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S e r i e E l a v u n d e r s @ k e l s e r

(Report on Morzwegiu?~ Fishery and ~Vlnrine I ~ z ~ e s t i g a t i o t z s f Vol. XYI. No. 9.

P u b l i s l ~ e d bx, t h e D i r e c t o r of F i s h e r i e r s

Notes on the energy

metabolism of the cod (Gadus callarias L.) and the coalfish (Cadus virens L.)

in relation to body size

2 9 5 7

A/S JOEIN GKIEGS BOKTRYICICERT, BERGEN

Inlroduction

T h e energy inetabollsill or oxygeil consuinptioll in fish has been

~vorkecl o u t for irlany species. Leiner (1938) ancl Black (1951) havc give11 good revie~vs o l the literature.

T h e r e seeins to be n o iilvestigation on the oxygen consumption 01 the cod anel the coalfish aiter the larval stage. Soirle pllysiological clata have been gatllered on these two species anel the present paper considers some results from these experiinents.

T h e work on which this paper is based was cal.riecl out at Fiskeri- direktoratets Havforskningsinstit~~tt i n Rergen.

-

i\'Patel-ials uncl 177 ethods

T h e fish ruhich were used in the experiirrents rilere all caught oLf the west coast of Norway. Tlre size range was lroin 2.8-1750 grains.

T h e coalfisl~ is by n o illeans a n experi~neiital animal. T h e epider- inis is easily dainagecl ancl lethal infections lolloxv. T h e cod, lioxvever, is a good experinleiltal animal. Adaptatioi~ to tlre experiinental con- ditions is goocl, making the lisll easy to rvoi k ~ v i t h .

All fish Itrere starved 3 to 4 clays before they .itrere placed in tlre experimental aquariuni. T h e experimental periocl lasted u p to 10 days. After the experirnents tlre fish were transferred to Iarge storage aquariums. T h e i r sur\~ival, foocl intake and behaviour in the following weeks were used as a n indication that conclitions had bee11 normal during the ineasurement 01 the energy metabolism. Data from fish which seemed to have been under abnormal conclitions were omitted.

Winkler's method, ~vitti the rnodificatioii introduced b y Krogh (1935), was used in estirnatiilg the oxygen consumecl. Tlre grespira- tion)> of the seawater was measuuecl, but the v a l ~ ~ e stayed inside the error linlit OC the analyses. T h e oxygen consuinption ol the fish was measured in closed aquariums.

1 2 3 4 5 6 7 8 9 1 6 ,

D a y s

Fig. I . Oxygen consumption of a cod of 1750 gr.

T h e tern1 ((basal metabolism)) is used in inedicine, but it is impos- sible to keep a fish in basal conclitions according to sucll a definition.

Iii most works about fish the external clata have been standarclised, but the measured metabolic. rate has not always been a standard metabolism. Tlie physiological conditiolls of the fish are very im- bortant. T h e energy nzetabolisnz o f fish zlnder standard conditions may be defined as a steady energy metabolic rate (inside the biological variation) over long t i m e intervals, zulzich can be clefined as a normal rate by the chemical and physical characteristics of t h e environnzent.

T h e experirneiital conditions were Ear different froin normal con- ditions, and therefore the oxygen consuinptioii of the fish was very high just after transfer to the experimental aquarium. T/Viiiterstein (1908) inentiolied this fact, which he co~isiclered himself able to sho~v by experiments.

T h e cod ancl. the coalfisli have a significalitly increaser1 energy

Fig. 2. Oxygen col~sumption of a cod of 280 gr.

iiietabolisril after being transferrecl to the experinieiltal aquariuln.

(Figs. I , 2, 3 a ~ i d 4).

I t seeins that tlie acclimatioil time is different for si~lall and large cocl. T h e sanrle was fotmcl on cocl ~ v l l i c l ~ were investigatecl ill different aquariums. T h i s pheiioinenoi~ is valicl for the coalfish as well. (Figs.

3 anc! 4).

'The clilference in tlie accliinatio~l time between large and sinall spe- c i i i i e ~ ~ s may have a coniiectioil ~ v i ~ t h bocly size in relation to the volnine 01' the expel-imental aquarium. However, measurements on sillall spe- c i ~ n e n s in a slnall aquariltm clo not confirin this. T h e i r accliinatioll time is as short as it ~ v o ~ ~ l c \ be in a large aquarium. I t is not possible t o give n geileral acclimatioil time for cocl and coalfish in the sailie

\\lay as Keys (1930 a) and 1,lTells (1932) have clone for I;'crndltlis !I(L,).-

vipinnis. TiVells (1982 p. 585) writes:

D a y s

Fig. 3. Oxygen consurnptiol~ of a coalfish of 1020 gr.

