Anda di halaman 1dari 12

J . Fish Biol.

(1985) 26, 127-138

Aerobic metabolic scope and swimming performance in

juvenile cod, Gadus morhua L.
Marine Laboratory, Aberdeen. Scotland & Department of Zoology, University of Aberdeen
Department of Zoology, University of Aberdeen, Scotland

(Received I 9 July 1983, Accepted 28 March 1984)

Juvenile cod (Gndus rnorhua) were made to swim in a tunnel respirometer to determine the
oxygen consumption during swimming at different speeds. Results were compared with
measurements of standard and active metabolic rates in static respirometers before and after
intense exercise. The oxygen consumption at maximum sustainable swimming speed was con-
siderably lower than the peak oxygen consumption following exhausting exercise. It is suggested
that these fish have a poorly developed system of aerobic (red) locomotor muscles which do not
normally make a major demand upon oxygen consumption. Apparent specific dynamic action
following feeding and repayment of oxygen debt following anaerobic exercise can each give rise
to greater rates of oxygen consumption. Following exhausting exercise there is a delay of about
I h before oxygen consumption reaches a peak level some 40% higher than the peak level
observed during sustained swimming.

As a fish goes about its daily activities in the natural environment, fluctuations
are observed in its metabolic rate (Priede & Young, 1977). The amplitude
through which the aerobic metabolic rate can vary is termed the metabolic scope
(Fry, 1947) and differs according to species, size, temperature and other factors.
The minimum observable metabolic rate is conventionally defined as the stan-
dard metabolic rate and corresponds to the oxygen consumption of a non-feeding
fish at rest (Beamish & Moorkherjii, 1964; Fry, 1971; Brett, 1972). The maximum
aerobic metabolic rate is usually measured as the oxygen consumption during
swimming at the maximum sustainable speed. This is referred to as the active
metabolic rate (Brett, 1964, 1965; Fry, 1971). Although metabolic scope is often
regarded as largely representing the difference between minimal and maximal
locomotor activity, feeding can also bring about an increase in oxygen consump-
tion. The processes of digestion, assimilation and biochemical transformation of
food material lead to an elevation of oxygen consumption referred to by Beamish
(1974) as apparent specific dynamic action (SDA). In a study on juvenile cod,
Soofiani and Hawkins (1982) showed that oxygen consumption due to apparent
SDA can rise to almost the active metabolic rate, taking as much as 98% of the
metabolic scope. Feeding to satiation seems to leave little metabolic scope
available for any swimming activity. Tracking studies on juvenile cod (Hawkins
rt al., 1974; Hawkins et al., 1980) and benthic feeding trout (-Holliday et ul.,
1974; Priede & Young, 1977) indicate that swimming speeds in such fish are so

*Present address: Dept. of Biology, College of Arts & Science, University of Shiraz, Iran.
0022-1 112/85/020127+ 12 $03.00/0 0 1985 The Fisheries Society of the British Isles

