the ventriclc and

rDrintains blood flow during venrricular cliastole.Thc clastic tissuc of the bulbus is vcry diffcrent nr slructurc li-(;nl thlt of thc ehsrica of :rrtcrics. Thc lvhole ofthc heart is encloscd in rhe visceral pericar

atrir.rm, rvhich lies ,'lorsal to the venrriclc. Thc arriunr has a rhin wail, and rnuscrilar tiabcculae rnverse thc lurnen rn : loose mesh$'ork. Thc endothelial liring x therelicrc Lrrqc in lrcr and rn sone specics has a phagocyric acrivity as pert of the rcticuloendolhclirl svstem. Contuction of rhc atrium

dial sac.rvhich is intirnatcly rssociateclrvirh thc cardiac suifacc. The p:rrict:rl or outer pcdcardial sac hncs the pericerdiel pericardial space is filled with scrous fluid, seperarctrittThc

fbrces the blood through valves into rhc venrricle. The ventriclc has a mrch thickcr wall rhan lhc Atrium. ancl in normal histoloeical sections only a mjnimal lumcn is appar cnt. There is a distinct outer compa.t layer of rnusclc and an inner spongy layer wirh numerous trabecr.rl:rc.Thc rhickness of the compact iayer is rel:rtcd ro rhe scope for activiry being :Lltrlrostabsen! in less active species such as plcuronectids_ Coronary vessels run ovcr the outside ofthe ventricle, sup-

ing rhc nvo nTernbranes. Thc clectrocardiograrrl (ECG) ofrhe releosr hcr t is usually sinilar to that ofothcr vertebr:rtcs (Fig.2.20), peirking at abour 70 rnv on the QRS wavc. The concluction velocirics lrc slorver than in nunmals. being tcmperarure dcpcDdcnt. f.i'pical vcntricular systolic prcssrires are 30 70 nrmHq (1.5 4 kPa). Heut r.rtcs (4 9 kl' a) and dirstolic 10 30 mmHg

plyine the conpact muscle, thc spongv muscie obtarning most

varv considcrably according ro temperature, falline es lorv ts 15 bcats/min for trour at 5'C.At mte is ebout 100/mrn15'C rhe maxirnurr heart

Arteries
Thc vcntral aorta runs fonv:d fion dre herrt nd tlisrributes blood to thc gills via the afferenr branchi,rl rr-teries. The

o's
rmVT 1s

Fig.2.19. Thc herrr ofanAdandc sxlmon showDg the nu\cuhr venricle (v), the fibroelastic bulbus ,rnd the dark. sofr atriLLn (r).

Fig. 2.20. A typical elecrroddiognm ofi teleos asncorded by e\terni contact cleclmdes.(B,vcou(esv ofDr R.L. Os\1,ald.)

24 arteries xfferent to the gills have a normal vertebrate arterial structure with tbree layers in the wall: adventide on the outside,media and intlma. The endothelium comprises {iattened ceiis, which can usually be distinguished or y by their dark-sBining nucleus' which bulgesinto the lumen. Contiguous cellsinterdigitateso that the endothelium forms a continuous surfaceThere is a membranebeneaththe endothelium,but this is fine basement visible only with the electron micioscope The intima is ussueand the mediais composedofelastictissue largelyelastic The wall of the laminaewith smooth musclecellsin beh)veen is highly resilient and can conffact !o make ventral aotta in aaljustments blood flow (Kir\ & Burnstock 1969) The outer adventitia is thin and composed mainly of collagen 6bres. The efferent branchial arteries join, dorsal to the pharynx' to form the dorsal aorta, the precise pattern varying in differFrom the first efferentbranchialsomeblood flows ent species. through the pseudobranch and thence to lhe eyes and cranium. Also in this region, arteries bmnch ventral !o the pharynx to supply the hyoideanand corcnary systems the dlop across gills so thar pressure There is a considemble in a typical mean blood pressure the ventnl aorta of trort is about 50 mmHg (6.5 kPa) and in *re dorsalaorta 25 mmHg (3 kPa).This is reflectedin the structureofthe arteriesin that the efferent vesselshave thinner walls with a smaller amount The dorsal aorta in 6sh can be of elastictissue and muscle. artery and vein in regardedas being intermediate bet'irveen structure. [n the dorsal aorra of lower teleosts,including salmonidsand clupeids,there is an elasticligement sfielched along the iength of the lumen of the dorsal aorta which can 'heart', automaticallyincreasingcirculacion act as an auxiliary to the musciesduring swimming movements(Priede 1975) Along the lengrh ofthe dorsalaorta there are lateralbranches the to the body musculature, viscerabeing suppliedmainly by the anterior mesentericaftery

PathologY Fish

Control of circulation
in changes stroke Cardiac output can be varied considrab1y' hcart rate volume being more pronounced lhan changesin The heart has a vagal inhibitory innervation and in some speciesan adrenergicstimulatory innerwxtion has rlso been demonstrated(Gannon & Burnstock 1969) The heart also respondspositively lo increasein venous return in accord ance with Starling's law Genenl increase in circulation during exercisecan be accountedfor by the action of circu in lacing catccholamineson alphareceptors various parts of the body. Vasomotor nerves have also been demonstrxted and, although it is apparentthat there are many inletspecific familiar many of the mechanisms in <lifferences the teleosts, ro rnammallrn phl,iologi'r' hrvr lhcir coLrnrerpart'in

Capillaries
ln mammals the capillary blood (trydtostatic) pressue opposes the capillxry across an equal blood colloidal osmotic pressure iow arreriai presswethis would seem !o bc wall. Due !o the are impossiblein frsh but the capillaries highly permeableso the wall is much lower than across that the osmotic pressure (Hargenset 41 1974) The interstitialfluid in other vertebrates has a high protein concenftation and, effectivcly, plasma runs quite freely through the capillary *dls Thus fluid balance in teieosts is firndamentally dill"erent from that in mammals so conccnftation in quite lergechanges plasrna that, for example, can be readily toleratcd

Lymph
The lymph drainage system of fish is very extensive,probably permeabiliry The lymph volurne ofthe high capi11ary because is about four times the blood volume (Wardle 1971) and its compositionis alnost identicalto thar ofblood plasmaln the main bulk of the myomeres the lymphatic circulation is rhe only circulation available since thete are no significant blood in vessels the white muscleThere are valious lymph propul 'lymph hearrs' along the length of major lymphatic sors or which aid lymph return during breathing movements vessels (Kampmeier 1969). A unique Gaturc of6sh circulatory systerNis the presence of a secondarycirculation, arising as narrow coiled rrtedal vesselsftom the gill primary vesselsand also ftom arterial lrerr sucha\ 'kin ;nd gur' Be'au'e supob ro \Jrious rurt.rce the majority of the blood cells are directed via the primery

Veins
are The veins offish, aswith other verlebmtes, relative\ indisrnainly of collagen The tensible and have wa11scomposed are major veins are large in dlameter,and pressures low' being iessthan 10 mmHg (1.5 kPa),although there is no evidenceof such asoccur in the sharks' negativeprcssures Thete is renal portal drainege through the kidneys, mainly from the caudal rcgion, and ftom che viscera there is a rypical vertebnte hePatic Portal systemValvesale not col non ln the leleostvenoussystem.

of TheAnatomy Physialogy Teleosts and circulation system at the bifiucations of the two systems. the blood in the secondarycirculation is normally oflower haematoctit and lower prcssure.Ils circulation time may rherefore be of lhe order of hours rarher dran the minutes & required for the pdmary circulation (Steffenson Lonholt 1992;Iwama & Farrell 1998).

25
blood and ; rcduccd considerably with increasc o:,1'genated in tenperaturc.Ar low lenpemtures rhe slope of the carbon curve n also grcater.The amount of dioxide dissociacion carbon dioxide,which can be carried by the blood, and the potenti;ll turnover at the gilLs is therefore grcater at low (Eddy 1971). temperalures

Haernoglobin

and gas ttansport

BLOOD COMPOSITION
Biood volumes of teleosts are small conpared with ,11 o*rcr classes of vcrtebrates, being in the region of 5% of body weighc.

Most teleostshave haemoglobin in their erythrorytes as do olher vertebratesSince blood tempenturcs are ofte4 1o1g nuch orTgen can be carried in simple soludon in the plasma and so certain polar lishes have no haemoglobin.There is up considerable veriation in fish haemoglobins, to four types occurring within an individual, each with its own character istics.Also, many 6sh have difi'erent haemoglobins at difl.erent stagesof development.Speciescan be adapredto different end acclimatizrtionto diferenvironmentaloq/gen tensions, drssociation ent tempentures entailsmoditcation in o>,rygen chamcteristrcs. in The Bohr effect,whereby increase Pco2 or reduction in pH lowers lhe afhnity for o:.fgen, occurs particulxrly in 6sh adaptedto conditions ofhigh oxygen and low carbon dioxide contcnt. Fish living in acidic watersof low oxygen content would not bencfit fiom e Bobr shift. ln many fish there is an additionalphenomenonwhere low pH lowers lhe total orygen-carrying capacityof the haenocurve.This so called globin aswell asshifting lhe dissociarion Root effect is unique to the teleosts.It great\ facilitates unlording of orygen to rhe tissues and is ofgreat importance in in sccretionofoxygen at high pressure the gasgland ofthe swim-bladder and in the choroid plexus of the cyc. The is at overall cxchangeofrespintory gases the gills and tissues gready influenced by the rate of loading and unloading of oxJgen !o the haemoglobin offish blood.ln the gills the pH is about 7.4 and here cherate ofoq'genation is approximately But four times fasterthan the rate of deo),1'genation. at low pH, q/pical of actively metabolizing tissue, the process is reversedand the rate of deoxygenationis 400 times faster than that of orygenalion (Forster & Steen 1969; Hughes & Koyama 1974). In the transport of carbon dioxide the turnover occun mainiy in the bicarbonatecomponent ofthe blood, and direct conbination with haemoglobin to fom carbamino compounds is small. this end the frsh crythrocyte containsthe To which facilitatesthe conversion cnzyme carbonlc anhydrase, Thc capacityfor carrying of carbon dioxide to bicarbonace. dran in carbon dioxide is signficandy higher in deoxagenaied

Plasrna
The composicion(mgl100 n ) of brcwn trout serrm is givcn by Wolf (1963) aschloride 424, sodium 358, megnesium2.3, potassium20.1, calcium 12.5, phosphorus 9.3, sulphate0.8 and whole blood glucose 77; fteezing point depressionis about 0.57"C.This is remarkablysimilar ro mammalianserum and indeed mamrnalian salincshavc bcen successfullyused for fish rissueculture. Frcg Ringcr solution is low in NaCI and KCl, is hypotonic and is tlnsuitablefor fish work. prorein concentrations lower *ran in man (7 g/ are Plasma litre), v:1ues from 1.68 to 6.19 g/litre havingbccn rccordedin differcnt speciesof teleosls. The imrnunological and othcr functions ofthc protcinsare broadly sirnilarto thosein higher remain to be vertebratesbut many interspecific difl-erences investigated.

Cellular cornponents of blood

The cellular components of 6sh blood differ from those of higher animals principally in relation to the nucleation of the red cells and the presenceof nucleated thrombocyles. These, rather than thc anuclear plateletsof higher animals, are the source of prothrombin. There are also differences between specieswith rcgard to particular elemenls and the rcles ofthe neutrophil and eosinophil ere lessobviots in the leleost.An excellent comparativereview ofthc blood cellsof tclcosts end their tinctorial properties is given by Yokote (1e82).

Erythrocytes
The teleosterydtocyte is similar in size,tinctoial properlies and ultrastructure that ofthc other vertebrates to but,like the avian and reptilian erylhloryte, ir is nuclcatcd (Fig.2.21).

Fish Pathology and alkaline phosphatase tests. (Jltrastructurally, the specifc gmnules are oval in shapc and exhibir a frbrous aPpearance. They bear a resemblanceto one of the lcss connnonly found gnnule q'pes ofmammalian neutrophils (Ferguson 1975a). Releasc of neutrophils into the blood, causinga neucophilia, is known to occur as a non-specific responseto a vadety of stressstimuli in mammals and fishes.This is prcbably mediated through the pituitary adrenal axN. The origin of teleost neutrophiis is most probably lhe tissueofthe kidney,though the spleenmay play haemopoietrc are ofteleost kidney,gmnuloblasts secn a minor role.In smears by in large numbers and may be characterized their histochemical prcperties.They are similar in morphology and stainingpropertiesto their counterpattsin mammalianbone is rnalloq the myeloblasts myelocytes.There litde informand ation on the lifespanof teleostneutrophilsbut they prcbably have a rapid turnover tine ofabou! 5 days,asin manlrrtals.

and white cels&om bloodsnear Fig. 2.21. Erythroqtes various = x1000. Leishman ofplaice. = b'rnphocytes,T ihrcmbocytes. L

and Numbers vary with species and are alsoaffectedby slress environmental temperature, but they usually range betlveen 1.05 x 106/nm3 and 3.0x10"/mmr- Immature erythrocytes, representapproximately1% of known as polychromatocytes, the total number and are rounder :nd biuish-greyin Giemsa Haemoglobin is, as in orher vellebmtes,the stained smearsmain vehicle for transport of oxygen and to a lessctextent where carbon dioxide,but unlike in mammalianerythrocytes, in anaercbicmetxbolism predominaces, the teleost erydnocyte, cell metabolismis primarily oxidacivephosphorylation, resuJting the production ofATP in

Monocytes
end cells,which, under Monorytes are partially differentiated will develop into mature cells of apprcpriale circumstances, the mononuclear phagocyte systembut are not capableof further division. ln teleostfishesthis systemis organizedasin other vertebmles,with circulating monocytes arising ftom renal haemopoietictissueand being readily able to take up a funchonal tissuercle. Monorytes ofEshesform ebout 0.1% ofthe circulatingleu in cocyte population,rhough they increase number for a short time (about 48 hours) after injection of foreign particulate matter like colloidal carbon. Morphologically chey are very similar to mammalian monocytes,which they also resemble a histochen1ically,possessing few 6lle scatteredgranules,which stainpositivelywith PAS and acid phosphatase. Uhra(truccurallythe cell membrrne i. thro\ n in(o pseudopodia and the chromatin of the eccentric nucleusdispersedmarginallyThe lysosonesvary in size and are usually prcm The Golgi rppantus is especially very electron dense. in fishes have been observedto takc up inent. Monocytes foreign particulatematerial such ascarbon and thorctrast,and where melanosomesare rcleased, in parhological processes Their powe$ of phagocylosis these are avidly phagocytosed. are,however,limited comparedwith those ofhigher animals.

