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Statistical Treatment of Animal Experiments

Author(s): John Wishart

Source: Supplement to the Journal of the Royal Statistical Society, Vol. 6, No. 1 (1939), pp.
Published by: Wiley for the Royal Statistical Society
Stable URL: https://www.jstor.org/stable/2983620
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Vol. VI., No. 1, 1939


[A Discussion, opened by DR. JOHN WISHART, at the meeting of the INDUSTRIAL

SOCIETY, November 24th, 1938. PROFESSOR R. A. FISHER, F.R.S., in
the Chair.]

I HOPE it will not be thought, because I have consented to open this

discussion, that I regard myself as an expert on animal experimenta-
tion, or that I have necessarily a great deal of experience of the
subject. The statistician must, as always, tread warily in dealing
with problems which are the acknowledged sphere of the practical
expert, but his advice on lay-out and statistical interpretation of
results may be useful, advice which should not be given without a
study of the practical problems involved. Nor could I have hoped,
in the very short time which has been available since the discussion
was planned, to study afresh the extensive literature on the subject,
in order to deal comprehensively with all animals, or with all experi-
mental problems. My purpose will be served if I am able to set the
ball of the discussion rolling.
My first contact with experiments of this nature was when, some
ten years ago, I examined the figures of certain trials laid out to test
the effect of treating grass-land with different grades of basic slag.
In some the effect was measured by the weight of hay taken off the
land, in others by the live-weight increase of animals fed on the
pasture. I recall one experiment where three, or at most four,
plots IO acres in size, each having a different treatment from the
others, were grazed by cows. Weekly weights of individual animals
were recorded, and the differences between initial and final weights
over the same time interval were averaged for each treatment, and
used as a measure of the effectiveness of that treatment. In a sense
this is not an animal but a crop experiment, but because the animal
was introduced to measure the effect, there were introduced at
the same time the difficulties which attend animal experimentation.
Chief of these was the large error involved, but there were others.
Practical considerations required the effective grazing down of the

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2 WISHART-Statistical Treatment of Animal Experiments [No. 1,

area, and should one particular treatment gain an ascendancy over the
others, it meant the introduction at a certain stage of an extra
animal. It was difficult, too, to fix in advance an integral number of
cows which could be depended on to graze the area effectively, while
mnixed grazing brought in other problems, and animals had a way of
falling sick and having to be replaced.
By that time satisfactory methods of crop experimentation had
been devised by Fisher, and the randomized blocks and Latin-square
lay-outs were becoming familiar, together with the essential principles
of good experimental design illustrated by these arrahgements. It
was natural, therefore, that animal experiments should be studied,
and criticised, in the light of these principles. From this point of
view the experiment I have just described was not satisfactory, since,
although live-weight gains were recorded for each cow on the plot,
there was no real replication of plots. The variation in live-weight
gain between cows on the same plot could not safely be taken as a
measure of the variation between cows in different plots; further,
the plots were bound to differ in inherent fertility, and a superiority
in pasture growth, as measured by its effect on animal growth, could
not be ascribed solely to the greater effectiveness of the particular
slag in supplying phosphatic fertiliser.
When it was suggested that the randomized block technique
should be applied to animals, various difficulties were raised. Thus,
it was stated that animals were more variable than field plots, too
variable, in fact, for small differences in growth to be detected.
Further, if this variability was to be reduced, all animals in one
experiment ought to be offspring of the same parents, and ought to
be of the same age and weight at the start of the experiment. This
limits the number of animals in a single experiment to the number of
young born at any one time, and even here it may be impossible to get
many of comparable weight. Thus the experiment can only be a
small one, and even if the standard error of a single animal's live-
weight gain is reduced by such a process of selection, the standard
error of the mean of any treatment will not, in general, be small,
beeause of the limited number of animals available. The expense of
animal experimentation is bound to limit the numbers capable of
being dealt with, thus keeping down the number of treatments to be
tested, and making an unsatisfactory experiment generally, since
while there may be several thousand unit plants of wheat in a single
plot of a cereal experiment, and several hundred unit roots on a plot
of potatoes or sugar beet, the unit with animal experimentation will
often be the single animal.
For these reasons progress in the direction of laying out animal
experiments which would permit of adequate statistical examination

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1939] WISHART-Statistical Treatment of Animal Expertiments 3

of the results has beell slow, and experimelnts have usually, been of a
very simple liature, comparable in fact to the duplicate plot experi-
ments that were in vogue prior to the randomized block era, except
that there was not always duplication. There have been exceptions
in the case of small animals; some of you may remember hearing
from Mr. Hartley the details of a fairly elaborate experiment on
poultry, at a recent meeting of this Section. I turn now to the series
of nutrition experiments on pigs- begun at Cambridge some two or
three years ago. These will serve as well as anything to illustrate
the methods involved, and they are experiments that I happen to
know something about. There may be others present who can deal
with cows, or sheep, or by contrast with small-scale animals such as
rabbits, or even rats and mice, while the point will not be lost sight of
that experiments on human nutrition deal with that very variable
quantity, the human animal. At an early meeting of this Section we
were given a contrast betweeni an experiment on rubber in Malaya
covering 2,ooo acres and one at East Malling, for which the total area
was only i/iOO acre. There may well be conitrasts as great on the
animal side.

