Prog. Neuro-Psychophormacof. & Hiol. Psych& 1987. Vol. 11, pp. 9-21 0278-5846187 $0.00 + .

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Printed in Great Britain. All rights reserved. Copyright 0 1987 Pergamon Journals Ltd.

STEREOTYPED BEHAVIOUR, HYPERAGGRESSIVENESS AND “TYRANNIC”

HIERARCHY INDUCED IN BANK VOLES (CLETHRIONOMYS GLAREOLUS) BY A
RESTRICTED CAGE MILIEU

GRETHE WRENSEN

Set. Hans Mental Hospital, Dept. E, Research Laboratory,

Roskilde, Denmark

(Final form, November 1986)

Abstract

Serensen, Grethe: Stereotyped behaviour, hyperaggressiveness and “tyrannic” hierarchy

induced in bank voles (Clethrionomys glareolus) by a restricted cage milieu. Prog.
Neuro-Psychopharmacol. & Viol. Psychiat. 1987, -11: 9-21.

1. This investigation was originally conceived as a further development of studies

made with the “amphetamine model” of psychosis. Since amphetamine itself seemed to
be the most important flaw in this model, it was judged promising to study abnormal
behaviour in animals elicited without the use of drugs.
2. Bank voles were placed in two different cage milieux, a restricted milieu approach-
ing usual laboratory conditions for experimental animals, and an enriched milieu
offering opportunities for satisfaction of natural behavioural (ethological) needs.
During an experimental period of 6 months normal behaviours (known from the
ethogram of the bank vole) and abnormal behaviours were recorded in eight 5-min
observation periods of each cage. Social structure (hierarchy) was studied by the
intruder method and social function (peaceful or tyrannic hierarchy) by inspection
of the subordinate voles for wounds.
3. Stereotypies, hyperaggressive behaviour and tyrannic hierarchy, replacing normal
behaviour and social structure, were seen frequently in the restricted milieu, but
only occasionally in the enriched milieu (differences highly significant).
4. It is concluded that these findings indicate possibilities for further development
of the basic studies of abnormal behaviour made by means of the “amphetamine model”
of psychosis. Opportunities exist for investigation of the interaction of milieu
factors with genetic , pharmacological and other factors relevant to the develop-
ment, prevention and treatment of abnormal behaviour and mental disease.

Keywords : abnormal behaviour, cage milieu, hyperaggressiveness, model psychosis, social

hierarchy, stereotypy

Introduction

Stereotyped, hyperaggressive and other abnormal behaviour elicited by amphetamine has

been studied for many years in this laboratory for comparison with psychoses, and this
work has proved useful i-a. in the study of antipsychotic drugs and for the development
of the dopamine hypothesis in psychiatry (Randrup et al, 1973, 1986, Marchais and
Randrup 1983).

9
10 G. Sorensen

The most important flaw in the “amphetamine model”, however, seems to be amphetamine
itself, and we therefore found it important to investigate stereotyped and other abnor-
mal behaviour elicited without the use of drugs.

It is well known that many species, e.g. zoo and farm animals, show stereotyped acti-
vities in restricted environments (Draper and Bernstein 1963, Berkson 1967, Fox 1965,
1968, Odberg 1976, 1978, 1981, 1986, Kiley-Worthington 1977, Sharman 1979, Ridley and
Baker 1982, Kennes 1984, Korsgaard et al 1985). Besides, the present author has often

observed this behaviour in several species of wild mice and voles in captivity.
“Stereotypy” has been defined differently by various authors, but usually denotes a
frequent and apparently aimless repetition of certain behavioural elements. Hyperag-
gressive and other abnormal behaviours are also reported repeatedly from observations
of various animal species in restricted and unfavourable environments (Ellenberger
1960, Calhoun 1962, Russell and Russell 1968, Davis 1979).

The experiments presented here were made with the bank vole. In experiments on milieu
effects, it is advantageous to work with small animals, because also an enriched (more

natural) cage milieu can be established in a limited space.

The specific objective of our experiments was to compare bank voles in restricted and
enriched environments, particularly with respect to development of abnormal behaviour
(individual, social and group behaviour).

Methods
Animals

Male bank voles (Clethrionomys glareolus) born in our laboratory and originating from
a stock caught in woods in Denmark and bred through several generations in the Danish
Pestinfestation Laboratory, Lyngby.

