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The Length-Weight Relationship and Seasonal Cycle in Gonad Weight and Condition in the

Perch (Perca fluviatilis)


Author(s): E. D. Le Cren
Reviewed work(s):
Source: Journal of Animal Ecology, Vol. 20, No. 2 (Nov., 1951), pp. 201-219
Published by: British Ecological Society
Stable URL: http://www.jstor.org/stable/1540 .
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[ 201 ]

THE LENGTH-WEIGHT RELATIONSHIP AND SEASONAL CYCLE IN


GONAD WEIGHT AND CONDITION IN THE PERCH
(PERCA FL UVIA TILIS)

BYE. D. LE CREN
FreshwaterBiologicalAssociation,Ambleside,Westmorland

(With 8 Figures in the Text)

CONTENTS
PAGE PAGE
I. INTRODUCTION . . . . . . 20I 4. THE SEASONAL CYCLE IN GONAD WEIGHT AND
CONDITION
2. THE ANALYSIS OF LENGTH-WEIGHT RELATION- (a) Gonad weight . . . . . 209
SHIP AND CONDITION . . . . . 20I (b) The weight of stomach contents . . 2I2
(c) Condition . . . . . . 212
3. THE LENGTH-WEIGHT RELATIONSHIP (d) Individual variations in relative con-
(a) Sources of material and methods of dition and gonad weight . . . 2I3
collection . . . . . . 205 (e) Discussion of the- seasonal cycle . . 2i6
(b) Analysis of data . . . . . 205 5. ACKNOWLEDGEMENTS . . . . . 2I8
(c) Results . . . . . . 207 6. SUMMARY . . . . . . . 2I8
(d) Discussion and conclusions. . . 209 REFERENCES . . . . . . . 2I8