Neutrophils
The term neutrophil, or polymorphonuclear lcucocyte, is drawn from human histology. Since the granules are not necessarilyneutral-saining, and the nucleus may not be multi lobed,in other species ofanimal lhe tenns heterophilor, in 6sh, 'type I leucocye', havebeen suggest-.d, in view ofits wide but usagethe term neuftophil will be used in dis descriptron. on Neutrophils have been identified in teleosts ultrastructural and histochemical grounds. Evidence of phagocytic activity, such as is found in mammalian neutrophils, is also available,and they are conmonly found at sites of inllamma r'or. (Figs 2.22 and 2.23). Neuhophils in fish are presentin about the samenumbersasin mammals(3-6 x 10rlnm3) bul they comprise a much smallerproportion ofthe blood leucoryte population (about 6-8% in fish comparedwith 60 70% in mammals). Morphologically, fuh neuaophils closely resemble their mamrnalian counterpats though the degree of nuclear polyThe histocheraical morphism in teleostsvaries considerabl)r charactersof plaice neuftophils have been extensively studied they berr closeresemblance by Ellis (1975)and in mos! respec6 to marninalian neutrophils, being positive with periodic acidacid Schift'(PAS), Sudanblack B and the benzidine-peroxidasc,

Thrombocytes
Thrombocytes arc responsiblefor blood clotting and are important in preventingthe lossoftissue fluids fron a surface

Ihe Anatomy ard Physioiogy Teleosis oi

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FishPathalogy

lO prh

Fig.2.23.B]oodcellsoft]rePlaice(continued).(u)Kidneysnerrshorvllgsagcsinthcdel'elopnenrofthenelltroPhil'prograuloblast(Pgb)] x1025 (!) Krdnev $reaf stagesfro thc srrnulolrlasrto Nxture granulobhst (gb);nuturc gnnulocrte (g); lvniphocvtes0) Lcnhnun (e) iegrrive (gb) gr"nulobl,isrr glanuloq'tc; (leb) late etvthiobixst grrnuloc,vtecontain,icid ptro,pr,"t"s" g,.nur",.The crythroq,tic scriesrrc strining granulesrpperr (gb) are ncgrtive,positil'cl-v uear Granuloblasts haemrtoxvlin x900. ($) Krdney terr Acid pliosFhxmsc counterstaiDed n.1,"'nth",'.,t*"gn.'locytes'sudanblrckBcounrcAtainedLeishnanXg00(\)Ki.]neysrrelr'Thegranu]ocyrcseriesfrolntheertl,v gnnuloblitst{gb)sninposilive]ywhilstthecrytlnocvticselies(cb.erytlrroL.1rv)artncgative'PAsDdhaeInatoxylnx900'(y)Kidneysnear shosl4sagesurthedevelopnenrotthecrythrccytespcb'proerlthrobhst;leb.letccll,titob]ast:Pec.pro-ervthroc}(.Leishnranx450, { F r o n lE h s 1 9 7 4 . )

n:rlian verleinjurn Thrombocytes are founcl in ail nontirn brates- Typically thev te elongatcd ce1ls'oftcn being terncd 'spindle cclls', fiough nosr conmonl-v. onc polc of thc cell is out inlo a point They clot rcadilv rnd if circ is not llrawn taken in thc prep3r:rtion of a blood snear the throrrbocytes mav crst olf most of thcil cytoplasn and lpperr '1s snall, dcrrselv staininq nuclei, surroundctl bv :r ninr'rle amourlt of

cytoplasm

lt is this

spent' lhromboclr'^ the lynphocyte

rvhich h'rs been When obscrved

fiequcntl-v confusccl with

rn the living st]le by plrtrsc-contrast rnicroscopy, a rcfi'rctilc vrcuolc can be secn ar thc basc of the polD[ed cnd of thc rhrcnbocyte, jusl lirterior to the nucleus Acid phosphrtase and PAS positive matcrial is zssoci:rtedwith that samc rcgrorl.

Ihe AnatonyandPhysiology Teleosts of The ultrastructure ofthe cytoplasmofthe teleosrthrombo cyle has a remerkable similarity to drat ofmammalian platelets (Ferguson 1975r). A labyrinth of interconnecting vacuoles ramifiesthmugh the cytoplasmand opensvia Gnestree the to exterior.Foreignparticlescan enter this labyrinthinesysten es in mammalianplatelets activephagorytosis probablynot but is involved. The dificulty in distinguishing 'speDt' thrornbocytes ftom lymphocyceshas led to much confusion regardingcounts of these cells.Unlcss thc tbrombocytes are preservedin their mature, intact, pointed or spindle forms, then differential counts oflhese celiswill not be reliable. In plaice blood, Ellis (1975)found, by a specificfluorescen! antibody technique,rhat the ratio of thrcmbocytes to lymphocyteswas 1.4 to 1, total numbersofthrombocyles ranging ftom 60 000 to 70 000/m#.

29 particles has been both claimed and denied by different

Basophils mastcells and

The basophilsof verlebntes are uncornmon granularlcucocontaining large basophilicmetachro rytes, characteristically matic granulessimilar to those of mast cells. The function of basophilsis not clear and though they contain 5-hydrorytrJptamine (5HT), in resemblance mast cells, lheir rela !o tionship to tissuemast cellsis not established. Like eosinophils they are affected hormonesfrom the adrenalgland:nd also by seem to be involved, in an as ye! undetermined way, in allergicand srress phenomena. The presenceofbasophils in 6shes, like that of eosinois, phils, claimed by some workers and clisput.^d others (see by review by Ellis 1975).Afirmativc reports of their presence liken them to lhe basophils nammals in their morphology of and stainingreactions.This hasnot, asyet, been implicated cell in any recognizeddefencemechanismin the fish. The cellsdcsignated nT cellsin fisheshave been idenrs ast cified solcly on rhe grounds that they have,in comnon with mammalian mast cells,a connective tissuehabitat and cyto plasmic granules which are basophilic and metachrcmatic. Recent wolk has, however,ako shown that the metachromatic granular cells prcscnt in the dermis of plaice skin, similarly contain 5HT. A propcrty offrsh mast cells obsened by rnany workers is the lability ofrhe cytoplasmicgranules. Roberts et al. (1971b) were able to stain the subepiderrnal mast cells of plaice orny aftet thc tissue was fixed in 4% paraformaldehyde and embedded in araldite,and Bullock et al. (1976) wcrc only ablc to stainthen in vital preparations (Fig.2.24).All agrccdwith the original conclusion of Michel (1923) that the mast cell granules of 6sh were ertremely soluble stmcnrres.Bucke (1972)found that the mast ccll granules ofthe goldish stained with dre pinacyanol erythrccyanate rnethod only after fixation in 10% bufferedformalin. In mammals the mast cells are mediators of anaphyh'ris. causingthe contraction of enooth muscle,dilation of blood vesels and increased vascular permcabiliry It is noc at all clear that this phcnomenon existsin 6sh, since attemptsto induce anaphylacric-type reactionsin fish have not generally been successful although occasional anaphylactoid reactionscan be observedclinically. In summary, although the presence ofcosinophils and mast cells in fishesis disputed,they cerrainly appear!o be prcsent in some speciesand probably ere prcscnt in all spccies.[n

Eosinophils
Eosinophilsare putativelyconsidered play a role in deGnce to nechanismsin mamrnalsby phagocytosingantibody/anrigen complexes. They rnay therefore have an important role in maintaining homeostasis during infccdon and are particular\ numerous when antigens are continually being released, as in pamsitic ihseases. manmals, eosinopbilscomprise or y In 1-3% ofblood leucorytes, though thcir numbersate rnodified by certain factorssuch ashormone levels. Eosinophils are chancteristically packed with large reftac tile granuies which havea high isoelecnicpoint, i.e. they stain with acid dyeslike eosin in alkaline medium. IJltrastructurally the gmniles of mamrnalianeosinophilspossess elecffon an denseaxial crystalloid, lhough this doesnot seemto be a constantfeatureoflhe eosinophils ofother vertebrates o<elenyi& Nemeth 1969). The literature concerning the presence and natffe of eosinophils fishes notorioudy conflsed,with many claims in is both oftheir presence and absence, often in the samespecies. They are normally reported to be rare in fish blood and most of the desctiptions of eosinophils in releostsrefer to the eosinopbilic granular cells found in the skin, haemopoietic and digestive tissues,which arc almost certxinly distinct ftom the true blood eosinophil. The only criterion for identifiing the eosinophil of fisheshas been the presence fairly large of eosinophiliccytoplasmicgranules. Fish eosinophils havebeen implicated in inflamrrrationand some reportsofphagocytic activity exist.For example, phago rytosis ofbacteria by eosinophilsin goldfish and guppieshas been reported, while phagocytosis thorctrast and c:tbon of

30 { l

PatholagY Fish onty a narrcw dm ofbasophilic cytoplasm ofthe ce11,leaving in which there are a few mitoqhondria and isolated ribosomes. In morphology the majority of circulrting smell lymphocytes aplear as inactive undi{lercntiatcd cells They circulaie in this form until stinulated into action by dlcir specificantigens.Alymphocyte is saidto be malure when it is

l''*F*t
'q*'*il-.-

competentto respondto ils antigenLymphocytescirculatethroughout the blood end lymph of rhe verlebrate body and congregatein orgns which filter body fluids. The number of lymphocytes in *re blood is noticeably lhe For instance, densiryof greatcrin fishesthan in mammals. in plaice is 48x103/mm3while in man it is only lymphocytes about 2x103/mm3. Thcre is great variabiliq' in thc rcported counis of lymphocyresin fish blood, such asto give risc to gravedoubts as Blaxhall and Daisley (1973) value,aJrhough to their diagnostic be thxt useful resultscan ncveriheless in their rcview suggest obtained. Much confusion ptobably slenx from inaccurate differentiationof lymphocytesand thrombocytes(seeabove)'

J+***.1; 'n*
r#**#td
P

! f *r,

ofa ofmastcelson undersurfecc Fig. 2.24. Wet Preparation red with 0 1%neurml in 0 85%salincx350 stained wtutingscJe, (fton Bulock etd/ 1976.)

HAEMOPOIETIC

TISSI'E

Sinceteleostfish haveno lymph nodcs and their boncs usually is haenopoietic cissue locatedin the haveno medullary cavity, strcrna of the spleen and rhe interscitium of che kidney To r ofthc liver, lesser extcnr ir is alsofound in the pcriportal areas lymphoid organ, and the specialized the intestinalsubmucosa the fiymus.

role of thesecells is coming to light mammalsthe defensive At the present rime the functional role of so only dowly. called eosinophilsend mast cellsin fish can only be infcrred and thet relationshipto ihe obvious eosinophilicgranule cell the and submucosae' EGC' wbilc probably ofteleost mucosae ciose,is asyet not defined (ll-eite,1998)

Renal haernoPoietic tissue

In the kidney the haemopoietictissueforms a support matrix fbr the nephronsof the posterior kidney bu! the antenor or heemopoieticThe blastcells headkidney is almostexclusively ere situated within a sflDma of rcticuioendothelie] tissue similar to that of the bone nurrow of thc mannral The through which blood endothelialcellsline numeroussinuses, for from the renal portal vein is passcd flltration oieffete cclls ofstennius and the and addition of ncw ones The corpuscles (adrenal) tissue (cortex and medulla) ere embedded internal within the haenopoietic tissue.Another cellular srructure, found throughout leleost haemopoietic tissue but not in centle (Roberts is higher verrebrates, lhe melanomacrcphage 1975b). centes vary in their degrec of organMelanomacrophage In ization, depending on sPecies. dre lowct teleoststhey are dislributed rhroughout thc haemopoictic of clusters dark cells

Lymphorytes
The lymphocytc is the celi responsiblefor rhe inrmune in Its response. propeties are discussed detailin the chaptcron and the prcsent descriptionis confined the inrmune process, of features *le cell.The morphology of to the haematological the lymphocyte is rcmarkably sirnilar throughout the phylum Vertebrata. They are usually and arbitr:ri\ separated into large and though they prcbably represen! for enali categories, referencq, different functional siatesof cels within populationsof cells rather lhan a differencein functional capaciry The avengc e fray differ betweenspecies, g-their sizeofsmall lyrnphocytes diametersaverage4.5 pm in plaice, 8 2 pm in goldfish and virtually the whole about 6 pn.I in man. The nucleusoccupies

TheAnatomy Physiolagy febosts and of

3I havea collar oflynphocytes. Circulatingmacrophages, replete with particulate matter, possibly microbial in origin, or metabolic waste products such as ceroid or haernosiderin, home selective\ on the melanomacrophage cenues,which can thereforebe considered metabolic dumps. as

The spleen
The spleen is the only lymph node like organ to be found in teleos! fish. It is dark red or black in colour nd in healrh usuallyhassharplydefined edges. is sitrated near the greater It Fig. 2.25. Melanomacrophage aggregation kidneyoftrout.ln in lower releosts melanomacmphrges ditruseand blackbrown in are cotouLH+Ix400. curvature of lhe stomach or the flexure of the intestine. Although usuallysingle, may in somespecies divided into it be !\,voor rnore smallerspleens.The spleniccapsule fibrousAnd is devoid of musclc and does not have the dense trabeculae extendinginto the tissucwhich are found in the mammalian splcen-In sorrrespecies pancreas locatedas a subcapsuthe is lar layer to the spleenbut in most fish the ruin elementsof the spleen are the ellipsoids,the pulp and the melanomacrophage centres. Ellipsoidsaie the thick-\ dled 6lter capillaries,which resultfiorrr thc division ofthe splenicarterioles. Each comprises a thick basement membrane-bound tube within which the vesselruns, usually eccentrically, sepamted ftom the membrane by a layer of sheathedcompartments. These contain erythrocytes and phagorytic cells and are capeble,of trapping large quanritiesofparticulate metter fmm thc circulation.Replete macrophages then migmtc fron ellipsoidsco the melanomacrophage cenrrcs(Fig.2.27) (Ferguson 1976). The splenic pulp consistsof sinusoidalphagocydc rissue similar to thar of the kidncy, in which latge numbers of red blood cells r,ray be held, and haernopoierictissue,which is supportedby argyrophilicfibres.Thisn mainly lymphopoietic but not exclusively so. Melanomacrophage cenfies, silrrilar to those of the kidney,occur and are usually located close to a vessel;they may evenbe investedby strands ofits extetna and usually have a fine reticulin limiring membrane.