The Camnbridge pig experimnent

The pig is in many respects a satisfactory animal to experillment

with. Relatively pure stocks are obtainable, and large famiilies are
the rule. Thus, particularly at a time when the Boards for Pig
Marketing and Bacon Development were set up, it was natural that
experiments should be unidertaken, not only to investigate the most
economical method of producing a satisfactory bacon pig in this
country, but also to study the experimiiental methods themselves.
The usual method has been the group-feeding trial, and I shall
illustrate this from an experiment set up alongside the individual-
feeding trial which will be described later. Three rations were tested:
A, a diet containing the amount of protein usual in farming practice,
B, a diet with an excess of protein, and C, a diet with a protein excess
about twice, by weight, that of B. Three pens were used, one for
each treatment, and each contailled ten pigs, both sexes beilng repre-
sented in the same proportions. The produce of a number of litters
was distributed as evenly as possible over the three pens; for example,
if six pigs were available from one litter, two would be allocated to
each pen. Care was taken to see that the animals were as homo-
geneous as possible, and approximately of equal weight at weaning
time. There were, of course, certain variations, but the three pens
were evened up so that the average weaning weights were approxi-
mately the same. After the pigs had been given a little time in which
to settle down to their rations, they were weighed individually, then

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4 WISHART-Statistical Treatment of Animal Experinments [No. 1,

at weekly intervals thereafter, the rationing proceecding according to

standard tables, until i6 weeks had passed. By that time some of
the pigs had reached 200 lb., and had to go to the bacon factory.
The remainder were kept until they, in turn, had reached 200 lb.,
but the experiment proper ended when seventeen consecutive weekly
weighings had been recorded for each pig, all covering the same
calendar period. The effect of the rations was measured by the
average live-weight gain per pig over the i6-week period, the figures
being 142-5, 141V0 and 132-7 lb. for rations A, B and C respectively.
It is doubtful how far any kind of statistical analysis on the individual
live-weight gains is valid. If, however, we may assume that we can
regard the ten pigs in any one pen as replicates of one another, then
it appears that the drop in live-weight gain with increasing protein
percentage in the ration is not significant. The standard error of
each pig's gain is 17-3 lb., or 12-5 per cent. of the mean, and that
of the above three averages is 5.5 lb., or 4-0 per cent. A point of
importance to the experimenter is the comparison of the efficiencies
of meal conversion with the different rations. In the present case
only the total meal consumption for the ten pigs is known, since the
food was provided in bulk to each pen, and while the amount of meal
per lb. of live-weight gain can be worked out for A, B and C separately,
no figures can be got for individual pigs, and no statistical analysis,
however invalid, is possible.
The standard error obtained is as low as we are likely to get with a
group-feeding trial, for the experiment was conducted with all
possible care. We thus see that if we are to be able to detect differ-
ences of the order reached in this experiment, something more
precise is necessary. For some time individual-feeding trials had
been advocated, and the opportunity was now taken to make a
comparison of the two methods, particularly as the group trial was
there to convince the farmer, who might well distrust the results of
trials carried out in a way that was not normal farming practice, but
who would probably be convinced if both methods gave the same

Individual-feeding trial

From the litters of each of five sows were selected six pigs, three
being hogs and three gilts. The six from any one litter were run
together in a pen, one hog and one gilt being given ration A, another
pair ration B, and the third pair ration C. Mechanical methods were
adopted to segregate the pigs at feeding time into small individual
pens, so that each pig had undisturbed access to its day's ration.
The experiment consisted of five pens, or thirty pigs in all. Weighings
were recorded as in the group-feeding trial. The experiment is of

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1939] WISHART-Statistical Treatment of Animal Experimtents 5

the randomized block type (for the rations were randomized in the
feeding-boxes), each block consisting of the pigs from one litter.
Thus pen (or litter) differences could be eliminated by the method
of the analysis of variance, showing the possibilities of this type of
experiment in cases where more pigs are needed than are available
fromn a single litter. The analysis follows the usual lines, and it is
possible to examine the effect of food, a possible sex difference, and
the interaction of these two effects, on any one of the variables.
These were: live-weight gain, meal consumption (known for each
pig), and a great number of post-slaughter measurements on the
carcase whose examination would reveal, if any, differences in
the conformation of the bacon pigs produced in consequence of the
differences in the composition of the food.
In the case of live-weight gain the differences were small, the
figures being I5-0, I46-4 and I42-6 lb. for A, B and C respectively,
with a standard error of 3-0 lb., or 2-I per cent. One very satisfactory
feature of the experiment was the fact that the accuracy was much
greater than that of the group trial; the standard error of each pig's
live-weight gain was 9-6 lb., or 6-5 per cent. of the mean, compared
with I7 3 lb., or I2-5 per cent. in the other case. The same story
is told by the means in both cases. Small as is the standard error,
the differences between A, B and C are not, however, significant on
the analysis of variance test, even when we note that nearly all the
sum of squares for the 2 degrees of freedom for food is contained
in the single degree of freedom component of " principal effect ",
which measures the drop from A to C. There is a way, however,
in which accuracy can be gained. The growth of a pig in any one
week obviously depends to a considerable extent on its weight at
the beginning of the week. Thus differences in initial age and weight
at the beginning may, and usually will, affect the comparisons. We
have no difficulty with age in the present case, but the limited
amount of experimental material makes it impossible to have the
six pigs in any one pen of the same initial weight. But this does
not really matter, as by means of the analysis of covariance we can
examine food and sex differences in live-weight gain after correction
for initial weight-i.e., adjusted to what they would have been,
knowing the regression of live-weight gain on initial weight, had all
pigs started at the same weight. When this was done it was found
that the food differences were still insignificant on the 2 degrees of
freedom, but when the single degree of freedom " principal effect "
was isolated, this was significant at the 5 per cent. probability level.
Thus, by a process of " squeezing," the drop in live-weight gain
from A to C was declared to be significant; the drop was 5-6 per cent.,
which figure had a standard error of 2-4, compared with 2-9 before

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6 WISHART-Statistical Treatment of Animal Experiments [No. 1,

adjustment for initial weight (without adjustment the drop was 5-7
per cent.). In neither case was there a significant difference due to
sex. It is of interest to observe that live-weight gain was signi-
ficantly correlated with initial weight (r = + 0.60), while in the
case of the group trial the correlation was not significant (r = + 0.36),
due perhaps to competition at the feeding-trough, the weakest
going, possibly literally, to the wall. This is a further point in
favour of the individual trial.
These results, together with the results of examination of the
rest of the data, were published two years ago.' The experiment
was found to be quite workable without too much trouble, once the
piggery had been built for the purpose. It was the pioneer of a
number of trials which have been conducted since, limited, of course,
to the same total number of pigs and the same number of treatments,
although the treatments themselves have varied. As far as I was
concerned, the figures then left the nutrition experimental stage and
entered on a new stage-that of the statistical study of growth.
As it is the statistical treatment of such experiments that is the
subject of our discussion (though this involves, naturally, the
question of lay-out), I shall ask you to bear with me a little longer
while I explain what I have since done.