1. INTRODUCTION A brief account is then given of the rather scanty


The present paper is an account of some of the data available on the weight of stomach contents.
investigations on the biology of the perch (Perca The seasonal changes in condition are then de-
fluviatilis Linn.) in Windermere, which are being scribed and, finally, some of the results are sum-
conducted in connexion with a trap-fishery experi- marized, combined and discussed as a picture of
ment (Worthington, 1950). This experiment is the seasonal cycle in the Windermere perch.
mainly a study of populations, but it has been In the statistical analysis of the length-weight
necessary simultaneously to investigate the general relationship the data for only one group of fish are
biology of the perch, particularly the growth and given in full (Tables i and 2) as an example of the
related aspects. The computation of a formula to method of computation used for all the groups.
express the length,weight relationship and provide Again, in the section on seasonal changes in gonad
a means of interconverting measurements of length weight and condition Figs. 2-7 are based partly on
and weight, revealed the relative complexity of the tables of data which are not published. The full
interrelationships of length, weight and condition. tables have been deposited with the Freshwater
Condition in turn was found to be correlated with Biological Association, from whom copies can be
the seasonal changes in gonad development and obtained.
growth, and the importance of the effect of stomach
contents on weight had also to be assessed. It was
decided, therefore, to combine these separate but 2. THE ANALYSIS OF LENGTH-WEIGHT
interrelated aspects in one paper. RELATIONSHIP AND CONDITION
The main part of the paper is devoted to the Data on the lengths and weights of fish have com-
questions of length-weight relationship and con- monly been analysed to yield biological information.
dition. A brief review of the fundamental bases for One or other form of such analysis has, in fact,
the concepts of length-weight relationship and become one of the standard methods employed in
condition and of some of the methods of analysis fishery biology. Often, however, the examination
of length-weight data precedes an account of the of length-weight data has become so stereotyped
application of the chosen methods to the present that confused thinking on its aims, the methods
material and its results. This is followed by an employed and the results obtained has resulted. It
account of seasonal changes in gonad weights. is not proposed to enter here into a discussion of the
202 Conditionin theperch
literature, but only to present an outline of the n times the increment in log length for the same
principal methods that can be used in the analysis period of time. Thus the length-weight relationship
of length-weight data before describing the applica- formula (i), besides providing a means for calcu-
tion of some of these methods in the present lating weight from length, and a direct way of
work. converting logarithmic growth rates calculated on
The analysis of length-weight data has usually lengths into growth-rates for weight, may also give
been directed towards two rather different objects. indications of taxonomic differences and events in
First, towards describing mathematically the re- the life history such as metamorphosis and the onset
lationship between length and weight, primarily so of maturity.
that one may be converted into the other. Secondly, The length-weight relationship may be expressed
to measure the variation from the expected weight graphically by plotting the observed lengths and
for length of individual fish or relevant groups of weights as a dot diagram on double logarithmic
individuals as indications of fatness, general 'well- graph paper. The points for fish having the same
being', gonad development, etc. Throughout this length-weight relationship will lie on a straight line
discussion the term length-weight relationship is with some scatter due to individual variation. This
applied rigorously to the first category, while the line represents the logarithmic form of equation (i)
tern condition is applied as a rigorous but general
log W=log a+n log L, (2)
term for length-weight analyses of the second
category. where n represents the slope of the line, and log a its
The length of a fish is often more rapidly and position. Changes in the value of n can usually be
accurately measured than the weight. Moreover, readily observed as changes in slope. If the scatter
back-calculations of past growth from scales, etc., is not too great, a line can be fitted by eye to each
usually yield data on length alone. Thus it is very range in length having the points on a straight line,
convenient to be able to determine a weight where and its slope measured. It is usually possible in this
length only is known, and occasionally it may be way to judge the value of n to one decimal place, an
useful to reverse this process. It has been found accuracy adequate for a preliminary investigation.
that the length-weight relationship of most fish can An accurate line can be computed from the same
adequately be described by a formula of the type: data by the regression method of least squares. Any
one line should be fitted only to that range of size
W=aLn, (I)
over which it is apparent that the fish have the same
where W=weight, L=length, a is a constant and length-weight relationship. This range, and the
n an exponent usually lying between 2-5 and 4-0 accuracy of the fits, can usually be more easily
(Hile, 1936; Martin, 1949). For an ideal fish which judged by straight line graphs on logarithmic paper
maintains the same shape, n=3, and this has than by drawing curves on arithmetic paper. It is
occasionally been observed (Allen, 1938). In the also important that the data from which the length-
vast majority of instances where length-weight weight relationship is calculated should not have
relationships have been calculated, however, it has been subjected to any selection for weight against
been found that the cube law is not obeyed and n - 3. length. For example, gill-nets may select the fatter
Further, most species of fish do change their shape as among short fish and the thinner among long fish,
they grow (e.g. Martin, I949) and so a cube relation- and thus lower the value for n, even though the
ship between length and weight would hardly be ex- means of length and weight may be unaffected
pected. It is, therefore, more logical as a general (Kipling & Le Cren, unpublished data).
basis for investigation to assume that probably Individual variations from the general length-
n ?3. It has also been found that while n may be weight relationship have usually been considered
different for fish from different localities, of more interesting than the length-weight relationship
different sexes, or for larval, immature and mature itself, and have been frequently studied under the
fish (different 'growth stanzas') it is often constant general name of 'condition'. In some cases the
for fish similar in these respects. The length-weight specific gravity of the fish may not be unity, and
relationship may thus be a character for the dif- variations in the specific gravity of the flesh of fish
ferentiation of small taxonomic units, like any other have been shown to occur (Tester, 1940) and their
morphometric relationship. It may also change importance in studies on condition has been dis-
with metamorphosis or the onset of maturity, as cussed by Kesteven (I947). Usually, however, in
has been shown for other relative growth ratios all but completely demersal fishes the density of the
(Frost, 1945; Huxley, 1932). Further, the exponent fish as a whole is maintained the same as that of the
n is the ratio of the logarithmic growth-rates for surrounding water by the swim bladder, and there-
length and weight, the increment in log weight for fore changes in weight for length are due to changes
any period of time (of reasonable length) being in form or volume and not specific gravity.
E. D. LE CREN 203
Such changes in condition have usually been First, there are those factors correlated with
analysed by means of a condition factor (or 'coef- length which affect the condition factor because the
ficient of condition', 'ponderal index', etc. (Thomp- fish does not in fact obey the cube law in its length-
son, I942; Hile, I936). This is calculated as a ratio weight relationship. The condition factor will vary
between the observed weight and that expected from with length itself according to the expression
the observed length. The basis of the expected
weight is that for an ideal fish in whose length-
KocL n-3. (9)
weight relationship formula (i), W= aLn, n= 3, and Thus length itself and any correlated factor will
thus obeys the cube law. Various types of condition affect the values of K. This means that, except in the
factor have been used, but in one of the original rare instances where n =3, the condition factors of
ones the condition was measured by the constant c fish of different lengths cannot be directly attributed
(equivalent to a in (I)): to features other than length. Further, factors such
as age, sex or maturity which may affect the value
W=cL3, (3)
of n, may in turn affect the values of K. Differences
therefore
w in mean K for fish from different environments may
c= L3. (4) be due to the fish exhibiting racial differences in
form, which will affect n and through it, K. Thus
As, however, c when so calculated is often an awk- differences in K attributed to environmental factors
ward decimal number, the average value of c found may in fact be genotypic.
by trial from formula (3) was incorporated into the Secondly, the values of K may be affected by
formula, and a new condition factor K found that selection in sampling. The effect of gill-nets on the
would vary about unity computation of length-weight relationships has
K= W (5) already been mentioned, they may also be selective
cL3' for condition factor (Farran, T936; Deason &
for example Hile, 1947).
K w Thirdly, there are those features usually associated
-O00427L3 (Menzies, 1920). (6) with K. General, long-term, features such as
environment, food supply and degree of parasitiza-
The value of c depends partly on the units used for tion may affect the fish's condition directly, or
weighing and measuring the fish; in formula (6) where K is correlated with length, via the growth-
these are pounds and inches. In instances where the rate and average size. Seasonal changes have
original value of c chosen was found to have only frequently been studied with the aid of condition
limited application, and K was found to average factors, which have been shown to be correlated
about some value approximating to, but not exactly with gonad cycles, rate of feeding, etc. Short-term
I, c has been further altered to a convenient round
cycles of alternating growth in weight and growth
number and often changed into its reciprocal in length have also been revealed by the use of
condition factors (Brown, 1946).
K L3W In view of this list of widely different factors that
for example can affect the condition factor, it is not surprising
that the interpretation of K is difficult and often
K= L30W f le, 1936) (8) leads to erroneous results. As an example a hypo-
thetical instance will be discussed. The population
(where L is in centitnetres and W in grams). In of a fish species is being studied for comparison in
this formula, which is widely used, c has ceased to two lakes. It is only possible to fish each lake on one
be equivalent to a in formula (i-) and no attempt is occasion, and gill-nets are used. Lake A is fished in
made to make K= I on average. All these formulae June and most of the fish are caught in a gill-net of
are based on comparison with an ideal fish, whose 2 in. mesh, and average 20 cm. in length. Lake B is
WocL3, and in subsequent discussion the term fished in August and most of the fish, which average
condition factor and its symbol K are applied only about 30 cm. are caught in a gill-net of 3 in. mesh.
to measurements of condition so derived. Condition factors are calculated for each fish
Differences in condition factor have been inter- (according to formula (5)) and averaged for each
preted as measuring various biological features such lake. For lake A, K=o-95, for lake B, K= I-03. It
as, fatness, suitability of environment or gonad is then concluded that lake B is a more suitable
development. The number of variables that can environment for the particular species. Actually the
affect the value of K is however considerable, and higher average condition factor in lake B could just
some of them will now be briefly discussed. They as easily be due to: (i) slight racial differences in
fall into three main groups. shape between the fish of the two lakes, (2) the
204 Conditionin theperch
greater mean length (due to growth-rate or mor- empirical formula (of type (i)) fails to measure any
tality-rate), (3) the fact that in August the fish are in change in form associated with change in length.
better condition after a summer of feeding than soon This is true, but the change in form or condition
after spawning in June, and (4) selection by the associated with length is succinctly and accurately
gill-nets. described by the value of exponent n, and, as Hile
In this hypothetical case further information on points out, the relationship between condition as
morphometric measurements, average size, growth- measured by K and length is given by the expression
rate, time of spawning, etc., might have ruled out
Koc L 3. (9)
many of the possible causes for the difference in K,
but probably would not have made it possible to With the relative condition factor, therefore, it is
attribute this difference to any one cause. It is thus possible to distinguish between and measure
clear that in using the condition factor great care separately the influences on condition of length
must be exercised to decide for exactly what purpose and other factors; whereas these are not readily
the information on condition is required, and separated when the ordinary condition factor is
whether the condition factor will yield this informa- used.
tion and give results undisturbed by the effect of For a more accurate analysis of condition, where
variables other than those being studied. Where the data are adequate, use may be made of the
other variables affecting K can be controlled or length-weight relationship formula itself. It has
eliminated, or where a biological feature can be usually been considered that the empirical length-
shown to be highly correlated directly with K, the weight relationship formula of type (i) can provide
use of the condition factor may be valuable. There little information on condition (Hile, I936). Unless
are, however, alternative methods of analysing the value of n in equation (i) is identical for the
condition which may be more suitable. groups of fish whose condition is being compared,
It is relatively easy to eliminate the effect on K the value of a in this equation gives no measure of
of length and correlated factors by calculating their relative condition, and is not comparable to c
a 'condition factor' based not on the 'ideal' length- in equation (4). Usually, as Hile (I936) points out,
weight relationship there is a negative correlation between the values of
W= cL3, (3) a and n. Where, however, it can be shown that n is
the same for two groups of fish, the values of a
but on an empirical, calculated length-weight obtained in separate length-weight relationship
relationship formulae calculated for each group will be a direct
W=aLn. (I) measure of their condition relative to each other.
The 'condition factor', in this case called the If it is expected that a large group of fish, con-
relative conditionfactor and designated by Kn to dis- taining a number of smaller groups the condition of
which latter it is desired to compare, is suspected of
tinguish it from the condition factor K based on the
being homogeneous in its length-weight relationship
cube-law, is calculated from the formula
(a logarithmic graph of the length-weight data
W
K aL
showing for instance one straight, but rather broad
Kn (Io) line of dots) then a length-weight relationship
formula (2) can be calculated for each subgroup.
In practice the length-weight relationship would The values of n determined are tested for homo-
first be calculated as the logarithmic formula (2), geneity, and if as was expected there is no significant
and smoothed mean weights, WV,for each length- difference between them, a pooled regression can
group computed from this log formula or read off be calculated for the whole group combined and the
an accurate graph. The relative condition factors values of a adjusted for this pooled length-weight
would then be calculated from the formula relationship. These adjusted values of a are accurate
w measures of the relative condition of the subgroups,
Kn= . (II) and the significance of the differences between them
can be subjected to accurate statistical test (e.g.
The difference between Kn and K is that the former Snedecor, I946; Goulden, I939; Mather, I943).
is measuring the deviation of an individual from the The computational labour involved in such an
average weight for length, while the latter is analysis of covariance is fairly large, but it is probably
measuring the deviation from a hypothetical ideal not much greater than that needed to calculate by
fish. The choice of which condition factor to use formulae (7) or (8) a K for each fish and then find
nlust be based to some extent on which of these two the mean value of K for each subgroup. Further
comparisons is the more relevant. Hile (I936) analysis, within the subgroups, can be carried out
argues that a condition factor calculated from an by means of the relative condition factor, calculated
E. D. LE CREN 205
from the pooled value of n and the value of a for relative to length have been included in the calcula-
each subgroup. tions of the regressions of weight on length.
The use of both the relative condition factor and Length was measured from the tip of the pre-
the analysis of covariance of the length-weight maxilla to the tip of the longest caudal fin ray
relationship as methods of directly comparing stretched out posteriorly. Length measurements
condition is confined to comparisons between fish were usually made to the millimetre below, but
which are homogeneous for n in their length-weight occasionally to the 2 cm. below. Age was determined
relationship formulae. Comparisons between fish from the opercular bone or from the length fre-
which have different length-weight relationships quency distribution for fish in their first two years
will normally have little relevance. If they differ in (Le Cren, I947). Age has been designated by the
size, differences between relative or ordinary con- number of completed years of life, but with the
dition factors will be difficult to interpret; if they birthday moved to I January. Thus a fish hatched in
are of the same length the weights themselves can June I945 will belong to the 0 group till i January
be compared. Comparisons between such groups I946, then to the I group till I January I947, and
of fish can perhaps best be expressed by graphs of so on. The sex and state of gonad was determined
lines on logarithmic scales representing their by internal macroscopic examination, for all but
respective length-weight relationships. running fish. This was usually easy for fish in their
To sum up this review, it may be concluded that second year and older, though in early summer it
the expression of length-weight relationship and the was sometimes difficult to distinguish between
measurement of changes in condition are two rather immature and recovered spent fish.
different but interconnected aims in the analysis of As the weights of the perch varied from I mg. to
length-weight data. The length-weight relationship I450 g., several balances were used in an attempt to
can best be expressed by a formula of the type weigh each fish to approximately the same relative
W=aLn (i), and is most conveniently calculated accuracy. The smallest fish were weighed in batches
and graphed in its logarithmic form (2). The of ten or more on an analytical balance to the nearest
condition factor K, where K= cW/L3 (7) being milligram. It was found impossible to obtain a wet
based on the cube-law, which rarely holds, is affected weight for these larval fish exactly comparable with
by length as well as many other factors. This makes the wet weight for larger fish, owing to the relatively
its interpretation difficult. The effect of length may large weight of any surface water, and the speed with
be eliminated by using a relative condition factor which this water and then the fish themselves dried
based on an empirical length-weight relationship. up. In practice surface water was removed with
The analysis of covariance in the log length-weight blotting-paper and the fish then weighed before
relationship may also be used for abundant data, they had time to dry up. Fish weighing between
where there is homogeneity of the exponent n in 0o and io g. were weighed on an ordinary chemical
the length-weight relationship. In any analysis balance to o-oi g.; fish between Io and 2oo g. were
exact ideas as to the aims are required, and the weighed on a 'Butchart' swinging arm balance to
method of analysis should be chosen so as to yield the nearest gram and for the largest fish a pan
the maximum unequivocal biological information. balance accurate to approximately I g. was used. In
all cases the fish were weighed intact with gonads
and stomachs, and damp with surface moisture, but
3. THE LENGTH-WEIGHT wiped clean of any adhering dirt.
RELATIONSHIP (b) Analysis of data
(a) Sources of material and methods of collection When a fairly large number of weighings from
The data used for length-weight analysis were different sizes of fish and different seasons had been
obtained from fish collected primarily for other collected, they were divided into a series of groups
purposes. These fish were collected in the years according to age, sex and maturity of the fish, and
I943-8, and were all caught in the north basin of the time of year, and were then plotted as a series
Windermere. They were collected mostly in traps of' dot diagrams' on double logarithmic graph paper.
(Worthington, I950) and seines, but also by angling These graphs revealed that the fish could be
and a few of the very large fish were caught in gill- divided up into a series of sex, age and maturity
nets. These latter are the only gill-netted fish that groups, in each of which the length-weight relation-
have been included in the determinations of length- ship could then be described by a formula of type
weight relationship, though some fish gill-netted in
log W= log a+ n log L. (2)
I948 have been included in the seasonal samples
used for determining relative condition factors. No Further, it seemed probable that the constant n in
samples likely to have been selected for weight the formula differed from one age, sex and maturity
2o6 Conditionin theperch
group to another, but that it was the same for fish coefficient was calculated from the residual sums of
of the same group caught at different times of the squares and used to calculate 95 % confidence limits
year. The constant a, however, varied with the and in a 't' test of the significance of the difference
season of capture. of the regression coefficient from 3. The residual
It was then decided to analyse the data with sums of squares for the 'within subgroups' regres-
accurate statistical methods and the fish were there- sion were used in a test of the homogeneity of the
fore classified first into groups according to age, subgroup regression coefficients. The regression
sex, maturity, and then into subgroups according for the means of the subgroups does not have much
to the time of year. The list of groups and seasonal value in this instance as the mean lengths were
subgroups is as follows: affected by arbitrary factors. The residual sums of
(i) Larvae Early summer
(2) 0 and I August, September, spring, July, August, September, October
(3) Female immature II April, May, June, July, August
(4) Female immature III and older April, May, June, late summer
(5) Female mature Spring, April and May (ripe), spent, late June, July, August, September,
autumn
(6) Male mature Spring, April (ripe), spent, late June, July, August, September, autumn