tissuc (Fig.2.25)-The degreeofmelanization varieswith age but at all agesthe pigment present is dark brcwn or black and has all *re biochemical and chcmical prcperties of melanin, although not necessarilylaid down on rhe characleristic melanosomes integumentalmelanin. of In higher teleoststhe amount of dark pigment presentin the melanomacrophage centresofspleen or kidney ofnormal fish is usuallyvery small,the majority of rheir pigmen! being much lighter in colour (Fig.2.26). Histochemically this is lipofuscin but Edelstein (1971) has produced evidence to suggestvery close chemical afinities between lipofuscinsor agepigments and melanin. The morphology of thc melanomacrcphagecentres of higher teleostsis also much more closely deflned.They are usuallynodular,with a delicateargyrophiliccapsule. nany In species they are closcly applied !o vascularchannelsand nay

The thyrnus
The thymus is a paired organ,an ovoid pad ofprimary lymphoid tissuesituatedsubcutaneously the do$al contilissure in ofthe opcrculum.It arises Gom prinordia associated wilh thc epidreliun of the pharyngealpouches.Lele (1932) made a compamtive study of thymic morphology in teleosts and found that its lifespan was very different in different species,
Fig. 2.26. Sectionthrcugh spleenofplaice showing chancteristic pale coloured focal nelanomacrophage centre.H + E x120.

involuting in lower teleosts before sexualmaturity but surviving, and even growing, for several years after maturity in

32

FlshPatholagy

rrr;rcrophagcs :re derivcd liorrr rlonoc-ites circuhtiug

in thc

b1oo.1rrrd their 1'rccursols: thus tLrere ere trvo ptpulrtions of nucrophaecs. one fired .urd thc other trcc to rllovc rLrorit Tbe cells of teleost lish rvhich :rrc corrsidc-redto tott4rrrsc thc llES. arc thc prcrnonoc,vrcr of the h.tctrropoictrc org;ttrs. the rnonocvtcs of the blood .rrrct1ymph, thc rnr.rophirges of ofthc

ffi'.i.fT,li't,iffi
Thcrt offish

l{$'..-tl*i,ii ,i,t"
o

loosc cornectivc tissue, thc ficc arrd fixei-l nlcrophatcs splecn .rnd kirtnq lc.' r . ' lr , r r - - u ', . . h , - r . l ' "

:urd, rrr man,v species,rhc lircd nrecrophages 'o t :. Do, ,r, o .rr' r' ofits :rnd thc atriurn bec.rrLsc

the kidncy becrusc ofits ,I'idin peculirrlv vulncrablc sitc. ate a mtnbcr

oi cliiti'rcnces Lretwccn the nlanD

arcl thc teleost u,ith rcgard to thc I{ES. l'hc limrrg oi drc . . .lr.r. rr''r.r .. .r hr:hlr .r ,li l . '. I r ,r r'\ 'D 1r' \\'hcrc'rs lr has no sudr xctiviq' rrr the lighcr auimrls ( F l g . l . l 8 ) . L y r l p h n o i l c s c l o l o t o c c u r i r f i s h . : r n c lt h e l i v e r ,

+;tffit$tlai

whosc Iiupficr

cc11sprovrrlc the larsest :rlcir ol phagoc,vtic virtudl)' irrerr ('ith r:j;rr cl to phago

ti\Lrc ir manm.rls. t

llit6iiffi X',;;*;S:,*;;:
lighcr tcleos|s. In \cctio| cords ele seen.rnct. r:rrely fbcrl cpithelirl corr-cspond to thc Hrssl's corpusclcs.

c,vtosisr|r the tcl.ost (Ferguson 1975,r).

itx,i

rnd eprthelioicl cclls. *'hicl-r are lbtrld iu teleost Girnt cc11s chronic infl.rmrlatorv lcslons. arc rlso considcrccl part of thc 'tnononuclc.rr nracrophrge sy\tern RES. or nore propcLlv the (V.rn Furth 197t1),sincc the,v erc lb,lnc'cl iiorn llsion of indi vidu.rl m:rcroph;rgcs in the prescncc of cerrain un.lcgreclable irirarrrs (Trrmrr 1975).Thc rnrto[ry oi thc various tisucs. rvhch poxes a phrgocvtic eorr\roneni, is devribcd rtnder

Fig.2.27. Se.rrcn olsplccn IIo,! Phic. nocuhrcd nnn!'erorsLy .r houfs prvioush srrh Indirn nk.Thc mk pltlcles hevebeer s L , { r p c dr i t h r r $ c $ ' i l l s o i t l e e l l i ; ' s o r J H.. + F x l i ) o

thcir ftspectivc orgrrr s-vslerls. An inlclestins liature oi drc nlrctophagcs of the telcost to tottn

thc thymus is arr regtegete of srn:r11

RES is thelr c.rpacitv. rvherhcr lixecl ol cilcuhting,

hrrrphocytcs with r fibrous c;rprule rnd trnc lg,wophilic \rLP porting cclls. Mecrophaecs ;ue vcrv nlLrlrerolrs rn the th,vmus of sonrc iish (e.g. /-o7rlrirs)mel alc t-ounrl rrr clo\c rssoci.ltion -lhc thttnus is thoughr to bc thc site of nith lvmphoc,vtes. irr11lr!rnocompc!cr)l f cell nr.rrurarion. L)ccilsion.i cpithelixl nesis which nrry

rggrcgrres orlcc they rre lcplctc UsLullv th._se.rggrcgatcs lre in rhe rlc.rs oi the nchnonracroph.rges oi thc illelrloPoicttc a tissuc\ (Robcr!! 19751) btt such rgerc'g:rtes r. rlso fbund.

'!+

THE RETICULOEND SYSTEM

OTHELIAL

(RES) rs rhe systcn ofphrgoThe leticulocnclotheli:rl s,vstcrrr cvtic cells, rvidcly dispersed throughout rcsponsiblc fut the rcnoval rn:rttcr ll onr thc rirculation. The critclr.r bv rvhich cclls are includcd irr th ll-ES rtc ligh phrgocltic rctn'itv rnd crplclw to corl(crlhrle vciqht :1n.i scs ot eYrdcrlce Fig. 2.28. Se(tlon drrough the rttiruD ofr phic. iricct(d rnll rrrnun ;crst cclls 6 hour prviouslyThc lircd phrgocytcsofth. hNe trken up thc yc:st and st,li! brislit red PAs xTanl thc bo.t1'. rvhich is of efii'tc celis rnd Puticulate

rcs.rtc such phagocvtoseri nutcri;rl.'fhe

surgcsts thrt irr mannr;rls an.t prob:rtrJ-r,telcost frsh. tissuc

The Anatony and Physiologyof Ieleasts

licqucntly pigmcnred, within or around chronic inflanuutory lesions. It is not kno\4.n \r'hether lhe piguenrs found in sr.rch regregates are of purelv exoscnous orisin, nor is it completely clcer why therc is so liequcntlv asociation with a collar of lymphocyrcs in mel.nin and

slrch aggregares. However,

rL'htcd pigmerts

are considcred ro play a defensivc rolc in thcir capacity fbr H2o. generation

rnany orgenismsr in (Edelstcin 1971).

THE RENALAND SYSTEMS

EXCRETORY

The regulationof thc inrernal body f,uid composirionof ish is a complex prccess. The skin is esentially imperuious in rdults but. as poinrcd out previously, fluxes of !.v?ter and ions rcrdily taLeplaceacross sills.Orher su ccsat which transthe tirs take placc are the gut wall and rhc kidneys.Thusit is rhe conrrol offluxcs in all rhlec organs-grlls, hidney and alimen, rary canal-that constirutes osn-roregularory cxcretory the and requirencnt offshes.

Fig. 2.29. Kidncyofrarrrbor rroutshorving s.h1re rrchncphric du.tsJoininqto torD urin!+ bladder (rrrowed).

dr.rimngrhe rail region tncl consritudnga renalportal systclt. Tbrs portal blood is rhlrsvcnous (Hickmrn & Trunlp 19ar9). Thc structurc of dre tclcosr nephron varies considerlbly bctwcen marinc, euryh:rlineand frcsh-rvater forms, mirroring rhe significint diflercnccsbenvccn dreir r:espcctivc funcrions. The lephron of rhc typical ftcsh-rvalertclcost conprises a rvcll vascul:rrizcd glomer-u1us (Fig. 2.30). r ciliatcd neck. trvo distincivc proxrnal segnents (one rvith a prorrircnr brush

The excretory kidney

Thc kidney of thc rcleostfish is a nixed organ comprising haenopoietic, rcticuloendothelial.endoctinc and excrcrory clcments.Thcflrst three funcrionsare Jl dealt rvith clservher.e .rnLl this section rvill be confined to consideration of the e\cretory component. The kidney oftelcosrsis usuallylocaredin a rcrroperitoneal posirion rp aeainsc ventralaspect the ofthe vertebralcolunrn. It is a lieht or dark brcwn or black organ nomaily exrending rhe length of rhe body caviry It is usually divided into anterior or head kidncy, which is largely composed of hacnopoietic elenrenls, and posterior or excrctory kidney.Although cmbq'ologicJly arrsing as a paired srructure,its aduh form \'arres rvirh spccies. ftom trvo separatc parallelorgans species in suchasthe anglerfish.rhrough varying degrccs ofatachmcnt, ro the cotnplete fusion found in salmonids. Thc ureters or :rrchirrcphic duds, which conducr urine lrom the colleding drcts to fie urinary papilla,may fuse ar aDylc.velanclrnay bc dilatecl, alicr sion,to folm a bladder (Fig.2.29).The urinary ducts open to the oursidepostedor to the anlls. Artcrial blood is supplicd ro rhe kidney by renal arceries ansing diredly fiom rhe aorta or frorn segmcntal vessels. Exccpt rn the smallnumber ofaglomerular species. thcseserve rhe glomerular capillay bcd and then drain into effcrentarte rioles-In marinc and euryhalinespecies peritubular capil dre laries also receive blood from caudal or segrnenred vesscls.

Fig. 2.30. Glomenl$ from thc nrld-kidDcy of conmon cxrp. pAS x600. (By co!rtcsy ofD. M.Yokore.)

34
boder and the second containing numerous mitochondria but a less devcloped brush border)' I narrow ciliated intermediatc segment, a dislal sesnent and a collecdns duct system Thc r,rarine nephron is rypically smallcr in slze 3nd consrsts ofglomerulus, neck segmcnti l\Jvoor three proximal segments, which constitute the major conPoncnt, an lnlermeoralc scgment occasionally found betwecn the first and second

Pathology Fish betwecn sPacc ofsalt in a specializcd liorn a high concentration ce11s, which are opcn !o lhe seaThe saltgr:l adjaccntchloride dient is producedby a vcry high densityofsodiur,r pumps,on thc membnnes, facing these intcrcellular spaccs(Fig 2 31)' The mechanisn of inward pumprng of s:lt in fresh-wrtcr r the thought to be csscntially samcasin amphibianskin can alternatebetwecn the nechanisrns EuqThalinesPecies accotdingto rcquiremcnls, fresh water and rnarine spccies of of the gloncrular apparatus somc undctgoesdcgenalthough eration in the marine enviromnent.Therc arc alsobasophilic to primordia. adjacenc glomcruli' in teleost nephrcns'which may bc the protoglomeruli rvhich will form new glomeruli, i[ reqLrlrcdOsmorcgulationis mrinly undcr endocrinc control via the posterior pituitary and interrcnal corticoid hormones and possibly the caudal neurosccretory syslen xnd the .juxtAglornerular apparatusof the kidney (L^g)er et al 1962)' Prolactinlikc horrnones apparenr\ control Na ef1lux from rhe gills but they also afcct the kidney and bladder wall' Corticoids also modify the ionic transport and perrneability chxmctciislics of different tissues.Thus mirny cndocrirre with scxual ma[lrrtiorl c:n secondarily changcs associalcd and osnoreguhtory mcchanismsln 'nadromous aflect ionic 6sh such assalnon rhe young fish hasto reacha certaln stagc

p!:oximal segncnts, and the collecting duct system The euryhaline teleost usurlly has a nephron combining thc sEucture ofboth qTpes,bcing similar ro that oflhe marine tclcost with, in :rddirion, a distal segment similar to that ofthe ftcsh water leleost. Certain marine and fiesh-warcr specics are rnonxlous nephron in rhar thcy do ]1ot possess glomeruli in the fhere rre rrenv rnJivrdual 'peci(' vrridtro'l\ in

ncphron structurc and the above descriptions provide only a broad summary. The major work on this field is that of Hickman antl Trurnp (1969) and this should bc consulted for n1orc detailcd comp:ralivc information-

Osrnotic and ionic regulation

In fiesh-warer the environmcnt is lrypo-oeronc and water tend: to passinto the body fluids thLough the gills and perby mcable sufaces of rhe pharyrrr' This is cornpensatcd thc volumes of d ute urine' so that A kidne)r producing large elomcrular frltrarion is of gteat importance. lost through rhc gi1lsand signifrcant Ions are alsopassively amounts arc lost in thc urinc The latler is mininized' by the Equilibriurr urine luving very low Na* and Cl concentrations is maintainedby acive uptakeofNa* and Cl by rhe gills and absorption from the food rhrough thc wall ofthe gut' ln the marine environment the blood has a relativelylow so osmotic Prcssure that, by passivediffusion, watci is lost through the gills and ions are gained.Urine is produced in small quantitieswith low or negligible glomerular frltr:tion The urine is approrimarelyisosmoticor slight\ hyporates. osmocicrnd conlainsmainly the divalcntions Mg'* and SOz-a' Largevolumesofsea-waterarc taken up by drinking to com water loss.Thedivalentions are generfor pensate the passive not takenup by thc gul so thcy apperrin the faeccsErcess a1ly monovalentions in the body are excretedthrough thc gills lhc gills in teleost Key slructuresin ionic exchangcacross which areimplicatedin pumping salt fish are rhc chloride cells, inwards in fiesh-water and outwards in seawatei Thcy are principally at locatedon both primery and secondlry lamellae, !wo, and are found in grcater the juncfion benveen the that salt 6sh.Ir hasbeen suggested in densitics marine-adapted pumping ftom the gills of marine fish rcsuits fiorn ditrusion

Fig. 2.31. Model for NaCl pumping acros thc gills m sea-water' Ep = ePrthelill cel cc = chloride cclls,srs= smoorh tubular svstem,

Ihe Anatarny Physiology feleosts and of of development involving drasric endocrine changes to become a smolt before being capablcof living in sea-water (Lengdonet al. 1984). lhc me,h;nxm. of ronic reg!l.rtionand o'motegulanon and their .ontrol arc particularly complex and the specialist tcxt of Maetz (1974) should be consulledfor further dctails. In pathological investigations should be noted that rhe it slrucluresinvoived in water and ion exchangecan undergo highly signilicant changesaccording ro the anilnal'sstate of development, compositionofirs aquaticenvironment and the endocrine factors.A.ll of thcse must be uxen rnro accounr when exar,riningclinical naterial fiom the gills, gut epithe lium and renal tissues. The tr:in nitrogenous end-products of metabolism arc ammonia and urea; both are highly soluble and erc readily expelled into the surrounding water, mainly by diffusion through the gills.Trirnerhylamine oxide occurs generally in fish and can be an important nitrogenousexcrerory componenr ofthc urine under marine conditions.