Examination of growth curves

I have always thought it a pity that more use was not made of
the regular weighings of the animals in such an experiment as this.
Ten years ago I fitted straight lines by the method of least squares
to the weights of the cows in the basic-slag trial, in the belief that
this would furnish a more accurate measure of the growth than the
difference between initial and final weights, and I turned now to the
study of the growth curves of the pigs. These appeared on graphical
inspection to be parabolic in character, with an upward curvature
-i.e., the weekly gains rose up to practically the end of the experi-
ment. Now, had the gain in weight been proportional to the weight
at the beginning of any one week, the logarithms of the weekly figures
would have been linear, but plotting revealed that this was not so,
for the graphs showed a downward curvature. I therefore began
with the fitting of second-degree parabole to the actual weights,
figures more easily interpretable by the practical man than the
logarithms, having ascertained graphically that such a parabola
appeared to be a good fit to the data. There were now two variates
for each pig, g, the average growth rate in lb. per week, and h, pro-
portional to the rate of change of growth in lb. per week per week.
Analysis of variance showed that on g the food difference was signi-
ficant on the single degree of freedom for " principal effect ", repre-

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1939] WISHART-Statistical Treatment of Animal Experiments 7

senting a gain in accuracy over the figures of live-weight increase.

The sex difference was not significant. Analysis of covariance on
initial weight w showed a further-gain in accuracy, for food differences
were now significant on the 2 degrees of freedom, and the sex difference
was found to be significant, the gilts having the greater mean growth
rate. The correlation between g and w was + 0-65. Although the
trend for A, B and C in the case of h was in the same direction as with
g, food differences were not significant, but there was a significant
sex difference, in that gilts had a greater rate of change of growth
rate than hogs. Covariance of h on initial weight showed no change
in the effects, the correlation being very small (r = + 0.04).
These results were given in a paper published in Biometrika this
year,2 but I have since returned to the attack, more especially as
further examination revealed that the cubic term of the curve fitted
to the weekly weighings, which I shall call i, was significant, while
the quartic term was not. Graphical examination of g and h against
w, and of h against g, revealed nothing abnormal. It seemed worth
while to see how far h appeared to depend on g, but analysis of
covariance showed that the two constants were not significantly
related (r + 0.20). In spite of this, however, the analysis of
residual variance after correction for g showed that the sex effect in
h fell below the 5 per cent. significance level. It would appear,
therefore, that it was the higher g in the case of gilts that was the
cause of the higher h, although the sex effect was not significant on
g alone. But there was an effect of initial weight on g, and this
suggested that the partial variablesg . w and h . w should be examined.
The following are the figures for the totals of IS pigs, for hogs and
gilts separately:
w q h
Hogs ... ... 617 138 2.43
Gilts .. ... 586 141 -9-73

Now, since rwA is negligible we have

rgh. V ( gh 2) approxim

0-20 0 26.
V\1 - 0.65 2)-

Thus it would appear that the significance of the sex effect for h
corrected for both g and w should be less than for h corrected for g
only. This was borne out by a multiple covariance analysis, which
reduced the sex effect to complete insignificance, although the
multiple correlation was not significant (R = 0 23).
Analysis of variance of the cubic term i showed a non-significant
sex difference, but very marked differences due to food treatments,

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8 WISHART-Statistical Treatment of Animal Experiments [No. 1,

far more significant than anything hitherto reached. The figures for
the averages of ten pigs each are:
A B c Mean Error
Mean i ... -000959 -000610 -000248 -0O00606 0 00109
Per cent ... 158.3 100-7 40.9 1000 18.0

The differences in size of the cubic terms between A and B, and

between B and C, are significant. Analysis of covariance of i on w
added nothing to the story, although the correlation was just
significant (r -0.43).
The result of all this is that we are now in a fair way to obtaining
a complete picture of the growth curves of the pigs under food and
sex differences, and can say with confidence that they are significantly
different. For the five hogs and five gilts, one from each pen,
given over to each of the three food treatments, it was possible by
working out the averages of the constants of the fitted curves-
namely, the mean and the linear, parabolic and cubic terms-to con-
struct curves for the " mean pig " in each case. These constants
are given in the following table, together with their standard errors,
obtained in each case from the analysis of variance tables. Note
that they are not the quantities g, h and i hitherto discussed (which
are the B, C and D of Fisher's treatment of orthogonal polynomial
fitting in Statistical Methodsfor Research Workers), but the quantities
a,l-12g, a2 = `ih I i, and, of course, ao, which are the imimiediate
result of the application of Aitken's method of polynomial fitting,3
and can be used directly, as indicated by him, for calculating the
Coefficients of Curves (actual weights)
aO a, a. al
Hogs A ... ... 109.47 4.803 0-0553 -0-00273
B ... ... 104.48 4 495 0.0568 -0-00146
a ... ... 106.97 4-478 0.0497 -0-00052
Gilts A ... ... 106X88 4-846 0.0641 -0-00302
B ... ... 108.71 4.793 0*0587 -0-00221
a ... ... 101.98 4.498 0 0593 -0-00096
Mean ... ... 106.42 4.652 0.0573 -0-00182
Standard Error ... 3-87 0.144 0 0037 0.00046
or 3-6% or 3.1% or 6.4% or 25.5%
The six growth curves are shown in the diagram, each being
displaced relative to the one on its left by the same constant distance
at the start, in order to avoid overlapping. The points on the
diagram are the weekly averages of the weights of the five pigs for
each sex and treatment. The curve was not directly fitted to these
points by least squares, but was drawn from the averages of the
constants fitted to the individual pigs.*
* It will be realized, as pointed out by Professor Fisher in the discussion,
that the curve so obtained is in fact identical with that obtained by a least-
squares fit to the weekly averages.

