55

THE GROWTH OF FISH

IV. THE EFFECT OF FOOD SUPPLY ON THE SCALES OF
SALMO IRRIDEUS
BY J. GRAY, F.R.S., AND S. B. SETNA, P H . D .
(Zoological Laboratory, Cambridge.)
(Received 15th April, 1930.)

(With Three Text-figures and Two Plates.)

CERTAIN scales of Salmonoid fish exhibit a series of concentric ridges or circuli,

the number of which increases with the size of the whole fish. Each year of
normal growth is recorded on the scale of a salmon by a concentric band which is
more or less sharply divided into two regions, (i) in which the concentric ridges are
relatively wide apart, (ii) in which the ridges are closer together. It is tolerably
certain that the first region is formed whilst the fish is in the sea, and the second
region whilst the fish is in estuarine or fresh water; roughly speaking the two regions
mark the summer and winter periods respectively. The factor or factors responsible
for the difference between winter and summer circuli are not fully understood.
There are two possibilities. The variation in width may be dependent on an inherent
rhythm upon which external factors may exert little or no control; or the variation
may be due to variations in the external environment of the fish. In view of the
high economic importance of the facts, curiously little experimental data are available
for a solution of the problem.
Cutler (1918), working with Pleuronectes platessa and P. flesus, altered experi-
mentally the temperature and food supply in the fishes' environment. Fish were
kept in "hot" and "cold" tanks, the latter being 50 C. cooler than the former, and
at each temperature the fish were fed once a day. The effect of food supply was
determined by a comparison of fish fed with excess of food twice a day with fish fed
sparingly every other day. Cutler's results, as summarised by Graham (1929),
were as follows:
Table I.
Tanks
Control Hot Cold Abundant Scanty

Length increment in cm. O"I 04 04 07 0-3

Sclerite number 9 ±2 10 ±1 9±i 13 ±3 8±f
Sclerite width in n (average) 9±* 12 ±i 8±£ IO±I 9±i
56 J. GRAY and S. B. SETNA
These data suggest that high temperature increases the sclerite width without
influencing either the growth rate or the number of the sclerites. Abundant
feeding, on the other hand, increases the growth rate and the number of sclerites,
without marked effect on the width. Cutler's observations are not readily harmonised
with those of Thompson (1923), Dannevig (1925) and Graham (1929), since these
authors found a marked correlation between sclerite width and growth rate. In a
state of nature, there can be little doubtthat the growth rate of fish during the summer
is greater than that during the winter. If the sclerite width is associated with
growth rate, it follows that wide sclerites will be formed in summer and narrow
sclerites in winter; at the same time the controlling factor might either be tempera-
ture (Cutler, 1918) or food supply (Thompson, 1923). The most satisfactory way of
attacking the problem is by the observation of fish under controlled conditions of
temperature and food supply. For this purpose trout form a satisfactory material,
they can readily be reared in captivity and the concentric ridges on the scales are
usually well defined.
The experiments here described were designed as a direct test of the effect of
food supply on the width of the circuli of Salmo irrideus; no attempt was made to
keep the temperature constant, although of course it remained the same for each
sample of fish under observation. The work was carried out at the Midland Fishery,
Nailsworth, Gloucestershire; to the manager of the farm, Mr F. Stevens, we owe
our sincere thanks for valuable co-operation and help.
It is significant to note, at the outset, that in no case which we have examined do
the scales of rainbow trout, which are fed by hand continuously throughout the
year, show any well-defined summer or winter zones (see PI. II, figs. 1, 2). This
fact seems to eliminate the suggestion that the periodicity of circulus width found
in other members of the Salmonoid family (when under natural conditions) is due
to an inherent rhythm over which the environment has no control. It also suggests
that temperature alone is not invariably a decisive factor.
For the present experiment, fifty rainbow trout {Salmo irrideus) were taken
from the yearling class on July 5th, 1928. These fish were approximately 20 cm.
long. Twenty-five of these fish were placed in a concrete tank (16 ft. x 6 ft. x 6 ft.),
whilst the remainder were kept in a similar tank below the first. The water supply to
the two tanks was derived from the surface of a lake measuring about an acre. The
amount of water that passed through the experimental tanks was 250 gallons per
minute. The top tank represented the starved tank. The fish in this tank were not
fed, but maintained themselves on the plankton available in the water. Although
the amount of food the fish thus captured was undoubtedly far below their maximum
requirements, they showed no symptoms of unhealthiness throughout the period of
experiment. Early in December 1928, the water entering the scanty tank was made
to pass through twofinescreens, so that the amount of natural foods that passed into
the tank was still further reduced. The fish in the lower tank were fed twice daily
with as much food as the fish would eat. They were given a diet of meat (raw and
cooked) together with biscuit and fish meals. This tank was well stocked with
natural foods (shrimps and snails).
 The Growth of Fish 57
The water flowing through both experimental tanks varied in temperature
according to the season of the year, although the extremes for the monthly averages
(Table II) were not as great as those to which fish are often exposed under natural
conditions. The daily temperatures were recorded; the maximum being 630 F.
and the minimum 34° F. The monthly averages were as follows:
Table II.
1928 °F. 1929 °F.