Cage Milieux

Restricted Milieu. Four males in a cage (370 x 220 x 150 mm) containing only bedding

of chaff in the bottom, foodpellets in the cover and a waterbottle.

Enriched Milieu. Three males in a larger cage (550 x 320 x 200 mm) offering opportu-

nities for satisfaction of ethological needs. The cage contains 1.a. a tubesystem in

the bottom, nest materials, hiding places, a system of twrgs allowing climbing, hazel-

nuts and many other kinds of food spread around.

In the experimental room there was artificial light from 6 a.m. to 8 p.m.

Experimental Procedure

Fourteen groups of voles were formed in the restricted milieu and 10 rn the enrxhed

milieu. The groups were formed, alternately in the two milieux when the animals reached
 Abnormal behaviour of voles in restricted milieu 11

the age of 25-30 days (subadult). Each animal could be identified by means of clippings
made in the fur. Seven animals died from various diseases during the experimental
period and, in addition, three voles (all in the restricted milieu) were killed for
ethical reasons, they were victims of hyperaggression. At the end of the experiment
there remained 51 voles in the restricted milieu with a minimum of two voles in one
cage and 25 in the enriched milieu (minimum two voles in one cage).

The experimental period was 6 months. After an initial period of 3 months, systematic
observations of behaviour were started by assessment of the social hierarchy in each

group with identification of the dominant vole. This was done by means of the intruder
method: A foreign vole of the same sex is placed in the group, and it is observed which
one of the group voles that attacks it. This one is named dominant (or alpha) and the
other voles showing no aggressiveness towards the intruder are named subordinates
(Sorensen 1980). For these observations, the voles in the enriched milieu were
transformed to smaller observation cages (the same size as those providing the restric-
ted milieu), because the opportunities for hiding in the home cages could have made a
very long observation time necessary. Bedding from the home cage was also transferred
to the observation cage providing the smell of home, which is important for the beha-
viour of the voles toward an intruder. The voles in the restricted milieu were observed
with the intruder in their home cages. The observation was continued until one or two
attacks on the intruder had occurred, this required between 5 set and 20 min
observation time.

The social hierarchy was judged “tyrannic” when one or more subordinate voles in a
cage were wounded. This was inspected weekly during the whole experimental period.

During the following three months all the cages (alternately a restricted and an en-
riched cage) were observed 8 times; each time the home cage was moved to an observation
table and touched to create a gentle noise. This method activates the animals without
frightening them so much that they hide for a long time. After a delay of 3 min, in
which the voles hide in the bedding, the cage was observed for 5 min, during which
period the voles come out from the bedding and perform exploration of the cage. Four
observations were made during the night period and four during the day period. In their
natural milieu bank voles are active both by day and night, but in the laboratory they
are most active by night. During the day observations there were, however, sometimes
improved conditions for observation of the subordinates in the restricted milieu, since
the dominants were then less active and therefore also less aggressive or
hyperaggressive. Normal and abnormal behaviours were recorded for each vole. The normal
behaviours (the ethogram) were known from previous studies (Serensen 1980).

The number of stereotyped behavioural acts were counted, they were quick and short
elements of behaviour usually repeated and readily distinguished from the normal
12 G. Sorensen

behavioural repertoire because of their automatic appearance. In some cases, however,

it was not quite clear whether a behavioural act should be regarded as stereotyped or
not. Such acts were counted separately and amounted to 8.5% of the surely stereotyped
acts. In Table 1 below they are counted as non-stereotypy. If they are counted as
stereotypy, however, or only excluded, this will make no essential difference in the
statistical evaluation (P-values becoming even more significant). Of the voles classi-
fied as non-stereotypic, seven from the restricted milieu and none from the enriched
milieu showed this type of behaviour.

Bouts of aggressive behaviour (threat postures, chasing, etc.) were regarded as part
normal behaviour, but when aggression became prolonged with many or prolonged fights or
biting, or blood stains appeared, it was recorded as hyperaggressive behaviour.

After the completion of these observations a lo-min observation of each group was
made in special observation cages. These cages were of the same size as those providing
the restricted milieu but contained only a thin layer of bedding, i.e. there were no
possibilities for hiding in the bedding. The material for the bedding was taken from
the home cage, thus providing the familiar smell, which reduces aggression. The obser-
vation cages were also without food and water. Finally, in order to see if the social
hierarchy remained stable throughout the observation period, the hierarchy and the
identification of the dominant vole were again checked by means of the intruder method.