low that the proportion of the energy budget attributable to swimming is almost
The purpose of this study was to examine the relationship between oxygen
consumption and swimming speed in juvenile cod G. morhuu.
A number of workers have studied the relationship between oxygen consump-
tion and swimming speed in various types of respirometer (Brett, 1964; Kutty,
1968; Farmer & Beamish, 1969; Beamish, 1970; Tytler, 1969). Despite recent
work on a number of species of marine flatfish (Priede & Holliday, 1980; Duthie,
1982) the most useful set of data for a marine 'roundfish' is that of Tytler (1969)
for haddock (Melanogrammus aeglejinus L.).
A preliminary study by Tytler (1978) on cod indicated that this species is
difficult to train to swim in a tunnel respirometer. We have measured the rela-
tionship between oxygen consumption and swimming speed in a small group of
individuals that would respond to such training. The results are compared with
estimates of maximum oxygen consumption obtained in a larger group of
individuals which were tested in a box respirometer following exercise to
(a) Tunnel respirometer
The respirometer used in this study was a modified version of Brett's (1964) tunnel
respirometer described in Priede & Holliday (1980). The whole tunnel assembly was
mounted on gimbals which could be tilted but in the present study the assembly was
retained in the horizontal position as a conventional tunnel respirometer.
The fish chamber was rectangular in cross section (299 x 148 mm) and 500 mm long.
Alternating black and white stripes on the outer surface of the swimming chamber pro-
vided visual cues for the orientation of the swimming fish (Griffiths & Alderdice, 1972).
Determination of oxygen consumption was carried out by closing the respirometer and
measuring the decline in oxygen tension using a polarographic oxygen electrode (1 BC,
model 500-501) and analyser. The oxygen tension was never allowed to fall below 80%
(b) Static respirometer for measuring maximum oxygen uptake
The open system respirometers for this part of the experiment consisted of simple
Perspex boxes approximately 61 in volume. U p to four fish chambers fed from a common
water supply could be set up simultaneously with a multichannel oxygen analyser
monitoring the outflow PO, of each chamber to give continuous records of the oxygen
consumption for each fish. The output was recorded on a data logger (Solartron type
3430BD) giving a printout at half-hour intervals. The rate of water-flow through the
system was adjusted so that ambient oxygen did not fall below 75% saturation. A detailed
description of the equipment is given by Soofiani and Hawkins (1982).
(a) Tunnel respirometry
The experimental fish, ranging in weight from 149-196.2g and length from
260-290 mm, were obtained from stocks in the Marine Laboratory, Aberdeen and then
were transported to the Zoology Department, Aberdeen University, where the experimen-
tal equipment was available. The fish were maintained in the aquarium, were acclimated
to the appropriate experimental temperatures (10' and 15' f 1 'C) for at least 4 weeks and
were fed to satiation every two days on a specially formulated diet composed from natural
food organisms (Soofiani & Hawkins, 1982). Each fish was deprived of food for a 36 h
period prior to being transferred into the respirometer. I t was anaesthetised with a

solution of benzocaine 25 mg 1- and length and weight measured to the nearest 5 mm and
0.1 g respectively, before transfer to the respirometer. The fish was then left for a period of
18 h during which a slow flow of 5 cm s - ' was generated, serving to orientate the fish and
to avoid oxygen depletion (Brett, 1964; Duthie, 1980). The fish chamber was covered by
black polythene to minimise the possibility of disturbance and excitation Brett, 1964;
Priede and Holliday, 1980).
Any elevation of metabolism caused by feeding (Beamish, 1964; Brett & Groves, 1979;
Soofiani & Hawkins, 1982) was thus eliminated by a total fasting period of 54 h.
The first swimming trial was conducted at a slow velocity to initiate swimming. In
subsequent trials the velocity of the water flow was increased in 5-10 cm s - steps from
the acclimation speed of 5 cm s-'.
Each step was maintained for 60 min and the maximum sustained swimming speed was
taken as the highest velocity which could be maintained by the fish for this period.
Immediately after each speed increment the respirometer was flushed through with
oxygenated water for 30 min. It was then closed for measurement of oxygen consumption
during the second 30 min of each speed increment.
Any fish that became disorientated or exhausted and unwilling to swim was allowed at
least 5 h rest at low velocity before resumption of swimming trials.

(b) Determinution of maximum oxygen consumption

The experimental fish ranged in weight from 684-129.9 g. They were acclimated t o 7",
lo", 15°C or 18°C ( f 1°C) for at least 4 weeks. Feeding and maintenance of the fish was
similar to that for the tunnel respirometry and they were deprived of food for 2 or 3 days
before the experiments. Each fish was then chased in a tank (123 x 69 x 28 cm) for a
maximum of 5 min until exhausted and could be removed from the water in a net without
struggling. The fish was then transferred to the respirometer and the rate of oxygen
consumption monitored until it reached its minimum steady level. This period usually
took 2 W O h.
For the purposes of direct comparison with the tunnel respirometry and other data the
oxygen consumption values were adjusted to a standard fish weight (Ws)g using the
formula described by Newel1 et al.; (1977) and Duthie (1980):

where VO, is the corrected oxygen consumption in mg 0, h- for a standard fish of

weight Ws g, W is live weight of experimental animal in grams, VO, (exp) is the observed
oxygen consumption of the individual mg 0, h-' and b is a constant. The generally
accepted value of 0.8 was used for the b value.

