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" DISPLACEMENT " BEHAVIOUR AND "PSYCHOSOMATIC" DISORDER
[DEC. 10,
ORIGINAL ARTICLES " DISPLACEMENT " BEHAVIOUR AND "PSYCHOSOMATIC" DISORDER S. A. BARNETT
THE object of this paper is to suggest a relationship between the " displacement behaviour " described by ethologists,* and what clinicians sometimes call " psychosomatic disorder." The argument leads to two general sets of proposals. The first concerns the necessity for a particular line of research ; the second suggests that the present terminologies of psychologists and psychiatrists are often a handicap on progress, and in particular that their interpretations should be strictly monist. Behaviour
Displacement behaviour is the performance of a behaviour pattern, in biologically inappropriate circumstances, when an appropriate action is prevented or the input to the central nervous system departs from the optimum. The behaviour does not directly serve the survival or reproduction of the individual or the species : biologically, it is irrelevant to the situation in which it is performed. Such behaviour has been mainly studied in fishes and birds (cf. Armstrong 1950, Tinbergen 1952), but it certainly occurs also in mammals. For example, a wild rat feeding, disturbed by another rat of a different species, makes " aggressive " noises and motions towards the interloper ; the latter flees ; the first rat then, instead of immediately resuming feeding, briefly grooms its face with its forepaws-an action which it would not ordinarily perform in the middle of feeding (Barnett and Spencer 1951). Deprived of the object of its aggression, the rat resorts to an alternative, and inappropriate, behaviour pattern. A slightly different example is that of wild rats during a series of copulations : at intervals both male and female may make a rapid to a feeding-point, feed hurriedly (and in a quite atypical manner) for a few seconds, and then of courtship and coitus resume the usual pattern (unpublished). In general, typical displacement activities occur in three types of situation. First, if an animal is interrupted in, or prevented from performing, a specific act, it may substitute another act, as in the example above of the aggressive rat. Second, seemingly irrelevant behaviour may occur if an animal has two incompatible types of behaviour open to it, as with herring gulls which resort to grass-pulling when the " appropriate behaviour could be either to attack or to run away (Tinbergen 1953). Third, the actual attainment of a goal-the performance of a consummatory act-may, strangely enough, be followed by irrelevant behaviour, as in the example above of the rats between copulations. Displacement activities are often incomplete actions, as in the nest-building movements of gulls or the grooming of rats ; but they need not be so. They are often hurried, as in the displacement feeding of rats. The so-called vacuum activities, in which an animal performs an action, such as a display, in the absence of an appropriate object, may possibly be responses to suboptimal stimuli ; they are comparable to displacement acts. Like them, they seem to reflect an overflow " of motor output. This point will be discussed in more detail later. Another phenomenon closely similar to displacement is the " redirection activity " , (Bastock et al. 1953), in which an animal directs the appropriate movement towards an inappropriate object. The authors quoted refer only to examples where the appropriate object is still present ;but a similar effect has been observed when
move
"
*
Ethology is the scientific study
6902
object is no longer available : a rat deprived of the object of its aggression may turn on a member of its own colony (Barnett and Spencer 1951). Bastock and her colleagues draw attention to the analogy to the " object of psycho-analysts. As Freud (1949) displacement
wrote :
ZOOLOGY, GLASGOW UNIVERSITY
Displacement
the
"
M.A. Oxfd LECTURER IN
1955
of animal behaviour.
We have found that instincts can change their aim (by and also that they can replace one anotherthe energy of one instinct passing over to another.
displacement)
The examples of displacement behaviour given above may be unconvincing to the critical reader who has not himself observed such behaviour. A critical attitude is very necessary in this field ; but fortunately, as we shall see below, good examples, quantitatively analysed, do exist. Learnt
Responses Resembling Displacement Behaviour
In the classical examples of displacement activity, the substituted behaviour always belongs to an alternative stereotyped (" innate ") behaviour pattern. This is reflected in the first definition of the term displacement activity " given by Thorpe (1951) : such activity is said to result from the activation ... of the action pattern belonging to another instinct " ; but alternative, wider definitions are given. For the present argument it is convenient to apply a wider definition, such as that given at the beginning of this paper. We can then consider whether there are learnt responses in mammals which, though very different from displacement responses of the usual sort, can properly be put under this general heading. Maier (1949) describes experiments in- which albino rats were faced with an insoluble problem in a Lashley jumping apparatus. Whatever solution they chose, they were "punished" in half of their jumps on the average. Moreover, they were forced to jump. Such rats tended to develop " behaviour-fixations " : they commonly, after some time, jumped, say, always to the right, regardless of the stimulus situation ; indeed, they did so even if the door was open and the food was visible on the left-hand side. Maier comments : "
It appears that the fixated group develops some kind of adjustment to the test situation and is therefore able to prevent
emotional tensions.
Fixated rats also showed less resistance to making a The detailed interpretation of Maier’s results remains difficult and to some extent controversial, but there is no doubt, from his and other experiments, of the reality of behaviour fixations " (Russell and Pretty
jump.
"
1951). Better-known examples of abnormal " learnt are those described under the heading of responses " experimental neurosis." In these, too, an insoluble problem may be presented, but the result is either an excessive undirected activity, or a general inactivity which may include refusal of food. The variety of these " abnormal " acts is considerable (cf. Russell 1950), and a fuller analysis is needed before the validity of the comparison suggested here can be assessed. However, despite the diversity of these responses, there seems to be at least an analogy to displacement behaviour in all these examples.: certainly, where there is a fixated response this provides a regular pattern of motor output in frustrating circumstances-i.e., in conditions in which a biologically appropriate response cannot be made. The
Consummatory
Situation"
displacement activity has been described substitute for an alternative activityif an animal is unable to perform or to complete a consummatory act, such as fighting, feeding, or copulation, it performs Usually,
as a
a
1204 other act instead. But this does not always apply. Moynihan (1953) studied nest-building movements and preening movements in gulls during incubation of the eggs. Nest-building movements were shown to increase some
if one or more eggs were removed, and the number of such movements in a given time increased in proportion to the number of eggs removed. Preening increased removed. An all when the were markedly only eggs of was the feature this work quantitative important expression of displacement behaviour: in this case, unlike many, subjective impressions could be excluded. (An earlier good example is that given by Tinbergen and van Iersel (1947) on fanning in Gaste1’osteus aculeattis.) Both types of movement were typical of displacement activities, in that they belong to stereotyped behaviour patterns, and that they occurred at inappropriate moments. They differed from the usual examples, however, in one notable respect : they were performed, not in place of a motor act, but when a specific sensory situation was altered. The substitute activity reflected the absence of what might be called a " consummatory situation " (cf. Thorpe 1954). The importance -of the afferent input is also shown by the work of H. Spurway and J. B. S. Haldane on the air-gulping of the climbing perch (Allabas testudineus). Haldane (in litt.) writes : Our fish got into a flap when they breathed N2. To men this certainly has not got an odd smell or taste. It makes you feel dizzy a few seconds after breathing it. Probably the brain is also affected by afferent impulses from the carotid sinus. The fish have done their consummatory act, but have not got the normal "feedback." This disturbs them much more than if they are mechanically prevented from carrying out the act.