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1939] WISHART-Statistical Treatment of Animal Experiments 9

A careful examination of the diagram will reveal the points of

difference that are brought out by statistical treatment of the growth
constants. Comparing the food treatments A, B and C, on the average
of hogs and gilts, it is seen that A has the smallest initial weekly
gain and C the greatest, B being intermediate. The last weekly gain
is, however, the same for all; A has curved up more than C, and has
reached its point of inflexion before the experiment ceased, while the
C curve behaves more like a parabola all through. In case it may
be thought that different initial and final weights account for these
differences, it should be added that interpolation of the figures for








A 8 C A B C


FIG. 1.-Growth curves of the "mean pig" for each treatment and sex
(actual weights).

the week after reaching 42 lb. and for the week before reaching i8o lb.
shows the same relative effects. Comparing hogs with gilts, averaging
for food treatments, we find that hogs start off with the greater
initial weekly gain and end with the smaller final weekly gain.
Hogs were, however, heavier at the beginning, and correcting for
this as before by interpolation, we find that the initial gains become
equal, though the final gains show no change. The curves showing
the two extremes in growth are those for hogs under the C treatment
and for gilts under the A treatment. These are side by side in
the diagram.

Examination of logarithmic data

Finally, because three constants have been found to be necessary
for the description of the growth of the pigs on actual weights, it is
interesting to see whether any economy in this respect is possible if
logarithms are used instead. If growth differences can be elucidated
by examination of one or two constants of the curves fitted to the
logarithms, that will represent a distinct advantage. Analysis of
the logarithms of a sample pig of the A treatment showed that
the linear and parabolic terms were significant, while the cubic

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10 WISHART-Statistical Treatment of Animal Experiments [No. 1,

and quartic terms were not. Accordingly the first two terms, which
we may call g' and h', were fitted to all pigs, and the usual analysis
of variance was worked out in both cases. The table for g' is as
follows, g' being calculated from the logarithms taken to two
places of decimals, and treated as whole numbers:

Analysis of Variance-g'
Variation D.F. Sum of Squares Mean Square
Sex ... ... ... 1 0-2811 0*2811 1-1435
Food ... ... ... 2 0-2101 0-1050 0.6511
Interaction ... ... 2 0-0329 0.0164
Pens ... ... ... 4 0.9115 0.2279 1.0386
Error ... ... ... 20 0.5710 0*02855

Total ... ... 29 2.0066

The sex difference was very significant, the mean for gilts being
distinctly higher than that for hogs. Further, the food differences
were significant, the trend being in the same direction as in the case
of the g values. The following tables summarize the results, apart
from the significant pen differences:

Summary of Results-g'
Hogs Gilts M[ean Standard Error
Mean g' . ... 4-091 4-285 4.188 0.044
Per cent. 97.7 102-3 100.0 1.06

A B C Mean Standard Error

Mean g' ... 4301 4.163 4.101 4*188 0-053
Per cent. ... 102.7 99.4 97.9 100.0 1.28

In the case of h' neither effect was significant. In view of the

clear-cut nature of the results, it was evidently not necessary to take
account of initial weights in a covariance analysis,* and the final
examination undertaken was the covariance of h' on g'. Contrary
to our experience of h and g, the regression was found to be significant
(r = -0 62), but adjustment of h' for g' did not alter the conclusions
already reached.
We conclude, then, that the simplest statement of the results of
statistical examination of this body of data is that treatment A has
given the largest mean proportionate growth rate, and C the smallest,
B being intermediate. Gilts have shown a greater mean proportionate
growth rate than hogs. All show a falling off to about the same
extent in proportionate growth rate as the period advanced. The
* The correlation coefficients for g' and h' with the logarithms of initial
weights were - 0-81 and + 0-47 respectively (both significant). The only
change on correction was that pen differences were now insignificant, showing
that differences in average growth constants for the pens were due to different
initial weights.

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1939] WISHART-Statisticat Treatment of Animal Experiments 11

following table gives the constants of the logarithmic growth curves

for the six " mean pigs ", in the same form as the previous table:

Coefficiets of Curves (logarithms)

a.' a. a, a3
Hogs A ... ... 199 49 2.122 -00424 -0-000021
B ... ... 197*47 2-034 -0-0368 -0*000712
C ... ... 198-72 1 981 -0-0368 -0-001241
Gilts A ... ... 197'70 2-179 -0 0397 -0-000121
B ... ... 199.23 2-129 -0 0425 -0-000018
C .. ... 195*56 2-119 -0 0389 -0-000970
Mean ... ... 198.03 2 094 -0Q0395 -0 000507
Standard Error ... 1.66 0-038 0-0025 -0-000357
or 0.8% or 1-8% or 6-2% or 70-4%
The six growth curves calculated from these constants are shown
in the secoind diagram, together with points which represent the