Statistical Analysis

The Mann-Whitney U-test was used for evaluation of measurement data such as the coun-
ted number of stereotyped acts (Table 1). For enumeration data such as the number of
.2
voles in various categories the chi test was used when the smallest expected value was
5 or above, if not the Fisher exact probability test was applied. All the P values gi-
ven represent a two-tailed evaluation of the tests.

Results

Stereotypy

Quantitative evaluations of stereotyped behaviours are shown in Table 1. The unambi-

gous result is that stereotypy occurred significantly more in the restricted than in
the enriched milieu. Also during the unsystematic observations made while cleaning
cages etc., stereotypy was observed very much more in the restricted cages than in the
enriched ones, where it was seen only exceptionally. Likewise, in the observation of

each group in special observation cages without possibilities for hiding in the bed-

ding, stereotypy was seen in all but one of the groups living in the restricted cages

but only in two of the ten groups from the enriched milieu; these were the same two
groups (and the same three voles) recorded with stereotypy in Table 1. Of the 36 voles
from the restricted milieu recorded with stereotypy in Table 1 thirty-one also showed
 Abnormal behaviour of voles in restricted milieu 13

stereotypy in the special observation cages, and of the 15 voles recorded with no

stereotypy, 12 also showed no stereotypy in the special cages.

Table 1

Stereotyped Behaviour in Restricted and Enriched Cage Milieux.

Results from Eight 5-Min Observation Periods

Restricted Enriched
cages cages P

No. of stereotyped acts

per vole
average 158 3.6 << 0.00006
range O-1081 O-69

No. of voles
Stereotypy 36 3
No stereotypy 15 22 << 0.002

No. of groups (cages)

Stereotypy 14 2
No stereotypy 0 8 < 0.0002

Qualitatively, five types of stereotypy were observed, each with several variants:
1. Backwards somersaults, this was the most frequent type.
2. Swaying forth and back in the rearing position against the short wall of the cage
opposite to the foodpellets.
3. Running forward along one half of one of the longer walls in the cage and retur-
ning by jump or somersault.
4. Crawling or rolling at the wire-netting of the sunk part of the lid that contains
the foodpellets.
5. Running following a fixed route (often round the circumference of the smaller ca-
ges providing the restricted milieu) with rearing in one particular corner.

Characteristic for all the stereotyped acts was their automatic appearance, the ani-
mals seemed “cut off from the outside world”, they did not react to stimuli from the
outside.

As mentioned in the Methods section, some behavioural acts could not be clearly re-
cognized as stereotypic and were recorded as doubtful stereotypies. These were:
a. Somersaults combined with much, and rather slow, crawling on the lid.
b. Hyperactive running not following a fixed route.

Many animals showed several of these types. Alternations of two stereotyped acts were
also observed. For instance, alternation of 1. and 2. was often seen. One vole showed 6
types of stereotypy: swaying (the most frequent type in this vole), rearing in a corner
without swaying, and backwards somersaults combined with much crawling on the lid,
14 G. Serensen

with only a little crawling on the lid, with just touching the lid and without touching
the lid,

As the stereotyped acts were repeated many times, they often left marks on the cage
walls (clear plastic). Thus stereotypy no. 2. produced a pattern similar to that made
by a windowscreen wiper in a car, and also stereotypies nos. 3., 4. and 5. gave rise ta
recognizable patterns of dirt on the plastic walls. Stereotypies nos. 1. and 5. left
characteristic patterns in the bedding of the cage.

Df the 14 dominant voles in the restricted milieu 12 were stereotyped (one other vole
was questionably stereotyped), and of the 37 subordinates in this milieu 21 were ste-
reotyped (four others quest~anably so). This trend is opposite to the expectation that
stereotypy might be observed most frequently in the subordinates, because of the stress
exerted by hyperagqressive dominants.

In preliminary experiments with breeding females in the restricted milieu, it vjas ob-
served in several cases that the stereotyped behaviour interfered with the care of the
young. The mother often moved the young to other places in the cage and sometimes per-
formed stereotyped activities carrying a young in the mouth. Such behaviours usually
led to the death of the young. This illustrates the severity of the abnormal behaviour.
These voles did not only perform stereotypies in their “spare time”; normal behaviour
was severely interfered with and to such an extent that it could have lethal effect for
the young.