It
reasonable to suggest that the concept of consummatory situation or state is more fundamental and more general than that of consummatory act. The latter marks the end of activity, or at least of a specific group of activities, such as those related to mating. But the performance of such an act must be accompanied by specific internal changes, such as those involved in filling the stomach ; and these changes, by evoking a particular afferent input, thereby alter central nervous activity and so the output to the muscles. seems
"
Kennedy (1954),
Instinct " in Man
in
a
polemical discussion of ethological
theories, suggests that the cessation of activity on the attainment of a consummatory act is "reflex "-i.e., due to an input of impulses from the periphery. Bastock et al. (1953), in the paper already quoted, ’suggest that "
consummatory stimuli," and especially proprioceptive feed-back, are important in what they call the " conof drive." (Since proprioceptive fibres make one-third to one-half of those in a nerve supplying a muscle, it is clear that they deserve a good deal of attention in the study of behaviour (cf. Lissnian 1950).) Disregarding the particular terminologies used in these two communications, each, from different points of view, suggests that we must look to the afferent input if we are to make a beginning on the physiological analysis of displacement behaviour. It is therefore worth while to consider a little further the subject of the influence of afferent impulses on certain sorts of behaviour. Here is an example (Hardy 1954) from a physiological study in which engineering and psychiatric terms are both used in a physiological
sumption
up
context : From the
point
of view of
servo
control of skin
tempera- ,
ture, it is worth noting that thermal sensations have strong when the average skin temperature is below 32° or above 34°C ....This " affect " of thermal sensation is probably one of the most powerful drives for. physiologic servo temperature regulation in the presence of external thermal stress. elements of
unpleasantness
or
pleasantness
in
A noxious stimulus of the sort described here evokes man either a general restlessness (analogous to
appetitive behaviour), or a learnt response which tends to reduce the unpleasantness. If there is no learnt response available, as in a young child, the restlessness may lead to learning-perhaps of the trial-and-error variety. In any case, the situation described in the quotation above is one in which a particular sensory condition induces and this activity ceases on the attainment activity, of a " consummatory situation "-in the example, a particular skin-temperature. If the analysis above is correct, it may serve to clarify the concept of instinct " as applied to man. In ethological writing the term instinct " is today commonly (not always) applied to a stereotyped activity. Depending on the author, the activity may be a single consummatory one ; or it may be a complex series of acts, of which the, "
"
consummatory one is the last. The term instinct, so used, doesnot apply to any internal agency in behaviour, " drive " may be used to refer to internal influences which lead to the activity. In man there is virtually no instinctive behaviour in the above sense. The infant breathes, sucks, and cries ; he also, perhaps, at 3-6 months, smiles in response to a particular " sign stimulus " (Spitz and Wolf 1946) ; but apart from reflexes, later behavioural performances are learnt. Yet human beings act most strenuously to achieve certain ends. These ends, regarded at the physiological level, are the attainment of certain sorts of neural input-of consummatory situations. The motor procedures which attain these ends are variable and learnt. Hence, to take an example from Freud, the word Trieb, usually translated as " instinct," refers to behaviour which ceases once a particular nervous input-e.g., from a full belly or empty seminal vesicles-has been achieved. Or, to put the obverse, the behaviour is activated by the input from an empty belly or full seminal vesicles. These examples must not be taken to imply that all such behaviour is activated peripherally, by the stimuli of the moment, or that it is innate." In higher mammals, especially man, with a vast mass of internuncial neurons in the cerebral cortex, it seems that learning-or at least previous experience--may lead to forms of neural organisation in which the activation of the motor output is largely within the central nervous system itself. Behaviour may then be both " spontaneous" and a product of learning. As Hebb (1949) has pointed out, it is this iliternuncial mass which makes possible the predominance of learnt behaviour in man, and his relative independence of the immediate sensory situation. Whitehorn (1952) approaching this topic from the point of view of a psychiatrist, has remarked that, despite the evidence from social anthropology for the cultural rather than the innate patterning of human behaviour, the term instinct is still retained by psychiatrists. He points out that the psychiatric use of the term represents a shift of meaning [from that of ethologists] to " inner driving forces correlated with biological needs," and that " this inner impulsive force is experienced as emotion." He further suggests that " emotion " is negatively correlated with instinctive behaviour : the less an animal’s behaviour is automatically determined by fixed neural patterns, the more it is necessary that powerful internal agencies should ensure that effective behaviour patterns are acquired (learnt) by each individual. Emotion in this context is the subjective counterpart of the neural changes due to a certain type of (unpleasant) sensory
though the term
"
"
"
input. Whether "
or
not
we
agree with this
emotion," Whitehorn’s analysis
hypothesis about brings out clearly
the notorious fact that the term " instinct" is what Ogden and Richards (1931) call a nomad : it wanders
1205
meaning to meaning. wanderings are not yet over.
from
Psychoneurosis "
and "
It is to be feared that its
Psychosomatic"
Disorder
Behavioural Cha.7zges In the virtual absence of instinctive behaviour in man, one might think it idle to look for displacement activities in human behaviour. The realisation of the importance of consummatory state," however, alters the picture. The analogy between displacement behaviour in other animals, and psychoneurotic behaviour in man, has indeed often been pointed out, notably by Armstrong (1950). Armstrong quotes a passage on hysteria in which Freud writes that in such states " we have an excessive innervation (in typical cases a somatic innervation), sometimes of a sensory, sometimes of a motor character, either as an excitation or an inhibition." This refers of course to the fact that the behaviour called hysterical covers a wide range, from outbursts of weeping to a functional paralysis or anoesthesia. Armstrong comments that both in displacement behaviour is a somatic and conversion hysteria " the performance in both the activity means of release from tension ; is incomplete and apparently detached from normal The same might be said of affective experience." obsessional behaviour. However, the fact that hysteria includes phenomena of inhibition, as well as oert activity, must make one hesitate to -regard it, in any simple way, as the human counterpart of displacement behaviour. As we shall see, this sort of question can be answered, or indeed effectively asked, only in neuro...
.
physiological terms. Psychoneurotic behaviour
in general is of course instinctive " : it does not consist of rigidly stereotyped activities common to a whole species or variety ; in a sense it is learnt. Behaviour, such as smoking or chewing, which is clearly analogous to displacement behaviour but which ought not to be labelled psychoneurotic, is quite obviously learnt. The same applies to the thumb-sucking of an infant. (Smoking and the rest could be labelled " abnormal," whatever that may mean, but there are other human responses, of biological value and in no possible sense pathological, which may have originated, in evolution, as displacement behaviour. Spurway and Haldane (1953) suggest that speech is a derivative (" ritualised ") of displacement breathing. Another form of communication, the movements of the facial muscles of expression, may well have originated in the same way, and have acquired increasing importance with the development of human not
"
perhaps
society.) a warning is perhaps required. It is sometimes overlooked, that human behaviour is so much more complex than that of any other species, that it is in some respects qualitatively This is evident in as well as quantitatively different. our use of symbols, not only in language but also indirectly. If a woman is sick of her husband, she may not say so or give any verbal indication that she is aware of it : but she may persistently vomit while her husband is at home. (I am indebted to Dr. R. D. Laing for this example.) Such a response can be interpreted according to a variety of systems. It could be described in neurological terms, perhaps with the addition of expressions derived from studies of conditioned reflexes ; it could be compared to a displacement response ; or it could be characterised in the more poetic or symbolic phrases (no sneer is intended) of Freudian analysis. All no doubt have some validity, but to say that psychoneurosis resembles displacement behaviour certainly does not " explain " it. The usefulness, if any, of the sort of comparison made here is of two kinds. First, if similar processes are found in man and other species, research possibilities
At this
obvious,
point
but
"
"
may be widened.