12 30


~~~~~~~/ /



:Fro. 2.- Growth curv

(logarithms of weights).

weekly averages of the logarithms of the weights for each sex and
treatment. The graphs bring out the differences in average slope
as between A, B and C on the one hand, and between hogs and gults
on the other.*
It is not suggested that all the above calculations are necessary in
any similar experiment. In fact, had we started with the logarithms
instead of the actual weights, it is unlikely that the latter would have
had the intensive examination which has been given to themt. But
the figures are presented as an illustration of attempts that may be

* Further examination has shown that the cubic terms fitted to the B and
a hogs, and the C gults, are significant. The values of a3' have been added to
the table since the meeting, and the diagram has been re-drawn. It would
appear, therefore, that working on the logarithms does not necessarily save
computational labour to the extent hoped for, if a comaplete specification of
growth has to be provided.

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12 WISHART-Statistical Treatment of Animal Experirments [No. 1,

made to represent the growth curves of animals under experimenta-

tion in a more complete way than has usually been done, by means
of relatively few growth constants. The imposed differences of
treatment in this experiment are small, but I understand that nutri-
tionists are interested in the change of form of the growth curve in
more extreme cases, with a view to relating these either to the food
supplied or to the subsequent conformation of the carcase. It may
be that similar methods to those outlined will prove useful in such
problems. They are, at any rate, open for discussion.

Co-ordinated Experiments in Pig Husbandry

I shall conclude by referring briefly to a development which may
be taken to be the direct outcome of the Cambridge experiment, and
for details of which I am indebted to Mr. Boaz. The ordinary centre
can only experiment with a limited number of pigs, and must resort
to group feeding. A number of co-ordinated experiments have,
therefore, been arranged to be carried out at research stations, farm
institutes and agricultural colleges in Great Britain and Northern
Ireland. Three treatments, one of which is usually a control, are
allotted to three pens, each having five or six pigs, which are group-
fed. Sets of three pigs are taken, from the same litter and of the
same sex and roughly the same weight when I2-I6 weeks old. Each
set is regarded as a " block ", and the pigs in the set are allotted at
random to the three pens under the different treatments. Thus each
pen contains one pig from each set. Regular individual weekly
weighings are made until the pigs reach bacon weight, and later
carcase measurements are taken at the factory. Statistical analysis
at each centre is of somewhat doubtful validity, but each centre can
be regarded as a replicate, and the data can be analysed as a whole.
At some centres there is effective replication of the sets of three
pens; at Harper Adams College, for example, there will be in the
present experiment eighteen pens of five pigs each, testing three
treatments in six fold replication. The results of this experiment
will be awaited with interest.

The CHAIRMAN said that it was not the Society's custom when a
discussion had been opened to move a formal vote of thanks to the
opener, so he would take the opportunity of expressing his thanks


1 Woodman, H. E., et al., 1936, J. Agric. Sci., 26, 546-619.

2 Wishart, J., 1938, Biometrika, 30, 16-28.
3 Aitken, A. C., 1933, Proc. Roy. Soc. Edinb., 53, 54-78.

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1939] Discussion on Animal Experiments 13

and those of the meeting to Dr. Wishart for an extremely interesting

address. He particularly wished to call attention to Dr. Wishart's
wise choice of subject. Animal experimentation was now awakening
from a condition of backwardness relative to agronomic experiments,
and Dr. Wishart had put before the meeting two or three carefully
described and simple experiments, both of the cruder and of the more
advanced types, upon which he would probably welcome detailed
Animal experimentation, which was going forward in many
directions, was a very wide subject. There were cases such as those
with which Dr. Wishart had dealt wherein the animal was regarded
primarily as a commercial product and experimentation was aimed
at producing that commercial product most economically. Dr.
Wishart had alluded in his paper to man as a subject of animal
experiment. The more earnest advocates of nutrition in these
times seemed. to believe that the juvenile human should be brought
up like a little pig, perhaps fearing that otherwise he would be brought
up like a little wolf, but the material difference, as between human and
animal experiments, was that the pig was expected to be turned
shortly into bacon, whereas it was hoped that the child would be
capable eventually of becoming self-supporting, and perhaps also
capable of self-control.
Finally there was that very large realm of animal experiment
in which the animal was used for purposes of assay, and the reaction
of the animal was a means of, for example, standardizing insulin or
carrying out operations of a comparable character.
He had been exceedingly glad to hear Dr. Wishart, from the
evidence he had given, reinforcing the practice of individual feeding,
which had been advocated now for many years by Crampton in
Canada and was, he believed, becoming universally adopted by
animal experimenters. He wished also to call attention to Dr.
Wishart's judicious treatment of growth curves, and particularly to
the fact that many statisticians, when faced with a growth curve,
would, at least until recent years, have been tempted to analyse the
variation in weight at a given age and later to try to combine the
evidence obtained at different ages, which latter process was hope-
lessly encumbered by the fact that, using the same animals, the
evidence to be combined was far from independent. In the method
which Dr. Wishart had used, each individual animal was made to
supply a series of constants which adequately described its growth
curve, and those constants might be treated like any other experi-
mental data, and rendered valid by the random sampling of the
He felt that in spite of the lucidity of the exposition, something
must have been kept back when Dr. Wishart, in speaking of those
growth curves shown in the diagram, said, " Note t-hat the curve is
not fitted to these points by least squares, but is the one determined
from the averages of the constants fitted to the individual pigs."
He could not see how that could be. He would expect to reconstruct
the least-square solution for the average weights by averaging the
least-square curves for the individual pigs.

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14 Discussion [No. 1,

DR. WISHART said the Chairman was perfectly right. The

least-square solution had been re-constructed in the process, and
the wording ought to be altered to make this clear.

DR. YATES also expressed his pleasure at hearing this opening

address by Dr. Wishart on a very important subject.
It had been said that it was all very well to use a good statistical
design with a field crop, but that hopeless difficulties were encountered
if the same thing were attempted on animals. He doubted whether
that statement were true. It was true that animal experiments
required more continuous attention. But against that was the
advantage of being able to repeat experiments at decidedly shorter
intervals, since with such experiments-and he had particularly in
mind pigs and the smaller animals-one need not wait for the next
season to come round.
Again, with feeding trials and similar experiments it was not
necessary to start the whole of an experiment at one time. Each
block could be started when a suitable batch of animals was forth-
Once a resort was made to individual feeding, a large number of
units could be treated separately, and that removed what had in
the past been one of the greatest difficulties in the way of adequate
statistical design and, in particular, of introducing anything in the
nature of a factorial experiment.
In general, therefore, he would say that animal experiments of
the ordinary type tended to be just as simple as agronomic experi-