Tests were carried out at two temperatures, 10°Cand 15"C, and the results are
shown in Figure 1. The data at 10°C were pooled from three fish and those at
1S'C from two fish. There was no significant difference between data for
individual fish within each group (Soofiani, 1983). Most fish did not readily
swim at slow speeds in the respirometer and showed a high variability in their
oxygen consumption. There was a strong tendency to dart to and fro in the
swimming chamber, in short bursts of activity. These cod were clearly not
inclined to hold station in a water-flow like salmonids. This restricted the work to
a few cooperative individuals. A regression line of the form:
log,, VO,=u+bU

200 r

10 o c

I00 0 T 04 0.8 1.2 1.6

Swimming speed (body lengths s-' )

FIG.1. Relationship between swimming speed and oxygen consumption at (a) 10" and (b) 15'C in Juvenile
cod. The horizontal dashed line is the adjusted minimum (standard) oxygen consumption for fish of
the appropriate weight (see text for details). Regression line equations and their statistics are given
in Table I. Arrows indicate the position of the maximum sustainable swimming speed.

is fitted to the data (Table I), where a is logarithm of the standard metabolic rate
and U is swimming speed in body lengths per second.
The horizontal dashed lines in Figure 1 show the adjusted minimum (standard)
oxygen consumption for fish of the appropriate weight (1 88.1 and 151.5 g at 10"
and 15°C respectively), obtained by the application of equation 1 to data derived
from separate experiments in a box respirometer (Soofiani and Hawkins, 1982;
Soofiani, 1983). These adjusted values of standard metabolism were 101 and
124mg 0, kg-' h-' at 10" and 15°C respectively, and fall within the 95%
confidence limits of the extrapolated values (Zar, 1974). Those confidence
intervals are: 904-106.7 at lO"C, and 119-3-147.0 at 15°C.
There is a significant difTerence between the slopes of the regression lines at the
two temperatures (P<O.Ol) and their elevations ( P < 0.001). These slopes 0.12 at
10°C and 0.21 at 15°C are relatively low compared with the range of values
TABLEI. Results of tunnel respiromztry experiments. Intercept values derived from extrapolation of the oxygen consumption-swimming speed
regression to zero activity

Mean Active rate Intercept
Temperature weight due to swimming (standard rate) swimming Swimming regression Fvalue Level of
speed significant
("C) (g) (mgkg-lh-') (mg kg-' h - '
(BI s-')

10 188. I 182.86 98.17 1.95 log,, VO, = I .992+0.12 U 37.77 P < 0.0005
15 151.5 322.08 132.43 1.44 log,, V0,=2.122+0.21 U 57.20 P < 0.0005

*BI s - =body lengths per second.


TABLE 11. Maximum oxygen consumption and details of the fish used in the static respirometry
experiments. The oxygen consumption is the average taken over 2 h starting 1 h after
placing the fish in respirometer

Weight Oxygen Consumption

Temperature (mgO, kg-' h - ' )

7°C 205 69.5 23 1.49 f 10.55

220 93.2 225.98 f 15.18
220 101.0 218.10f 11.21
230 125.0 241.19 f9.09
Mean k SD 218.75+ 10.31 97. I8 & 22.88 229.1 1 f 14.00
10°C 215 85.0 279. I 1 f8.71
205 68.0 285.1 1 f 17.90
220 92.0 280. I7 f9.90
Mean fSD 213.33f7.64 81.67f 12.34 281.36f 11.78
15°C 200 72.0 445.14k 15.00
210 91.0 437.48 f 12.23
210 80.0 451.39f14.56
Mean fSD +
206.67 5.77 8 1 f9.54 444.67 f6.97
18°C 225 103.7 371.03 f6.81
230 129.4 417.37f 18.63
Mean fSD 2273 k 3.54 116.55+38~17 394.20f27.96

reported for other fish (Beamish, 1978). The active rate of oxygen consumption
was 182.9mg kg-' h-' and 322.1 mg kg-' h-' corresponding to maximum
sustained swimming speeds of 1.95 and 1.44 at 10" 15°C temperature.