The proposal is then, in effect, a that similar internal " mechanisms " underlie all the various phenomena compared. Secondly, the attitude to particular aspects of human behaviour The objective, nonmay be changed for the better. ethical, approach to behaviour attempted by ethologists is superior to a subjective moralistic one, both heuristically and humanely. Yet if a person is said to have a mental " disorder, for many-even for some doctorsthis amounts to a moral stigma or aesthetic flaw, such as is not felt to exist in the person with, say, peptic ulcer or tuberculosis. (It is perhaps instructive to realise that a great variety of so-called perversions," from masturbation to fetishism, have been observed in various animal species. Homosexuality, in particular, is a widespread phenomenon among vertebrates.) There is, indeed, on these grounds alone, a case for revising the terminology of what, in my view, should be called behaviour disorders but which at present are usually described as mental illnesses. (It is a strange fact that in psychiatry the term " behaviour disorder ’’ is reserved for a single " disconduct group of antisocial activities--the orders which are also lumped under the head of psycho-
hypothesis, namely,
"
"
"
"
pathic personality.") Soinatic
-Responses examples
In all the
output
so
from the central
far
given,
nervous
there is
an
altered
system. Instead of the
"
normal " output, another is substituted. The alternative motor pattern may be "instinctive" or not; it may be alternative to an active pattern of behaviour or to an inactivity ; it may itself be an inaetivity-an inhibitory effect. All these statements apply also to the so-called psychoIn these, the substitute somatic disorders of man. output is not to the somatic musculature, but is as a rule autonomic. There is, however, no part of the body and no organ which is outside the scope of these disorders. This is not surprising, since every part of the body has a blood-supply partly under autonomic control and carrying hormones of which the secretion is influenced by the autonomic nervous system. Wolff (1953) gives an excellent review of these condí tions. An example is the pattern of response of the stomach to the ingestion of food : increased bloodflow, increased motility, and secretion of acid. These may all occur, quite inappropriately, in conditions of stress. Another familiar response to stress is diarrhceahyperactivity of the gut muscles. Wolff comments that the person -with violent diarrhoea may be outwardly calm, sweet-mannered and serene.’A third standard example is asthma, in which the response is an inappropriate contraction of the smooth muscle of the lung. Alexander (1952a) describes these responses as follows :
biologically appropriate
or
"
Whenever the expression of competitive, aggressive, and hostile attitudes is inhibited in voluntary behaviour, the sympathetic adrenal system is in sustained excitation. The vegetative symptoms result from the sustained sympathetic excitation, which persists because no consummation of the fight or flight reaction takes place in the field of coordinated voluntary behaviour, as exemplified by the patient suffering from essential hypertension who in his overt behaviour appears inhibited and under excessive control. Likewise in migraine headache, the painful attack may terminate within a few minutes after the patient becomes conscious of his rage and gives open expression to it.
It is easy to see that, in one aspect, the psychosomatic responses fit quite readily into a Pavlovian framework : they can be interpreted in terms of conditioned reflexes, once it is understood that a conditioned reflex is an exceedingly complex phenomenon (cf. Hebb 1949). In each instance the primary response is innate," and would ordinarily be called reflex. In the innate character of the response, these disorders show a closer similarity to the ordinary displacement behaviour of "
1206 birds and fishes, than is shown by typical " psychoneurotic " behaviour. They are, however, different in three obvious ways from what we may call classical displacement activities.
First, they are very variable within the species. There is evidence, as might be expected, of genetically determined differences in the ways in which individuals react to stress : in one man a given stress may induce, say, diabetes, in another similar conditions may lead to hypertension. This is perhaps yet another example of the exceptional polymorphism of the human species
(cf. Haldane 1949). Second, the environmental causes of these disorders is much more complex and diffuse. Even when there is an immediate precipitating cause, it acts on a central nervous system already prepared by previous experience. This of course reflects the size and complexity of the human nervous system, with its enormous accumulation of connecting neurons. Third, as already mentioned, the output is not somatic motor, but is visceral. Here again, evidently, we have a reflection of a distinctively Primate (especially human) evolutionary development-namely, the importance of the cerebral cortex, and especially of the frontal lobe, in controlling autonomic output. Fulton (1949) writes : I am inclined to regard the recognition of the frontal lobe as the primary autonomic centre of the forebrain as perhaps the most significant disclosure of recent years in the field of scientific medicine, for it affords at long last a sound physiological background for that still shadowy subject of psychosomatic medicine." Despite these differences, the psychosomatic disorders and the psychoneuroses on the one hand, and displacement activities on the other, all seem to exemplify "
general
feature of
complex nervous systems-that, excitatory state which would ordinarily produce a motor discharge, and if the motor output cannot take place in the biologically " normal " fashion, then it will follow some other path. The other path a
if there is
may be
"
rigorously
an
innate " or fixed for the
"
learnt," but, if learnt, may be
particular
The Problem of
"
individual.
Nervous
Energy
"
T enninology An early
discussion of " nervous energy " was that of Freud. As Ernest Jones (1953) points out, Freud in his early psychiatric works wrote in terms of " an economics of nervous energy." (At that time, too, Freud adopted a notably " psychosomatic " approach to his subject. In interpreting anxiety neurosis as a of sexual consequence deprivation he points out that increased respiration-rate, palpitation, sweating, and congestion occur both in coitus and in anxiety states. As Jones shows, the history of Freud’s thinking is of great interest for the study of ideas on the relation between " soma " and " psyche.") What, then, is the nature of the central state responsible for the motor discharge ? This is as crucial a question for ethology as for psychology. Whether the central state is in turn due to a specific neural input, or whether it is in any sense autonomous, there can be no doubt that it exists. Lorenz (1950) interpreted the variation in intensity of " innate behaviour patterns " in terms of reaction specific energy." This expression has been extensively criticised, and Thorpe (1951) proposed the use of the term " specific action potential " (S.A.P.), later modified to " specific action potentiality," as an alternative. The definition is : "
" The state of the animal responsible for its readiness to the behaviour patterns of one instinct in preference This specific readiness to all other behaviour patterns. diminishes or disappears when the consummatory act of the discharged instinct has been performed."
perform
The advantage of this term, so defined, is that it is objective, unambiguous (provided that terms such as instinct " are defined with similar rigour), and non-
"
committal on the nature of the internal state involved. The last feature might however be regarded as a disadvantage, since it glosses over the major the nature of the central state of problem-namely, " readiness." Whatever terminology is adopted, if the suggested relationship between displacement behaviour and certain human disorders has any significance, it should lead to some improvement in our understanding of behaviour. 1 suggest that it can do this in two ways.
for Research (1951) has remarked that a major gap in neurophysiology is our ignorance of how the output of the cerebral cortex is organised. For mammals it is of course just this output which is crucial for understanding Possibilities Halstead
the behaviour discussed here. The need for links between neurophysiology and ethology has been urged often enough (cf. Darwin 1872, Tinbergen 1951), but the links are slow in the forging. We have seen that there is a general rule that, if one motor output is blocked, for whatever reason, there will be discharge along another pathway. Can physiologists make this notion more precise, in terms of the actual functioning of neurons ? One of the great difficulties is to decide on a line of attack. What experiments can be devised to establish specific relationships between neural processes and behaviour! Up to now, there has been more success in relating hormonal effects to specific behaviour patterns, but we know almost nothing of how the hormones influence the central nervous system-or, for that matter, how the nervous system influences endocrine secretion. Nevertheless, it seems possible that this is a field in which advances might soon take place, given sufficient The phenomena of "stress " (cf. Selye 1950) effort. have already been mentioned in connection with psychosomatic responses. They have been studied especially in man and laboratory mammals, in which they involve a complex tangle of relationships between the endocrine glands, the hypothalamus, and the cerebral cortex. Selye has described a " general adaptation syndrome," involving a characteristic set of changes in endocrine and other organs, and leading sometimes to psychosomatic " disorder. Despite controversy on detail, we have the beginning of a foundation for the physiological study of these disorders (cf. Russell 1953). Parallel studies come from the psychological side, apparently uninfluenced by developments in ethology. but with a bearing on them. Stellar (1954) has put forward a hypothesis that " motivated behaviour" is quantitatively dependent on the activation of hypoHe uses Morgan’s term " central thalamic centres. motive state" as his counterpart of the ethology’ He suggests that, if the hypothalamus is indeed S.A.P. concerned in the control of all types of motivation," this may help to account for " interaction among motivations." Brain function is likely to be more complex than this suggests (cf. Fessard 1954), but in so far as it is valid, it has obvious implications for the understanding of displacement behaviour in vertebrates. Psychologists have also carried out experimental work with results which ought to be incorporated in any general picture. Maier’s work has already been mentioned. Miller (1948) hat> shown that white rats may show generalisation (of response), or displacement ... from one drive to another." The physiological side has been emphasised here. because it seems essential, for real progress, that a physiological analysis of behaviour should be developed. But this does not imply that the study of overt behaviour should be neglected. There is also needed, among much else, a thorough study of the whole range of substitute responses throughout the vertebrates at least : such a survey should preferably be carried out in conjunction with physiological studies. "
"
"
1207
Laitgiiage
and
hiterretation
In this efforts of
endeavour, it will be necessary to join the people working in widely different fields. The gulfs between psychiatry, psychology, ethology, and physiology are both wide and deep, and the attempt to bridge them is liable to bring on fits of dizziness. An urgent necessity is for those who practise one discipline to learn the language of the others. It would perhaps be naive to hope for a unified terminology, at least in the next few decades, but some simplification might be attempted. Psychiatrists often use words in ways which bewilder outsiders and perhaps repel students. Here, for instance, are two statements by an authoritative follower of Freud-Franz Alexander (1952b) : the uncoordinated movements of the not yet subordinated to any utilitarian goal and probably have the sole function of erotic discharge. Erotic phenomena have a playful quality-in fact, all play activities are erotic in nature.
