267

THE DAILY RHYTHM OF ACTIVITY OF THE

COCKROACH, BLATTA ORIENTALIS L.
I. AKTOGRAPH EXPERIMENTS, ESPECIALLY IN RELATION TO LIGHT

BY D. L. GUNN
From the Zoology Department, University of Birmingham

(Received 2 January 1940)

(With Five Text-figures)

I. INTRODUCTION
IN the last few years there has been an increasing interest in animal rhythms.
Various aspects of the subject have been reviewed by Szymanski (1920), Fox (1923),
Hoffmann (1926), Richter (1927), AmirthaUngam (1928), Stein-Beling (1935),
Scott (1936), Gunn et al. (1937), Pie'ron (1937) and Kalmus (1938a, b). This paper
is concerned with the daily rhythm in the locomotory activity of the cockroach,
Blatta orientaUs L. Those who are familiar with this insect in its natural haunts
are agreed that it comes out and is active only at night. Can this rhythm of nocturnal
activity be reproduced under experimental conditions and can it be controlled
experimentally? Szymanski (1914) and Wille (1920) showed that it could be
reproduced but they did not attempt to control it.

II. APPARATUS AND METHODS

Szymanski (1914) designed several kinds of apparatus, which he called akto-
graphs, for recording animal activity automatically on a smoked drum. I have
used two practically identical aktographs, one of which has been described in detail
(Gunn & Kennedy, 1936). The animal chamber is 20 cm. long and 10 cm. wide;
when the animal walks along the length of the chamber, the chamber tips and an
attached lever makes a nearly vertical mark on a smoked drum (Gunn, 1937, Fig. 5).
Movements of the animal across the chamber are not recorded, so that one complete
vertical mark on the record represents a walk of at least 20 cm. but probably not
usually as much as 30 cm. A spiral drum was used, rotating once a day, so that
about 4 days' records came on to one drum paper. A time marker showed hours.
In the principal experiments, only recently moulted adult cockroaches were
used. Nymphs were kept until they moulted, so that prolonged experiments could
be carried out with individual animals before they died of old age. Only males
were used. With certain specified exceptions the work was done in a constant-
268 D. L. GUNN
temperature room at 25-5° C , the normal variation being a short cycle of about
+ 0-5° C. This is a suitable temperature for behaviour experiments on this species
(Gunn, 1934).
In all the experiments, beneath the perforated zinc platform on which the
animal walked there was moist common salt, which has an equilibrium relative
humidity of 75 %. About 30 g. of fresh carrot was put in for food every 3-5 days.
This enabled experiments to be carried on with single individuals for as long as
15-20 weeks but, of course, the humidity was disturbed by the carrot. It would
naturally be higher near the carrot than elsewhere, and probably higher before the
carrot had dried than later on.
When the carrot was renewed, the animal was removed from the chamber, the
excess salt solution and faeces pipetted away and more salt was added. The animal
was led gently during this exchange and not agitated, and after a few weeks it
walked into the removal tube and back into the aktograph, when the time came, in
most docile fashion. A few individuals made a temporary escape and had to be
handled roughly, but that only occurred early in their case histories.
Thus, apart from the isolation from its kind which was suffered by the cockroach
during these experiments, the living conditions were good. I believe that an adult
life as long as 22 weeks (19 November 1938 to 24 April 1939) at 250 C. has not
previously been recorded for Blatta orientalis (see Rau, 1924).
All the apparatus was roofed in and curtained off in a corner of the constant
temperature room, and lighted by its own 40 W. electric lamp. This lamp gave
a light intensity of about 20-30 metre-candles (1 f.c. = 10-76 m.c.) inside the
aktograph, with a narrow shadow at one end and another behind the carrot. This
light was controlled by a clockwork switch kindly given by Mr Robert McDowell,
and it could thus be automatically turned on and off regularly at pre-determined
times. When this light was off, it was too dark for the dark-adapted human eye to
read the scale of the light meter. In the main room, daylight was always excluded
and a 200 W. ceiling lamp was on day and night.
Thus the temperature was uniform and the light on the aktograph was kept
uniform or varied at will. The humidity varied, but with a rhythm of 3-5 days as
the carrot dried up and was replaced, and with no daily rhythm. The external factor
which was not controllable was mechanical stimulation-pressure change, noise and
vibration in general. The stands of the aktographs rested on 4 cm. of sponge rubber
on a slate slab built into the wall, so gross vibrations must have been rare. The
results obtained show that vibration was not a factor of overwhelming importance.
The chamber had a glass lid sealed on with vaseline, so draughts were completely
absent, and any chemical stimulation from outside must have had a 3-5 day period.
The only other factor which might have affected the activity of the cockroach
was the presence of the experimenter and of occasional visitors; these potential
disturbances occurred only during the solar daytime, and the complete inversion
of rhythm obtained experimentally shows that they were not important. The
apparatus normally required no more than a few minutes attention each day, with
an hour or so each fourth day.
 Daily Rhythm of Activity of the Cockroach 269