Table I1 shows the details of the fish and the mean maximum values of oxygen
consumption for each individual at each adaption temperature. A high standard
deviation shows the great variability in the data. Figure 2 shows the time course
of change in oxygen consumption during recovery from exercise. The peak in
oxygen consumption occurs an hour or two after activity has ceased. Recovery
time increases with temperature requiring about 42 h for recovery at 18°C. It
seems that during activity oxygen consumption is sub-maxima1 and a peak is only
reached during the repayment of oxygen debt.
In order to allow a reasonable comparison between individuals used in these
experiments, the oxygen consumption values have been adjusted to an arbitrary
standard fish weight of 150g using equation 1. Table 111 shows the adjusted
values of the respirometry results. Values for the minimum (standard) oxygen
consumption from Soofiani and Hawkins (1 982) are included in Table 111. These
can be used to calculate metabolic scope, i.e. the difference between maximum
and minimum oxygen consumption. This is shown graphically in Figure 3. It
can be seen that maximum oxygen consumption and maximum scope occurs at
15°C. It is also noticeable that the maximum oxygen consumption values are
significantly higher than the active rates obtained in the swimming experiment.

300 I


4001 I

Ftc. 2. Oxygen consumption following exercise to exhaustion at (a) 7". (b) lo", (c) 15"and (d) 18°C for four
different individual fish. The expected standard oxygen consumption rates for the fish (weight cor-
rected) are indicated by the dashed horizontal line. Expected ' active ' rates at 10°C and 15°C based
on the tunnel respirometry are indicated by the dashed and dotted lines. The weight of each
individual fish is given and the expected standard and active rates are all corrected to that weight.

The most remarkable finding in this study is that at maximum sustained
swimming speed in the tunnel respirometer the rate of oxygen consumption was
below maximum oxygen consumption. During recovery from exhausting exercise
the oxygen consumption could be 40% higher than that observed during
sustained swimming. Peak values of oxygen consumption attributable to SDA
TABLE 111. Maximum and minimum, active and standard (intercept value) rates of oxgyen consumption in Juvenile cod. Values given have been
adjusted to an arbitrary standard fish of weight 150 g and shown below. Scope for activity, which is the difference between maximum and minimum
oxygen consumption is also given

Oxygen consumption (mg 0, kg-' h-')

Temperature Minimum rate

(-C) Maximum Standard Active Scope
rate mean wt. rate rate
observed k SD Corrected

7 2 10.06 60.15 91.08 i7.38 75.83 - - 134.19

10 249, I5 48.10 132.72k 13.15 105.72 102.71 191.32 143.43
15 393. I 1 44.08 159.03 15.63 124.48 132.95 323.36 268.63
18 374.80 58.40 149.08 f5.83 123.45 - - 25 1.35

-- 300


2 20c



) ,


10 15
Temperature ("C)
FIG. 3. Relationship between oxygen consumption level and temperature, adjusted to an arbitrary
standard fish weight of 150 g. W--U =the maximum oxygen consumption recorded after
exercise to exhaustion. 0-0 =the minimum resting oxygen consumption in unfed fish.
c2-C =the highest oxygen consumption recorded during sustained swimming in the tunnel
respirometry experiments. 0-0 =the intercept derived from extrapolation of the oxygen
consumption-swimming speed regression to zero velocity.

following feeding to satiation (Soofiani & Hawkins, 1982) also exceed the active
metabolic rate observed in the tunnel respirometer; 259.8 versus 182.9 mg kg- '
h- ' at 1O'C and 397.6 v 322.1 mg kg-' h -' at 15°C. These differences are far
greater than can be explained by any margin of experimental error.
The increase in oxygen consumption during sustained swimming must largely
reflect metabolism in aerobic red muscle. Wardle (1977) states that the Gadidae
have only small proportions of red aerobic swimming muscle. If this is so, the
increase in oxygen consumption during swimming must be small.
In an extensive study of proportions of red and white muscle in marine fish,
Greer-Walker & Pull (1975) however give a figure of 17% red muscle for cod
which is relatively high. This measurement is made from transverse sections of
the tail and does not make a precise allowance for the fact that in cod, red (slow)
and white (fast) fibres are arranged in a mosaic mixture of both types of fibres
(Greer-Walker, 1970). Sidell (1980) used a biochemical method of determining
the total bulk of muscle in goldfish (Curassius uurutus); this may be a better
method to use. The proportion of red muscle can vary with exercise Greer-
Walker & Pull (1973) and temperature (Sidell, 1980), and there is evidence of
variation between different stocks of cod (Love et al., 1974). All these aspects
need to be critically examined before general conclusions can be drawn.
The slope (6) of the relationship between the logarithm of oxygen consumption
and swimming speed (Table I, Fig. 1) is low, 0.12 at 10°C and 0.21 at 15°C.