35
and specializations suit spccificdicts,arc oftcn vcry distincro tive. Overall length ofthe digestiverube is a mxjor differcnce, *rat of herbivorousfishesbeing vcry much longer than that of carnivorousspecies. Othcr specializatiofi,related to diet, including dentirion, prcsenceand numbe$ ofdiverricula and evenin somespecies complctc absence ofstomach, are alsoof considerable taxonomic signilicance(Fig. 2.32). The swim-bladder, which is derived embryologicallyfrom rhe upper digestivc tracr, bur has no digestivefunction, is considercdasa separate system.

Mouth
The mouth and buccalcavity arc sharcdby thc rcspiratoryand digestive systems. Their digestivc funcrion is confined to selection,seizureand oricntation offood for transferto the stomach.Details ofthe rangc offeeding mechanisms usedby fish are rcviewed by Alcxander (1981).Chewing and predi gestion,found in manrrnals, not usually a function of the are mouth of the teleost except in a few highly developcdhcrbivorous species. The mouth end pcriorel rcgions fie well

THE DIGESTTVE SYSTEM

The digestive tract

cndowed with sensorynervc cndings and teeth, which vary greatly in location, morphology and mmber The lining of the buccalcavily consists ofa str:tificd mucoid cpirhelium on

Teleost feedon a vcry wide varieryof substrates that fish so a drick basemen!rnenrbrancwith a very condenseddermis differenccs oral srructures digestive in and rract.adaptxtions binding it to bone or muscle.

Fig. 2.32. The abdominalcontentsof a minbow trour. a top predatory carnivorewith a very short l.uestine.

FishPathologY

36 Oesophagts
and vcry muscr'rlar is The oesophagus usuafu a short' straight ofthe stomach lts tub. p",riog fto- ,h. mouth !o lhe catdia with mucous cclls epithelial lining is usually well endowed folds into which its *a *rir, ^rrd t;e extensive longitudinal of awkward inner surface is thrown, allows for easyswallou'ing are blind diverthere ln food particles- some tropical species teeth or sacs) oesophageal (oesophageal dcuia

Stomath a sigmoid' higbly disThe stomachvariesin size'It is usually lt is highly mus tensible sac\vith numerous fol& in its lining from the siiatcd the cular and the cardiader,rarcates change tbe smooth muscle muscle of the anterior digestivetr:rct to are a number of layersof muscle' occurring distally There tissuc mucosa,and rhe layersof conneciive with a r,rr-rscularis with large numbers of adjacent to rhis are often endowed mucosa itself is very eosinophilic granule cells The gastric ofthe folds Some hxses mucoid, with numerousglandsat tile for rcasoD this no are species agastric; particular\ convincing el al (1975) have but anomaloussituationis available Kapoor for consumption of suggcst.d that it is a modification co allow which mus! be passed i"r"g'. "-oorru of indigestiblc ballast quickly through the digestlve tract' from the distel' Pyloric caeca, blind-ending diverticula and liom the anterior pylo.i. uriu. rcgion of lhe stomech but notably in the intestrne, ate found rn many specres or more Their histo salmonids, where they may number 70 featurcs resembie those of the logicai and histochemical intestinerather than the stomach'

ofa regionofthe intestinc ofthe Pyloric Fiq.2.33. Secdon cells:c = circr ar musclc; ,,,i0.* ".",. t = eosioophilicgranule lnins H + E x240 ccllsofnrucosal M = mucous digestivetmct of Physiologicalchangcstakc place in thc periods of starvation' ,n"r-ty,p".i., of telcost duing cyclical Thcse changesare particular\ well Agr"rion o. spawningsuch as the species and anadromous m"i.d in catadromous salmon' whcre Europcan eel and the Atl:ntic and Pacific shrunkcn and datkint..iind ;ola. flattcn, cells become into rirc epithclielnecrosrs end lhere is oflen cxtensive scaining tenaciousmucoid intestinal'ontent

The liver
organ ln wild fish it is The teleostliver is a rclativcly large and lighter brown in herusually reddish brown in carnivores may be ycllow or even bivores but et cefiain times of year it generally contain off-white. ln farmed frshcs, where diets in colour than in the higher levelsoflipid, it is usuallylighter be a localizedorgan efuivalent wild spccimenThe liver may or may' in somc spccresihave in the anterior abdomen or are which extend the lcngth o1'the abdorncn processes ln some speciesit is a clos.ly "pptied to lhe other viscera but in organ in the form of a hepatopancreas' .o-pooti orgao ic orher,thc pancre,r, r 'epamte rn of {ish liver dift'ers Oorn the mammalian The histology of the hepetothat there is far les ttndtnry for disposition ancl rhe rypical Portal triads of thc cytcs in cords or lobules which are irre_ mammali"n live. ate no! apperent Sinusoids' polygonel hepatocyrcs'are gularly distributcd betwecn the cells with very f"*". in ,ru-b., ".d are lined by codothelial cells ate not found in prcminent nuclei Functional Kuppfer

Intestine
to diet' the Atrhough its relative length may vary accordrng tube' which doesnot rncrease intestineofmost frsh is a simple lt may be straght' in diameter to form a colon posteriorly of the abdominal sigmoid or coilcd, depending on the shape epirhelium' overlay.i'trr. u n", a simple, mucoiil' columnar often richly endowed with eosinophilic ing " ,tbmrl.o,, muscuiaris rnucosa and grl.r.tl" ."[, and limited by a dense Rodlet cclls arefrequendyseenin ibroelastic layer (Fig 2 33) ' the lining of the intestrne

Rectum
*rall than that of the The rectum has a thicker muscle r,rucigcnic lt is capableof intestine and its iining is highly corxiderabledistension

TheAnatonyandPhysiology leleosts of

37

Fig. 2.34. Section rhrough liverofnormaltroutshowing sinusoi& bemeenmuralia ofpolygon: hepatocytes. x110. H+I

1t

the lining of the sinusoids. This was first demonstratedby Varichak in 1938 and has since been adequatelyconfirmed (Ellis et al. 1976) but descriptionsofso called Kuppfer cells ba.ed .o1elyon rrorphologicel crrterraconLJnue rppear ro (Hinton & Pool 1976).The sinusoidalIining cells are fenes trated and overlie the space of Disse which is the zone betwecnsinusoidcellsand hepatocytes containsmicrcvilli and ftom both, rs well asnumbers of fat storagecells,the cellsof Ito. Hepatocytesare polygonal and have a drstinctivecenffal nucle$ with densely stainingchromatinmarginsend a prominent nucleolus (Fig. 2.34).The hepatocytes oflen swollen are wrth gJycogen neulr2lfJr when nutrrBonr, eren mrrgin or ally lessthan ideal and during cyclical starvationphasesrhe cells may be shrunlen and the entire liver loaded with yellow ceroid pigrnents. The biliary system also differs ftom that of rnammals in that intracellularbile canaliculioccur which eventuallyanastomose to form typical bile ducts. The bile dtcts fuse and ultimately form rhe gall bladder, which contains the greenish yellow bile which is conductedto the intestinevie the conmon bile duct.The gall bladderlining is transicional epirhetium which olien contains rodlet cells. Haemopoietic tissue,complete rvith melanonracrophagecentres,is firund iD varying amounts around rhe larger vesels of the liver, and where a hepato pancreasis present this invests the larger branches of the heoatic porul verna e glrndulJrexrerna.

Fig. 2.35. Section ofnornal rainbow rrcut pancrcas.The darkly stainingexocrine pancreaticacini surround the pancreaticducts (PD). The ch:racteristicpaDcrearic cellsare ar bonoD ofpicrure. fat I = isletsof Langerhans. + E x70. H

pyloric caeca, a subcapsular as investmentoflhe spleenand as an exteinal layer around the hepaticporral vein. The acrnarstructureof rhe exocrrnepen(rcatrr' tr:\ue r. very similar to that of the mammal xnd is cornprisedof cells with a very dark b:sophilic cyropiasm. ln actively feeding fish thesecontain large numbers ofbright, eosinophilic,secretory granules.Thepancreaticduct usuallyjoins rhe common bile duct somcwherealong its length.The.endocrine components of the pancreas, isletsof Langerhans, the consistofa number of Iightly capsulated, poorly staining structurescolnprised of thc srnallfusiform o, B and 6 cells-The sizeofislet cellsrnay vary wich scason and,in somespecies, there is one major islct, known as dre Brockman body (Fig.2.35).

NUTRITION, GROWTH

METABOLISM

AND

The pancreas
The pancreatic tissueis more variablein locacion, even within a single species, than rhe other abdominal viscer:.The mosc common sitesfor it are asscaltered islandsof searetorytissue interspersedamong the fat cells in rhe mesentery of lhe

Most knowledge of food requirementsfor 6sh is basedon work on salmonand trout in artificial culture.More recendy, however, information conccrning the dietary requirements of omnivorousand herbivorousculcuredspecies such :s carp and cilapiahas become availablefor comparison {auncey 1982; Jauncey 1999). The main energy foods are carbohydrates, proteinsand fats,asin all other vertebrates.

38 Carbohydrates
as offish can ucilizecarbohydrate wcll species Alrhough rTrany aslipid as an energysourcc,at levelsup ro 25% ofthe diet, it is generallynor an inportant dierary corrponent. Essentially, that of a diabeticmanrnlel. fish encrgy metabolismresembles hypcrThus the ingestion of glucoscwill produce persistent hours-This slow glucosernetabolismis glycaemiaovcr nrany pertiy due to low catabolic activiry in tissuessuch as livcr' muscle and kidney. lnsulin, of fish origin, will lower blood but levclsand pronole transfbr!o tissues, it scemslhat glucose insuljn is not calleduPon to modulateblood glucosclevelsin is rcleost fish.While in man,malsrapid glucosehomcostasis ro mainlain bmin function, in {ish higher glycogen required unnecessery. this responsc rcscrves thc bnin may n-rake in levelsremain unchangedfor In starvedfish, liver glycogen which indicatcsthxt the oKidationofother subup to 22 days, over lhe mobilization and hydrolysis slratestekesprecedence of glycogen and the oxidation ofglucose. Thus the capecity of fish to metabolizc glucosc aerobicallyis lorv relarivc to mammals.It aiso appearslbal thc nervous tissue'which uses rather ghrcosease primary fuel, may rely on gluconeogencsis formct proces 1shighcst when the The than glycogenolysis. dict is high in protein and ]orvestwhen protein is low and is carbohydratc high. It would seem that lish, which are predomin:ntly carni vores,use non-essentiaiamino acids and lipids as preferred Whcn dietary carbohydnre levclsare low, sourcesof energy. blood glucoseand glycogen rcservescan be mrintained by gluconeogenesis.
Tabl 2.1 Mean essentisl $1nio

Patholagy Fish
acid requirements, based on data belonging to the

from various species ofreleost 6sh, nainly

Ddilt tueight-rc|aled ttl t ner1." t fuE/kr Lrtly Right/dat)

.1.311.0
Histidlne

1.7X0.2 2.610.5 ,1.010.6

3.1i0.7 3.110.7 0 610.2 .1.110.1

a rangc fron-r420 to 766 g/kg live weight gain, rvhich is not dissimilarro tcrrcstrialfarmed animals. Although meny abiotic factors influence metabolism and thus growth (seelatcr).asyet therc is no evidenccthat f:ctors such as tcnperatute or salinity affect the relativesizc of the protein conponent of lhe diet. which have bccn studied so far rcquirc the Thosc species arrino acids (EAA) as tcrrestrialfarm animals essential samc (Table2.1).

Lipids
Fish require lipids as a sourceof energy and to maintain the With lower structure and function of cellular mcmbranes. than homeotherns, 6sh udlize lipids with temperatures body lorv mciting points. Many of the conrmercially imPortant msrine species fccd on crustacean zooplankron' which faity acidsin dre forr,r contain high levelsof polyunsatumtccl As of wax esters. a consequenceof this diel, fish such as herting and capclin iay down large lipid depotsin the livcr and muscle. The constituent lipids are oils consisting of fatty acidsof the ro3 triglyccridescontaining polyunsaturated largeamounrsofoils, which rnakeup ofsuch Possession series. 10-20% of the body weight, means that lipids rathcr than As arc carbohydrates the Dain cnergy reservc. a rule, body Jipid contcnt is inversclyproportlonal io body watcr conteni; in thrsrs .ru'eful rnde' of G'h condicion 'omc circum'lrnLe' ln gencral an increasein rhc non prolcin componcnts of the diet leadsco improvcd prolein utilization Diets conhin ing between 10 nd 20lo lipid by weight, opdmize ptoteln utilization, although conrmercial saLmonid diels now fre quently contain up to 30%.Alsothe protein-sparingaction of lipid, Excess lipid is more e{lectivethan that of carbohydrate.