III. TOTAL ACTIVITY PER DAY

In summarizing the results, the records of complete journeys along the aktograph
chamber were counted for each hour of each day; parts of journeys were either
ignored or reckoned as whole journeys, according to their length. Two animals
gave very long experimental records. They were supplied from Sheffield by
Dr Kenneth Mellanby as nymphs, kept at a variable temperature about 250 C. in
normal daylight and darkness until their last moult, and then used in these
Weeks

20 10 20 10 20
1938 Nov. Mar. Apr. 1939
Fig. 1. Daily activity, measured with the aktograph, of two cockroaches from just after the last
moult until death. Indications for both parts of the figure: Alt. = alternating light and darkness,
12 hr. each per day; Alt. noise = rattling noise for la hr. per day, in continuous darkness; Light
and Dark = continuous light and darkness respectively; the spots near the top of the figure indicate
the dates on which the food was changed, etc.

experiments. The distance they walked in a day varied from six to 1300 journeys,
with an average of about 150, corresponding very roughly to 1^-300 m. per day
(average about 40 m. per day). Fig. 1 shows the relation of this daily activity to the
age of the animals and the conditions of light.
It is easy to read several long-period rhythms into these two records, but such
rhythms do not coincide for the two animals, and in any case cannot be correlated
with any external changes. There are, however, two points of resemblance: for
about a week after the cerci of the two cockroaches were cut off (12 December)
both animals were rather inactive; in the second place, in each case impending
jBB-xviiiii *"
270 D. L. GUNN
death was signalized by a few days of tremendous activity about a fortnight before
death.
It is surprising to find that light had little, if any, effect on the total activity
per day. Thus, for these two animals, omitting the periods which were exceptional
because of amputations or approaching death, the average number of journeys per
day for the whole period was 146 (4876 hr.). The average for alternating light and
darkness was 156, for continuous light for periods of upwards of a fortnight at a
time it was 158 and for continuous darkness it was 125 journeys per day. The figure
for continuous darkness is for one animal only, and that towards the end of its
life. For the first month in continuous darkness the average is 145 journeys per day,
so it would not be proper to conclude that continuous darkness has any depressing
effect.
It can be stated, however, that prolonged illumination under these experimental
conditions has no great depressing effect on activity. Neither did the onset of
continuous light have any observable effect (Fig. 1). The rise in activity on one of
the two occasions when continuous light was replaced by alternating light and
darkness can be correlated just as well with recovery from the depressing effect of
cereal amputation. Light, then, appears to have no effect on total activity per day
but, as is shown below, daily variations in light certainly have a great effect on the
time of day at which the activity occurs.
In passing, it may be placed on record that in these experiments one animal
walked very approximately 3^ km. (14,500 journeys) in 100 days and the other
5^ km. (22,000 journeys) in 155 days.