ments, and definitely more so than the sort of problems which were
encountered when dealing with long-term experiments in agriculture
concerning crop rotations and so on. On the other hand, it was
certainly true, because of the larger size and cost of the individual
unit, that animal-breeding was in general more difficult than plant-
Much had been said, at various times, about the high variability
of animals. Certainly a single animal-a pig, for instance-was
variable, but from Dr. Wishart's figures, and from similar figures
obtained in experiments at Rothamsted, it would be seen that in
fact there was not much to choose in this respect between one pig
and one agricultural plot. They both tended in a well-designed
experiment to give an experimental standard error of the order of
iO per cent.
The utility of factorial design in animal experiments was only
now beginning to be appreciated. As an example of this type of
design he might instance an experiment on four factors carried out
at Rothamsted on the feeding of young pigs. The factors were:
large versus small amounts of green food, rationing versus ad lib.
feeding, comparison of pigs permitted to exercise in a field each day
versus those confined to pens, and coarse versus finely ground meal.
This gave i6 treatment combinations. of which 8 were assigned to
each pen or group of pens, the four-factor interaction being con-
founded. 64 pigs in all were used, each block of i6 being started
at an interval of about four weeks, Individual feeding was used,

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1939] on Animal Experiments 15

and litter differences were eliminated by the arrangement. Thus

information was provided on all four factors simultaneously, and at
the same time possible interactions between the various treatments
were investigated. Each pig was in effect used four times over,
with a corresponding gain in overall efficiency, quite apart from the
information on the interactions.
Such an arrangement was, of course, only possible if a reasonably
large number of experimental units were available. This was one
of the great merits of individual feeding. Another advantage was
that proper estimates of experimental error were available from
even a small number of pens. The comparison of the individual
weights in a group-feeding experiment could never furnish such an
estimate, not only because all the pigs in the same pen might suffer
some common damage, such as disease, but also because of competi-
tion within each pen. Lastly, individual feeding might be expected
to increase accuracy considerably, since it eliminated competition
at the feeding-trough.
There were other possibilities in the design of animal experiments
which were not available in agronomic experiments. One of these
was the rotation of treatments on the same animals. Mr. Wake
Simpson had recently given a paper to this Section on certain
experiments he had carried out on school-children, in which he had
made use of this principle.. The objection to rotation of treatments
was that the effects of the treatments might persist from one period to
the next, so that the results might be complicated by residual effects.
Use might also be made of concomitant information such as
initial weights, inilk yields over a preliminary period, and so on.
The statistical process here involved was known as the analysis
of covariance. There were, in fact, many possible ways of increasing
the accuracy of an experiment: it was the job of the statistician
to incorporate these in the design, and not to attempt to use them
when the experiment was completed, but whatever refinements of
design were adopted, proper replication and randomization were
essential. It might seem unnecessary to emphasize this, but a
most cursory examination of the literature would show how often
these cardinal principles were neglected, and how often through their
neglect the whole of the work was rendered valueless.
In conclusion, a brief description of a grazing experiment at
present in progress at Rothamsted might be of interest. The
problem was that of evaluating the residual effects (on the pasture)
of feeding cake to grazing cattle. It was a difficult problem experi-
mentally, and had been made more so by the agriculturists because
they insisted that practical grazing conditions had to be reproduced,
and that to do this each plot must be at least five acres in area.
In the Rothamsted experiment there were nine five-acre plots in
three blocks of three plots. In each year one block was grazed by
cattle, those on one of the plots being fattening bullocks to which
cake was fed. In the subsequent winter one of the other two plots
of the block received nitrogenous fertilizer. On the third plot
nothing was given. In the two subsequent years mixed grazing was
adopted, with the intention of measuring the effect of the treatments

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16 Discussion [No. 1,

on the herbage by the differences in live-weight gain. It was also

envisaged that samples of the grass might be cut and the yields
evaluated quantitatively. The remaining two blocks were treated
similarly at one-year intervals, so that all three phases of the rotation
were represented.
Whether the experiment would ever be capable of providing the
hoped-for results remained to be seen-Dr. Yates had grave doubts
on the matter. But it was at least interesting to note that the
variability of the animals in the first year (which was a bad one
from the grazing point of view) was by no means as large as was
feared. From the comparison of the increases of animals on the same
plot the standard error per plot due to the variability of the animals
was found to be only about 6 per cent. of the increase in weight.
This error was of course not cumulative from year to year. Another
interesting result that had already emerged was that mixed grazing
was more efficient from the point of view of live-weight increase
than grazing by cattle alone.

MISS TURNER said this was her first appearance in the Society,
and she expressed her thanks for being allowed to be present. She
wished to speak on some problems which might be likened to the
long-term experiments of which Dr. Yates had spoken. Her own work
had been done in Australia, and related mainly to sheep. In
Australia the natural pastures were deteriorating, and one method
of improving them was to plant English grasses and fertilize them
annually. Some graziers had raised the objection that; although
this increased wool production, it caused coarsening of the wool,
with consequent decrease in value. It was necessary, therefore, to
find out what effect the improvement of the pasture had on the wool,
and whether such an effect was cumulative. It was also important
to determine the effect of improved nutrition, such as that provided
by improved pastures, on lambs whose growth had been checked at
various stages by feeding on poor pastures. Both these might be
described as long-term experiments.
Another point was the use of the group trial. Miss Turner
agreed that individual feeding was essential for a laboratory trial,
but thought another kind of trial was necessary in Australian
conditions. Pigs could be fed in pens either individually or in
groups, and when thus feeding them in groups one was approaching
quite closely to actual husbandry conditions. But feeding sheep
in pens was getting nowhere near actual farming conditions in
Australia, and the grazier was far more likely to be convinced by a
trial which did approach such conditions. Experimenters had to
convince both themselves and the grazier by translating any trial
from laboratory conditions to the field. She would describe one
large experiment with cattle which was being carried out by the
Australian Commonwealth Council for Scientific and Industrial
A certain cattle disease in Queensland was known to be a deficiency
disease, and preliminary experiment had shown that the feeding
of certain supplements was successful in preventing it, 13ut it was

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1939] on Animal Experiments 17

obviously necessary to extend such an exp

into the field. About 3oo animals, varying in age, sex and breed,
were collected from various sources. It was decided to run a
randomized block experiment, arranging the animals so that in each