In
child
early infancy
are
Such statements may have
biologist they suggest a of language : what does a
a valid meaning ; but to lack of rigour in the use " erotic " mean in these
sentence’? It is not suggested that clinicians will derive much ilirect benefit, as clinicians, from watching rats or seagulls, or from reading about them. But some intellectual stimulation may result, and perhaps a more critical approach to theory and a more rigorous terminology. In particular, one fundamental conclusion arises from the argument presented here-that a monistic frame of reference is the only one which satisfies the needs of communication about behaviour. We are obliged - to speak of the animal body, including that of man, as a material entity, of which the existence can be fully explained in terms of the obsetvable, physical processes going on in it. This excludes any reference to a mind " in the sense of an independent agency or separate mode of existence (cf. Young 1950). This statement is not intended as an excursion into theology, or even epistemology. Nor does it deny the reality of consciousness. Our consciousness is tied firmly to the integrity of certain parts of our brain : we are conscious when certain neural processes are occurring, and unconscious when they are not. When we say, colloquially, that we have something "in mind," we refer to our conscious thoughts ; but there is no reason why we should reject the notion that these thoughts are reflections, or aspects, of brain function. It is assumed that the aim of ethology, of which psychology is a part, is the fullest possible understanding and consequently prediction and control, of behaviour. To improve understanding, we have to devise ways of communicating our observations and theories about behaviour. At present, however, dualistic modes of speaking are so deeply embedded in our language, that a man may explicity support monism, and in the same breath use a dualistic terminology. The obvious example is the persistence of the division of illness into " mental " and physical." We can see-this very clearly with the " psychoneuroses " and the " psychosomatic disorders." As their names imply, the mind or psyche isheld to be involved in both : the first, however, are commonly regarded as strictly mental " disorders, while the second are, by definition, diseases in which the manifestation is somatic." The distinction, however, disappears when we find that the two kinds of condition may alternate or co-exist in the same individual : a man with persistent pulmonary tuberculosis, given a phrenic crush which improves the condition of the lung, develops a conversion hysteria instead (Day 1946) ; similarly, anxiety may be without any manifestation in talk or other behaviour when there are " somatic " indications of stress, although the somatic state disappears when anxiety is removed. .
"
"
"
"
From this point of view the common practice of using the term " neurosis," instead of psi/cAoneurosis," is appropriate ; but " neurosis " should include, not only conditions such as anxiety states or obsessional behaviour, but also " adaptive bodily changes with repression or denial of anxiety and other disturbances in feeling, with no verbalisation of conflicts except after considerable enquiry " (Wolff 1953) ; in other words, it should include the psychosomatic disorders, and all these disorders might more reasonably be called neurogenic." The has been suggested, not very term " somato-somatic for this seriously, approach (Schorstein 1954). Russian physiologists use- the term " cortico -visceral " to refer to the effects of emotion on smooth muscle or glandular secretion. (It would be wrong, no doubt. to pursue etymology But it is remarkable that the terms much further. anxiety " and " angina " both have the same origin, which they share with the Latin verb angere, to choke. There are other instances of monism in etymology.) A difficulty in the approach suggested here is that one can palpate a spastic colon, while the neural changes of an anxiety state can be inferred only from general behaviour or from speech. A psychologist or physician, accustomed either to the somatic or the " psychic " frame of reference, finds difficulty in switching to the other. This example further underlines the desirability of psychiatrists, psychologists, and ethologists learning each other’s languages. Another, related, difficulty is that the way of speaking adopted here is, to some people, distasteful or even odious : to them it seems inhuman. This attitude deserves respect, since it often reflects a real and necessary concern with the needs of the whole human personality. The question remains, however, whether the formulations suggested in this paper help towards the successful application of knowledge to human ends. The ends to which knowledge is put depend on the motivation of scientists, physicians, and others, not on their prose style. It is not suggested for a moment that only doctors use imprecise or obscurantist terminologies. The example Drive " of instinct has already been mentioned. remains an instance of a mendicant word-one which goes around begging for a meaning (Ogden and Richards 1931). The whole of learning terminology, despite the efforts of Thorpe (1950) and others, is in a most unsatisfactory state (cf. Haldane 1954). This state of affairs will not be remedied by logical meditation or semantic analysis alone. Greater precision must depend on improved understanding, and this will develop only from further experiment. The proposals put forward here for future research and for more rigorous interpretation are therefore interdependent. "
"
"
"
-
"
"
"
Summary and Conclusions Disturbed behaviour, in which one reaction pattern replaces another, more " appropriate " one, is widespread in vertebrates. It
occurs when an activity is interrupted prevented (i.e., in conflict or in frustrating circumstances) or when a particular sensory situation is not achieved (e.g., when the number of eggs in a nest is reduced, or when the skin-temperature is outside a certain, optimum, range). In the study of animal behaviour (ethology), responses of this sort are called displacement behaviour. These responses have been mainly described as substitutes for other activities (usually for consummatory acts such as breathing, eating, or copulating), but it is suggested that the immediate cause of such behaviour must always be some deficiency in the afferent input to the central nervous system. All activity is accompanied by -a (and proprioceptive interoceptive) input, and this is presumably responsible for the, normal cessation of "
or
"
"
"
1208
activity which occurs when a consummatory performed.
act has been
In human behaviour there is an almost complete absence of innate " behaviour in the sense of unlearnt stereotyped reaction patterns. Human behaviour of the kind which is said to arise from " instinct " is infinitely variable (apart from reflexes) on the motor side ; but it tends to achieve particular sensory inputs (consummatory situations). If such ends are not attained, behaviour may be disturbed: psychoneurotic behaviour-e.g., in hysteria or obsessional statesis analogous in certain respects to displacement behaviour in birds or fishes. Psychoneurotic behaviour is, however, not innate, but in a sense learnt. In this it resembles the behaviour fixations developed by, for instance, rats in frustrating situations. In these examples one motor output is substituted for another. The same applies in the " psychosomatic disorders of man, in which an autonomic discharge, resulting from some form of stress," produces a condition such as asthma. It is suggested that the distinction between psychoneurotic and psychosomatic disorder is unjustified ; that both might properly be called " neurotic or neurogenic " ; and that the distinction arises from a conventional mind-body dualism which is now a handicap in the interpretation of behaviour. It is further suggested that displacement behaviour and neurotic responses of all kinds may represent a general feature of the functioning of nervous systems-namely, that if there is an excitatory state which would ordinarily produce a motor discharge, and if the motor output cannot take its " normal " form, then it will follow some other pattern. This proposal may be regarded as an hypothesis. It requires investigation and clarification
BIOPSY OF THE RECTUM IN ULCERATIVE COLITIS
"
"
GEORGE LUMB M.D. READER
IN
"
"
WESTMINSTER
MEDICAL
SCHOOL;
CONSULTANT PATHOLOGIST
R. H. B. PROTHEROE B.A., M.B. Camb.
"
"
Lond., M.R.C.P.