IV. DAILY RHYTHM OF ACTIVITY

Each of the two animals dealt with above was first tested in alternating light
and darkness, with darkness between 18 and 06 hr. G.M.T. Fig. 2 (a, d) shows that
high activity did not occur at all parts of the day indifferently, but that in this
experiment it was concentrated into thet hours of darkness. The average figures
shown in the graph fairly represent the results for the separate days, except that
a few isolated bursts of activity during the period of illumination do not show.
Similar teats were carried out with the dark period lasting for 12 hr. after 11 and
04 hr. G.M.T. daily (Fig. 2, b, c, e, f); in these cases the animals had been kept in
continuous light for at least a fortnight before the new rhythm of light was started.
In all six experiments (Fig. 2) there was one continuous period of high activity
each day and the greater part of this period occurred during the hours of darkness.
The average figures shown fairly represent the individual days' records. Thus
in Fig. ze high activity is shown as starting between 08 and 09 hr., about 3 hr.
before the light was turned out; this actually occurred on 7 out of the 11 days,
while on 2 of the other days activity increased 1 and 2 hr. respectively before
darkness fell.
In general, high activity started during the first hour of darkness. There were,
however, two exceptions to this in the six tests; in one (e) it started 2-3 hr. before
 Daily Rhythm of Activity of the Cockroach 271
dark and in the other (d) there was no great increase of activity until the third hour
in darkness. At the other end, it was usual for activity to reach a fairly low level
several hours before the light came on again. That is to say, activity is brought
to an end by some cause other than the change from darkness to light, and it does
not necessarily start when darkness begins. Nevertheless there is a clear correlation

38 AM 2

38AM3

Hours of the day (C.M.T.)

Fig. 2. Hourly activity of two cockroaches in alternating light and darkness (iz hr. each per day).
The hours of darkness are indicated by diagonal lines. In order to reduce the six parts of the figure
to uniform size, for each experiment the average number of journeys was worked out for each hour
of the day, for the 7-17 days, and each average was expressed as a percentage of the sum of the
averages.

between the rhythmical occurrence of darkness, whenever that starts during the
day (04, 11 or 18 hr.), and the rhythm of high activity. In fact, by suitable adjust-
ment of the time of darkness, the high daily activity of the cockroach can be made
to occur when one pleases. Mellanby (1940) has obtained a similar result in a study
of cockroaches in a natural infestation. Fig. 3 a shows that this rhythm starts
without delay as soon as continuous light is replaced by alternating light and
darkness (see below).

V. ABOLITION OF THE RHYTHM

Since the activity rhythm is related to the rhythm of light and darkness, it might
be expected that in continuous light the activity rhythm would disappear. Practically
speaking, it does, but not at once (Fig. 36). The rhythm previously established by
18-2
272 D. L. GUNN
alternating light and darkness persists for a time but, within the first few days,
complete hours of the active period pass with no activity at all, while bursts of
activity occur increasingly frequently at other times. The process of averaging the
results from several experiments conceals these effects (Fig. 36), but they can be
seen clearly in the record of a single test (Fig. 4). Continuous darkness (Fig. 4)
has much the same effect as continuous light.

Time
Fig. 3 a. The initiation of rhythmic activity immediately the alternation of light and darkness begins.
Average number of journeys in each hour in four experiments which were put together irrespective
of the time by the clock; the onset of darkness (diagonal shading) on the fourth day was made to
coincide for the four experiments.

Time
Fig. 3 b. The slow disappearance of the daily rhythm of activity in continuous light. In the first
week shown there was an altematiot of light and darkness. Average of four experiments put
together as in Fig. 3 a. Diagonal lines indicate the dark periods.

After several weeks in continuous light or continuous darkness, the record of

a single day's activity no longer gives any indication of the previous rhythm. Even
so, the averaging of the daily records of one individual long kept in constant light
or darkness reveals some trace of rhythm (Fig. 5). In a selected group of 12 con-
secutive hours averaged over a fortnight the activity may be twice as great as in the
other 12 hr.; but in the proper rhythms, such as are shown in Fig. 2, the difference
is of a different order, the journeys in the active period usually numbering at least
six times as many as those in the inactive period.
 Daily Rhythm of Activity of the Cockroach 273
This residual rhythm, if it may be so called, is not detectable by simple
inspection of the records, but appears only when the numbers of journeys are
averaged for a week or more. It has been found in all the records for continuous
light or darkness. It is characterized by the activity being above average in a series
of consecutive hours, but only slightly above average. Thus in the four periods in
continuous darkness shown in Fig. 5, the selected groups of 12 hr. contained
69, 59, 59 and 64% respectively of the day's activity. This active period was
28. i. 39 30 Jan. 1 Feb. 3Fe
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5 Feb.
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Fig. 4. The disappearance of the daily rhythm of activity in constant darkness. Individual record
from animal numbered 38 AM 3. During the first week (top) there was an alternation of light and
darkness (iz hr. each) and from 04 hr. on 4 February onwards there was continuous darkness.