block they were uniform for age, sex, breed, the station from which
they were purchased, initial body-weight, growth curve during a
preliminaryperiod, and size. Body-weight always presented a problem,
since the weight of the same animal might vary at differenit times
of the day, and the animals were therefore run together for nine weeks,
being weighed weekly. The initial weight was taken as the mean
of the nine weighings, and the growth curve over the nine weeks was
also considered. Size was measured in terms of height, length and girth,
since if body-weight alone were considered a large cow in poor con-
dition would not be differentiated from a small cow in good condition.
The main trial consisted of three groups, two receiving supplements
and one being the control. It was hoped that in addition to analysing
any mean responses to treatment it would be possible to sort out any
sex differences or breed differences. In the preliminary experiment,
which had given an indication of successful treatment, no control of
variation such as the use of randomized blocks had been attempted,
and by their introduction the difference necessary for significance
had been considerably reduced.

DR. S. BARTLETT said that at Shinfield they were accustomed to

dealing with a variety of animals for experimental purposes but he
himself was particularly concerned with dairy cattle of all ages. He
thought the question of whether to adopt group or individual
feeding for dairy cattle was still of controversial interest. With
mature dairy cows the matter was not important because it was
easy to treat the cow as an individual both in regard to intake of
food and output of milk, and live-weight, but it was sometimes
difficult to arrange individual-feeding experiments for young cattle,
and at Shinfield, until recent years, the group-feeding method was
always used. For experiments on the winter feeding of young
stock it was usual to let the animals run loose in a yard, half a dozen
or so together, and feed as best they could. During the last year
or so, however, individual feeding had been introduced, the
practical difficulties having been overcome by tying up the
animals during the day so that all their food was taken individually,
and letting them loose at night for exercise and so-forth. This plan
worked remarkably well, both from the practical and the experi-
mental point of view. The experiments were now of the random-
ized block type with as many treatments and replications as the
number of animals permitted.
Dr. Wishart had mentioned a comparison of the variability of
group and individual feeding. He could safely say that the standard
error of individual live-weights in young cattle on individual feeding
was less than half of what it was when they were on a group-feeding
system. With cattle, therefore, they showed perhaps slightly more
improvement in this respect even than Dr. Wishart had found with

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18 Discussion [No. 1,

An argument often put forward in favour of group feeding for

young stock was that it imitated practical conditions, but this
argument needed qualification. Under practical conditions a farmer
who had a group of animals in a shed, and found one of them being
badly bullied and half starved, would probably place the animal in
another pen. This would be good, or at least relatively good, farming
practice, but it would spoil an experiment.
He considered it paid to tie up animals for feeding during the
winter whether they were under experiment or not. An improvement
in these animals was noticeable over animals running loose, in mean
live-weight gain as well as in uniformity.
The amount of bullying of certain animals in group feeding
varied, and he strongly suspected that this bullying propensity was
influenced by nutrition. This would suggest that there were other
factors to be taken into account besides live-weight gain. The
grazing experiment which Dr. Wishart mentioned also brought to
mind all sorts of other factors. While he had tried to show that
individual feeding was a great advance from the point of variability,
he believed that some of these other factors could only be tested by
group feeding. It would be well, therefore, not to assume that
because individual feeding gave less variability than group feeding
the latter must be given up. Group feeding was still useful in
measuring certain factors.
He was interested in Dr. Wishart's growth curves, which appeared
to place the animal experimenter ahead of the crop experimenter,
because the former would now utilize the interim weights. Here he
would like Dr. Wishart's opinion. In a winter feeding-experiment
the cattle were weighed on three successive days both at the beginning
and again at the end of the experiment. During the course of the
experiment it was proposed to weigh the animals each month on
three successive days. One alternative might be to weigh them
one day each week. Which was the better method for use
with Dr. Wishart's group-curve technique? It was probably a
question of whether the greater accuracy of three weighings was
more important than the danger of missing the exact point of
inflection of the growth curve which might occur with monthly
He had talked over the matter with a colleague of his who had
experimented with rats and it appeared that in rat-feeding experi-
ments the question of group feeding seldom arose. Individual
feeding was invariably used and was superior in every way for
nutrition experiments.

DR. M. S. BARTLETT had one or two comments to make which

arose fairly directly out of the paper. The first was in connection
with design, and bore on the last section of the paper dealing with
Mr. Boaz's experiments. It was sometimes difficult to obtain
enough animals at any one centre, and it was therefore hoped to
conduct instead co-operative experiments with a rephcation at each
centre. That point was brought out in a discussion opened by the
late Mr. Gosset in 1936, who said that this might be all right if the

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1939] on Animal Experiments 19

centres were at research stations, but i

variability in farming practice between centres might cause differ-
ences in the nature of interactions in the same way as in series of
field experiments. But in spite of that difficulty, Dr. Bartlett
thought there was still a greater chance of usino each cenitre as a
separate replication than in the case of field experiments. With the
latter all the nitrogen required, for example, might be obtainable from
the soil, whereas in animal-nutrition experiments differences were
likely to be of a smaller order, for the animals would always need
a definite ration with so muLch protein, so much carbohydrate, and
so on.
A difficulty that sometimes confronted the statistician in the
analysis of experiments (not only in animal experiments) was that
arising from several sets of observations. If the experimenter gave
his own instructions on what should be tested, there was hardly any
difficulty (though it might perhaps have to be pointed out that all the
information available in the experiment was not being extracted);
otherwise it was a separate problem in itself to consider what to do
with the data. Sometimes one could arrive at a compromise. In
long-term dairy-cow experiments at Jealotts Hill, the mean weekly
milk yield during the experimental lactation period was taken.
The farmer was mainly interested in this, and it also probably
gave a fairly efficient analysis of the differences between the lactation
curves due to treatment. In other cases one might have to depart
from the sort of thing the experimenter himself would select, and
try to reduce the observations for any treatment to one or two
constants. It was interesting to observe that by doing this for the
live-weight figures Dr. Wishart had established a clearer-cut difference
between the food treatments than had been obtained originally from
the straightforward live-weight increases.
There was still a slight statistical difficulty, even if one had cut
down one's observations to one or two constants, in that, strictly
speaking, it was not fair to isolate one analysis from others being
done at the same time. It might therefore be relevant to summarize
Dr. Wishart's analysis in regard to the joint significance of all the
fitted constants, which he would do by writing down the following