Protein
for Protein requiremcnrs frsh are linkcd to grcwth and gonad developrnent-In fact therc is a cle:r linerr relationship betwecn daily protein requirementand specificgrowth fot x wide rangc offish.This implies that the utiljzation ofprotcin is relatively constant and is independent of the feeding i.e catcgory, carlivores,omnivoresand hetbivorcs 45 Dietary proceinrequirementfor fish mngcsbec'iveen and 70% of the grossenergy content ofthe dict. Comparedwith high. On the other hand,for farmcd homeothcrms,this seems body tempera horneotherm to rnaintain a constAntelevated is ature,thc total encrgy requirementfor maintenence higher than that of an ectolherm of similar size This diference will to lead, consequently, a highcr propottion of non prctein componentsin che diet. If daily protein requiremcnts energy offish have are relatedto flesh produclion then all catcgories

TheAnatomy Physiology Teleasts and af however, crn cluse mortaliry associated wirh fatty livers, particularlyin salmonids. Polyunsaruratcd fatty acids of rhe 0)3 series have becn shown to be csscntial componentsof rhe dier of fish.A deficicr,cy can lcad to ccssarion ofgrowth, caudd fin crosion,fatty liver,c:rdiac myopathyand the developmentofthe shock syndromc.Whcn linolenic acid (24:6c03) addcd ro rhe diec.at is lcvcls of around l% by rvcight or 2.7% of the total energy contcnt, growth and improved lood conversionis stimulated. A comprehensive rcview of rhe lipid nutririon of fishes is by Bell (1998). sivcn Frcsh-waterspecies achievehighestrvcight gainswith diets containing borh co3and (06 scricsfarty acids-Thelafter series is more typical of terrcstrial organismsand m:ry icflccr thc origin oflood in fresh-rvater fish. It is inrerestini notc that to dietary requirements ofa given fish species closelymatch thc body composition of that species caught from thc rvild. and presunrably feeding on a totally nacuraldiet. There ale. as night be anticipated, non-cncrgy food rcquircncnts lbr minerals and vitamins.These requirements xre broadly similar to rhosc of higher animals. and rhere is a graduallyaccumularing body ofinfornation on the effeclsof rherr deficicncyin the diet.

39
changc but also by the r:!te and dircction of thc change. Abrupt and substantial increase even u'irhin the nornel telnperaturerangecan be lethal duc to a metabolic overshoorand a failure in the trxnsport of respiratory gases. smaller A decrease ternpeialurcis lcss rraumaticrnd hasbeen fotnd in to be usefulfor handling and ttansporting{ish.sinceactivity is suppressed aDd chc metabolic ovenhoot teduces oxygen consu1nption disproporrionately. Ifthe lemperaturcis increased grrdually (1"C per day) fish can be acclirnacized a wicle mngc of tcmpcntures, often to span ng 20 30'C, dependine on the species(Fig. 2.36). Within this ranee lhe melabolic mle doubles,apprcximately, lor cach 10'C rhe in tenperature.Also, within this ranec. there is an optimum tempcrrture for the speciec. which ir is rt p-rfornrng r'rosf cfiLr(ntl\. Ar rhr. oprimrrn ren,perrrure. conversioneffciency,growth and swirlming perfornrnce are nraximal.For cach species optirnum temperaturc rclarccl the is to lhe lenperature reginlcn ofih natunl cnvircnment. As temperaturcincrc$cs, the level of spontaneous accivity alsoincreases, this is reflectedin swimrning pcdormlnce and and active metabolic lare. both of which rncrease to the up optinum tcmperatrre,aiier r,vhichthcy flanen out and even lually crash rt thc upper lethal temperature. nost spccics In thc curtailment of the increases Duximal active nletabolic itr

Metabolic

rate

The measurcmcnt ofmetabolic rate,primarily asolll,genconsumption, prcvides estimates the energy requirementsof of fish, fbr both husbandry and watcr managcmcnt.Also, as aqualicectotherm, fish arc very responsive ch:ngesiD their to cDvironnent. which is reflicted in changes merabolicrate. in T\us rhe rrer'urcmcntof orygcn consllnrptiorr J .en\rflvc rr method of establishingthc relative importance of various environmentalfactors.As a result of rhe variabiliry of mettbolic rneasurements standard thc metabolic ratc is tekcn ,s an arbitrarynon-ectiveleve1 which is extnpolated ftom the rela '',vrrnmrng .peedand o',ygen. on\un.phon. tron.hrp berween mcasurcdafrerat leasr24 hours fasting.Thisstandard rate can thcD be usedto eslimateminimal nuintenancerarionsand the influcnce ofenviromnental factors(for reviewsseetsrett 1962; 1971);Brett Groves & 1979).
I

28 24

100min

E<*t,

*.^trF.1 '!o- -- 1...-o-

E20

upperlethal temp.

Fte
E t z -9

".,*Try-o------4
lower lethal temp.

; 0
4

Ltr-.-d.-.-----

i .-'ffi-a? --too t-a1

4 8 12 16 20 Acclimaiion tsmperature

'/ .' -rJ - ' ' '..i)/ min

2A "C

24

Abiotic factors inJluencing netabolic rate Tefiperutrrc, Thc cctotherm body temperaturc follows

Fig.2.36. The efec! ofthernul acclinadon ot1 upper rnd lower lctlEl rcmperrtluesofthe sockeye salmons shown as a serierol tolemncepolygons.The iDner polygon indicaresthe renpenrure tolenDce at which 50% ofthe samplesurviveson unLinited ti le of cxposure.Thetwo outer polygonsrefer to 1000 and 100 nnnutes of exposure.Temper3tures selected fish acclimatcdbcrs'een.l rnd b]' 20"C :rte :rlsoshoivr.

ambient tenpemture very closely,except in thc large eame fish such astuna and marlin.The metabolic responses rem to pcratufe are governednot ody by the e:<tent lemperaturc of

40
mte is due to the rcduction in rhe concentrationof dissolved o>rygen lhe lemperaturerises. es Swimming per{ormanceand active metabolic rate will often continue to increaseabove above optimum conditionsifthe Po, ofthe water is increased air saturationlevel. Fish have been showr to adjust their metabolic rates io compensate, a limited extent, for long-term changesin to in body temperarure. Thus *re differences metabolic ratesof individualskepcfor long pcriods at either end ofthe thermal coefii Ienge arc lessrhan that predictedby the temperacure for cient (Q16).Thecellularbases this therr,ralacclimatization with dif: are thought to involve switching between isozymes fering thermal propertiessuch as substreteafinity. There is elso cvidcnce for the opening of alternarivemetabolic pathenergy-releesing weys to supplement temperaturc-sensitive reactions.LoFlemperature acclimatizationproducesrcduc tion in the mclting points of membrane lipids, through thc incorporation ofpolyunsatunted fatty acidsto maintaininter"-,l i-*--,,t". ---L^-,.^

FishPathology

at which these restrictions occur is called the incipient linitillg leml, above which the fish is totally independent of orygcn tension. Below the incipient limiting lcvel svimming activity

and active metabolic rate decrease with decrease in orrygen tension. When active melabolic mte and standard rate arc an oxygen

identical the fish cannot move widrout incuring

actility, which corredebc and has rcached the letel oJno excess sponds to the critical level. Benveen the level of no exces activiry and the incipient limiringlevel is the zone oftolerence

in which a frsh can survivc indefimtely buc at a reduced pace. The short range of orJgen tensions between the level of no excess activity and the lethal level where the stendard rate is

elevated and no activity is possible is rcferred to as the .zone o,f reiirldfire.In this zone ofresiscance metabolism begins to show anaerobiosis with production oflactic acid and free shortchain fatty acids in muscle and livcr tissue.In the zone ofresistance mosr fish are unable to sul'vive indefinitely. Acclimatization to low oxJgen tensions does not appear to alter the incipient limiting or no excessactivity levels but does in some species extend the zone ofrcsistance. From Table 2.2

Oxlger,

although lhere is Fish are fundamentally eeroLres,

it is clear that few species arc independent ofo:.1'gen tensions much below air satumtion levels. In the Salmonidae the level of no excess rctivity at optimum temperacures is when the

weeks in hypoxic evidenceof cypdnids surviving for several In conditions,bymaking useof anaercbicmelabolicpathways. genemlit can be saidrhat orf'gen is a limiting frctor in that it governsthe metabolic rxte by virrue of its opemtion within rhe metabolic chain.Thc availabilityof orygen in the aquatic environmcnt is low, due to rts rclatively low diffusibility and low solubiliry which is furthcr reduced by increase in lemperarure and vtinrq. ln ion\(quence oxl,gen i. more commonly a limiting factor in water than In altspecies that the standard mte It has been shown for several of oxygen consumption is constant over a wide r:engeof oxygen lensions fiom atmospheric I1o2 down to a critical tension.This critical tension,which varies lvith tcmperature and species, marks the point where the shndard rate increases rapidly as thc lethal level is reached,after which there is e the crash-Theindependentzone reprcsents range of ambient Po, in which rhe 6sh arc able to extract suflicient orygen withou! increasing ventilation ofthe gills.The rise in scanthe cost of dard rarebelow the critical levelindicatesthe increased venriiating gjll' u hen orlgen becomcs LmJtrng. rhe of Although the metabolic responses 6sh to low orrygen tensions under standardconditions indicatc the maximum it extent to which they can toleralelow orTgen tensions, does not show the exlent to which the normal activitiesare constrained.In actively moving 6sh aclive metabolic rate and maximun swinrming speedrarc reduced at orJgen tensions tension higher than thc cdtical level.The oxrTgen considerably

water is about 30% air saturation (fbr review see Hughes

1973). Catbon dinxide. Dissolvedcarbon dioxide,on irs own, has

liftle effect on the standardmetabolic m!e. However high che maximum active mte of oxygen Pco2 valuesdo depress consumption (Fig. 2.37).Thc relationshipbetween Pco2 and and in genenl the active metabolic rate is species-specfic sensitivityis reduced at higher acclimatizationtemperatures. AJso there appearsto be an interaction between increasing Pco2 and decreasing Po2 to augment the reduction of lhe rrrr\rmumlcHver.lteo[ oxJgenconsumpljon. ammonia Afifiot id. Fish are very intolerent of dissolved (NH3). Concentrationsas low as 0-1 mg/litre arc harmful. However, in water below nertral pH ammonia er.ists mainly and Table 2.2 The relationship betveenort genconsumption bodyweiglt for sevenl species offrsh
Sp&tri b ldlla

0.73 0.73 0.78 0.iIJ 0.85

los ), = 0.73 log a+0.36.1 logI:0.73logr+0.364 loC 1 = g.tt ,o* "-,,., na togl=0.881o9F0.847 logl= 1 . 0 4 l o g{ 1 1 . 9 1 0.52 logI=0.85logr

tl 13 20 10 20 15

The Anatomy PhystologyIeleosts and of

o air saturatlon (F0,160 mmHg) x40% air saturalion(fr" 64 mmHg) '30% air saturation (Fo,48mmHg)

E25 E ; R20 = .t c 10 Partialpressureol Cq (mmHg)

Fig. 2.37. The influence ofincrease ofitor on the active rcspiratory metabolism ofthe commoncary:t 25'C at threep:rrrial pressures oforfsen. (Frcn Beamish 1964.)
o

30 Satinity(%)

Fig. 2.38. lnfluence ofincrease sdinityon lhe weeklyfood in intrke.expressedr percentage 3s ofthe fresh weightofuinbow trout at 10oC.

in the non-roxic ionized form, a processenhxnced,in thc presenceof carbon dioxide,by the formation of alnnonium carbonare. Unless pH is significantly .lkalinc, therefore, ammonia rvill not be a mejor inllucncc on mctabolic mte. However,in recircularionsystensbiodegmdationof arrxronia can pioducc harnful nitrites. Saliritl. The influenceofsalinity on the netAbolisn offish

logY=logd+rlogX where Y is the oxygen consumption in mg/hour; X is the freshweight ofrhe fish in grams;l is chc cxponcnt ofweighc; and a is a constantequivalentto rhc oxygcn consumplion of l fish rvcighing 1 i.This equation is shown in Fig. 2.39 as a linear relarionshipbctween the logarichrnof weight and the logarithm of oxygen consumption,rvith b the dope of the

is only relevantto those eurJhalinespecics which are able ro toleratea certainlange ofenvironmcntalsalinitics. Many ofche conxrercially important species such as the Salmonidaeand Angu lidae llll into this categoryThe interesrin the effects of salinity on the physiologyoflhese spccies comesfrom rcports that juvcnilcs tnnsferred into higher salinities show indeased growth. However,there is litde evidencethat wirhin thc salinity range 0 28 g/lirre the metabolic rate 01 conveision effcienciesare significanrlychangedThere are reporrs that rhc metabolic cost of locornotion et maximal cruising speeds is minimxl at the iso-osmotic point of 11.6g/litre when rhe osmotic gradient is zero. However, no such eflect has been der,Tonstraled under standard conditionsand the only explan ation for improved growch ar higher saliniriesis that lherc is An incrcasc feeding rate (Fig.2.38). in

500 active 100

standard

Biotic Jactors inJluendng ffietabolic rate Body size. Sn'r:llfishhavea relacively higher metaboticrate

'r

510

50100 weight (g)

1000

than large 6sh and this genenl rule appliesboth intra- and interspecifically. The relationshipbetween body weight and orygen consunption is an cxponential funccion which is normrlly f(prci nrrd by rh. followingequ:rion

Fig.2.39. Relationshipbctreen standard and activer.ates ofo'f,gen consumptionand weight ofjuvcDne Or.orli,nd?,r',.,'Pa 15'C.The at 'l' \,rlues for standardand actrve rares are 0.78 and 0.97 rcspecrively.

42 linc. When I is unity the metabolic rate is directly propor tional to body wcighc, but with vrlues less than unity the netabolic rate of small fish is highcr th:Lnin largc fish. It has bccn shown for a number of species fish that thc , value of under standard conditions at 20'C is within rhe mnge 0.73 1.04with a meanvalueof0-83The weight exponcnt appearsto increasein value with decrease tenperature even wirhin the same species.This in phenomenonxppears be intrespccifcsincethere are no sigto nificant difcrencesin b valuesevenbelween uopical xnd polLr species.Swinming activity also tends to push the value of ir upwards.In sockeycsalnon rhc weight erponen! increases from 0.78 under standard conditionsto 0.97 when mtximally activc. Thc a values under difering environmental conditions prcvidc an interesting intraspecific colrparison. At thc optimum lempemcurcsfor many speciesthe d vxlues arc rcmarkablysimilar,suggesting thereis an optimum level of thrt energy cxpenditure which, through thennal acclimatizarion, .pecie' haveevolvedto e\plori rhcrf rrmpcrJrurc rnJrvr.:uaJ reginen efficiently. Thus tropical,tempemteand polar species have similar metabolicratesfor their particularthermal r:ange. (Orherwise tropical frsh would tend to btrn themselves up while polar species would take too long to developand grorv) Masculat actit ity. Spontaneous swimming adiviry of

Fish Patholagy standardrate. This correlateswith thc incrcase in orygen consumption whcn frsh are induced to swilr thrcughout the (Fig.2.40).However,rhese rangeofcruising speeds periods of intenseaciiviry are of short duration and swimming activiry, * hen 'p"e.rd over che24 hour p.rroo. increrc' orlgen .on ( r u m p t r o rl e s s h J nn v r ( eL h f . r : n d a r d a r cr r o p r i m u mr e r n r peratures.In the lolver end of the tenperarure nnge rhc roudnc lcvcl ofmerabolisnr is even less.