For each of these subgroups a regression was squares for the means regression has, however, been
calculated for the logarithm of weight on the used as a test of the significance of the difference
logarithm of length, by the method of least squares. between the subgroup means, when they have been
The lengths were grouped in 2 mm. groups for the adjusted for the general regression. This test is
smaller fish and 5 mm. groups for the larger, while therefore a test of the significance of subgroup
the weights were grouped in equal-sized logarithmic differences in weight corresponding to the grand
groups, varying from o-o6 to 144 g. in size. Fre- mean length, or in fact a test of seasonal variations
quency distributions of lengths, and even more of in relative condition. Two further tests are possible
weights, from fish tend to the Galton-MacAlister relative to the means regression. One tests the
distribution rather than the normal, although the significance of the difference between the pooled
lengths are usually very near the normal. Thus the regression coefficient and the means regression
logarithms of the weights are approximately coefficient, and the other tests the significance of the
normally distributed. deviation of the subgroup means from their own
When this series of regressions had been cal- regression. As the means of the subgroups are
culated, it was clear that in general the results affected by arbitrary selection of the length of fish
agreed with the approximations obtained graphically. caught in the samples from different times of the
The data were then analysed further for covariance year these tests have little meaning in this analysis
(Snedecor, I946, pp. 3I8-29; Goulden, I939, of length-weight relationship. A summary of the
pp. 253-9; Mather, 1943, pp. 119-28). The regres- tests of significance for the female fish is set out in
sions for each group of subgroups were tabulated Table 2, as an example of the tests carried out for
with their sums of squares and regressions; and then each group. In each case Bartlett's test of homo-
the sums of squares were calculated for: (i) the scedasticity was also carried out to verify the
' pooled'. regression within subgroups, (2) the means validity of the 'F' tests.
of subgroups, and (3) the total for the whole group. When the analysis of covariance had been com-
As an example of these calculations the skeleton data pleted for each group, the values of the adjusted
for the mature female group are shown in Table i. means for each subgroup were calculated. This
The series of data obtained for each group by this gave for fish of the grand mean length the smoothed
analysis of covariance provided a set of statistical mean weight at each season of the year and was
tests of significance. thus an accurate measure of seasonal change in
From the residual sums of squares for each sub- relative condition. The significance of these dif-
group were calculated the standard error and 95 % ferences between the adjusted means had already
confidence limits for the regression coefficient for been tested as described above.
each subgroup. The 'pooled' or 'within subgroups' Further, accurate graphs were drawn on double
regression is the best estimate of the regression logarithmic paper for each group of fish using the
coefficient for the whole group, as unlike the 'total' regression coefficient b for 'within subgroups' and
regression (given in the last line of Table i), it is not the adjusted value of a for the subgroup with the
affected by the arbitrary differences in mean length maximum value of a. From these graphs smoothed
between the fish in the different subgroups. The mean weights (1') could be read off for any length.
standard error of the 'within subgroups' regression In certain cases these smoothed mean weights were
E. D. LE CREN 207