Tyrannic Hierarchy and Hyperaqqressive Behauiour

A dominance hierarchy (one dominant per group) existed in all the groups in both
milieux. In most cases it could also be seen that the hierarchy was “linear”, i.e. from
the degree and amount of defensive behaviour made by the individual subordinates, it
appeared that one of them was second in rank (8) and another one was lowest in rank
(~1. The intruder experiments and observations of aggression during the eight 5 min ob-
servation periods shomed that the hierarchy remained stable throughout the observation
period; the vole that was dominant at the beginning of t.his period remained so at the
end in all cages except two. In one of the two exceptional cages the dominant
(a) became ill (see footnote in Table 2) and in the other (an enriched cage) it died.
In both cases the former B became et. in some cages the hierarchy became tyrannic, this
happened significantly more often in the restricted than in the enriched milieu as
shown in Table 2.

A total of 18 subordinate voles were hurt in the tyrannic hierarchies, 17 in the

restricted milieu and one in the enriched milieu. These vales were wounded at the hind

part of the body or at the tail, three of them (all in the restricted milieu) so
severely that they had to be killed during the experiment for ethical reasons. In some
cases the tail was totally bitten off.
 Abnormal behaviour of voles in restricted milieu 15

Hyperaggressive behaviour as defined in the Methods section was performed only by do-
minant voles. It was looked for in the 5-min periods of systematic observation, but was
seen only in the restricted milieu as shown in Table 2. Sometimes the hyperaggressive
behaviour dominated the whole observation period, and some observations had to be in-
terrupted (by addition of more bedding (hiding) material) in order to save subordinate
voles from being killed. In some cases the aggression was directed at one particular
subordinate, in other cases at several. In one cage it was observed that the dominant
attacked the lowest ranking subordinate first and then the two others in ascending rank
order. In 7 of the 8 cages where hyperaggressive behaviour was observed, the hierarchy
was judged tyrannic.

Table 2

Tyrannic Hierarchy and Hyperaggressive Behaviour

in Restricted and Enriched Cage Milieux

Restricted Enriched
cages cages P

No. of groups (cages)

Tyrannic hierarchy
(21 subordinate vole
wounded) 1oa lb
c 0.005
Peaceful hierarchy
(no vole wounded) 3 9
No. of dominant voles
Hyperaggressive
behaviour observedC 8 0
co.01
Hyperaggressive
behaviour not observedC 6 10

aIn one additional cage an atypical tyrannic hierarchy developed. Towards the end of
the experimental period the CI vole became ill and the B vole attacked and wounded it.
After this the former B became a, and during the same period the hierarchy shifted from
Beaceful to tyrannic.
cThe tyrannic hierarchy in this group was highly atypical, see case story in text.
During the eight 5 min observation periods.

Sexual behaviour (a special type of following sometimes with subsequent mounting or

attempt to mount) occurred occasionally among these male voles. In three voles it deve-
loped into hypersexual activity (repeated many times). This happened in the one en-
riched cage with tyrannic hierarchy and in two restricted cages also with tyrannic hie-
rarchy. All the three hypersexual voles were subordinates and all three directed the
activity selectively at one of the other voles (in the enriched cage and in one of the
restricted cages at the dominant). When this one performed stereotyped behaviour such
as backwards somersaults, the mounting could not be completed, but many attempts were
made.
16 G. Smensen

Case story. In the only group in the enriched milieu with tyrannic hierarchy the
hierarchy was also very atypical. The cx vole in this group was not hyperaggressive. The
life of the group was dominated by the extreme hypersexuality of the 8 vole, directed
at the o. The 8 vole spent much time chasing the third vole, the W, away from the
CL, and just in such a situation hypersexual and hyperaggressive behaviour may be mixed.
Even though subordinate voles almost never show aggressive behaviour, in this group it
was the 8 vole and not the cx that injured the third, the w vole.
The 6 vole was very abnormal. It was both hypersexual (only against the a), hyperag-
gressive (only against the w), very stereotyped and hyperactive. In periods where B was
totally occupied by performing stereotypies, o dared to approach ~1, which was friendly,
and these two could lie together and perform the friendly social activities listed in
Table 3. But most of the time w had to live an isolated life because of 0. In short
observation times it is then seen that two voles (CY and B) live together, while w
“isolates itself”. This may lead to speculations about animal models of autism. In this
case, however, after prolonged observations (hours) of the daily life of this group, it
must be concluded that it was another, very abnormal vole, the B, which was the cause
of the almost totally isolated life of W. w itself appeared in all respects quite
normal.