Specifc rlynamic actiotl and starvdtioh Specific dynamic adion (SDA) is the terrn used for rhc incrcascd merabolic rate fbllowing thc ingcstion of a meel. It is the enetgy usedin digestion.assimilarion, growrh, deamina tion ofrnrino rcrds rnd the ') nr-he.r, nirrog.rou' ercretory oi products. The energy released SDA is rclatcd to thc com as ponentsofthc dict. SDA is lighest for prorein ar 30% of lhe total energy content of the dict. Lipid and carbohydrare have SDA costsof 13 ar,d 5% rcspctivcly.The proportions ofthese componentsin the diet will affectthe contriburion of SDA to the tocal nctabolic rare.Since most conulercially inportant specics carnivores, arc with a high tequirement for protcin in the diet. SDA will bc high. The traditional view is that SDA represcnts energy cost ofprocessiDg thc food and n therefore compedtivewilh growth. ln other words bcst dicrs ire rhose which have the lowcst SDA per energy intake. However, Jobling (1983,1985)hasproposedcheoppositevielv that SDA is the encrgy expenditureassociated with srowth and is rhcrcfore inleractivc.His alternariveview is thxt diets prcmoting

aquariumfish hasbeen shown to hevea pronounccd eft-ect on oxygen consumplion.The ma-rin,umroutine rate (the orygen , orsumprionar norIn,rll(vcl\ of 'porrrnc.ur ryrovemenr In rnoderate confrnement) can bc as high as ten times the

1000
!/ DUU

100 4 0 ; 30 E, 20 E)
'v i

100

E
llJ

0.5

1.0

'1.5

2.0

Swimming speed(bodylengths/s)
Fig. 2.40. Relationship between netrbolic nte and sninming speedfor 1 + ye3r group haddock at 5'C and 15oC.Nore rhet rhe ordinate has a logarrihmic sca1e.

TheAnatamy Physiology febosts and of good growih will induce high SDA.Thus dietsrich in protein promote grcwth in fish. SDA n ofparticular importance in farmed carnivorcs, with a high or1'gendemand,sincefeeding at the upper end oftheir remperature rangecan resultin lethal overdemand orygcn. for thc Gill pathology and reduced water supply rvill exaccrbate si!uation. Star-vation been shown to decrease has standard metabolic rate by 25-50% ofthe non-scarved rate in 7 300 days. Rapid decrease characterizes eady slages starvation, the of followed by an exponentialdecay. Even for short periods of star-vation (1-2 day$ the standard rate can fall by 10%. Finally,changing day length end the onset of sexualmatu rity and reproductivebehaviourcan changcboth the standard and dle me-ximalrcutine metabolic levels.Differcncesin [he order of 30% have been found benveen the rcsiden! and migratory phases somespecies white 6sh.Unfortunatcly of of ic is extrenely difiicult to separatc the colrrbinedeffcctsof out photoperiod and temperature upon the physiologyoffish and the role that endocdneshave on metabolhm.

43 bladder. They vary in size from small strands of tissue in the juvenile to iargewhite flabby organsapproaching\2y" of the total body wcight. Therc is a nain collccting duct for gcnital secrctions ftom thc testis-the vasdcfcrcns which conducts mature sPermatozoa an cxcrctory mcatus at lhc urlnary to papilla. The testisitself comprisesa seriesof cubules blind sacs, or rhe seminiferous rubules, which are lined with spermatogenic (or seminiferou$epithelium.The process maturation of the of male gameteinvolvesthe multiplicarion of spermatogonia, or sperm molher cells,which develop from the spermatogenic epirhelium to fbrm spermacocytes. Many of theseeventually undergo a meiotic division to become the haploid spermalo zoa.Spermatozoa attachto thc surfaccofthe pyriform, nourishing cells of thc seminiferousepithelium known as Sertoli cells, until readyfor release. Secrctionoftestosterone, male the sccondarysex hormonc, is by thc intcrstitial cclls ofthc tcstis, locaccdin the fibroussupportiDg tissucor clscirr thc bescircrlt nembrane cif the seminiferous tubules-

The ovary

THE REPRODUCTIVE

SYSTEM

The femalegenital tract ofthe celeost vrrics in structurefrom the simple cluster of ovarian follicles found in thc lower 'pecies. releost rhe very complexorgenlound rn vrvrp,rrous to Thn not only produceseggsbut aiso acts es a spernatozoa store,a vaginaand a uterus where the young crrbryos can bc nourished. The mature ovariescan representas much as 70% of the rotal body wcight. They arc suspendcd from thc abdominal wall by a mesenteryand usually appearas a sn,all cltsrer of ninute omogc whitc sphcrcs in thc immaturc fish. Thc primary ovarian cells arc the ovarian follicles.These line a hollow cavity or potential cavity which has a very colrplex scrics of folds in its lining. Ova arc passcd into this cavity as they marure.In the higher teleosts the ova are pass-.d directly ro cheoutsrde an ovrducr. rhe nore pnn'rhve.pecre\ vra but such as the salmonicls passthe eggsinto a fold of mesentery which ukimately ruptures and relersesthe eggsdirectly into the abdominalcaviry for evacuation the genital opening. via Oogonia, the cellswhich are beginning !o mature,are sur rounded by a singlelayer of small epithelial cells:nd it is this aggregate ova and epitheJialcells which is known as the of ovarian follicle. The epidrelial cells grow as rhe ovum grows and are separated rtom it by a gradually thickening hyaline capsule, zona pellucida. the These granulosa cells,as they are known, are responsible nourishing thc ovum end sccrcting for its yolk. Should an ovun degeneralebefore ovulation they

The teleost fish show greater divenity in their reproductive pattcrns than does any other group in the animal kingdom. Although most specieshave male and female sexes, hermaphro(development ditism and bi-sexuality occur. Parlhenogenesis liom an unGrtilized ovum) and gynogenesis (development liom an ovum stimulated to divide by penetration from a sperm which does not conrribute gene$ are alsorecorded,either in *re wild or in the labomtory (Purdom 1972).Eggsand sperms may bc discharged into thc waccr for cxtcrnal fcrtilization or copulation may take place, resulting in either discharge of fer tilizcd eggsor viviparousreleasc young fish (Hoar 1969). of Young 6sh may be hatchedin nestsand protectedby nulc or femrle,hatchedftom eggsdeposited redds, in released inro the plankton liom floating eggs or even held in the nrouth of a Parent. An understanding of leleost reprcductive analomy and physiologyald is pathophysiology particularlyimportant in is cultured species, where *re egg production and larval stages are normally the most criticrl in cerms of economic as well as biologicalcficiency ofthc system(Brcnage & Roberts 1995).

The testes
The testes are paired organs, suspended by mesenceries from the dorsal abdominal wall, alongside or below the swinl

Fish Pathalogy cxcept of6sh rcserrrble thoseofother species The neurones which :]re thar therearesomc,suchxsthe Mauthneriangroups, very large comparedto those of mammals.Supporting cells, and uricmglia),are thc neurcglia (:srrocyres, oligodendrocytes grey and elsopresent.CNS tissueis divided into the ciassical white malter consistingof nuclei of neurcnes:nd neuroglia, (Aridns Kappers respectively and myelinatedaxonalprocesses, et al. 1960). Thc brain and spinal cord are protected by a single primitive rneningeallaycr, thc ncninx prirnitiva, enclosing cere brospinalfltid (CSF)producedby the chomid plexuses.These glomcrulus-like, invaginations the vcntriclcsare of ependymal, often in very differcnt sitcswithin the brain, cornparedwith the mammal becauseof the differenccsin infolding of the Fig. 2.41. Sectionihrcugh ovaryofl:inbow trout showingprnury . I n d . e . o n d ro u . ) , e . P O p r i m " r y o o . \ 4 e : S oe . o n d r ^ D = macrophages ofooclte degeneration.+ E x60. H at site oocytei M releostbrain. in The roots of rhe spinal ncrvcs,cspccialJy rhe region of the donal rcot ganglix, are ustatly ovcLlaid by clusters of eosinophilicgranularce11s, which are norphologically similar in invade the degenerating cell before themselvcs turn being In invaded by rnacrophages and nelano-macrophagcs. m:ny speciesseveralgenemtionsof ova may be found in differenr stages developmenr(Fig.2.41). of The fecundity of teleost 6sh varics. Those fish which merely liberate their eggs into the water, usually prcduce fbr considerable numbe$, to compensatc the lack ofcare.Thc mature comnon cod, for instance,can prcduce some nine million eggsper season, whereaslhe Nile rilapia,which cares for irs ofiipring by mouth brooding, produces only a few thousand, several in brcods, over an annual cyclc (Maclntosh & Little 1995). to thosc frequently observed in the teleost intestinal and other loose connectivetrsslres. submucosa

The brain
The teleost brain is similar in its besic components to the but with nany diffcrencesin forrn brain of higher anirnals, and cornple;iry For easeof description it is usually divided into five divisions comprising, from thc anterior: the relencephalon,thc diencephalon, the mesenccphalon,rhe metencephalon or celebelhm and the medulla oblongata (Fis.2.12).

THE NERVOUS SYSTEM

The nerwoussystem of the teleost extends throughout d1e body as an interconnectingsystemofintegr:tion centresand the coll nunication pathways: neuronesand their axonal and processes. The largest concentrationsof nervous dendritic tissueare in the bmin and its poslerior extension,the spinal cord, and rhesetogether comprise lhe central nervoussystem (CNS)-The peripheral neNous system(PNS) comprisesthe out from fie CNS and their nerue endingsor neruespassing organsof specialsense. Only a part of the nervous function is under conscious control the nerves serving rhe striated myolomel muscles end voluntary crxnial muscle. Regulation of heart beat, chromatophores,gill respiratory movementsipcristalsisand rhe other functions of smooth muscles are controlled by as autonomic componcntsofthe syslem, in highcr spccics.

Ttlenrephalon
The telencephalonor forebnin is rcsponsiblefor olfaction and for aspefisof colour vision, riernory eDd rcproductive and feeding bchaviour The olfactory bulbs, sjtuated in the nostrils, arc dirccdy connected ro the telenccphalon via axonsfrom thc telencephalonproper,known asthe olfactory rracl. itsclfhxs no obviouslateralventriclc bu! The relencephrlon The is usually divided into pars vcntralis and pars dorsalis. olfactory component of the dorsalis, dominant in lowcr widr cvolurion:ry level so that in highcr teleosls,decrcascs teleostsit is a higNy differcntiated centre proccssingmuch morc than simply olfactory information. ln fact ablarion of telencephalon in actinopterygians causcs disruption of a wide range of behaviour, including startlc reflexes,lexrned behaviour and reproductivebehaviour (Fig. 2.43).

TheAnatonyandPhysiolagy Te/eosts of

Fig. 2.42. The brain of rn Adantic salnon (x2). S = spinalcordi O = opric lobesiC = cerebelluni I I = opt;c nerve:T = telencephalon.

cutaneous sensatlon IasIe

a

culaneous sensation

culangous sensation lateral line

olfaction skeletal musdes headandpharynx muscles eye muscEs viscera

Fig. 2.43. Generalizedfish brain showing fLrnctions ofdif|erent resions.Stippled recrangles:iegions ofgeneml integntion. 1,lexrning, appetitivebehaviour.attentioni2, sensorycoordinatioD, notor intcgration; 3. postural conrrol and autononric regulationi4. alerrnrl ni.chansms:5. honeostatic and appentivecoordination;6. motor coordination.Eferenrs:A. retlculospiul rracrjC and D cranial nerver 5.7,9, 10, l1i I, cranialnerves3. 4. 6.Afferenls: 1, spinocerebelar tract; 2, spinoretlcular tr:rcti3, cranial nencs 7. 9. 10: ,1.cnDi,l neFes 0. 5,7, 9, 10; 5, cnnial nercs 7,9, 10i 6. nene 8i7, nerve 2j 8, nerve 1. (Redruwn Fom LamrDs 1981.)

Diencephalon The diencephalonis very variablcin form but n usuallysmall and subdividesinto thrcc distinct components.rhe epichrla m u ' . t h e l h J l l m u .a n d r h c h l p o r h r h m u. The epithalamus consists ofrhe pineal body,rvhich is a light receptorwith possiblcendocrinetunctions.and rhe habenular nuclei, which ser-ve coordinate oulputs from pineel and ro telencephalon the thalamus.Thisdorsalregion ofrhe rhal to amusalsohacneurallinks with thc tetina and thc optic lobci. The thalamusis very complex in structlrie and homologics with higher species cvcn bctwccn tcicoscspecies often or are

erceedinglydifficuJrto infer.It hasa numbcr ofnuclci whosc sizes vary considerably with spccics.Ingcncraltcrms it can bc stated that thc vcntirl prrts of the dicnccphalon function mainly rs corrclation centres fbr sensory nrputs sLrchas
, - , . r ^ r i ^ - ^- , 1 ^ l r - - r ; ^ The hypothalamus is morc rcadily defincd and usually relatively large in fishes. It apperrs to comprise mainly nuclei rcsponsiblc for coordination of forebrain stimuli and lareral

line impulses. An area conrrolling fbedin!: behaviour has been locared in the inGrior iobe of rhe hyporhalamus rvhich and sccms

receives both olfactory and gusratory information

Pathology Fish to have notor control over the jaw muscles involved in or fceding.Thc neurohypophysis pln nervos:!is a downward pouching ofrhc floor ofthe hypothalanus,the infundibulum, carrying a-xonalffacls of thc Preop[ic grnglionic neuroncs The.e nrct' mr) be mrs.ivcrr 'pr\ nrngttme in rolnerPciie\' as but are not divided into supreoPticand p:rravenfiicular in higher vertcbr:tes. Inmediarely bchind the infundibulum a lies the saccusvasculosus, choroid plexus responsiblefor fluid secretion. cerebrospinal enlarged gustxtory elements derived frol-r their very wc1l supplyfrorn the medulla developcdtastereceptors.Autonomic with the laiter to the oculomotor and vagusncr-ves, is restricted bed, heart' stomach dlid,thc branchialvascular supplying,ir?lel The intestinc is not usually innervrted by and swim-bladder. Nervous rcgulationofrcsPirationis vested the par:asympatheric. ofthe medulla ofmotor neurones in an i1l-deinedaggregation the which coordinates ventil centre, knorvn esthe respirarory ation rhythrnswithout higher influence (Shclton 1970). The nerve supply to taste receptois extensivciystudied nutrirional ald fishery significance-is of because ecological, located within the medulla, as are lhe acoustic fibres,with sound perception fibres in the pars superior end vcstibular ftinction in the parsinferior.Thc medulla alsoactsasa cenffc The Maurhner cells,two vcry for control of chromatophores. larg;encurones,1ic in the mcduth at the levcl of rhe VIUth ventromedialiyright down rhe nerve rcot,but lheir axonspass spinalcord :rnd help to coordinaleswimiins Inovements.