Table i. Data for regressionsof log weight (y) on log length (x) for maturefemales, showing method of
computationand analysis of sums of squares(s.s.) and degrees of freedom
Regression Degrees
s. products coefficient s.s. due to Residual of
Subgroup s.s. x s.s. y xy b regression s.s. freedom
Spring Io-6099 i2-62806 3 64674 3 437I I I253425 0-0938i 93
April-May I-o8443 13-40003 3 7I86I 3 42909 I2-75145 o-64858 392
Spent I-26846 15-39107 4-33807 3 4I995 14-83598 055509 257
Late June 0-37000 4-31748 IP24122 3135465 41i6386 OI5362 II9
July 0-20429 2-43482 o-69505 3-40227 2-36475 0?07007 51
August o040923 4-67813 P335927 3 32153 4-5I486 o0i6327 122
September 0o4I593 4-66059 IP37450 3 30464 4 54223 O-II836 I04
Autumn 2-87077 33-59050 9 74773 3-39553 33-09871 0-49I79 358
Total 2'29459 1496

Total within 7-684Io 9I'I0068 26zI2II9 3 39938 88&79585 2-30483 1503


subgroups Difference O-OI024 7

Total between means 1-10438 9-46914 30o8322 2-79I8I 8&60776 o-86138 6


of subgroups Total 31i662I 1509

Total 8-78848 I00-56982 29-20441 3 32303 97'04713 3'52269 I510


Difference 035648

Table 2. Analysis of variance for data in Table i


Sums of Degrees of
Source squares freedom Variance F P
Due to total regression 97-047I3 I 97-04713 63,000 < OOI
Between regression coef- 001024 7 0-00147 0-95 > 0-05
ficients within subgroups
Difference between 0-35648 I 0-35648 233 < OOI
'pooled within subgroups'
and 'means' regressions
Deviation of means from o086138 6 0-14356 94 < 00I
means regression
Residual 2-29459 I496 OOOI53
Total I00-56982 I5II

used to calculate relative condition factors, (K.) by weights of fish from hatching till they were 3 0 cm.
formula (i i).. Mean relative condition factors were long. The regression formula found was
then calculated for small samples of fish caught at
log W=o-68969+3 59i63 log L.
different dates especially in the 'spring' and
(autumn' when the adjusted mean weights for the The length of newly hatched fry was found to
subgroups for the whole season were inadequate to average 6 mm., and although accurate weighing was
trace detailed changes in condition. very difficult, several batches gave an average
weight of i mg. per fish. The regression calculated
(c) Results for the larvae, because of the difficulty of weighing
The results of the analysis of the length-weight and the bulk weighing of the fish is not based on
data will be presented group by group followed by data quite comparable to the regressions calculated
a summary of the combined data. for older fish. It will be seen, however, from Table 3
Larvae. The graphical plotting of the data re- that the regression coefficient of 3-59i63 for the
vealed that the few points for weights of larval larvae is significantly greater than that obtained for
perch lay off the line taken by the young fish of any other group, and is significantly different from
between 3 and I4 cm. in length and formed a fairly 3.o0
distinct larval length-weight regression. A regres- O and I. Between the time they reached a length
sion was therefore calculated for all the lengths and of 3-0 cm. at an age of about z months, till the end
208 Conditionin theperch
of their second growing season, the perch seemed to larger than those for the three later months, but the
form a homogeneous group with their weight varying test of homogeneity showed that this difference is
with approximately the cube of the length. Regres- not significant. Bartlett's test showed, however,
sions were calculated for 0 group fish in August and that the II group females were heteroscedastic,
September, and I group fish in the spring, July, there being a significant difference in the variance
August, September and October. These regression of the subgroups. The cause of this remains obscure,
coefficients varied from 2-5983I to 31i8333 though but it means that the tests of significance based on
most of them were very near 3-0, and the differences variance ratios may be erroneous. The 2-year-old
between them were not significant. The pooled females may include fish which will remain im-
regression coefficient was 3-O I7, and it was not mature, and which thus may make the group