between 03 and 15 hr. in the first two and between 12 and 24 hr. in the last two
periods. During the latter, there was a motor making a continuous rattling noise
between 06 and 18 hr.; but it seems unlikely that this could have affected the
animal, for the aktograph was well buffered and the cerci had been cut off. Hair
sensilla on the cerci have an auditory function (Pumphrey, 1940).
The excess over 50% shown by the activity during the 12 hr. period of greatest
activity is statistically significant. The active period did not tend to fall into the
same part of the day in different experiments, but most of the experiments were
not carried on for long enough to allow the period of slightly higher activity to
 Daily Rhythm of Activity of the Cockroach 275
In one experiment lasting 4 days, the 12 hr. darkness was followed each day
by only 8 hr. of light, so that darkness came on 4 hr. earlier each day. On the
first 2 days activity started 2 hr. after dark, on the third day 4 hr. after dark and
on the fourth day 8 hr. after dark. Thus the onset of activity did occur earlier
than previously, but it did not shift as much as the onset of darkness.
A number of individuals were first tested in continuous light (250 C ) ; of these,
three trapped specimens which had been kept in an open dish in an ordinary room
did not show any rhythm from the start. Perhaps the rise of temperature to 25° C.
upset the internal "clock" of these animals. On the other hand, of seven animals
which had been bred from the egg inside a dark oven at 250 C , three showed a
clear rhythm in continuous light, two a doubtful one, one none at all, while the
remaining animal was quite inactive. No doubt the oven was not quite light proof,
while all manipulation during rearing—cleaning, feeding, etc.—was carried out in
daylight hours, so these experiments cannot safely be used as evidence about the
method of acquisition of the tendency to show a rhythm.

VII. DISCUSSION
Szymanski (1914) with Blatta orientaUs and Wille (1920) with Blatella (Phyllo-
dronria) germanica did ten aktograph experiments each, each with one animal at
a time and lasting 24 hr. Both found an average of 3 hours' activity, starting at
15.30 hr. at the earliest and ending at 00.30 hr. at the latest. The temperature varied
around 200 C. In my experiments at 250 C. the active period lasted longer than
this, being seldom less than 6 hr. Szymanski also recorded a short period of activity
at about 06 hr. in the morning, and because of this second period he regarded the
cockroach a9 polyphasic (Szymanski, 1920). Truly polyphasic animals, like mice,
have many short periods of activity each day. My experiments indicate that the
cockroach is really monophasic.
Neither of these authors attempted to analyse the cockroach's activity rhythm.
A number of authors, on largely or entirely theoretical grounds, have considered
that diurnal changes of temperature and humidity provide both the stimulus for
the rhythm and the evolutionary reason for its existence (Picard, 1912; Necheles,
1927; Crawford, 1934). It may be that factors other than light can be used to
control the rhythm, but so far light has been found to be potent and other factors
impotent in a number of cases (Kalmus, 1938a, b). I hope to investigate these and
other questions at a more suitable time. For example, can rhythms of periods other
than 24 hr. be set up? Will a short pulse of half an hour's darkness each day suffice
to maintain a rhythm? Would rhythm9 occur if there were no correlation between
danger and light in the life of the individual ? What is the internal clock, especially
for a persistent rhythm? Does the light act through the compound eyes?
Szymanski (1914) carried out two experiments which are of considerable
importance. In one of these, half the aktograph chamber was shaded and half
lighted all the time, while in the other, half was warmed and the other half not
warmed. In both experiments, during its active period the cockroach walked
276 D. L. GUNN
indifferently in the two halves of the chamber. During this time, the well-known
responses to light and temperature seem to have disappeared. Mellanby (1940)
found, however, that in a natural infestation—in a less limited space than is
available in an aktograph—cockroaches did not emerge into a continuously lighted
room. They may have emerged in other rooms nearby or they may have been active
inside the cavities in the wall.
Esterley (1917) showed that certain marine plankton copepods tended to come
to the top of a long vertical tube in the evening, even though kept in continuous
darkness at an even temperature for 2 days. This accentuation of geo-negative
behaviour occurred with a diurnal rhythm even in the absence of external clues.
Broadly speaking, it is thus to be expected that many animals which show a
diurnal rhythm will behave differently according to the phase in which they are
tested. Most of the work on behaviour is done during the daytime, and this work
on the cockroach suggests that the nocturnal phase can be investigated without
doing more than check experiments at night. In the cockroach, at any rate, the
rhythm can be reversed by a suitable alternation of light and darkness and then
for a few days the rhythm persists almost in full in continuous light. Experiments
on behaviour during the active phase can then be done in the daytime during these
few days. In the experiments described here, the institution of a new rhythm was
always preceded by a period of continuous light; this may not be necessary.