Mesencephalon is The mesencephalon relativelylarge and an:tomically subdivides into thc optic tectum, which providesthe roof ofthe third ventricle,and lhe tcgnenlum, which is irs floor' Usually the optic tccttm is a relalivelylargecomponent and is divided by a longicudinal furrow into nvo globular sffuctlrres,the corpora bigemina.lt is particularlyconcernedwith reception and coordination of optic nerve inputs, which reach the tn lectum efler complete crossingover at the opcic chiasrna. hasa cortex confish with well fbrmcd eyesthc optic tectum taining cellswith various visurl functions.For example.clls xrrangedal righl anglesto the tcctal with dendritic prccesses for surfaceare thought to be responsible positionalfixing and detail in thc visualfield.Anorher group ofcells with processes orientatedparallel!o lhe tectal surfaceseemto be involved in the pcrcepdon of movemenl.

The spinal cord

The spinal cord oflcleosrs extends the length ofthc body and lermin;rtes, in ligher teleosls, in an cndocrine structure' lhe urophysis. Grey and whire matter in the tclcost spinal cord is well demArcrted' increasing in complexiq' wilh cvolutionary levcl, although the r'i'vo dorsal horns of grey malter are fused They contain nunrerous large motor neuroncs in both the dorsal and the ventral horn tissue.Thc dorsal and vcntral roots of chc spinal cord do not have a demarcarion of notor end sensory nerve fibres as in highcr conhin vertebralcs: both tracis a mjxture of nerve fibres. Major featurcs of the cord

The cerebellum or metencephalon The cercbellumof teleos!frsh variescol iderablyin size and with It morphology between species. is gener"lly associated and balance and cootdination of prrcprioceprive rcception a it In stir,ruli. most telosts hastwo components, vesdbulolater_ thc vestibular aPpaalis ftasal) lobe, which receivesscimuii frorn ratus and leteral line inputs, and the corpus cerebeili,more which rcccivessensorystimuli via the spinal dorsallysituated, cord fiom eruemitiesand prcpriocepton.The sizeofthe basal to lobe is rc'lated thc degreeofdcvelopment ofthe lateralline

tmcts xre the very largc ventrcmedial axons. the Mxuthnerian rxons ftom the medull!.

Peripheral nerves
There are ten cranial nerves scl'ving both scnsory and ntotoq voluntary and involuntary fundions of the head and. in the case ofthe vagus, pxm-sympathctic s[pply to the main visccml organs also-

Medullaoblongata
The rnedullamergeswirh the spinal cod without any distinct mainly four columnsofnerve fibres. demarcation.Itcomprises the visceml sensoryand motor and the somaticsensoryand dorsaland ventraltractsalsoforr,r *re roots motor ffacts.These frbrecomponent to X.In cyprinidsthe raste ofcmnial nervesv the of the medulla is evertedto encompass very considerably

Special sense organs The eye

Thc teleost eye is rcrnarkablysimilar to the eye of all orher (Fig.2-42).The sclera,lhe outer fibrous cort' has vertebrates innerv.ted by insertionsfor three pairsofoculomotor rrruscles or rhe rh|fd craniJ nervr Jnd ha. cartilaginou, o\\cou( sup-

TheAnatony Physiology Ieleosts and of ports.The cornea,the extern;rlwindow of rhe eye,is similar in its refiactiveindex to water. Its layerscomprisc an epider ma1 conjunctiva, a bascncnt membrane (Bowman's mempropria and an internal brane),a dermally derivcd subsrantia membasemenc membraneand endoihcJial layer (Descetret's brane and endothelium).Thc teleostlens is not lenticular in shapebu! usually completely spherical.It protrudcs parti:11y 'fish through the iris to prcvide a very wide angleof view (a eye'lens).Theshort focal lcngth ofthc cye neans that the fish dcgrcc of hasa large fleld ofview and deprh offocus.A sma11 accommodationis achievedby moving thc lcns towardsthe lentisn.usclc(Fig.2.44). In retina,by the action of rhe retractor many species lens,or the cornea,is tinted. the nenvork of Thc choroid vessels ofthe eyeforr,r a subscleral capillaricsfor nourishment of the rerina.They are associated

47 rvith thc largc choroid gland,a nctwork ofcapillarieswhich is lclive in oxygen secretionrnd whose function is considered ro be relatedto ensuringa high lcvel of oxygenfor the retina, although it alsohasblood-nonitoring functions.Thc orrygen secretionmechanismis the same as in thc gas gland of the srvim-bladdcr,so ccrtain syndrcmcs nay becone apparent in simultaneously borh organs. in The ocular humours have no! been rnvestigated any detail but, although there are considerablc diffcrences in viscidity between species, they can usuallybe comparedwith the aqueousand vitreous ofhigher animals. The iris is virtually fixed, having very poorly developed Thc sphincter and dilator n,uscleseven in advancedspeciestcleostrctina,thc light-scnsitivctissue, organizedasin olher is vertebretes,with trensparentnervous elemenls innermosl,

Fig.2.44. Diagnmnutic representation ofthe teleosteye.(Af.crwa s 19'12.)

48 overlayingrhe rcd and conc receptorcellswith the black pigMos! shsllowwaler fish havecolour nented layerperipherally. vision buc conesand colour vision are absentin frsh living ar depths where absorprion of long-wavelength light by lhe waier colurnn produces low intensity blue-green mono chlome illumination (Munz 1971). The visual pigments, accordingto lhe visual rhodopsins, vary in specftalsensicivity envircnment of the frsh.The optimum wavelengthof lighr tendsto match that which is best transnitted by the weter in to which the fish lives.Thus oceanic fish are most sensrtlve light with a blue bias, while the eyes of ftesh-water frsh respond best to light with reddish tint. Some 6sh have two types of visual pigment, the prcpofiions of which can vary or seasonally during migmtions betweensea-and licsh-water.

Fish Pathology with a tapetun, of sensory hairs similar lo the mcchano receptorsof the latcral line. The senson arc stimulated by the endolymph,which movementsof the fluid in the canals, to and thc stinuli are passed the indicateangularaccelemlion, tract. cerebellumvia the acoustico-lateralis ofthree inter-connccting Thc otoLithorgan usuallyconsists known asthc menbranous labyrinth (the utriculus, chambers, sacculusand lagena).The semicircular canalsinsert in the utriculus. In each chamber are loceted otoliths, distinctively shapedwhice calcifed'stoncs' ovcrlying the sensoryepitheliurr, which consistsof sensorycells which are again very sinilar !o the latcral line neurolrasls. Thc membranous labyrinth is alsofilled with endolymph,and movcncnt of the otolirhs, which are denser than the endolymph, ovci lhe providesstinulation by the force ofgraviry and sensorytissuc, low-frequency sound vibretions. The otolith organs arc and change important in the perceptionofsound, accelcration in equilibrium.

The labyinth
dcvelopmentofthc anlerior The labyrinth is an evolucionary with line, fonmng a complex sensoryorgan associated lateral 'hearing'. [t consists two of maintenanceof equilibrium and and the otolith organs canals connecledparts:ihe senricircular (Fig.2.45). The semicircular canals comprisc three fibous serrrilunlr tubcsembeddedin the skull.The endsofeach canelinserrinto the la\rinthine cavirywhich is expandedat one end to form small ridges the sphedcalampullawhich containsthe cristae,

The lateral system line

in AJthough h:' origin..rnJ ncrvou\corrponenr. iomrrron ir with the la\rinth, the laleral line is a distinct organ, found The main componcnts are the only in lower vertebrates. but there :Irc dso w.'11 pairedlateralline canals in somespecies developedcontrguoushead canals. The canalis a groovein rhe ftunk ofthe fsh, on eachsidc, with a bony suppott and an inleiumentalcover which is

semicircular cana|s

punctuated by scquential pores along its length- The mechanotcccptors locaredbasallyin the canal,alccrnatine are 'neuromxsts' stimulated erc These with the position of pores. by transferofmooon in the cxternal ,r,ilierto the watct inside dre receptorsof the the canal,which mechanicallydisplaces ncrircnasts.These consist of pyriform reccptor cells with x bundle of sensoryharr-like stuctures which exlendsup into the gclatinous cupula (Fig. 2.46) (Flock 1971). Lateml linc to neuromxstorgansarc very sensitive water patticle disPlacement such as that induced by neardeldsound of frequencies that somc fish are caPable up to 200 Hz. k hasbcen suggesled up ofdeteding the vibrations of acriveprey over distances to 32n. The nerve supply to lhe lateral line nerve, which is paralleland dightly medial to the canalitseli is mainly derived &om the vxgus(Xtb cranialnerve).

senses OlJactory gustatory axd

The olfactory organs are paired pics on the snout with e single
F i g . 2 . 4 s . T h e l d b y n n , ho r g a no r " r y p L . a 1 r e l e o v .

by opening traversed a cuspofskin dividing it into an anteriot

TheAnatany and Physialogy leleosts of

49

.:.i'r:ili-il
S,t '

Thc slrstirtorv olgrns

thc tastc buds

arc lbund both on

the oLrrer sLrdaceofthe lips. he,rcl,b.tbcls rncl fins rncl on the gil1r-*cr-s, gil1 rrchcs rnd rnouth. Indeed rn somc spccies the.v rrray bc found splcrcl ovcr thc cntirc body sudicc. Norlr ally innervatcd by branchcs of dre VIIth, ncrvcs, thc buds .rrc conposcd scgment-shaped cells lornrine IXth rnd Xth cr:rnial

of clusrcrs of a sphere. They

clongated consist of

rcceptor. \upportin!! mcl bas:rl cells. Ther-e iue rlso 1n.rn_v ftcc ncLrrones in the teleost epiclernis.

SWIM-BLADDER Structure
The gas filled su'in bladrlcr is .r conspicuous (up to 71" of bodv votunrc) r)cl ch:!rrctcristic lcrtulc of thc visccra of

t c l c o s r 6 s h ( F i g . 2 . . 1 7 ) .l r s p r i r r a r y f u n c t i o n i s r s a b u o y . r n c v rrr('chrnisln sincc thc teleost body has r specific qlaviw 107% rrnd 105% ofthat officsh:rnd sea-rvater respectivel_vlt is rlso

used loL sound and pressur-crcccprion and in sorrc spccics is cquippcd \i'ith ci urrmine rrrusclesfor sourrd prodrrctior.Thc sn'im bladder is absent in nany botton livirg spccics. \\'hcrc

ncutlal buol.:rncv is not necessxry,:rnd in sonre f'.rstsrvinuning pc'1.rgic species.ivherc rt rvould incL.easc drag bv incrcasing the surf:rce area, Irr thc lish lrrvr divcrticulun thc srvnr-bl.rddel develops es a dolsal

oi thc forcgut. so that nr the ;rdult nmny struc-

turel features of the digcstivc trrct .rrc ro:rincc1. Histologicelly it consists ol m'o nr.ril laycrs: a runica irter r.r, rvhich lines the
Fig. 2.46. The litenl ...1.- " /'t| hne orerr. ArThe "1,n H | Ddibulxf (.3: cDrl Dd

gas spacc, .rnct .r tunic.r erterna. The tunicr mterrla has ;r transitrlr.rl cpithclial layer ovedying a nuscularis nlucosa and ir submucos.r of loose, vescuhr aorlncctivc tissLlc-Thc tunic,

-t.

microgrrph of $r{rcc burbot

o{senso5

c;.idrchum of r crml or!:rn of tlic hlr burdlcs iuth kinocrLia (k) lacing

ln ce,,tr,: lre nlo rnsoF

;r opfoslte dircctions. Sterlocili,L (t are L.L'r)Jles ofnr.reasnrg length. {A b\' courtesy of Dr G.R. Crdr)eri ! bv coLrrtes! ofDrA. Flock.)

externa c_onsists ofan ooter st'tosr bcncath rvhich lics r tor.rgh fibr-ous hler rrr l'hich rusclc ]rrcl chstic comn('ctrvc ussuc 'rrc founcl. Thc cmbr-yonic conncclion betu,'een the gut rnd the su.irnbl.rrlder is rerlined rs a pneonirtic duct ir nrny of the rrrore -o. . l.^.por,iy I rJ rr rl, r 1pF of \' I I ' r r r ri r r t 'phvsostones bhdder are lelerrecl to as . ln nrost ofthe spin_v ravcd fish the functional pne rnatic drct is lost: thc dored or 'physoclstous srvinr blirdder. Lr both plrysostones rnd plryso clists the su'im bhddcr' h.rs .r rviclc r:rrrgc of Lnorphohgic.rl variation rcl:ued to hrbitrt rrd bchrviour (Fig. 2.-18). anv M ph-vsostomatous sl.nl blaclclcrsh.rve nvo charlbers sep.u.rteci

inlet.urcl postclior cxh.rust.Su.nmrnrg and breathnrgrllorv p;rsagc oflarer through rhe sacs, passineover the olfactor,v epitheliurn.u.hich n raned tu a seliesoiverv vrscul.rrfoldr or lingcn rvhich increase surfaceater,r1 sensotvtisue. the The ecual olircrorn tisuc cor'rsists oflbcal groupsofr.:cc1r t,or cellssurroturclecl nrucoicl.lrrdcililted colunu;r cpitheby liurrr.Axons lron thc olfactory bulb collect lrom the b:rses of lecepror ce11s forur rhc olf:rctory trrcts to thc rclcn to cephalo . ln rrrirD\, species thc loosc conncctivc tissucof thc subepiclc'rrlal tissuc of thc nral rnucosrL hc.n'ilv cndorvcd is with eosrnophilicgLanulccclls.

by r di.rphrrgrn. TlT e .urtcrior chmrbcr is thcn .rssocirtcd rvith grr rcccption:rrld rctcntioD and consequently has:r thicker *all.The postcrior chrmbcr: bcnrg involved in gas reabsorp-

don. has i thin runica intern.r

50

Fish Pathalogy

Fig. 2.47. The swim-bladderofthe ninbow trout (arrcwed).

f,fltr eg 98
D
ofswim-bladders from r Fig. 2.48. Diagrammatic rePresentation variery ofshallow rurer 6shes They are drawn rs though thc with the stiPpledarearepresenting extent ofthe gas transparenr, secretingconplex and the brcken lines the sphinctersepanting secretory aod absorbent charnbers or areasofthe swim-bladder' A,The soby:an exampleofa singlegasgLrndwith r large oval area B.The gadoid pattetn: a single gasgland and oval sphincter'c,The wruse pattern: double-charnbered swim-bladder with a four-part gas ofthe chambers gland-D,A varianr olc in which rhe surfacearees E,The be alteredby musculardisPlacenentofthe sPhincter' can gurnard: a multilobulate gas gland in a double-chambered bladdet r.The perch: a multilobulate gas gland wirh an oval sphincter'

ffir-------r V-t
V \-"sSYz

m
\{,

TheAnatanyandPhysiology Teleosts of

51 Frsh rvith swim-bladders can perceive relative pressure changesequivalencto lessthan 0.5% of the ambient hydrostetic pressures, while those without can only detect pressure changcs betwcen 2.5% and 10% (Tyder & Blaxter 1973).