1 2 3 4 5

D a y s

Fig. 4. Oxygen co~lsulnption of a coalfish of 230 gr.

L o g b o d y w e ~ g h t in g r a m

Fig. 5. Energy-metabolism of cod and coalfish in relation to body weight.

(Both species lumped together).

<<Indeed I should expect that with very active, nervous fishes seve- ral clays ~voulcl be required for thein to reach a normal level of meta- bolism, ~vliile o n the other hand, quiet, sedentary fishes may do so in a few liours.,

T h e results in the present work show that )the acclimatioil time is not species specific. Tlie pheiloinenon is more complicated, and con- tributiiig tactors must be sorrgl~t in each single individual and aniong the external stiinuli.

T h e high oxygen consu~nptioil in the cod and tlie coalfish just after being trailsterred to another acluariu~n is kno~vn. S~uldnes (1956) outlines the situation ~vlien these species are transported alive in

~vell-boats.

E ~ z e ~ g y - ~ ? ~ e t a b o l i s ~ ~ ~ i1.1 relation to b o d y size

Oxygen consumption in relation to bocly s i ~ e has lollg beell dis- cussed. One ot the difficulties is tlie unit of body size, because this is important in the comparison of clilferellt animals. Zeu~then (1947)

I inds

r

units.

T h e r e is little ot this rvorl\ o n fish. Cron2leiin (191 1) writes that he finds from the experiments ol Kiiauthe (1898) that the oxygen tonsumption in CyPri~llrs sp. is proportional to tlie surface of the body. Linclsteclt (1914) reaclles the same results on Tznca v l t l g a n ~ . P i ~ n t and Joilgbloecl (1946) empllasize that the oxygen consumption in fish is proportional to the bocly surface. Keys (1930 a), Vis~vanathan and Tanipi (1 952), ailcl We1 Is (1 935), llave rnacle investigat ions on lisll i n velatioil to body ~veight lor almost tlie xvhole size-uange ol the species.

I n the present n~orl. the coilclitioiis have been staudarclizecl. T h e results are plotted (fig. 5). Keys (1930 11) has found a three phase curve lor E'rtnclzclus I ~ a ? u i p i ~ z ? z i ~ . Eliassen (1952) has founcl the same for Atten7in salillcr ancl Zeutllen (1947) tlte same for Mytillrs ~ d u l i ~ . I n a later ~ v o r k Zeuthen (19.33) belie~res he tindc tlle saliie phenome- non in other groups o l the aiiiirial kingdorn iroin tlie results of othei authors ancl liis own experiments.

Statistics

\.\ihen tlie values origliially iouncl in the present investigation are tiansferred to a logarithinic scale, there seeins to be a clear linear legression bet~veen the ~veiglit 01 the fish ancl the oxygen consumptioil.

T h e corielation-coelficiei~t Tvas fount1 to be 0.99 £01 the ~ v l ~ o l e nlaterial (30 fish). This sigiiiiicant coefficient close not exclude the possibility ol a clittereiice i oxygen--~ons~unptloil ¹¹¹ ielation to the weight I~et~veeii sillall fish and the bigger ones.

T h e pal-amcter lourlcl, can not prove that the oxygen consumption

~ n c r e a ~ e s 111 linear proportioi~ to increasing sveight. As a l~ypothesis the 11 smallest tish tvere tested as one group and tlre 19 biggest fish a5 another group. (From the tliagrani there seems to be a natural gap between these two groups). T h e correlation coefticients found were 0.97 and 0.94 respectively.

If'esting t l ~ l e e difierent ccxielation t o e l t i ~ i e n t s against each othei i ~ a s not becil clone, ns no s ~ n l p l e ciitelioil is I,t~ol\n. 011 acc onnt ol thc hillall a m o ~ ~ n t ok material n o iurtliei statistical methocls Ilave been iiivolved, and it ~volilcl in any case have heel? rlif lictllt to get signi- l icant answers to the p~ oblems.

T h e regressioii-lniec lo1 tlic tlilec i ~ i e n t ~ o i i e d g i o ~ l p s liave, how- e l e r , bee12 cal~ulatect, and, as x\~ill be seen iroin he diagram, there ls

a slight clitlere~~ce bet~veeii the rise of the I I-group ancl the 19-group.