approximate but simple x2 table for the food effect (adjusted for
initial weights).

d.f. Direct analysis ILog. analysis

9 ... ... 2 7'12 13 28
h.g ... ... 2 0.45 0.51
i.gh ... ... 2 8 77 1.23

ghi ... ... 6 1634 15 02

Note on the above Table.-Any item could have been tested exactly by the
z test. For testing the last line, see, for example, Proc. Camb. Phil. Soc., 30
(1934), 327-340 (339). For the approximate x2 test for the last line, as used
here, see Proc. Camb. Phil. Soc., 34 (1938), 33-40 (380). The other items were
chosen to make the table additive. It should be noted that (i) the x2 test is
leas accurate for these items, since the multiplying factor used to get the best

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20 Discussion [No. 1,

x2 approximation does no
(ii) caution should be used in interpreting the significance of any partial effect
such as h.g (h adjusted for g), since the adjustment implies using covariance
with g, which is itself varying significantly between treatments. Nevertheless
the entire table still serves as a useful summary of the significance of the food
effect. It was amended to include the cubic term for the logarithmic analysis,
which was not completed by Dr. Wishart until after the meeting. The
significance level of the total x2 nearly reaches P 0-01 for the direct analysis
but is only about P = 0 02 for the logarithms. It is interesting, however, to
note the large contribution to x2 for the logarithms from the linear term. The
cubic term in this analysis was found just significant by Dr. Wishart, but does
not provide any extra significance when tested jointly with the other constants.

MR. F. J. DUDLEY said that his chief interest was in poultry

experiments, but from time to time he had examined figures obtained
at Harper Adams Agricultural College in connection with pig
experiments. It was of some interest to consider the results of what
might be called a uniformity trial, carried out some years ago by
Dr. Crowther. Fifty pigs were divided into five lots of iO pigs each,
in such a manner that the lots were made as closely comparable
as possible, attention being paid to breeding, weight, age, and sex.
The IO pigs of each lot were housed in two sties, the pigs in one sty
being made comparable with those in the other. Throughout the
experiments all the pigs were on the same treatment. Taking as
the criterion the total gains in live-weight, the effect of making the
sties comparable at the outset was, as might have been expected, to
yield a very high variance within the sties and a significantly lower
variance between sties. The variance between the sties in the lots
was not significantly different from the variance between lots. A
further point was that if the pigs had been distributed at random,
the difference between the means of two lots of iO pigs each could
not have been considered significant unless it had been greater than
I 2 per cent. of the meani.
It was considered that the most satisfactory method of conducting
a group-feeding trial in which all the pigs in a sty fed at the same
trough, was to regard a group of pigs as the experimental unit, rather
than the individual pig. The prospect of being able to test only
differences greater than I2 per cent. was not at all encouraging, but
the low variance between the means of lots of iO pigs in the uniformity
trial, giving a standard error of only 3 I per cent. of the mean,
suggested that the method adopted to make the lots as even as
possible at the outset might be used.
The further requirements after the groups had been formed were
(1) adequate replication to improve the accuracy of the treatment
comparisons, and, what was more important, to provide an estimate
of error by which to judge the significance of treatment differences,
and (2) to assign the treatments to the different groups at random.
For what he described as reasons of good husbandry, the practical
man preferred to keep the large pigs separate from the small pigs,
and attempts had been made to introduce replication through
groups of pigs of different sizes on each treatment. In one such
experiment 6o pigs were first divided into four groups according to
initial weight, while in all other respects the groups were made as

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1939] on 4nimal Experiments 21

comparable as possible. The I5 pigs of the heaviest group were

then divided into three sub-groups, each to receive a different type
of management. The pigs of the other size-groups were treated in
a similar manner. The standard error of a treatment mean in this
case was 4-5 per cent., based on 5 degrees of freedom, after eliminating
variation between size-groups and correcting for differences in initial
group weights by linear regression.
Soine caution was necessary in taking as an estimate of error the
interaction between treatment and size. In a similar experiment
where there were three size-groups and a comparison was made
between two levels of protein in the ration, it was found that the
superiority of the one ration over the other was smallest for the heavi-
est pigs and greatest for the smallest pigs, while it was intermediate
for the pigs of medium size. It was impossible to say whether this
was significant or not, because of the lack of replication of the size-
treatment groups, but it seemed to be generally accepted that in
proportion to its size the small pig required more protein than the
large pig.
A serious drawback to the success of group-feeding trials at a
single centre was the limitation of accommodation for experimental
purposes. In the majority of the experiments at Harper Adams
College the number of pigs had been between 30 and 40, but in some
cases as many as 6o pigs had been used. The consequences of this
limitation were that the experiments had been of a simple type, the
number of treatments and replications was restricted, and the estima-
tion of error was usually based on so few degrees of freedom that only
large treatment differences could be shown to be significant.
There seemed little doubt that group-feeding trials could serve a
useful purpose in pig-feeding investigations, provided the experiments
were properly planned and carried out on a sufficiently large scale.
One means of meeting the latter requirement and at the same time
extending the range of conditions under which treatments were to
be tested was to co-operate with other centres of a similar character.
A certain amount of co-operation of this kind had already taken
place, and Dr. Wishart had given some details of a co-operative
scheme which was at present in progress. Incidentally, the part
which Harper Adams College was taking in this investigation was the
largest pig experiment carried out at that centre. From what he had
learned regarding the arrangements the experiment at that station
would effectively provide its own estimate of error.
He had been particularly interested in Dr. Wishart's remarks
concerning correlation between initial weights and total gains in
weight. His observations from the limited data he had examined
were that when the initial weights ranged from 35 lb. to 6o or 70 lb.
there was a positive correlation which was well marked. When the
initial weights ranged from 6o lb. to go lb. there appeared to be no
correlation. Further, among a group of pigs where the initial
weights ranged from 42 lb. to 96 lb. the regression of total gain on
initial weight was not linear, the medium-sized pigs gaining slightly
more than the larger and considerably more than the smaller

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22 Discussion on Animal Experiments [No. 1,

DR. WISHART, in reply, said that he would be inclined to recom-

mend weighing on one day in each week instead of taking three suc-
successive daily weights in each month. The latter process would
give a more accurate determination at any given time, but if growth
curves were to be fitted the accuracy of each individual weight was
not so important as the number which could be obtained.
In conclusion, he expressed his gratification at the way in which
his paper had been received. He was glad that the discussion had
widened out and dealt with different animals, different methods,
and different problems; also that there had been a pleasing absence
of criticism of his own contribution.

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