Beamish (1978) reports values of 0.1-0.6 in studies on different species of fish;

0.3-0.35 however would be considered to be a typical value for teleost fish.
Low b values are often reported in studies where there is evidence of stress or
hyperactivity at low swimming speeds resulting in high values for standard
metabolic rate. The close correlation between our estimates of standard metabol-
ism extrapolated from the regression line and independent values from quiescent
fish suggest that this error does not arise in this study.
It is possible that as swimming speed increased an increasing proportion of the
work was carried out anaerobically by the white muscle fibres. Beamish (1968)
found evidence for this in cod and Duthie (1982) found a similar effect in several
species of flatfish demonstrating elevated lactate levels in the muscle as evidence
of anaerobiosis. Oxygen consumption cannot be taken as a suitable method of
estimating the energy expenditure. The maximum sustained swimming speed in
these fish is low. We observed a clear limit to how fast these fish would swim in
the respirometer. If forced to exceed this limit they would respond with a burst of
activity clearly involving the main myotomal fast white fibres. The fish would
dart forward, hit the grids of the swimming chamber and then be forced against
the rear grid by the water flow. A struggle would ensue as the fish tried to
re-orientate itself and the experiment would be curtailed. The fish was apparently
trying to use a burst and glide mode of progression (Weihs, 1974), but obviously
this would not work within the confines of a respirometer. Beamish (1966)
reports a sustained swimming speed of 2.6 body lengths per second for cod
353 mm long but Wardle (1977) found that larger cod 0.7-0-8 m long could only
sustain speeds of less than 1 body lengths per second. He suggests that there is a
general reduction in swimming performance with size in the Gadidae. For fish of
the size used in these experiments a swimming speed higher than we observed
might be expected. It is possible that these fish which had been kept in an
aquarium for some time without exercise may be poorer swimmers than typical
wild cod. Bams (1967) and Brett (1958) both observed a reduction in swimming
performance in artifically reared or aquarium stocks of fish.
Using acoustic tags attached to individual fish the swimming speeds of juvenile
cod in Loch Torridon off the West Coast of Scotland have been measured in an
extensive series of studies (Hawkins et al., 1974; Hawkins et al., 1980). G. Smith
(unpublished) reports a mean swimming speed for these fish of 0.09 body lengths
per second. This level of exercise is probably not sufficient to maintain a high
swimming performance capacity. This may only be true for the juveniles and the
fish may develop improved performance when they move to offshore feeding
Despite possible uncertainties regarding the condition of our fish in relation to
wild stocks, it seems realistic to assume that the active oxygen consumption
observed in the tunnel respirometer corresponds to peak activity of the red muscle
fibres. It is a little surprising that there was no evidence during swimming of
repayment of oxygen debt in further elevation of the oxygen consumption rate. It
is possible that repayment of debt is inhibited by adverse distribution of blood
flow during exercise. In the exhaustion experiments it is noticeable that peak
oxygen consumption only occurs about 1-2 h after the bout of exercise. (Fig. 2).
Many reef dwelling and benthic fish display low levels of swimming activity
and yet may feed at high rates and grow fast. In such fish the demands of SDA