T h i s might imply that the oxygen consumption by increasing weight, dose iloe increase to the same exten1 arliong big lid1 as among snlall ones.

Ttre present work on tlie oxygen conslmlption in the cacl and th(.

coallisll s11orvs that there is a c o ~ l n e c t i o ~ l bct.tveerl the body size ailtl the acclima~tion time required tor reaching a ccnormal,, level of meta1x)- lism alter being transferred to anohel- aquarium. (Figs. 1, 2, 3 ailci 4).

Tire oxygen consumption tlncler s~andarcl coiiditioi~ sho~vs that tllei-e is a clear l i ~ l e a r regressioil between the two variables used. However, [he significant coeflit i e i ~ t does not exclucle the possibility ol a cli ffe- rence in oxygen-con~~lmptioll in relation to tlte ~veizht het~veen stnall Jiuli and bigger ones. (Fig, 5 ) .

L,ITERATURE CITED

BLACK, E. C., 1951. Respiration in fishes. Publ. Ont. Res. I,nl). 71 pp. 91-11 1.

CRONHEIM, TV., VON, 191 1. Gesanltstofl'x~echsel cler Kaltbliitigen Wirbeltiere, im be- sonderen cler Fiscl~e. 3. Firh. XV, pp. 319-369.

EI~IASSEN, E., 1952. The energy-metabolism of Artenzin salina in relation to body size, seasonal rllythms, and different salinities. Univ. Bergen Arb. no. 11, pp. 1-17.

I ~ N A U T H E ? K., 1898. Zur Kenlitnis des Stoffwechsels cler Fische. Pfliig. Arch. ges.

Ply.riol. 75, pp.

Iceus, A. B., 1930a. The nleasurement of the respiratory exchange of aquatic animals Biol. Bull., CVoods Hole., 59, pp. 187-1 98.

- 1930b. A study of selective action of decreased salinity of asphyxiation on the Pacific Killifish, F~ndulus parui/)innis. B~rll. Scrip/~s Imt. Ocennogr. tecf~., 2 , 1Jp 4 17--490.

I ~ K O G H , A., 1935. Syringe Pipets. Precise deter~nination of oxygen in water by syring-e pipets. Industr. fiagng. Chem. (Anal.) 7, pp. 131-133.

LEINER, M., 1938. Die Physiologic der Fischatmung. Akademische Verlagsgesellschaft M. B. EI.

LINDSTEDT, PH., 1914. Untersuchungen uber Respiration und Stoffwechsel xion Kalt- bliitern. 2. Fiscfz. l4., pp. 193-245.

I'ulv~, A. and J. JONGBLOXD, 1946. O n factors influencing the gas-exchange in fish.

Arch. neerl. zool. 7, pp. 1-15.

S U N D N E ~ , G., 1956. Transport ol live fish. In press.

VISWANATI-IAN, R. and P. R. S. Tarnpi, 1952. Oxygen consumption ancl viability of C/tnnos c/zanos (Forski%) in relation to size. P~oc, Inrliatz Acad. Sci. XXXVI, pp.

135-159,

WELLS, N. A., 1932. The importance of the time element in the determination of the respiratory metabolism of fishes. Proc. nut. Acarl. Sci., Wash. 18, pp. 580-585.

- 1935. Variations in the respiratory metabolism of the Pacific Killisfih, Fin- dulus parvipinnis due to size, season ,and continued constant temperature.

Physiol. zool., 8, pp. 318-336.

WINTERSTEIN, H., 1908. Beitrage zur Kenntnis der Fischatmung.

Pj'liig. Arch. ges., Physiol. 125, pp. 73-98.

ZEUTHEN, E., 1947. Body size and metabolic rate in the animal kingdom. C. R. Lab.

Carlsberg, Ser. chim. 26.

- 1953. Oxygen intake as related to body size in organisms. Qztart. Rev. Biol., 28, pp. 1-1 1.

F I S I C E R I D 1 R E I < T O R r \ T E T S S K R I F T E R S E R I E H A V U N D E R S B K E L S E R Reborts on No? zwegian Ijishe?y and fifa7ine Investigations

C O R R I G E N D A

1701. 11, NO. 9.

Page 7, Fig. 5, on the abcissa for Log ¹¹¹¹ O2 j e ~ hour read Log ml 0, per hour (+ I ) .

Val. 13, No. 5.

Page 3, line 5, for Sclzonlnncler reacl Scholaader.