July 56-1 January 44-0

August 48-2 February 4 3- °
September 460 March 45 4
October 50-0 April 460
November 470 May 51-0
December 446

For an examination of the scales, Winge's (1915) method was employed, the scale
being mounted, on a microscope slide, in glycerine. An eyepiece micrometer was
focussed on the centre of the scale and the number of ridges in the direction of the
anterior longitudinal radius read off, together with the distances apart of successive
ridges. The chart shows the total number of ridges and the variations in spacing of
the ridges in the anterior quadrant. The scales for examination were removed
in each case from the same region (shoulder), and the extent to which they were
magnified was the same throughout. The fish were placed in the tanks on July 5th,
1928; the first scales were taken on September 18th, 1928. Scales were removed
from fish selected at random from each tank. Charts were prepared and represen-
tative scales were preserved in 10 per cent, formalin. Material of this type was
collected at monthly intervals, care being taken to avoid undue disturbance to the
other fish in the tanks. Individuals used for scale examination were not replaced in
the tanks.
In order that an experiment of this type should yield useful results it is essential
that growth should occur at a measurable rate in each tank. If the amount of food
present in the scanty tank falls below a critical level, the fish will not grow since the
available food is all required for maintenance: on the other hand, the food supply
must be definitely lower than in the tank in which food is abundant. However
carefully the conditions are controlled, there is always a marked variation in the
ability of individual fish to grow on artificial or natural diets, and this variation is
accentuated when food is scarce. The twenty-five fish placed in the abundant tank
increased in length from approximately 20 cm. in July 1928 to 30 cm. in November
1928, and to 33 cm. in March 1929. These fish under normal hatchery conditions
would have reached an average length of 29 cm., although a significant amount of
variation might be expected. It is, however, quite clear that an abundance of food
in the experimental tank resulted in a very rapid growth rate. In the scanty tank, on
the other hand, it is impossible to give even approximately comparable figures. The
fish under these conditions are best divided into three categories: (i) those which
grew comparatively steadily and attained in February 1929 a length of 26-5 cm.;
58 J. GRAY and S. B. SETNA
(ii) fish which grew very much more slowly; (iii) fish which showed little or no
increase in size, particularly during the latter months of the experiment. Marked
differences of this type might well be expected, and probably indicate a variation in
the ability of the fish to obtain the food available.
An examination of selected scales revealed quite definite data. PI. II, fig. i, shows
a scale typical of thefishas placed in the tanks in July 1928; PI. II,fig.2, shows a scale
from a fish reared under hatchery conditions precisely similar to those of fig. 1, but
killed on January 15th, 1929. These two scales may be regarded as typical of fish
reared under standard conditions of feeding. It will be noted that, although the
latter fish was two years old, the scale exhibits no well-marked seasonal variation in
the spacing of its ridges; the spacing tends to increase towards the periphery of the
scale, in spite of the fact that these rings were formed during the winter months.
The contrast between these scales and some of those obtained from fish fed with
abundant food is quite definite. A definite percentage of scales removed from the
abundantly fed fish at approximately monthly intervals from September 1928 to
March 1929 show that the peripheral rings are markedly wider apart than those
nearer to the centre of the scale, and are clearly wider than those characteristic of
standard hatchery conditions. PI. Ill,figs.6,7,8, and Text-fig. 1 are typical records of
such scales from the hand-fed fish. It is, of course, difficult to determine precisely
those ridges which were laid down after the period of abundant feeding began, but the
change from normal width to wider widths is usually fairly abrupt and can be safely
assumed to represent the period of rapid growth under the experimental conditions.