generally exceed any demands f o r oxygen from the locomotor muscles (Warren &
Davies, 1967).
We suggest the metabolic scope in cod and other such fish has evolved t o
accomodate peaks in oxygen demand following feeding rather than peaks during
locomotor activity associated with aquisition of food. T h e metabolic scope can
nevertheless be fully utilised during repayment of oxygen debt following violent
We are grateful to the former Director of the Marine Laboratory, Aberdeen, Mr B. B.
Parrish for the use of laboratory facilities during part of this study, and to Dr A. D.
Hawkins, Dr P. Tytler and G. W. Smith for helpful discussions during the preparations
of this manuscript. We also thank Miss Hilda Stewart for her help in preparing the
Bams, R. A. (1967). Differences in performance of naturally and artificially propogated
sockeye salmon migrant fry, as measured with swimming and predation tests. J .
Fish. Res. Bd Can. 24, 11 17-1 153.
Beamish, F. W. H. (1964). Respirations of fishes with special emphasis on standard
oxygen consumption. 111. Influence of weight and temperature on respiration of
several species. Can. J . Zool. 42, 177-188.
Beamish, F. W. H. (1966). Swimming endurance of some Northwest Atlantic fishes. J.
Fish. Res. Bd Can. 23,341-347.
Beamish, F. W. H. (1968). Glycogen and lactic acid concentrations in Atlantic Cod
(Gadus morhua L.) in relation to exercise. J . Fish. Res. Bd Can. 25,837-851.
Beamish, F. W. H. (1970). Oxygen consumption of largemouth bass Micropterus
salmoides in relation to swimming speed and temperature. Can. J . Zool. 48,
1221-1 228.
Beamish, F. W. H. (1974). Apparent specific dynamic action of largemouth bass, Microp-
terus salmoides. J . Fish. Res. Bd Can. 31, 1763-1769.
Beamish, F. W. H. (1978). Swimming capacity. In Fish Physiology (W. S. Hoar, D. J.
Randall, eds.), Vol. 7, pp. 161-187, New York: Academic Press.
Beamish, F. W. H. & Moorkherjii, P. S. (1964). Respiration of fishes with special
emphasis on standard oxygen consumption. I. Influence of weight and temperature
on respiration of goldfish, Carassius auratus L. Can. J . Zool. 42, 161-175.
Brett, J. R. (1964). The respiratory metabolism and swimming performance of young
sockeye salmon. J . Fish. Res. B d t a n . 21, 1183-1226.
Brett, J. R. (1965). The relation of size to rate of oxygen consumption and sustained
swimming speed of sockeye salmon, Oncorhynchus nerka. J . Fish. Res. Bd Can. 23,
1491-1 50 I .
Brett, J. R. (1972). The metabolic demand for oxygen in fish, particularly salmonids, and
a comparison with other vertebrates. Respir. Physiol. 14, 151-170.
Brett, J. R. & Groves, T. D. D. (1979). Physiological energetics. In Fish Physiology (W. S.
Hoar, D. J. Randall and J. R. Bretts, eds.), Vol. 8, pp. 279-352. New York:
Academic Press.
Brett, J . R., Hollands, H. & Alerdice, D. F. (1958). The effect of temperature on the
cruising speed of young sockeye and coho salmon. J . Fish. Res. Bd Can. 15,
Duthie, G . G. (1980). Physiological responses of flatfish to temperature: A comparative
study. PhD Thesis, Aberdeen University, U.K.
Duthie, G. G. (1982). The respiratory metabolism of temperature-adapted flatfish at rest
and during swimming activity and the use of anaerobic metabolism at moderate
swimming speeds. J . exp. Biol. 97,359-373.
Farmer, G. J. & Beamish, F. W. H. (1969). Oxygen consumption of (Tiulpia nilotica) in
relation to swimming speed and salinity. J . Fish. Res. Bd Can. 26,2807-2821.
Fry, F. E. J . (1947). Effects of the environment on animal activity. Univ. Toronto. Stud.,
B i d . Ser. 55, 1-62.
138 N . M. SOOFIANI A N D 1. G . P R I E D E

Fry, F. E. .I.(1971). The effects of environmental factors on the physiology of fish. In