PATHOLOGY,
SENIOR
IN
REGISTRAR
PATHOLOGY,
AND
RESEARCH FELLOW
IN ULCERATIVE COLITIS
WESTMINSTER
HOSPITAL,
LONDON
RAPID autolysis has hitherto greatly impeded study of the morbid histology of the gastro-intestinal tract. For this reason post-mortem material is of very limited value for detailed study, and examination of surgical specimens also has its limitations because of the degenerative changes that occur before fixation is complete. The stomach and the rectum, however, are accessible for biopsy and the gastric mucosa has been carefully studied with the biopsy gastroscope. Palmer (1954) has used this method extensively and reviewed the published reports. Rectal
biopsy
has been used almost
entirely
for
diagnosing
by neurophysiologists. In general, there is need for more effective communication between ethologists, psychologists, physiologists, and psychiatrists, and for the development of research in which - physiologists and students of behaviour
collaborate. Given such combination, it should be to devise possible experimental programmes which would lead to a rapid advance in the understanding of behaviour. I am grateful to Prof. T. Ferguson Rodger and his colleagues of the Department of Psychological Medicine, Glasgow University, and to Prof. J. B. S. Haldane, F.R.S., and Dr. H. Spurway, for illuminating discussions on the subject-matter of this paper. Especial acknowledgment is due tq Dr. A. B. Sclare for reading and criticising the manuscript. ADDENDUM
After this paper was written Kortlandt (1955) published a long and valuable discussion of the concept of instinct, with many references to the relationships betweenethological and psychiatric theories. I have also been allowed to see, in advance of publication, a paper by Morris in which an interesting aspect of the connection between autonomic function and behaviour is presented. Both papers can be recommended to anyone interested in the topics I have discussed. REFERENCES
Alexander, (1952a) Psychosomatic Medicine. — (1952b) In Alexander and Ross’s Dynamic Psychiatry. Chicago. Armstrong, E. A. (1950) Symp. Soc. exp. Biol. 4, 361. Barnett, S. A., Spencer, M.M. (1951) Behaviour, 3, 229. Bastock, M., Morris, D., Moynihan, M. (1953) Ibid, 6, 66. Darwin, C. (1872) The Expression of the Emotions in Man and
Fig. I-Age-distribution
—
cases
of ulcerative colitis.
Hardy, J. D. (1954) Harvey Lect. 49, 242. Hebb, D. O. (1949) The Organization of Behaviour. New York. Jones, E. (1953) Sigmund Freud, Life and Work. London ; vol. i: Kennedy, J. S. (1954) Brit. J. anim. Behav. 2. 12. Kortlandt, A. (1955) Arch. Néerl. Zool. 11, 155. Lissman, H. W. (1950) Symp. Soc. exp. Biol. 4, 34. Lorenz, K. Z. (1950) Ibid, p. 211. Maier, N. R. F. (1949) Frustration. New York. Miller, N. E. (1948) J. abn. soc. Psychol. 43, 155. Morris, D. Behaviour (in the press). Moynihan, M. (1953) Ibid, 5, 58. Ogden, C. K., Richards, I. A. (1931) The Meaning of Meaning London.
F.
Animals. London. Day, G. (1946) Lancet, ii, 703. Fessard, A. E. (1954) In Brain Mechanisms and Consciousness (edited by J. F. Delafresnaye). London. Freud, S. (1949) An Outline of Psychoanalysis. London. Fulton, J. F. (1949) Functional Localisation in the Frontal Lobes and Cerebellum. Oxford. Haldane, J. B. S. (1949) In Genetics, Paleontology, and Evolution. Princeton and Oxford. (1954) Behaviour, 6, 256. Halstead, W. C. (1951) In Cerebral Mechanisms in Behaviour. New York.
in 150
Russell, R. W. (1950) Brit. J. Psychol. 41, 95. (1953) Int. J. Psycho-Anal. 34, 1. Pretty, R. G. F. (1951) Quart. J. exp. Psychol. 3, 151. Schorstein, J. (1954) Personal communication. Selye, H. (1950) The Physiology and Pathology of Exposure to -
-
Stress. Montreal.
Spitz, R. A., Wolf, K. M. (1946) Genet. Psychol Monogr. 34, 57. Spurway, H., Haldane, J. B. S. (1953) Behaviour, 6, 8. Stellar, E. (1954) Psychol. Rev. 61, 5. Thorpe, W. H. (1950) Symp. Soc. exp. Biol. 4, 387. (1951) Bull. anim. Behav. 9, 34. (1954) Nature, Lond. 174, 101. Tinbergen, N. (1951) The Study of Instinct. Oxford. (1952) Quart. Rev. Biol. 27, 1. (1953) The Herring Gull’s World. London. van Iersel, J. J. A. (1947) Behaviour, 1, 56. Whitehorn, J. C. (1952) In Alexander and Ross’s Dynamic —
—
—
—
—
Psychiatry. Chicago. Wolff, H. G. (1953) Stress and Disease. Springfield, Ill. Young, J. Z. (1950) Doubt and Certainty in Science. Oxford.
ORIGINAL ARTICLES " DISPLACEMENT " BEHAVIOUR AND "PSYCHOSOMATIC" DISORDER S. A. BARNETT
THE object of this paper is to suggest a relationship between the " displacement behaviour " described by ethologists,* and what clinicians sometimes call " psychosomatic disorder." The argument leads to two general sets of proposals. The first concerns the necessity for a particular line of research ; the second suggests that the present terminologies of psychologists and psychiatrists are often a handicap on progress, and in particular that their interpretations should be strictly monist. Behaviour
Displacement behaviour is the performance of a behaviour pattern, in biologically inappropriate circumstances, when an appropriate action is prevented or the input to the central nervous system departs from the optimum. The behaviour does not directly serve the survival or reproduction of the individual or the species : biologically, it is irrelevant to the situation in which it is performed. Such behaviour has been mainly studied in fishes and birds (cf. Armstrong 1950, Tinbergen 1952), but it certainly occurs also in mammals. For example, a wild rat feeding, disturbed by another rat of a different species, makes " aggressive " noises and motions towards the interloper ; the latter flees ; the first rat then, instead of immediately resuming feeding, briefly grooms its face with its forepaws-an action which it would not ordinarily perform in the middle of feeding (Barnett and Spencer 1951). Deprived of the object of its aggression, the rat resorts to an alternative, and inappropriate, behaviour pattern. A slightly different example is that of wild rats during a series of copulations : at intervals both male and female may make a rapid to a feeding-point, feed hurriedly (and in a quite atypical manner) for a few seconds, and then of courtship and coitus resume the usual pattern (unpublished). In general, typical displacement activities occur in three types of situation. First, if an animal is interrupted in, or prevented from performing, a specific act, it may substitute another act, as in the example above of the aggressive rat. Second, seemingly irrelevant behaviour may occur if an animal has two incompatible types of behaviour open to it, as with herring gulls which resort to grass-pulling when the " appropriate behaviour could be either to attack or to run away (Tinbergen 1953). Third, the actual attainment of a goal-the performance of a consummatory act-may, strangely enough, be followed by irrelevant behaviour, as in the example above of the rats between copulations. Displacement activities are often incomplete actions, as in the nest-building movements of gulls or the grooming of rats ; but they need not be so. They are often hurried, as in the displacement feeding of rats. The so-called vacuum activities, in which an animal performs an action, such as a display, in the absence of an appropriate object, may possibly be responses to suboptimal stimuli ; they are comparable to displacement acts. Like them, they seem to reflect an overflow " of motor output. This point will be discussed in more detail later. Another phenomenon closely similar to displacement is the " redirection activity " , (Bastock et al. 1953), in which an animal directs the appropriate movement towards an inappropriate object. The authors quoted refer only to examples where the appropriate object is still present ;but a similar effect has been observed when
move
"
*
Ethology is the scientific study
6902
object is no longer available : a rat deprived of the object of its aggression may turn on a member of its own colony (Barnett and Spencer 1951). Bastock and her colleagues draw attention to the analogy to the " object of psycho-analysts. As Freud (1949) displacement
wrote :
ZOOLOGY, GLASGOW UNIVERSITY
Displacement
the
"
M.A. Oxfd LECTURER IN
1955
of animal behaviour.