Table 3. Summary of regressionsof log weight on log length for each group of fish, and the combinedtotal
Significant
Homogeneity b signi- difference
of ficantly between
No. Range in Range in Regression regression different adjusted
Group of fish length (cm.) weight (g.) slope, b within group from 3 means
0 and I 667 3-0-I4-o 0o22-28 3.0I2 Yes No Yes
V immature II 462 9'5-15-5 7-40 3-I95 Yes Yes Yes
V immature III I3 II0-oI85 I I-70 3?053 Yes No Yes
and older
S?mature I512 II'5-43'5 II-1250 3-399 Yes Yes Yes
d mature 640 I0-0-44-5 7-I450 3'28I No Yes Yes
(due to August)
Total 34I2 3'0-44 5 0'22-I450 3-i65 No Yes Yes
Larvae 323 o-65-2-95 O-OOI-0-25 3'592 Yes
d immature 35 8'5-26-5 5-I80 3'243 Yes

Table 4. Average dates of collection, numbersof fish and adjusted mean weightsfor
seasonal subgroupsof 0 and I fish
Percentage
Average No. of Adjusted of maximum
Subgroup date fish mean weight weight
0, Aug. I i Aug. 85 3-536 97.1
0, Sept. 5 Sept. 9I 3-642 I00*0
I, Spring i8 Apr. 209 3.o84 84.7
I, July 2 I July io6 3-468 95*2
I, Aug. i i Aug. 29 3-538 97.,
I, Sept. ii Sept. I35 3 493 95 9
1, Oct. 26 Oct. I2 3 327 9I 4

Total - 667 3 379 92-8

significantly different from 30o. The adjusted sub- heterogeneous. The adjusted means were very sig-
group means differed significantly from each other, nificantly different from each other (Table 5).
thus showing that there were significant seasonal After August, when they began to develop their
changes in relative condition. The adjusted means ovaries in preparation for their first spawning the
and mean dates of collection for each subgroup are next spring, the maturing II females were classed
set out in Table 4. with the mature females.
Immature females. At first all immature female Some older females were found to remain im-
fish were grouped together, but the trial plotting mature. Some of these may be fish which mature
indicated that most of the 2-year-old females late, others may never mature. However, they were
probably had a distinct length-weight relationship treated as a separate group. The pooled regression
from those that were older. The immature females coefficient is 305292, and this is not significantly
were thus divided into two groups by their age. different from either 3-0 or the regression coefficient
The I + and II + females had a pooled regression of 3'I9506 obtained for 2-year-old immature fe-
coefficient of 3-I9506. The regression coefficients males, but it is significantly different from that for
for the subgroups April and May were considerably mature females. There was no significant difference
E. D. LE CREN 209
between the subgroup regression coefficients, but the ordinary condition factor. It might have been
the adjusted means varied significantly (Table 6). possible to split up the groups still further and carry
Mature females. This was the largest group, and out a more detailed analysis of covariance, but it is
details of the analysis of its length-weight relation- probable that the data that were available would not
ship have already been discussed as illustrating the have warranted the extra computational labour
method of analysis, and set out in Tables i and 2. involved.
The pooled regression coefficient was 3-39938; this The length-weight regressions for the various age,
is significantly different from 30o and from all the sex and maturity groups are shown as lines on
other groups. There was some difference between a logarithmic graph in Fig. i, and the successive
subgroup regression coefficients, they ranged from changes in slope will be noticed in this figure. The
3-30464 to 3-437Ii but these differences were not regression coefficient for the older immature females
significant. The whole group shows significant is very similar to, and not significantly different
heteroscedasticity with Bartlett's test, but when from, that for the young immature fish of both
the 'spent' subgroup is omitted the result loses its sexes. This suggests that the change from this
significance. The spent fish were rather variable relationship to that of the mature fish is correlated
owing to the inclusion of many fish that were part with maturation rather than age. The older imma-
recovered and some only just spent. The adjusted ture females may retain the length-weight relation-
mean weights show considerable and significant ship of the young fish because their gonads have
variation (Table 7). not matured.
Mature males. The pooled regression coefficient A detailed comparison of the length-weight
is 3128063 which is significantly different from 3-0 relationship of perch from different lakes is not
and from the other groups except for the 2-year-old possible as different authors have used different
females. There was no great variation among the methods of measurement and analysis. Perch, on
subgroup regression coefficients except for August the whole, would appear however to have a length-
which was 2-95847 and the spring which was weight coefficient greater than the cube, although
3-48383 but based on only ten fish. The whole series this is not invariably the case (e.g. P. flavescens
were significantly heterogeneous (P < o-oi), but Mitchill in Lake Michigan (Hile & Jobes, I942)).
when August is omitted the remainder are homo- Alm (1946) discusses the condition of perch from
geneous. The data for August consist of only a few different lakes, and although there appears to be
samples over a very limited range and thus may be quite a number of exceptions there is apparently
atypical. The heterogeneity has been ignored in the a tendency for the faster growing populations of
further analysis of the results. The significant varia- perch to have a higher average condition. The
tion among the adjusted mean weights is very Windermere perch would seem to be intermediate
similar to that found for mature females (Table 8). in condition to the Swedish examples Alm quotes,
Immature males. The lengths and weights were but perhaps to have a higher value for the length-
recorded for thirty-five immature male perch of weight coefficient n.
2 years old or older. They gave a regression coeffi-
cient of 3-24253 which is not significantly different 4. THE SEASONAL CYCLE IN GONAD
from that for mature males. WEIGHT AND CONDITION
(d) Discussion and conclusions (a) Gonad weight
Kesteven (i947) discussed the possibility of using The total weight of the perch as recorded included
the analysis of covariance in the treatment of length- the weight of the gonads. As these change in size
weight data from fish, but the present work was with the season, and when ripe may constitute
started before his paper was published. As no other a considerable fraction of the total weight, records
use of the method so far appears to have been made, were made at different seasons of the gonad weights
the present analysis functions to some extent as a of small samples of fish. It was also hoped that these
trial of the method. It' has revealed information weights would provide evidence of the season cycle
which would not have been shown if ordinary con- in gonad development.
dition factors or a single length-weight relationship The perch were opened ventrally and the single
regression had been used. It is clear from the results ovary or both testes dissected out, an operation that
described above, that no single regression will could usually be accomplished in a few seconds.
adequately describe the length-weight relationship The gonads were then weighed complete. At first
for the perch. The fact that the length-weight a rough chemical balance was used, and the gonads
relationship coefficient n was considerably greater were weighed to the nearest o-i g., later a more
than the cube would also have complicated the accurate chain type balance was used and weights
analysis of seasonal changes in condition by means of were recorded to o-oi g. For the largest fish the
2I0 Conditionin theperch