Table III. Distribution of circulus width for peripheral circuli.

Number of circuli measured on each scale: 20
width Abundance of food Scarcity of food
in n Total Total
A B c D E F A B1 C, Fi

5° 2 5 1 8
45 1 3 1 5 3 13 .
40 3 8 b b 5 9 37 1 1 2 1 5
35 5 1 1 3 2 1 13 4 1 2 5 2 2 16
3° 2 4 4 3 3 20 7 10 7 4 7 1 36
25
20
I
2
b
2
2
4
4 1
2
'9
10
5
1
4
4
3
5
2
5
5
4
3
8
22
2
15 . 0 1 1 2 4 l8
10 0 2 2 1 1 6
Total 20 20 20 20 20 20 120 20 20 20 20 20 20 120

A.M.== 35 ±8 A.M. ==26 ±6

Taking the outer twenty ridges on each selected scale as revealed by the Winge
charts, the variation can be expressed by the graph in Text-fig. 2. It will be noted
that, although the total number of fish observed was small, the effect of the food
supply upon the spacing of the ridges is sufficiently clearly marked to leave little
or no doubt of the conclusion to be drawn, or of the fact that wide rings can be laid
down in winter months. In order to compare these results with those obtained
 The Growth of Fish 59
i i i I i i i i

D WV
\

11111111 111111111 111111111 111111111 111111 11 ir

Text-fig, i. Curves of scales showing the successive widths of circulus from measurements of in-
dividuals kept in the "abundant" tank. Each minor subdivision of the ordinates represents 5/1;
each subdivision of the base line indicates a circulus.
6o J. GRAY and S. B. SETNA
from the "scanty" tank it is necessary to select from the latter each of the specific
types mentioned on p. 58. Firstly, we may consider the scales from a fish which
grew steadily on the reduced food supply. PI. Ill,fig.9, is a typical scale of such a fish
killed on February 12th, 1929, when it had attained a length of 26-5 cm. With one
exception the distance between successive ridges are all less than 40 ft, the average
width being approximately 22 /*. It will be noted that the peripheral rings, on the
right side of the scale, are markedly narrower than the average. The scales of fish
(from the scanty tank) which grew more slowly exhibit the same phenomenon,
viz. the peripheral rings are narrow, and are of the order of 20 /x or less (see Text-
figs. 2 and 3 and PI. II, figs. 3, 4, 5). It will also be noted that the outer rings on the
scales of slowly growing fish are often incomplete, and are more irregular in form
than those characteristic of well-fed individuals.
401-

Scanty food U-Abundant

30 food

20

10

10 15 20 25 30 35 40 45 50 55
Width of ring in /t
Text-fig. 2. Showing the distribution of concentric rings of varying widths at the periphery of fish
fed with abundant and scanty food respectively.

Although the population concerned is not as large as might be desired, the

facts seem fairly clear, (i) Abundant food can effect an increased growth rate, and
during this period the ring width in some scales is increased, (ii) The formation of
wide rings can occur during the winter, (iii) Scarcity of food entails a reduction of
ring width. It is also noticeable that several of the fish in the abundant tank spawned
during the winter, but the fact is not recorded by any disturbance of ring formation.
The results of the present experiments obviously differ from those obtained by
Cutler (1918), but are in closer harmony with those of Thompson (1923), Dannevig
(1925) and Graham (1929), all of whom recorded a correlation between growth rate
and sclerite width. It should not be inferred, however, that temperature is without
its effect. Hathaway (1927) has shown that the ability of fish to feed at low tempera-
 The Growth of Fish 61
"ures is markedly less than at higher temperatures; in other words, although food
may be present in abundance, the amount actually absorbed may be greatly reduced
at a low temperature, and in this way the growth rate will be affected. In addition,
temperature may affect the growth rate in another way. The food which is actually
J I I i I I I II I I I I I I I I I I I I I I I I I I MIL
r r r TTT i i i

i i i
Text-fig. 3. Curves of scales showing the successive widths of the concentric rings from measure-
ments of individuals kept in the " starved " tank. Each subdivision of the ordinates represents 5/x.

