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org/ on April 4, 2018

The metabolism of Psammechinus miliaris eggs 557

Shapiro, H . 1935 Biol. B ull. Wood's Hole, 68, 363.
Shearer, C. 1922 Proc. Roy. Soc. B, 93, 213.
Tang, Pei-Sung an d G erard, R . W. 1932 J . Cell. Comp. Physiol. 1 , 503.
W arburg, O. 1915 Pfiiig. Arch. ges. Physiol. 109, 324.
— 1925 Biochem. Z . 164, 481.
W arburg, O., K ubow itz, F . and Christian, W . 1931 Biochem. Z . 242, 170.
W hitaker, D. M. 1931 J . Cen. Physiol. 15, 183.
— l 933a J ' Cen. Physiol. 16, 475.
—• 1933 & J-Gen. Physiol. 16, 497.

598.65:611.013.16

Visual stimulation and ovulation in pigeons

B y L. H arrison Matthews

{Communicated by F. H. A . Marshall, —
Received 27 September 1938)

I t has long been known th at the female pigeon, unlike the carefully
selected domestic hen, does not ovulate spontaneously. Harper (1904)
showed th at in pigeons ovulation occurs not less than 8 days after the
introduction of the male to a female when both are ready for mating. He
further recorded th at two female pigeons when confined together may both
take to laying eggs. The act of mating therefore is not necessary to induce
ovulation, the mere presence of a companion sufficing. This exteroceptive
stimulus to ovulation is usually regarded as exerting its influence through
the pituitary.
The experiment described in this paper was designed to show whether
the stimulus is visual, auditory, olfactory, tactile or a combination of two
or more of these alternatives. The essential part of the experiment carried
the confining of two females together a stage further. Two birds were
confined in adjacent cages separated by a sheet of glass, and single birds
were confined by themselves but provided with a mirror. The birds were
thus supplied with companions to which they had no tactile access.
Wooden cages with fronts of half-inch mesh wire netting were used.
They measured 1 8 x 1 4 x 10 in. Some of them were joined together in
pairs, the partition between them being movable, so that it could be
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558 L. Harrison Matthews

removed or a sheet of glass substituted for it. Others were provided with
slides at the back so th at a sheet of mirror could be slipped in to cover the
back wall.
The birds used were dealers’ ordinary stock, mongrels showing a pre­
dominance of the common blue rock. They were placed in solitary con­
finement for a month, as a preliminary to the experiment, so th at the
effect of any previous stimuli might pass. They were then confined as
follows:
Double cages.
(1) Male and female with no partition (control).
(2) Male and female with glass partition.
(3 ) Two females with glass partition.
(4 ) Two females with no partition.
Single cages.
(5 ) Female alone, with mirror.
(6) Male alone, with mirror.
(7 ) Female alone without mirror (control).
Owing to the difficulty of ascertaining the sex of mongrel pigeons a
mistake was made with some of the birds. One of those in cage no. 3
turned out to be a male, so th at this cage merely duplicated cage no. 2.
No. 6 was confined as a female, but was found to be a male after the
experiment had started. Cages nos. 1 and 2 were kept in a different
building from the rest. All the cages were arranged so th at none of the
birds could see any of those in other cages.
The results obtained are shown in Table I, where the dates when eggs
were laid are shown.
I t will be seen th at eggs were obtained from the control pair in 9 days’
time, and from other birds after longer intervals. The female of the control
pair was isolated on 12 June. She laid again on 14 and 15 June, but
thereafter produced no more eggs up to the end of July when the experi­
ment ended. The control female, shut up by herself with no mirror, did
not lay at all up to 12 June. On th at day she was placed in a double cage
with a male, and laid on 23 and 25 June. The male (6), which was mistaken
for a female and confined with a mirror, soon started calling and displaying
in the usual manner of pigeons (cf. Whitman 1919) to its mirror image.
The results obtained show th at the stimulus which causes ovulation in
the pigeon is a visual one. The ovulation of birds separated from the male
by a sheet of glass, or confined alone with a mirror, shows th at the stimulus
is not tactile. The absence of ovulation in the control, which was denied
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Visual stimulation and ovulation in pigeons 559

sight of another bird but was in the same room with the others, shows th at
the stimulus is not olfactory or auditory. Were it either, or both, she
would have laid. Her subsequent ovulation when confined with a male
showed her to have been a normal bird which lacked only the visual
stimulus for ovulation.