It is extremely unusual that a vole in a group lives isolated. Even in the most
tyrannic hierarchies in the restrlcted milieu the animals slept close together (inlcu-
ding the wounded vole and the one that had wounded it) and had many friendly interac-
tions in spite of everything.

Normal Behaviour

In order to study abnormal behaviour in a species of animals, it is necessary to have

basic knowledge of the normal behaviour of that species. Those animals that do not show
the investigated forms of abnormal behaviour in this experiment cannot automatically be
characterized as normal unless they actually show normal behaviours (they might be
apathetic, showing no behavioural activity at all).

Therefore, in every 5 min observation period, all behaviour of each vole was recor-

ded. Because the behavioural repertoire of the bank vole is large, the normal beha-
viours were not recorded quantitatively. It was only recorded if a behaviour was
sent or not.

The behaviours observed are listed in Table 3; they correspond to the ethogram of the

bank vole studied earlier (Sorensen 1980).

 Abnormal behavior of voles in restricted milieu 17

Table 3

List of Normal Behaviours of the Bank Vole

in Restricted and Enriched Cage Milieux

Normal behaviours observed

Behaviours possible in both restricted and enriched milieux:

Social behaviours
Oominant behaviours
Aggression, short bouts of threat postures, chasing and attack
Other behaviours characteristic of dominant voles:
Hindleg scratch, the flank glands are scratched by a hindleg, a behaviour of scent
marking
Displacement digging, violent digging, displacement activities are shown by anima IS
either thwarted or in conflict
Dancing, small tripping staccato-steps, often combined with pilo-erection, a sign
of excitation
Pendulum sniffing, the head is swung rhythmically from side to side, a conflict
behaviour
Defensive behaviours, listed in the order of increasing aggression inhibiting signal
strength
Intention movements of defensive posture, i.a. one foreleg lifted
Defensive posture, upright on hindlegs
Submissive posture, head raised higher than in defensive posture so that the throat
is shown
Flight
Boxing, two voles rearing, mutual boxing with forelegs.
Behaviours which may form parts of friendly social interaction
Sniffing at another vole
Grooming another vole
Sleeping together
Ordinary following
Sporadic sexual contact, a special type of following sometimes with subsequent moun-
ting or attempt to mount
Individual behaviours
Behaviours which may form parts of exploration
Walking
Runnino
Rearing
Crawling on the wire-netting of the lid
Sniffing around in the cage
Hidinq
Buried in the bedding of chaff
Hanging in the wire-netting of the lid
Lying under the sunk part of the lid, containing food-pellets
Sitting visible
Eating
Drinking
Sleeping alone
Digging
Grooming
Displacement grooming
Freezing
Table 3 continued on the next page
18 G. Ssrensen

Running in panic, one vole, but may spread to all in a group, often followed by
freezing

Behaviours possible only in the enriched milieu:

Climbing in twigs
Gnawing bark of twigs, some voles also eat it
Searching food (apple, carrot, sunflower seeds, corn, peanuts, hazelnuts)
Opening hazelnuts
Cracking of seeds
Hoarding
Running through the tubesystem
Nest-building, using the materials provided (paper and three types of glass jar)
Marking behaviours on glass jars (smooth surfaces are preferred for marking). Particu-
larly two types of marking behaviour gave rise to the secretions visible on the glass
surfaces. These were hindleg scratch with activation of the flank glands (performed by
dominants only) and urine marking (performed mostly by dominants)

During much of the observation period the voles in the enriched cages were occupied
by normal behaviours, in particular such behaviours that were made possible by the en-
richment of the milieu. Thus it was ascertained that the voles made use of the opportu-
nities of this milieu. Gnawing of bark was prominent with some voles, hoarding with
others. The dominants showed the forms of dominant behaviour not constituting direct
aggression, which forms we regard as contrasts to hyperaggressive behaviour and as
characteristic of a peaceful (functional) hierarchy.
In the restricted cages, however, much time was occupied by stereotyped and hyperag-
gressive behaviour. Because of a hyperaggressive dominant vole, many subordinates i.a.
quite normal ones had to spent much time hiding in the cage bedding or by hanging con-
stantly in the wire-netting of the lid. For this reason, among others, normal beha-
viours were seen only sporadically in the observation periods. When there was an oppor-
tunity, however, the voles utilized the time for normal behaviours, so that all the
behaviours listed in Table 3 were observed.