We have found that instincts can change their aim (by and also that they can replace one anotherthe energy of one instinct passing over to another.
displacement)
The examples of displacement behaviour given above may be unconvincing to the critical reader who has not himself observed such behaviour. A critical attitude is very necessary in this field ; but fortunately, as we shall see below, good examples, quantitatively analysed, do exist. Learnt
Responses Resembling Displacement Behaviour
In the classical examples of displacement activity, the substituted behaviour always belongs to an alternative stereotyped (" innate ") behaviour pattern. This is reflected in the first definition of the term displacement activity " given by Thorpe (1951) : such activity is said to result from the activation ... of the action pattern belonging to another instinct " ; but alternative, wider definitions are given. For the present argument it is convenient to apply a wider definition, such as that given at the beginning of this paper. We can then consider whether there are learnt responses in mammals which, though very different from displacement responses of the usual sort, can properly be put under this general heading. Maier (1949) describes experiments in- which albino rats were faced with an insoluble problem in a Lashley jumping apparatus. Whatever solution they chose, they were "punished" in half of their jumps on the average. Moreover, they were forced to jump. Such rats tended to develop " behaviour-fixations " : they commonly, after some time, jumped, say, always to the right, regardless of the stimulus situation ; indeed, they did so even if the door was open and the food was visible on the left-hand side. Maier comments : "
It appears that the fixated group develops some kind of adjustment to the test situation and is therefore able to prevent
emotional tensions.
Fixated rats also showed less resistance to making a The detailed interpretation of Maier’s results remains difficult and to some extent controversial, but there is no doubt, from his and other experiments, of the reality of behaviour fixations " (Russell and Pretty
jump.
"
1951). Better-known examples of abnormal " learnt are those described under the heading of responses " experimental neurosis." In these, too, an insoluble problem may be presented, but the result is either an excessive undirected activity, or a general inactivity which may include refusal of food. The variety of these " abnormal " acts is considerable (cf. Russell 1950), and a fuller analysis is needed before the validity of the comparison suggested here can be assessed. However, despite the diversity of these responses, there seems to be at least an analogy to displacement behaviour in all these examples.: certainly, where there is a fixated response this provides a regular pattern of motor output in frustrating circumstances-i.e., in conditions in which a biologically appropriate response cannot be made. The
Consummatory
Situation"
displacement activity has been described substitute for an alternative activityif an animal is unable to perform or to complete a consummatory act, such as fighting, feeding, or copulation, it performs Usually,
as a
a
1204 other act instead. But this does not always apply. Moynihan (1953) studied nest-building movements and preening movements in gulls during incubation of the eggs. Nest-building movements were shown to increase some
if one or more eggs were removed, and the number of such movements in a given time increased in proportion to the number of eggs removed. Preening increased removed. An all when the were markedly only eggs of was the feature this work quantitative important expression of displacement behaviour: in this case, unlike many, subjective impressions could be excluded. (An earlier good example is that given by Tinbergen and van Iersel (1947) on fanning in Gaste1’osteus aculeattis.) Both types of movement were typical of displacement activities, in that they belong to stereotyped behaviour patterns, and that they occurred at inappropriate moments. They differed from the usual examples, however, in one notable respect : they were performed, not in place of a motor act, but when a specific sensory situation was altered. The substitute activity reflected the absence of what might be called a " consummatory situation " (cf. Thorpe 1954). The importance -of the afferent input is also shown by the work of H. Spurway and J. B. S. Haldane on the air-gulping of the climbing perch (Allabas testudineus). Haldane (in litt.) writes : Our fish got into a flap when they breathed N2. To men this certainly has not got an odd smell or taste. It makes you feel dizzy a few seconds after breathing it. Probably the brain is also affected by afferent impulses from the carotid sinus. The fish have done their consummatory act, but have not got the normal "feedback." This disturbs them much more than if they are mechanically prevented from carrying out the act.
It
reasonable to suggest that the concept of consummatory situation or state is more fundamental and more general than that of consummatory act. The latter marks the end of activity, or at least of a specific group of activities, such as those related to mating. But the performance of such an act must be accompanied by specific internal changes, such as those involved in filling the stomach ; and these changes, by evoking a particular afferent input, thereby alter central nervous activity and so the output to the muscles. seems
"
Kennedy (1954),
Instinct " in Man
in
a
polemical discussion of ethological
theories, suggests that the cessation of activity on the attainment of a consummatory act is "reflex "-i.e., due to an input of impulses from the periphery. Bastock et al. (1953), in the paper already quoted, ’suggest that "
consummatory stimuli," and especially proprioceptive feed-back, are important in what they call the " conof drive." (Since proprioceptive fibres make one-third to one-half of those in a nerve supplying a muscle, it is clear that they deserve a good deal of attention in the study of behaviour (cf. Lissnian 1950).) Disregarding the particular terminologies used in these two communications, each, from different points of view, suggests that we must look to the afferent input if we are to make a beginning on the physiological analysis of displacement behaviour. It is therefore worth while to consider a little further the subject of the influence of afferent impulses on certain sorts of behaviour. Here is an example (Hardy 1954) from a physiological study in which engineering and psychiatric terms are both used in a physiological
sumption
up
context : From the
point
of view of
servo
control of skin
tempera- ,
ture, it is worth noting that thermal sensations have strong when the average skin temperature is below 32° or above 34°C ....This " affect " of thermal sensation is probably one of the most powerful drives for. physiologic servo temperature regulation in the presence of external thermal stress. elements of
unpleasantness
or
pleasantness
in
A noxious stimulus of the sort described here evokes man either a general restlessness (analogous to
appetitive behaviour), or a learnt response which tends to reduce the unpleasantness. If there is no learnt response available, as in a young child, the restlessness may lead to learning-perhaps of the trial-and-error variety. In any case, the situation described in the quotation above is one in which a particular sensory condition induces and this activity ceases on the attainment activity, of a " consummatory situation "-in the example, a particular skin-temperature. If the analysis above is correct, it may serve to clarify the concept of instinct " as applied to man. In ethological writing the term instinct " is today commonly (not always) applied to a stereotyped activity. Depending on the author, the activity may be a single consummatory one ; or it may be a complex series of acts, of which the, "
"
consummatory one is the last. The term instinct, so used, doesnot apply to any internal agency in behaviour, " drive " may be used to refer to internal influences which lead to the activity. In man there is virtually no instinctive behaviour in the above sense. The infant breathes, sucks, and cries ; he also, perhaps, at 3-6 months, smiles in response to a particular " sign stimulus " (Spitz and Wolf 1946) ; but apart from reflexes, later behavioural performances are learnt. Yet human beings act most strenuously to achieve certain ends. These ends, regarded at the physiological level, are the attainment of certain sorts of neural input-of consummatory situations. The motor procedures which attain these ends are variable and learnt. Hence, to take an example from Freud, the word Trieb, usually translated as " instinct," refers to behaviour which ceases once a particular nervous input-e.g., from a full belly or empty seminal vesicles-has been achieved. Or, to put the obverse, the behaviour is activated by the input from an empty belly or full seminal vesicles. These examples must not be taken to imply that all such behaviour is activated peripherally, by the stimuli of the moment, or that it is innate." In higher mammals, especially man, with a vast mass of internuncial neurons in the cerebral cortex, it seems that learning-or at least previous experience--may lead to forms of neural organisation in which the activation of the motor output is largely within the central nervous system itself. Behaviour may then be both " spontaneous" and a product of learning. As Hebb (1949) has pointed out, it is this iliternuncial mass which makes possible the predominance of learnt behaviour in man, and his relative independence of the immediate sensory situation. Whitehorn (1952) approaching this topic from the point of view of a psychiatrist, has remarked that, despite the evidence from social anthropology for the cultural rather than the innate patterning of human behaviour, the term instinct is still retained by psychiatrists. He points out that the psychiatric use of the term represents a shift of meaning [from that of ethologists] to " inner driving forces correlated with biological needs," and that " this inner impulsive force is experienced as emotion." He further suggests that " emotion " is negatively correlated with instinctive behaviour : the less an animal’s behaviour is automatically determined by fixed neural patterns, the more it is necessary that powerful internal agencies should ensure that effective behaviour patterns are acquired (learnt) by each individual. Emotion in this context is the subjective counterpart of the neural changes due to a certain type of (unpleasant) sensory
though the term
"
"
"
input. Whether "
or
not
we
agree with this
emotion," Whitehorn’s analysis
hypothesis about brings out clearly
the notorious fact that the term " instinct" is what Ogden and Richards (1931) call a nomad : it wanders
1205
meaning to meaning. wanderings are not yet over.