Buoyancy adjustment
Neutral buoyancydepcndson maintaining r constantvolume in a flerible, gasfilled, buoyancy chambcqirrcspcctiveof the depth ofthe fish. ln physostomes with access the water/air to intcrface, inflation is producedby swallowingair which is thcn fbrced via the pneumatic duct to the swimbladdcr. physoIn clisc, and rhosephysoslomes with no access the water/air: to interfacc,inllation is by the reiease gasfrom arterial blood of passing through a gland situatedin the tunice interna of the anterior ventral area of rhe swin-bladder Gas reabsorption occurs when a capil1aryplexus (the oval) arising from the dorsal aorta is exposed to swim-bladder gas.The oval is an impervious muscular diaphragm which contrcls the area of plexusand henceabsorption.The conponents ofthe exposed gas are mainly orTgen, nitrogen end carbon dioxide but rhe proporcionsare,in many cascs, differenr6om thosein air.The partial pressurc oxygen can bc ashigh as 176 atmospheres of in some species ofdeep seanucrurid. [n physostomcs such as the cyprirndsthc swim-bladdercontainspure nitrogen,while crrbon dro'.ide r': andTyder 1976). mo-e rrrirble componenr occurring (for further information seeSteen 1970 mainly in physoclists

Sound production
A.lthoughthey do not possess laryn-\,some ish can producc a sounds by rubbing the serratedsurfacesof special skeletal con,ponents. Meny spccics of the families Sparidae, Tetrodontidae and Holocentridac producc high-pitched sound by grinding thcir tccth, but thc vibration ofthe srvinbladder wall by special musclesprovidcs thc grcatcst rcpetoire of noiscs or calls produced in fish. The mernbers of the Tiiglidae arc wcll knowr for their sound production. In fact, thc crcpusculer chorusesof the searobins causedgreat confusion amongsroperators of anti stbmarine acoustic echo location deviceson board warshipsof the US narry during World War II, unril the target was eventually recognized (Moulton 1956).

THE ENDOCRINE

SYSTEM

The endocrine system of frsheslus thc sarnc basic com-

Sound and pfessure reception

Sound and pressure receptionare similar,in lhat sound propagation through water involvcsprcssure oscillations wcll as as water particlevelocity and displacement.Thus flexiblc gasthc filled swim-bladder which responds!o pressurechangesby changing volume is an obvious potcntiel pressure receptor. Thc swim-bladders of many speciesare found to have direct or indirect linkage with the perilymphatic systemofthe inner Although near sound sources can be perceived by fish without swim-bladders, bone conduction, otolith vibrx by tion or laterel line response, they are insensitiveto distant sound sourcesabove 400 Hz. Fish with swim-bladdcrs but without connections with the inner ear (e.g. gadoid$ can produce good conditioned responses frequcncics below at 520 Hz, whereas the clupeids,ictalurids or cyprinids, with their direct connection, may have frequency rangesof per ception betlveen 13 and 4000 Hz.The presence ofweberian ossicles pushesche uppei liequency responseto as high as 5000 Hz. As with buoyancy,any uncompensatedchange in depth wili allect rhe sensitiviry of sound perception by changing the volume of the swim bladder (Chapman & Hawkins 1973).

ponents es that of higher vcrtcbratcs but, bccaurc of rhc considcrable differences in environmental consftaiDtsand evolutionary developrnentexperiencedby the celeosts, has it many distinctivefeatures (Matty 1985).Thc nost striking dif ferenceis the number ofendocrine sffuctureswhich hxve no apparcnr analoguc thc mamrnals.These in includethe urophysis, the corpuscles ofStannius and possiblythc pscudobmnch-

The pituitary
lhc rclcostpiruirrry as in :ll vertrbratrs. comprise. rwo ernbryologicallydistinct components.These are the neuro hypophysx, which grows down from the dicnceph.lon of thc brain, and the adenohypophysis, which originates as en upward budding of the cctodcrm of thc cnbryonic buccrl caviry The two ftise together with their rcspcctive mescnchy mal vascular supply to fonn :r cornposite endocrine glend enclosedaboveby thc dicncephalonand laterally and below by a bony cupula, the cella turcica-Extracts of hypophys--al rissucarc ftequendy used to stimulate ovulation in ct trred fishesand the proceccion affordcdby thesestructures makesits rcmoval ftom cadaverssomewhat dif[cult. The teleostneurohypophysis much simpler than charof is mamrnalsand comprises a stalk of nervous tissue with an

52
enlergementat its tip, which forms the core of the complete gland.The stalk is composedof the axons of neurosecretory whose cell bodies are located in the iypothalamic neurones, nuclei. Thc :denohypophysis secrctesa vatiety of prctein or peptide hormones and is usually divided anatomically into a pars intermedia and pars distalis lt is composed of a number of different cell types with a variety of linctoial propeties' ThJ precise correlation of the different cell types with their specifrchormones has not yet been completely worked out & berweenspecies and may dift-er @a11 Baker 1969) The pituitary hormones ofteleostscan be divided into tlvo groups: those which stimulate activiry of other endocrine and adrenal,and those influorgens,e.g. the th)'roid, gonads, such as the movements of processes encing physiological dermal melanophores, osmoregulation, metabolism and glowth. A motc detailed description of the teleostpituitary and its hormones is presentedin the excellent revievr by Matty (1985).
t

Fish Pathology

'

Fig. 2.49. Folliclesofthyroid tissuein the comective tissuearound rs the venu:alaorta.Colloid (C), the thyroid hormone secrcBon, *rce folicles. H + E x70. present in rhe lumen of

The thyroid gland

The teleostthyroid is similar in basicstructureto that of the mammal, and the t\roid hormone, which has a stimulatoty is efFecton many metabolic processes, a similar iodinated thy roxine to rhat ofhigher animals(Gorbman 1969)-Thethyroid foilicles arc, as in the mammal, usually round to oval with low cuboidal epithelium and PAS-positive colloid secretion, but a very important point of differentiation is the anatomically diffuse distribution ofthe follicles,which varies considerably lnsteadofbeing or betlveenspecies evenbetween individuals. located vsithin a disctete capsulethey are distributed through out the connec[ve tissueof the pharyngealareaor even,in around the eye,ventral aota, hepaticveins and some species, renal haegropoietictissue(Fig.2.49).This diversityof disrribution hasled to a number ofreports ofsuch folliclesasbeing but although th]'roid tumours do occur' most such neoplastic reports are probably merely of nornally ectopic follicles Control of thyioxine and tri-iodothl'ronine is through pituitary thyroid stimulatinghormone (TSH). Both are transproteins Unlike the case ported in the blood,bound to plasma in mammals, thyroid hormones do not produce a calorigenic metabo in rcsponse fish,but seefr to influence carbohydrate of responses lisrn and the mobilization oflipid reserves.These 6sh to thyroid horrnones are very dependent on the nutti_ phocoperiod,and selinity' tional state,ambient temperature, Recent work has shown that thlroxine prornotes growth in juvenile fish by stinrulating appetite.

Adrenals (interrenal and suprarenals)

The compact adrenal-typcendocrine gland comprising both cortex and medulla is only found in a few teleostgroupssuch an Normally, in teleosts adrenalcortical equiv asthe sculpins. alent, the, interrenal, a series of strands of Jighdy staining cuboidal eosinopbiliccells,is situatedin the anterior kidney, through passing with malor blood vessels ofren in association the are:r (Fig 2.50). The steroid hornones of thc adrenal

Fig.2.50, Section of anterior kidney of rainbow trout sho{ing strands ofpale suining adrenal corkx cells and the single nodule of neurosecretory neurones forming rhe adrenal medllla.The stronu of haemopoietic tissue is also wel endowed with individual dark cells.H + E x110. brown melanomacroPhage

Ihe Anatorny Plrysiology febosts and of cortex include gluco- and mineralocorticoids, and theshave very irnltrL fun.tron' to tho'e rn rhe hrgherarumals. Thc adrenal medulla, the chrornafin tissue Go called because its stainingrcaction to chrcmic selts), variablein of is location. It may be found accompanying the sympathetic gnglia, in clumps becweenanterior kidney and spine or, as indicated above,in close contact wilh the inlerrenal tissue, within the head kidney [t secretessympathomin-retic sub stances such as adrenaline, associated wirh imrnediate suess responses. perticuler, increase in blood levels of [n catecholaminescauses hyperglycaemia and increases the functional areaofthe gills for gaseous and ionic exchange. The endocrine pancreas

53
secreteinro the centre ofthe cluster (Fig.2.51).They secrete a glycoprotein hormone called teleocalcin, blocking absorption ofcalcium by the gills,so they ere in some weys akin to the parathyroids of higher animals. They also have othcr propertiessuch assecrctionofpressor substances possible and involvernent in osmoregulation,but they do not seem to secrete stercid aompounds.

The distribution of the pancreatictissuevaries considerably with specics. Thc cndocrine component is also varied, with small islets of Langerhansscatteredthroughout lhe lissue, in the salmonidsor anguilliforms.In the higher teleoststhe endocrine tissueconsists ofa small number ofscatteredislets and a large compacr islet which varies in sizewith the stage in the life cycle xnd is known asthe Brockrnan body (Epple 1969).They have a delicateflbrous capsule within which are lhe three types of isle! cells,cells which produce glucagon, cells producing insulin and cells of unknown function (Fig.2.52).Thereis considerable changein idet sizeat spawn ing and senility and with dietary changesbut rhere are aiso reported seasonaldifferences in the prcportions of the
.1itr"- -r."ll h,h..

IJltimobranchial glands
All vertebrateshave the capacity to rcgulatc their serum calcium and in fish this is achieved the activity ofthe ulti by mobranchialglands, cordsofpolygonal cells\ingjusr venrnl to the oesophagus within rhe septum separatingthe sinus venosusftom the abdomen (Copp 1969).The organ is an embryonic fifth gill arch and correspon& in function to the pamthyroid in manrmals.

The corpuscles of Stannius

The corpuscles Stanniusare usuallypaired whitish clusrers of ofendocrine tissuenormally locatedrerroperitoneally the on suface ofthe kidney.Thelargeclear endocrinecellsappearto

Insulin causes hypoglycaemiabut 6sh do not exhibit the rapid blood glucoseclearancercsponsetypicei of menlmals. Insulin release linked to blood glucosclcvcls but only to is enablerelativelylow icvcls to lcave thc extracellularspaceto

Fig. 2.51. Sectionthmugh the mid-kidney ofninbow trout passing through a corpuscleof Stannius(CS).H + E x35.

Fig. 2.52. Islet ofl-ogcrhms cclJs a\e pancrcas h ofthe cormron carp.A = alphacells;B = beti ce1ls; = delta cells. azocarnnre D AT G x800. (By courtesyofDr M.Yokote.)

Pathology Fish function ascellularfuel. lt may be directedto oxidativecleat ar,ce of glucosebui not to glycogen dcposition.Insulin has from amino acidsand been found to inhibit gluconeogenesis to reduce the rurnovcr of liver protein. The prime role of insulin may be to conserve prctcin and arnino acids and proomote tissuedeposition. to Glucagon actsantagonistically insulin in that it increases [t blood glucoseby liver glycogenolysis. also stimulatesthe incorpomcion of amino acids in thc ljver and stimulates gluconeogencs$,

The pseudobranch and choroid body

but where Thc pseudobranchis noc prescnt in all celeosts presentit is a rcd, gill like structurederivcd from the first g l arch and attachcdto the incernalsuface ofthe operculurn lt consistsof parallel blood cepillariessupported by cartilege connection with has rods.The pseudobranch a dircct vascular the choroid ofthe eyc,which is composedof similar arraysof capillaties alternatingwith rows ofslendcr fibroblast-likecellsa Those frsh which do not possess pseudobranch (e g Anguillidae) invariably alsolack a choroid rete.Although it is considcredto have an cndocrine and rcgularory function as well as a hyperoxygenationiunction for the rctinal blood supply,*resc are still ro be defined in full.

The urophysis
Thc caudal endocrine secretory struciure is only found in obscurebut sharks and bony fish.lts function is still somervhat ils anatomy has been well descdbedby Bcrn (1969) It is a small whitish ventral expansron of the spinal cord, at its postedor end.It is invesledwith a mcninx,like the rest ofthe supply draining to rhe cord, and has a very extcnsivevascular It renal portal sysleDr. is comprised largely of large neuro ccllswilh pollnnorphicnuclei,whosenon-myclinated secretory to st3lkanalogolls axonsextcnd ftom the cord,in : urophyscal adjacentto terninate in pallisadcs rhat ofthe hypophysis.They capillarywalls ofthe neulo-hacmalcomplex, akjn to thoseof thc hyporhalamus. Thc utophysealhormones are pcPtides which are concerned principally with osmoregularion,although one acts specificallyon the smooth muscle of the reprcductive lract (Bern & Ntshioka 1993).

The gonads
As well as their obvious gamelogcnic funccion, the teleost hormones,which havea gcnemlizedeffbct gonadsalsosecrete Their outpu! is conholled by the on a wide range of tissues. 'conductor of ftom rhe pituitAty,lhe ouqut of gonadotropins pro and androgens the endocrine orchestra'.Thc oestrogens duced in the gonadscauscthickening ofskin, colour changes, kypes and swclling of rhe ofbrccding tuberclcs, developn-rent urogenitalarca, alsoheve aar rcaching effectson Androgensand ocstrogens Andrcgensstimulipid and protein mctabolism. carbohydrare, promoting growth by increaslatc protein anabolicprocesses, ing the protein and RNA content oflivcr, kidney and muscle. Oestrcgcns tend to incrcase body reservesof lipids and of initiatc the synthesis vitellogcnic proterns.