absorbed by the fish is utilised in two ways, firstly for the maintenance of existing
tissues, and secondly for the formation of new tissue. If a fish absorbs a unit quantity
of food (a) at a low temperature, we may suppose that a definite fraction (xj) is
required for maintenance, leaving the remainder (a — xx) for the production of new
tissue. If the temperature be raised, the rate of metabolism increases so that Xi is
62 J. GRAY and S. B. SETNA
increased—to x2—leaving a smaller proportion (a — x2) for the purposes of growth.
As shown by Gray (1928) for the embryonic cycle of the trout, the effect of high
temperatures is to produce larger organisms for each standard diet, although the
time required for their growth is increased. As far as one can see, under natural
conditions, the growth of post-embryonic trout will be influenced by temperature
in several ways: (i) a higher temperature may induce a more rapid formation of
nutritive organisms—and hence increase the quantity of food available; (ii) it will
increase the capacity of the fish to capture the food; (iii) it will increase the rate at
which the food is converted into new tissue; (iv) it will decrease the proportion of
acquired food which is available for growth. The net result will depend on the
equilibrium which exists between these various effects.

SUMMARY.
1. Salmo irrideus fed continuously throughout the year exhibit on their scales
no seasonal periodicity in the distance apart of their concentric ridges or rings.
2. Similar fish fed with abundant food form on a proportion of their scales
abnormally wide rings even during the winter months.
3. Fish fed with limited diet develop abnormally narrow rings.
4. The width of the rings is probably ciosely associated with growth rate.

BIBLIOGRAPHY.
CUTLER, D. W. (1918). Journ. Mar. Biol. Assoc. 2, 470.
DANNEVIG, A. (1925). Rep. Norwegian Fish. Invest. 3, 6.
GRAHAM, M. (1929). Fishery Investigations, Ser. 2, 2.
GRAY, J. (1928). Brit. Journ. Exp. Biol. 6, 125.
HATHAWAY, E. S. (1927). Ecology, 8.
THOMPSON, H. (1923). Fisheries, Scotland, Set. Invest. 5.
WINGE (1915). Medd. Komm. Havunsdersegelser Fisk. 4, 8.

EXPLANATION O F P L A T E S .
PLATE II.
FIG. 1. Scale characteristic of a fish placed in the experimental tanks in July 1928. Length of fish,
20 cm.
FIG. 2. Scale of a fish of the same age as experimentalfishbut fed under standard hatchery conditions.
Fish killed on January 15th, 1929. Length of fish, 29 cm.
FIG. 3. Scale of a fish removed from scanty food conditions on September 18th, 1928. Length of
fish, 23 cm.
FIG. 4. Scale of a fish removed from scanty food conditions on December 15th, 1928. Length
offish, 25-8 cm.
FIG. 5- Similar to the above, but killed on February 12th, 1929. Length of fish, 202 cm.
PLATE III.
FIG. 6. Scale of a fish fed on abundant food from July 1928 until September 18th, 1928. Length of
fish, 30 cm.
FIG. 7. Similar to Fig. 6, but killed on November 17th, 1928. Length of fish, 325 cm.
FIG. 8. Similar to Figs. 6 and 7, but killed on March 12th, 1929. Length of fish, 32-5 cm.
FIG. 9. Scale of a fish which grew steadily on scanty diet from July 1928 to February 12th, 1929.
Length of fish, 26-5 cm.
JOURNAL OF EXPERIMENTAL BIOLOGY. VOL. VIII, PLATE II.

Fig. i. Fig. 2.

Fig- 3-

Fig- 5-

J. GRAY AND S. B. SETNA—THE GROWTH OF FISH (pp.55-62).

JOURNAL OF EXPERIMENTAL BIOLOGY. VOL. VIII, PLATE III.

Fig. 6. Fig. 7.

Fig. 9.

J. GRAY AND S. B. SETNA.—THE GROWTH OF FISH (pp. 55-62).