Table I. D ates of egg-laying

Birds placed in solitary confinem ent 2 February. E xperim ent started 3 March
1 2 3 4 5 7
control $ w ith $, no
<? ¥ <? glass ? <J glass ? ? ? m irror m irror
12 Mar. 15 Mar. 29 Apr. 23 Apr. 14 May
13 Mar. 17 Mar. 1 May 25 Apr. 16 May
30 Mar. 28 Mar. 19 May 27 Apr. 27 May
1 A pr. ' 30 Mar. 23 May 29 Apr. 28 May
16 May 1 Ju n e 1 Ju n e 8 May 9 Ju n e
18 M ay 4 Ju n e 10 May 11 Ju n e
3 Ju n e 15 Ju n e 19 May 24 Ju n e
18 Ju n e 20 May
? isolated 4 Ju ly 22 May Placed w ith
12 Ju n e 23 May 12 Ju n e
14 Ju n e 2 Ju n e 23 Ju n e
15 Ju n e 4 Ju n e 25 Ju n e
No fu rth er 18 Ju n e 9 Ju ly
eggs to end 10 Ju ly
of J u ly

Although visual stimulation from a companion of the same sex, or even

from a mirror image, is sufficient to produce ovulation, it would appear
th a t the response of the female to the behaviour of the male is also self
stimulatory. When the birds are denied access to each other, or are of the
same sex, the pattern of the mating behaviour cannot be completed, and
the lower level of stimulus produced is reflected in the longer interval
before the appearance of the first eggs.
I t is claimed th at this experiment lends further support to the ex­
planation of sexual display in birds and other animals brought forward by
Marshall (1936), who concludes th at display does not produce a conscious
sexual selection as proposed by Darwin, but is essentially a mutual pitui­
tary stimulation. The visual stimulation here shown to exist in pigeons
when confined with a single companion, or merely a mirror image, is
carried to a higher level in those birds which nest in colonies. Fraser
Darling (1938) showed that a definite minimum number of pairs is a
requisite in any breeding colony of certain species of birds that nest
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560 L. Harrison Matthews

gregariously, in order to produce the threshold stimulus for ovulation.
Visual stimulation has also been demonstrated in pigeons by Patel (1936),
who showed th a t unless the male can see the female sitting on the eggs
during incubation, he secretes no milk from the crop gland after the eggs
are hatched.

The writer wishes to express his thanks to Dr Richard Clarke, who

kindly found accommodation for the experimental birds a t the Clifton
Zoological Gardens, and to Keeper Victor Jones who looked after them.

R eferences
D arling, F . F raser 1938 “ B ird Flocks and th e Breeding Cycle.” Cambridge.
H arp er, E . H . 1904 Am er. J . A nat. 3, 349.
M arshall, F . H . A. 1936 Croonian L ecture. Philos. Trans. B, 226, 423.
P atel, M. D. 1936 Physiol. Zool. 9, 129.
W hitm an, C. O. 1919 “ The B ehaviour of Pigeons.” W ashington.

612.014.445:612.461.I
616—006.4:547.681

The photodynam ic activity of the urine of mice

treated with blastogenic hydrocarbons*
B y J. C. Mottram
From the Radium Institute, London, and The Mount Vernon
ospital, Northwood
H

(Communicated by J. W. Cook, F.R.S.— Received 15 October 1938 )

In view of the fact th at a number of blastogenic agents and hydrocarbons

have been found to make the skin of mice photosensitive and also to have
photodynamic action on Protozoa (Mottram and Doniach 1937, 1938),
examinations of the urine and faeces of mice either painted or inoculated
* The term “ blastogenic ” is used to denote a tum our-producing agent, irrespective
of th e character or m alignancy of th e tum ours produced. All of the compounds so
described in th e present com m unication have been shown to give epitheliom as in
mice, and w ith m an y of them sarcom as have been obtained by subcutaneous in ­
jection.