from
Psychoneurosis "
and "
It is to be feared that its
Psychosomatic"
Disorder
Behavioural Cha.7zges In the virtual absence of instinctive behaviour in man, one might think it idle to look for displacement activities in human behaviour. The realisation of the importance of consummatory state," however, alters the picture. The analogy between displacement behaviour in other animals, and psychoneurotic behaviour in man, has indeed often been pointed out, notably by Armstrong (1950). Armstrong quotes a passage on hysteria in which Freud writes that in such states " we have an excessive innervation (in typical cases a somatic innervation), sometimes of a sensory, sometimes of a motor character, either as an excitation or an inhibition." This refers of course to the fact that the behaviour called hysterical covers a wide range, from outbursts of weeping to a functional paralysis or anoesthesia. Armstrong comments that both in displacement behaviour is a somatic and conversion hysteria " the performance in both the activity means of release from tension ; is incomplete and apparently detached from normal The same might be said of affective experience." obsessional behaviour. However, the fact that hysteria includes phenomena of inhibition, as well as oert activity, must make one hesitate to -regard it, in any simple way, as the human counterpart of displacement behaviour. As we shall see, this sort of question can be answered, or indeed effectively asked, only in neuro...
.
physiological terms. Psychoneurotic behaviour
in general is of course instinctive " : it does not consist of rigidly stereotyped activities common to a whole species or variety ; in a sense it is learnt. Behaviour, such as smoking or chewing, which is clearly analogous to displacement behaviour but which ought not to be labelled psychoneurotic, is quite obviously learnt. The same applies to the thumb-sucking of an infant. (Smoking and the rest could be labelled " abnormal," whatever that may mean, but there are other human responses, of biological value and in no possible sense pathological, which may have originated, in evolution, as displacement behaviour. Spurway and Haldane (1953) suggest that speech is a derivative (" ritualised ") of displacement breathing. Another form of communication, the movements of the facial muscles of expression, may well have originated in the same way, and have acquired increasing importance with the development of human not
"
perhaps
society.) a warning is perhaps required. It is sometimes overlooked, that human behaviour is so much more complex than that of any other species, that it is in some respects qualitatively This is evident in as well as quantitatively different. our use of symbols, not only in language but also indirectly. If a woman is sick of her husband, she may not say so or give any verbal indication that she is aware of it : but she may persistently vomit while her husband is at home. (I am indebted to Dr. R. D. Laing for this example.) Such a response can be interpreted according to a variety of systems. It could be described in neurological terms, perhaps with the addition of expressions derived from studies of conditioned reflexes ; it could be compared to a displacement response ; or it could be characterised in the more poetic or symbolic phrases (no sneer is intended) of Freudian analysis. All no doubt have some validity, but to say that psychoneurosis resembles displacement behaviour certainly does not " explain " it. The usefulness, if any, of the sort of comparison made here is of two kinds. First, if similar processes are found in man and other species, research possibilities
At this
obvious,
point
but
"
"
may be widened.
The proposal is then, in effect, a that similar internal " mechanisms " underlie all the various phenomena compared. Secondly, the attitude to particular aspects of human behaviour The objective, nonmay be changed for the better. ethical, approach to behaviour attempted by ethologists is superior to a subjective moralistic one, both heuristically and humanely. Yet if a person is said to have a mental " disorder, for many-even for some doctorsthis amounts to a moral stigma or aesthetic flaw, such as is not felt to exist in the person with, say, peptic ulcer or tuberculosis. (It is perhaps instructive to realise that a great variety of so-called perversions," from masturbation to fetishism, have been observed in various animal species. Homosexuality, in particular, is a widespread phenomenon among vertebrates.) There is, indeed, on these grounds alone, a case for revising the terminology of what, in my view, should be called behaviour disorders but which at present are usually described as mental illnesses. (It is a strange fact that in psychiatry the term " behaviour disorder ’’ is reserved for a single " disconduct group of antisocial activities--the orders which are also lumped under the head of psycho-
hypothesis, namely,
"
"
"
"
pathic personality.") Soinatic
-Responses examples
In all the
output
so
from the central
far
given,
nervous
there is
an
altered
system. Instead of the
"
normal " output, another is substituted. The alternative motor pattern may be "instinctive" or not; it may be alternative to an active pattern of behaviour or to an inactivity ; it may itself be an inaetivity-an inhibitory effect. All these statements apply also to the so-called psychoIn these, the substitute somatic disorders of man. output is not to the somatic musculature, but is as a rule autonomic. There is, however, no part of the body and no organ which is outside the scope of these disorders. This is not surprising, since every part of the body has a blood-supply partly under autonomic control and carrying hormones of which the secretion is influenced by the autonomic nervous system. Wolff (1953) gives an excellent review of these condí tions. An example is the pattern of response of the stomach to the ingestion of food : increased bloodflow, increased motility, and secretion of acid. These may all occur, quite inappropriately, in conditions of stress. Another familiar response to stress is diarrhceahyperactivity of the gut muscles. Wolff comments that the person -with violent diarrhoea may be outwardly calm, sweet-mannered and serene.’A third standard example is asthma, in which the response is an inappropriate contraction of the smooth muscle of the lung. Alexander (1952a) describes these responses as follows :
biologically appropriate
or
"
Whenever the expression of competitive, aggressive, and hostile attitudes is inhibited in voluntary behaviour, the sympathetic adrenal system is in sustained excitation. The vegetative symptoms result from the sustained sympathetic excitation, which persists because no consummation of the fight or flight reaction takes place in the field of coordinated voluntary behaviour, as exemplified by the patient suffering from essential hypertension who in his overt behaviour appears inhibited and under excessive control. Likewise in migraine headache, the painful attack may terminate within a few minutes after the patient becomes conscious of his rage and gives open expression to it.
It is easy to see that, in one aspect, the psychosomatic responses fit quite readily into a Pavlovian framework : they can be interpreted in terms of conditioned reflexes, once it is understood that a conditioned reflex is an exceedingly complex phenomenon (cf. Hebb 1949). In each instance the primary response is innate," and would ordinarily be called reflex. In the innate character of the response, these disorders show a closer similarity to the ordinary displacement behaviour of "
1206 birds and fishes, than is shown by typical " psychoneurotic " behaviour. They are, however, different in three obvious ways from what we may call classical displacement activities.