Table 5. Average dates of collection, numbersof fish and adjusted mean weightsfor
seasonal subgroupsof II immaturefemales
Adjusted Percentage Percentage
Average No. of mean of maximum of August
Subgroup date fish weight weight weight
Apr. 23 Apr. 46 I4.9 8I-9 85.2
May 2oMay 70 I6-I 88 5 92- I
June 2I June 55 i8'2 I000 I04^0
July 20 July ii6 i6-9 92'9 96-7
Aug. iI Aug. I75 I7.5 96-2 b00'0

Total - 462 I6-9 92'9 96-7

Table 6. Average dates of collection, numbersof fish and adjusted mean weightsfor
seasonal subgroupsof III and older immaturefemales
Adjusted Percentage
Average No. of mean of maximum
Subgroup date fish weight weight
Apr. 2 IApr. 36 25 4 86-I
May i8 May 50 z6-o 88-I
June 2I June 24 29-2 9910
Late summer 5 Sept. 2I 29-5 I00*0

Total - 3I 27.0 9I.5

Table 7. Average dates of collection, numbersof fish and adjusted mean weightsfor
seasonal subgroupsof maturefemales
Adjusted Percentage
Average No. of mean of maximum
Subgroup date fish weight weight
Spring 2i Feb. 95 50o6 88-5
Apr. and May 2I Apr. 394 55-4 96-9
Spent 22 May 259 46-5 8I.3
Late June 2I June 121 52-0 9019
July 22 July 53 55'9 97.7
Aug. 8 Aug. I24 56-5 98'8
Sept. 14 Sept. Io6 57-2 Ioo-o
Autumn 25 Oct. 360 53 o 92,7

Total - 15I2 52'85 92 4

Table 8. Average dates of collection, numbersof fish and adjusted mean weightsfor
seasonal subgroupsof mature males
Adjusted Percentage
Average No. of mean of maximum
Subgroup date fish weight weight
Spring 5 Feb. I0 I8-9 86.3
Apr. 22 Apr. I6i 19-2 87-6
Spent 25 May 46 I8.5 84.4
Late June 21 June 82 2I'0 95-8
July 20 July I2 21'0 95-8
Aug. I2Aug. I53 21-7 99g0
Sept. 8 Sept. 56 21*9 I0oo0
Autumn 23 Oct. 20 2I-6 98-6
Total - 640 2zo6 94-0
E. D. LE CREN 2II
gonads were weighed on a 'Butchart' balance to the type were drawn for adequate samples of fish from
nearest gram. In every case an accuracy of at least different seasons. The data for mature females in
5 % was obtained. In the case of the smallest February and October each gave a horizontal band
immature fish, however, an accurate dissection and of points on the graph. The points for very large
weighing of the minute gonads was not attempted. fish weighing about i kg. also lay on the same
horizontal band. The data for both ripe and spent
1000 fish in May in each case also showed no definite
tendency for an increase or decrease in gonad: body-
500 _
weight ratio with weight of fish. Data for other
months, and for immature females, were less
extensive and less conclusive. In the case of male
fish, data for February, April, August and Sep-
100 / _ tember were plotted, and in each case were not
inconsistent with the conclusion that the gonad:
body-weight ratio is constant at any one season for
any size of fish. It can therefore be concluded, that
although there is some individual variation in the
gonad:body-weight ratio, it tends to be constant at
10 any one season for all sizes of fish of the same sex
and state of maturity.
Once it was clear that the gonad: body-weight
ratio did not alter with size of fish, mean gonad:
body-weight ratios of samples of fish that varied in
0-5-~~~~~ size could be used to trace the seasonal changes
in this ratio. The data for mature fish are plotted in
Fig. 2. It will be seen that the ovary has a minimum
size in mid-summer when it is quiescent. In August,
however, it begins to increase in size, and this
increase proceeds regularly through the winter and
spring till the spawning time in May. At this time
0.1
the gonad averages about 20 % of the body weight.
The freshly spent fish have empty ovaries weighing
0.05-
about 3 % of the body weight. By the middle of
ino Lengt cetmte
June these ovaries have shrunk to about i % of the
body weight, at which ratio they remain till August.
There is considerable variation among the different
0.01
samples in the average gonad: body-weight ratio,
0-005-
especially in the spring, but this is almost certainly
due to phenological differences from one season to
the next. In I947 in particular, after an unusually
severe and late winter, the ratios in May were
approximately equivalent to those for April in
0.001
0-5 1.0 5 10 50 other years.
Length in centimetres The mature male fish differ from the females in
Fig. i. The length-weight relationships of the perch.
several ways. Development of the testes starts in
The regression lines for log weight on log length are August, but by October the testes have reached
given for larvae, 0 and I group, mature females and their maximum size, weighing about 8 % of the
mature males. The regressions are those forthe seasonal body weight. They then remain this size through
subgroups with the maximum relative condition. the autumn, winter and spring, till April. The
average gonad weight then decreases through April
First, for each fish whose gonads had been weighed and May till it is only I-2z% of the body weight.
a gonad: body-weight ratio was calculated and Quantitative data for the summer are nearly non-
expressed as a percentage. It was then decided to existent, but from visual observation it can be con-
determine whether this ratio altered with the size cluded that the testes remain small and quiescent
of the fish. Accordingly 'dot diagrams' were con- through June, July and most of August. As the
structed of the logarithm of the gonad: body-weight testes begin to be ripe early in April, the apparent
ratio plotted against body weight. Graphs of this fall in their weight from the beginning of April may
212 Conditionin theperch
be due mainly to inevitable loss of milt with handling ratio for all the forty-eight fish was 0-52 %. Indi-
in capture and examination. These data on seasonal vidual ratios varied from o-o to 2 I %; there did not
changes in the testes weight are very similar to seem to be any significant difference between the
those obtained for Perca flavescens Mitchill in mean ratios for samples from June, August or
America by Turner (I919), who also studied the September; or any correlation between the ratio
histological changes in some detail. and the size of the fish, though the data available
The data for immature fish are very limited, but are not really adequate to determine these features.
suggest that the gonad:body-weight ratio is rela- The perch in Windermere are known to feed more
tively constant at about o 5 % for immature females heavily in summer than in winter (Allen, I935), but
and does not change with size or season. This ratio even if they did not feed at all in winter, these
~Z I X i [ -
TI I I
10 o _ > Males
8 0 4A Q

6 0A

0~~~~~~~~~~~~~~~~~~~
no o I ?1IXIII 1 11 ,-1 1 1
0
~~~o24 0
'
1943
0
o 22 ?0 Females 01944
620 eS A~0 0 1945
~~18 ~ ~ ~ ~~)4t-1'. o~~~~~~A1946
~~~~16 0 x~~~~~~~~~~~01946
o14 .1948
12
1p4 - l l | , +
A~~~~~~~~~~~~~~~~~
1 0~ ~ ~~~~P
o 8eban Mar Ar Maoue Jl u Sp.O o e

Fig. z. Seasonal changes in the gonad weight as a percentage of the total body weight. The larger symbols are the
means for samples of five or more fish with the vertical lines indicating 95 %oconfidence limits. The smaller
symbols represent the means of samples of less than five fish.