Discussion

Stereotypies

Stereotyped behaviour of bank voles (Clethrionomys glareolus bred from a stock caught
in the neighbourhood of Edinburgh, Scotland) in a restricted cage milieu, but caged
singly, has earlier been studied in some detail by Ddberg (1981, 1986) and by Kennes
(1984). Qualitatively, these authors report the same types of stereotyped behaviour as
we have observed: somersaults (“looping”), “windowscreen wiper”-like swaying in the
rearing position (“ruitenwissers-stereotypic”), and running forth and back following a
fixed route. In addition, however, they observed jumping (often in a corner of the
cage) and in their experiments, this was the most frequently occurring type of
stereotypy. It was sometimes seen in alternation (combination) with running forth and
 Abnormal behaviour of voles in restricted milieu 19

back. In preliminary observations the present author saw much jumping stereotypy in
caged bank voles caught in the North of Sweden (Norrland), while bank voles from the

South of Sweden (Sk&e, neighbouring Denmark), observed at the same time in identical

cages and in the same room, showed only little jumping. The jumping stereotypy may thus

be a characteristic of certain, geographically located strains of bank voles (a more

general description of differences between the bank voles from North Scandinavia and
those from South Scandinavia and Central Europe is given by Hansson, 1985). Quantita-
tively, Odberg (1981, 1986) and Kennes (1984) found, as in our experiments, large
individual differences with high frequency of stereotypy in some animals.

Causative Milieu Effect

The high frequency of stereotypy in some individuals, replacing normal activities,

and the severe distortion of social behaviours seen in our experiments indicate that
the restricted milieux elicit severe behavioural pathology in the bank voles.

Stereotyped behaviours of various types and abnormal social behaviours have been ob-
served in many other mammalian species in captivity, and it is the general belief of
ethologists that these behaviours are caused by the restricted milieux, which do not
offer opportunities for satisfaction of natural behaviour needs (see the references in
the Introduction). Our experiments provide some more definite experimental confirmation
of this belief.

Possible Generalizations to Other Mammalian Species and Man

With due consideration of the differences between these observations in many species,
we may contemplate the possibility that they represent a behavioural homology in the

zoological evolutionary sense. Could this possible homology be extended to the species
of man? Stereotypy and abnormal social behaviours (including hyperaggressive and
hypersexual activities) are certainly known in man. They constitute parts of various
mental diseases, in which milieu factors may well play an etiological role. In view of
the wide variations of the available observations in both man and animals, more speci-
fic comparisons between the species cannot be considered at the present stage, however.

With respect to the stereotyped behaviour elicited by stimulant drugs, a zoological

homology seems to exist between all mammalian species including man. The stimu-
lant-induced stereotypies have been compared with those seen in the functional psycho-
ses, schizophrenia, depression and mania. These comparisons have proved useful for cer-
tain investigations in biological psychiatry as mentioned in the Introduction. Possib-
ly, the study of abnormal behaviours, elicited without the use of drugs, may have po-
tential for further advancement. One reason for the relatively few reports of studies
of milieu-induced abnormal behaviour compared to the reports on drug-induced abnormal
behaviour may be that some of the most freuqently employed experimental animals, labo-
20 G. Serensen

ratory strains of mice and rats, seldom show milieu-induced stereotypies. In contrast,
these are shown by many other species and strains of mice and voles.

Conclusions

The experiments show that the bank voles develop stereotypy and abnormal social beha-
viours in a restricted cage milieu, and that it is possible to avoid these behaviours
in captivity by an enrichment of the milieu, which can readily be established in the
laboratory. Possibilities therefore exist for further and more detailed laboratory
studies of the milieu factors and also of interactions of these factors with genetic,
pharmacological,neurochemical, and other factors relevant to the development,
prevention and treatment of abnormal behaviours and mental disease.

Acknowledgements

The author wishes to thank Gjmva Brendstrup's fund and Asta Alverde's fund for finan-
cial support, and Dr Axe1 Randrup for his continuing interest in this work.

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Grethe Serensen
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