Fish Physiology (W. S. Hoar, D. J. Randall, eds.), Vol. 6, pp. 1-98. New York:
Academic Press.
Greer-Walker, M. (1970). Growth and development of the skeletal muscle fibres of the
cod (Gudus morhua L.). J . Cons. perm. int. Explor. Mer. 33,228-244.
Greer-Walker, M. & Pull, G. A. (1973). Skeletal muscle function and sustained
swimming speeds in the coalfish (Gadus virens L.). Comp. Biochenz. Physiol. 44A,
Greer-Walker, M . & Pull, G. A. (1975). A survey of red and white muscle in marine fish.
J . Fish Biol. 7,295-300.
Griffiths, J. S. & Alderdice, D. F. (1972). Effects of acclimation and acute temperature
experience on the swimming speed of juvenile coho salmon. J . Fish. RPJ.Bd Can.
Hawkins, A. D., Urquhart, G. G. & Smith, G . W. (1980). Ultrasonic tracking ofjuvenile
cod by means of large spaced hydrophone array. In A Handbook on Biotelemetry
and Radio Trucking ( C . J. Amlaner, Jr., & D. W. MacDonald, eds), pp. 461-470.
Oxford: Pergamon Press.
Hawkins, A. D., Maclennan, D. N., Urquhart, G. G. & Robb, C . (1974). Tracking cod in
a Scottish sea loch. J . Fish Biol. 6,225-236.
Holliday, F. G. T., Tytler, P. & Young, A. H . (1974). Activity levels of trout in
Airthrey Loch, Stirling and Loch Leven, Kinross. Proc. R. Soc. Edin. 74B,
3 15-331.
Kutty, M. N. (1968). Influence of ambient oxygen on the swimming performance of
goldfish and rainbow trout. Can. J . Zool. 46,647-653.
Love, R. M., Robertson, I., Lavety, J. & Smith, G. L. (1974). Some biochemical
characteristics of cod (Gadua morhua L.) from the Faroe bank compared with those
from other fishing grounds. Comp. Biochem. Physiol. 47B, 149-161.
Newell, R. C., Johnson, I. G. and Kofoed, L. H. (1977). Adjustment of the component of
energy balance in response to temperature change in Ostrea edulis. Oceologiu, 30,
97-1 10.
Priede, I . G. and Holliday, F. G. T. (1980). The use of a new tilting tunnel respirometer to
investigate some aspects of metabolism and swimming activity of the plaice,
Pleuronectes platessa L. J . exp. Biol. 85,295-309.
Priede, I. G. & Young, A. H. (1977). The ultrasonic telemetry of cardiac rhythms of
wild free-living brown trout (Salmo trutta L.) as an indicator of bioenergetics and
behaviour. J . Fish Biol. 10, 299-3 18.
Sidell, B. D. (1980). Responses of gold fish (Carassius auratus L.) muscle to acclimation
temperature: alteration in biochemistry and proportion of different fibre types.
Physiol. Zool. 53,98-107.
Soofiani, N. M. (1983). Growth and energetics of captive and freeranging juvenile cod,
(Gadus morhua L.) PhD thesis, Aberdeen University, U.K.
Soofiani, N. M. & Hawkins, A. D. (1982). Energetics costs at different levels of feeding in
juvenile cod, Gudus morhuu L. J . Fish Biol. 21,577-592.
Tytler, P. (1969). Relationship between oxygen consumption and swimming speed in the
haddock (Melunogrammus aegle3nu.v). Nuture 221,274275.
Tytler, P. (1978). The influence of swimming performance on the metabolic rate of gadoid
fish. In Physiology and Behaviour of Murine Orgunisms. (D. S. McLusky, A. J.
Berry, eds.), pp. 83-92. Oxford: Pergamon Press.
Wardle, C. S. (1977). Effects of size on the swimming speeds of fish. In Scule Eflects in
Animul Locomoiion (T. J. Pedley, ed.). pp. 299-31 3. New York: Academic Press.
Warren, C. E. & Davis, G. E. (1967). Laboratory studies on the feeding bioenergetics
and growth of fishes. In The Biological Basis of Freshwater Fish Production (S. D.
Gerking, ed.), pp. 175-214. Oxford: Blackwell.
Weihs, D. (1974). Energetic advantages of burst swimming of fish. J . Theor. Biol. 48,
2 15-229.
Zar, J . H. (1974). Biostatisticul Analysis. Englewood Cliffs, New Jersey: Prentice-Hall