of 0o5 % is approximately equivalent to that of the figures are adequate to show that, relative to the
quiescent mid-summer ovaries of mature females. seasonal changes and individual variation in total
Although the evidence is not sufficient to be con- weight, the weight of food in the stomach is not
clusive it suggests that this ratio begins to increase in important. The weight of material in the remainder
normal females in their third August at the same of the alimentary canal may nevertheless have some
time as the mature females. begin to develop for significant influence on the total weight, but visual
spawning next spring, but remains constant through- observations suggest that the weight of the intestinal
out life in those females that still remain immature. contents would tend to vary rather less than that of
the stomach contents.
(b) The weight of stomach contents
In order to determine how much of the individual (c) Condition
and seasonal variation in relative condition might The seasonal changes in relative condition are
be due to the weight of food in the stomach, the given in outline by the adjusted mean weights for
stomach contents of a few samples of perch seined the seasonal subgroup regressions for weight on
in the summer of I 948 were weighed. The per- length. These are presented in Tables 4-8. These
centages of the total body weight contributed by data have been amplified by calculating relative
the stomach contents were determined for each condition factors for individual fish and finding the
fish. The average stomach content: body-weight mean of these for each sample of fish from the two
E. D. LE CREN 213
years 1945 and I948. These years were chosen to of about 78-80 %. By 20 June the condition of the
give a comparison between one year and another, spent fish has increased considerably and this
and because they provided the most ample sources increase is continued through July, till by August
of data, but a few additional samples for the three the relative condition is about 99 % of the maximum
spring and three autumn months from other years in September. After the middle of September it
are also included. These means are plotted (with falls sharply and by the middle of November it has
their 95 % confidence limits) in Figs. 3-7. reached an average value of about 9I %. It then
For the 0 and I group fish data from the adjusted continues to fall slowly through the winter and early
mean weights of the regression only are available. spring.
It will be seen from Fig. 3 that the 0 group fish have The mature males present a picture similar to the
a somewhat higher relative condition in September mature females, except that the loss of relative con-
than the I group fish a year later. There is a con- dition on spawning is not so great (Fig. 7). There is
siderable gap in the data between September and evidence that the fall in condition from December
the following spring, but the low value of 84-7 % to March is somewhat steeper than in the females.
for the middle of April indicates that there must be There appears to be a similar sharp rise in April
a considerable fall in condition over the winter. followed by a general decrease through the latter
A subsequent rise in the early summer is indicated part of April and most of May as the male fish
by the higher values in July and August, but the spawn. This spawning process appears to take
detail of this increase cannot be followed without several days for an individual male fish, unlike the
further material. As in the other groups the con- females who probably lay all their eggs in one
dition falls again in October, and presumably spawning act. After spawning the males increase in
decreases throughout the autumn and winter. condition rapidly and then more slowly throughout
In the case of the II group immaturefemales the the summer to a maximum in September. The
adjusted mean weight for June was thought to be autumnal fall in condition does not begin until late
unexpectedly high, and as it was based nearly in October.
entirely on one sample of fish, it was ignored, and
the value for August taken as the basis (of i000%) (d) Individual variations in relative
for calculating the relative condition factors. The condition and gonad weight
points for the adjusted regression means in April The data on relative condition and gonad weight
and May and the sample means for that period, that have just been discussed have been based on
show that the relative condition is low at the mean values obtained for a number of fish. There is,
beginning of April but then rises rapidly through however, considerable individual variation in both
May. The July adjusted regression mean may be relative condition and percentage gonad weight. As
aberrantly low, and the relative condition probably a measure of this individual variation the coefficient
increases slowly through June, July and August. of variation has been calculated for a few repre-
After August the II group immature females begin sentative samples. For relative condition factors
to develop their ovaries and join the mature female the values obtained varied between 6 and I 7 %;
group. while for ovary weight as a percentage of body
Relative condition factors for the few females weight the coefficient of variation ranged between
that remain immature though three or more years Io and 28 % and for testis weight from 8 to 36%.
old are plotted in Fig. 5. Data are somewhat The higher values were obtained at the spawning
scanty, except for April and May, but it will be seen time or in the autumn when both the relative con-
that these fish follow a similar pattern to the II dition factor and the gonad weight were undergoing
group females. Condition probably falls slowly rapid change. As might be expected the greater
through January, February and March, but rises individual variation shown at such times is probably
in May and June. In summer the maximum is due more to individual differences in the time of
probably reached early in September followed by spawning or development of gonads than to inherent
a fall through the autumn. individual variations in condition or gonad weight
For the maturefemales the data are more adequate maintained throughout the season, though the
and are shown in Fig. 6. The condition is fairly latter does occur to some extent. This greater
constant with perhaps a slight fall through January, variation at times of change is also shown by the
February and March. In April it rises sharply, and greater range of the 95 % confidence limits for mean
at the beginning of May some samples of ripe fish condition factors in April, May, June and October
have a mean relative condition factor of nearly than at other times as illustrated in Figs. 4-7.
IOO %. The ripe fish from later on in May, however, The individual relative condition factors for
have much lower mean condition factors. Samples a number of large samples from August, September
of newly spent fish have mean relative conditions and October, the latter part of the growing season,
J. Anim. Ecol. 20 14
I~ ~ I I -

0 and I

0
100 -

0-0
oR // N

C~~ ~ ~I ~ ~~~~I N

. 90 / N N

cs~~~~_* ~~~ regression


Adjusted means _
80

I_ __ I

Jan. Feb. Mar. Apr. May June July Aug. Sept. Oct. Nov. Dec.
C ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~I
Fig. 3.

_Group 11females

o tAdju June
Adjusted meanst
regression s
Nov. Dec.
Jan. Feb. Mar. Apr. May July Aug. Sept. Oct.
80
Group 11 females if Fig- 3. :~~~~~A1945

L 80

00

0 c
Jan. Feb. Mar. Apr. May
May June July
July Aug.
Aug. Sept.
Sept. Oct. Nov. Dec.
Dec.
Fig- 4-
E. D. LE CREN 215

Mature females }

100

IA 194
r- I II I I I I I I I

80 I 0

C:

8 IN
o 1 LIAly
-u 90 T

Adjusted regression means

I 1945~A A94
t .~~~~~0
1948

b 100
Jan.
Jan. Feb.
Feb. Mar.
Mar. Apr. May June
June July Aug. Sept. Oct.
Oct. Nov.
Nov. Dec.
Dec.
Fig. 7.
6.