VIII. SUMMARY
1. In an aktograph at 25-5° C , at upwards of 75 % relative humidity and with
food present, the average locomotory activity of the cockroach per day does not
depend on whether there is continuous light for weeks, or continuous darkness, or
a daily alternation of light and darkness.
2. When temperature and humidity do not vary during the day and other
factors are kept as constant as possible, the cockroach's activity can be largely
concentrated into any desired half of the day, simply by suitably adjusting the time
of onset of the half-day's darkness. A rhythm can thus be set up, so that the main
activity occurs at the same hours each day.
3. This activity rhythm persists for some days in continuous light or con-
tinuous darkness, but eventually activity becomes much more evenly spread over
the whole day, leaving only a slight residual rhythm which is unrelated to the
previous conspicuous one. A new conspicuous rhythm can then be started at once
by alternation of light and darkness.
4. There are indications that animal responses to physical stimuli may depend
to a considerable extent on whether the animal is in the active or the inactive phase
of its daily cycle. A method is suggested for making it possible to study the
nocturnal phase during the daytime.

I have enjoyed the pleasure and benefit of discussing this work with Dr Kenneth
Mellanby and of comparing his unpublished results with mine.
 Daily Rhythm of Activity of the Cockroach 277

REFERENCES
AMIRTHALINGAM, C. (1938). J. mar. biol. Ass. U.K. 18, 605-41.
CRAWFORD, S. C. (1934). Quart. Rev. Biol. 9, 201-14.
ESTERLEY, C. O. (1917). Univ. Calif. Publ. Zool. 16, 393-400.
Fox, H. M. (1923). Proc. roy. Soc. B, 95, 523-50.
GUNN, D. L. (1934). Z. vergl. Pkysiol. 20, 617-25.
(i937)- J- exp- Biol. 14, 178-86.
GUNN, D. L., JENKIN, P. M. & GUNN, A. L. (1937). J. Obstet. Gynaec. 35, 839-79.
GUNN, D. L. & KENNEDY, J. S. (1936). J. exp. Biol. 13, 450-9.
HOFFMANN, R. W. (1926). Handb. norm. path. Physiol. 17, 644-58.
KALMUS, H. (19380). Riv. Biol. 24, 191-225.
(19386). Z. vergl. Pkysiol. 25, 494-508.
MELLANBY, K. (1940). J. exp. Biol. 17, 278-85.
NECHELES, H. (1927). Chin. J. Physiol. 1, 143-56.
PICARD, F. (1912). Bull. set. Fr. Belg. 46, 235-47.
PIERON, H. (1937). J. Psychol. norm, pathol. 34, 398-412.
PUMPHREY, R. J. (1940). Biol. Rev. 15, 107-32.
RAU, P. (1924). Trans. Acad. Sci. St Louis, 25, 57-79.
RICHTER, C. P. (1927). Quart. Rev. Biol. 2, 307-43.
SCOTT, W. N. (1936). Trans, roy. ent. Soc. 85, 303-30.
STEIN-BELING, I. VON (1935). Biol. Rev. 10, 18-41.
SZYMANSKI, J. S. (1914). Pfltig. Arch. ges. Physiol. 158, 343-85.
(1920). Z. allg. Physiol. 18, 105-62.
WILLE, J. (1920). Biologie und Bekfimpfung der deutschen Schabe (Phyllodromia germanica L.)
Pp. 140. Berlin: Paul Parey. Beiheft zu Z. angew. Ent. 7.