i eaiecniinepesda ecnaeo
Figs.3-7.easoalurchanges
forrege Matnubreup
males {h en fsmlswt hi 5%cniec h aiu.Teajse
iisaegvn uv en a

for egrssin sugrops nd te mansof smpls wth teir95 cofdjsenreglmt ressogiean.s uva
been drawn by eyethrough these points
2I6 Conditionin theperch
were also examined for correlation between con- nearly all the difference between the seasonal curves
dition and growth in length. It was suspected that for condition for mature and immature fish is due
those fish that were growing slowly in length as to the gonad-weight cycle in the former. For
indicated by a small 'plus' growth for the current example, in the mature females the nearly constant
season, might also be growing slowly in weight and level of condition through the winter, as contrasted
so be in poor condition. Further, those fish that with the continued fall in the condition of immature
were making the maximum increment in length fish, is due to the increase in ovary size offsetting
might be doing so at the expense of weight and also the fall in body weight. The rise in condition in
be in poor condition. On examining the data, April is further due to the continued increase in
however, it was found that although one or two ovary weight concurrently with a slight increase
samples showed a very slight suggestion that such in body weight.
a correlation existed, the overall picture gave no This slight, and unexpected, rise in body-minus-
indication that there was any correlation between gonad-weight in April before spawning has been
growth increment in length and relative condition examined in rather more detail by calculating
factor. condition-minus-gonads for a series of samples
(e) Discussion of the seasonal cycle from I944 and I947. They confirm the suggestion
It is to be expected that there will be a seasonal from the general Fig. 8 that there is a slight rise in
cycle in condition with a high level of condition condition-minus-gonads in April followed by a
during the late summer and a low level at the end of slight fall, possibly due to the final increase in gonad
the winter. The immature fish (O and I group, II weight just before spawning being made at the
females, and III and older immature females) show expense of body weight. A similar tendency is
this cycle with a minimum in March and April, apparent in the males as well as the females. As,
a rapid increase in May and June, a maximum in however, there is considerable individual variation
August and September and a rapid fall in October it is doubtful how significant these relatively small
and November. changes are. It is impossible with the present data
These groups, and the mature males but not the to elucidate them fully.
mature females, also show a high value for condition It appears then that the loss of weight on spawning
in June and low in July, relative to the expected is almost but perhaps not quite entirely due to loss
trend. This may be because most of the data for of ova or milt and not general 'condition'. That some
these 2 months comes from a few samples mainly sacrifice in general body weight is made towards
collected in one year ( 945) and may thus be aberrant. the build up of the ova is, however, indicated by
On the other hand, the younger fish were probably the high rate of fall and subsequent low level of the
feeding mainly on zooplankton (Allen, I935); and condition-minus-gonads through the winter. It is
Smyly (unpublished data), studying the food and significant that all the groups of fish, except the
growth of 0 group perch in Windermere, has found II group females, have their lowest level of condition-
that in some years there may b'! a shortage of littoral minus-gonads about the middle of April and the
zooplankton about the middle of July. Thus the spring increase in condition starts at about the same
apparent July fall in condition might possibly be time. In the II group females, however, it seems to
due to a temporary shortage of food for the smaller, start at about 3 weeks earlier.
plankton-feeding fish. In the mature males the development of the mass
The effect of the actual weight of the gonads on of the testes takes place in about 6 weeks between
the relative condition and its seasonal changes can the middle of August and the middle of October.
be seen in Fig. 8, where the seasonal curve for The curve for condition-minus-gonads of the males
condition is plotted with and without the percentage falls off steeply earlier than in the other groups, but
gonad weight subtracted. For convenience the the condition with gonads is maintained at a high
lower curve may be termed the 'condition-minus- level for longer into October than is the case for the
gonads', although it has been calculated indirectly females. Thus the development of both the testes
from the data for average condition, and gonad and the ovary is made partly by maintaining a total
weight as an average percentage of body weight, body weight for length higher than normal and
and not from calculating the condition of fish partly at the expense of the condition of the rest of
weighed after the removal of their gonads. the fish. It is significant that a large proportion of
It will be seen from Fig. 8 that the curve for the the food available for growth, and the growth
mature females minus gonads is very similar to potential of the mature fish, particularly the females,
that for the immature females 3 years old and must be devoted each year to the annual production
older, but that the seasonal fluctuation is rather of gonad products.
greater. Again the curve for the mature males It is hoped to provide some detailed evidence of
minus gonads is similar. It appears therefore that the season when growth in length takes place in
E. D. LE CREN 2I7

100
Mature males

90

80

100 _

Mature _>
-females _

90

80

100

Ill + immature females

90

Jan. Feb. Mar. Apr. May June July Aug. Sept. Oct. Nov. Dec.
Fig. 8. Diagrammatic seasonal curves for relative condition with and without gonads. The solid black represents the
gonad weight; the upper edge of the black the condition with gonads and the lower edge the condition-minus-
gonads.
2i8 Conditionin theperch
a later paper. It would seem, however, that the upon the statistical section. Finally, I am deeply
immature fish grow in length from early in May until indebted to Miss Charlotte Kipling upon whose
early in October, but that growth is most rapid in statistical acumen and computational perseverance
June, July and August. The mature fish do not much of this paper depends.
begin to grow in length until after the spawning
season; probably about the middle of June and when 6. SUMMARY
the relative condition has reached 90-95 %. If the
start of the growing season for length is correlated i. The methods of analysing length-weight data
with some stage of the spring increase in condition, from fish are reviewed. Emphasis is laid on em-
it would then be expected that the annual growth of pirical formulae of the type W=aL", and the
immature fish, as compared with mature, might limitations of the conventional condition factor.
benefit from the longer growing season of the former. z. The length-weight relationship of perch of all
The females do grow more in their second year than sizes was determined from a series of regressions of
the males, but the older immature females grow at log weight on log length, and an analysis of co-
about the same rate as mature females of the same variance. Relative condition factors were calculated
age. These older immature females have in fact for individual fish from smoothed mean weights
a spring increase in condition at about the same obtained from the regression lines.
time as the mature females. It cannot be concluded 3. In length-weight relationship it was found that
however, from the evidence at present available, the perch could be divided into a series of six
that the seasonal changes in condition are the sole groups corresponding with age, sex and maturity.
factors controlling the growing season for length, Each group was generally homogeneous within
although it would seem reasonable to suggest that itself throughout the seasons, but usually differed
a fish would recover the weight lost over the winter significantly from the other groups. Relative con-
and in spawning before starting again to grow in dition was found to vary significantly with the
length. season.
4. At any one season the gonad weight is a
constant percentage of the body weight for fish of
5. ACKNOWLEDGEMENTS all sizes. The seasonal changes in gonad weight are
Most of the fish used for this paper were collected described and differ somewhat for the two sexes.
by Mr G. J. Thompson and his assistants, and Stomach contents weigh up to 2% of the body
Miss M. Hullock helped to prepare the figures. weight in summer.
Mr H. C. Gilson, Dr Winifred Frost and Mr 5. There is a regular seasonal cycle in condition
W. J. P. Smyly have made helpful comments on which is at its maximum in September and minimum
the manuscript, and I also wish to thank Dr Ralph in early spring. The different seasonal changes in
Hile for his criticisms and general interest in the condition between mature and immature fish can
first part of the paper. Dr N. L. Johnson and Mr largely be accounted for by the cycle in gonad
N. W. Please have kindly read and commented weight of the former.

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