First, they are very variable within the species. There is evidence, as might be expected, of genetically determined differences in the ways in which individuals react to stress : in one man a given stress may induce, say, diabetes, in another similar conditions may lead to hypertension. This is perhaps yet another example of the exceptional polymorphism of the human species
(cf. Haldane 1949). Second, the environmental causes of these disorders is much more complex and diffuse. Even when there is an immediate precipitating cause, it acts on a central nervous system already prepared by previous experience. This of course reflects the size and complexity of the human nervous system, with its enormous accumulation of connecting neurons. Third, as already mentioned, the output is not somatic motor, but is visceral. Here again, evidently, we have a reflection of a distinctively Primate (especially human) evolutionary development-namely, the importance of the cerebral cortex, and especially of the frontal lobe, in controlling autonomic output. Fulton (1949) writes : I am inclined to regard the recognition of the frontal lobe as the primary autonomic centre of the forebrain as perhaps the most significant disclosure of recent years in the field of scientific medicine, for it affords at long last a sound physiological background for that still shadowy subject of psychosomatic medicine." Despite these differences, the psychosomatic disorders and the psychoneuroses on the one hand, and displacement activities on the other, all seem to exemplify "
general
feature of
complex nervous systems-that, excitatory state which would ordinarily produce a motor discharge, and if the motor output cannot take place in the biologically " normal " fashion, then it will follow some other path. The other path a
if there is
may be
"
rigorously
an
innate " or fixed for the
"
learnt," but, if learnt, may be
particular
The Problem of
"
individual.
Nervous
Energy
"
T enninology An early
discussion of " nervous energy " was that of Freud. As Ernest Jones (1953) points out, Freud in his early psychiatric works wrote in terms of " an economics of nervous energy." (At that time, too, Freud adopted a notably " psychosomatic " approach to his subject. In interpreting anxiety neurosis as a of sexual consequence deprivation he points out that increased respiration-rate, palpitation, sweating, and congestion occur both in coitus and in anxiety states. As Jones shows, the history of Freud’s thinking is of great interest for the study of ideas on the relation between " soma " and " psyche.") What, then, is the nature of the central state responsible for the motor discharge ? This is as crucial a question for ethology as for psychology. Whether the central state is in turn due to a specific neural input, or whether it is in any sense autonomous, there can be no doubt that it exists. Lorenz (1950) interpreted the variation in intensity of " innate behaviour patterns " in terms of reaction specific energy." This expression has been extensively criticised, and Thorpe (1951) proposed the use of the term " specific action potential " (S.A.P.), later modified to " specific action potentiality," as an alternative. The definition is : "
" The state of the animal responsible for its readiness to the behaviour patterns of one instinct in preference This specific readiness to all other behaviour patterns. diminishes or disappears when the consummatory act of the discharged instinct has been performed."
perform
The advantage of this term, so defined, is that it is objective, unambiguous (provided that terms such as instinct " are defined with similar rigour), and non-
"
committal on the nature of the internal state involved. The last feature might however be regarded as a disadvantage, since it glosses over the major the nature of the central state of problem-namely, " readiness." Whatever terminology is adopted, if the suggested relationship between displacement behaviour and certain human disorders has any significance, it should lead to some improvement in our understanding of behaviour. 1 suggest that it can do this in two ways.
for Research (1951) has remarked that a major gap in neurophysiology is our ignorance of how the output of the cerebral cortex is organised. For mammals it is of course just this output which is crucial for understanding Possibilities Halstead
the behaviour discussed here. The need for links between neurophysiology and ethology has been urged often enough (cf. Darwin 1872, Tinbergen 1951), but the links are slow in the forging. We have seen that there is a general rule that, if one motor output is blocked, for whatever reason, there will be discharge along another pathway. Can physiologists make this notion more precise, in terms of the actual functioning of neurons ? One of the great difficulties is to decide on a line of attack. What experiments can be devised to establish specific relationships between neural processes and behaviour! Up to now, there has been more success in relating hormonal effects to specific behaviour patterns, but we know almost nothing of how the hormones influence the central nervous system-or, for that matter, how the nervous system influences endocrine secretion. Nevertheless, it seems possible that this is a field in which advances might soon take place, given sufficient The phenomena of "stress " (cf. Selye 1950) effort. have already been mentioned in connection with psychosomatic responses. They have been studied especially in man and laboratory mammals, in which they involve a complex tangle of relationships between the endocrine glands, the hypothalamus, and the cerebral cortex. Selye has described a " general adaptation syndrome," involving a characteristic set of changes in endocrine and other organs, and leading sometimes to psychosomatic " disorder. Despite controversy on detail, we have the beginning of a foundation for the physiological study of these disorders (cf. Russell 1953). Parallel studies come from the psychological side, apparently uninfluenced by developments in ethology. but with a bearing on them. Stellar (1954) has put forward a hypothesis that " motivated behaviour" is quantitatively dependent on the activation of hypoHe uses Morgan’s term " central thalamic centres. motive state" as his counterpart of the ethology’ He suggests that, if the hypothalamus is indeed S.A.P. concerned in the control of all types of motivation," this may help to account for " interaction among motivations." Brain function is likely to be more complex than this suggests (cf. Fessard 1954), but in so far as it is valid, it has obvious implications for the understanding of displacement behaviour in vertebrates. Psychologists have also carried out experimental work with results which ought to be incorporated in any general picture. Maier’s work has already been mentioned. Miller (1948) hat> shown that white rats may show generalisation (of response), or displacement ... from one drive to another." The physiological side has been emphasised here. because it seems essential, for real progress, that a physiological analysis of behaviour should be developed. But this does not imply that the study of overt behaviour should be neglected. There is also needed, among much else, a thorough study of the whole range of substitute responses throughout the vertebrates at least : such a survey should preferably be carried out in conjunction with physiological studies. "
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1207
Laitgiiage
and
hiterretation
In this efforts of
endeavour, it will be necessary to join the people working in widely different fields. The gulfs between psychiatry, psychology, ethology, and physiology are both wide and deep, and the attempt to bridge them is liable to bring on fits of dizziness. An urgent necessity is for those who practise one discipline to learn the language of the others. It would perhaps be naive to hope for a unified terminology, at least in the next few decades, but some simplification might be attempted. Psychiatrists often use words in ways which bewilder outsiders and perhaps repel students. Here, for instance, are two statements by an authoritative follower of Freud-Franz Alexander (1952b) : the uncoordinated movements of the not yet subordinated to any utilitarian goal and probably have the sole function of erotic discharge. Erotic phenomena have a playful quality-in fact, all play activities are erotic in nature.
In
child
early infancy
are
Such statements may have
biologist they suggest a of language : what does a
a valid meaning ; but to lack of rigour in the use " erotic " mean in these
sentence’? It is not suggested that clinicians will derive much ilirect benefit, as clinicians, from watching rats or seagulls, or from reading about them. But some intellectual stimulation may result, and perhaps a more critical approach to theory and a more rigorous terminology. In particular, one fundamental conclusion arises from the argument presented here-that a monistic frame of reference is the only one which satisfies the needs of communication about behaviour. We are obliged - to speak of the animal body, including that of man, as a material entity, of which the existence can be fully explained in terms of the obsetvable, physical processes going on in it. This excludes any reference to a mind " in the sense of an independent agency or separate mode of existence (cf. Young 1950). This statement is not intended as an excursion into theology, or even epistemology. Nor does it deny the reality of consciousness. Our consciousness is tied firmly to the integrity of certain parts of our brain : we are conscious when certain neural processes are occurring, and unconscious when they are not. When we say, colloquially, that we have something "in mind," we refer to our conscious thoughts ; but there is no reason why we should reject the notion that these thoughts are reflections, or aspects, of brain function. It is assumed that the aim of ethology, of which psychology is a part, is the fullest possible understanding and consequently prediction and control, of behaviour. To improve understanding, we have to devise ways of communicating our observations and theories about behaviour. At present, however, dualistic modes of speaking are so deeply embedded in our language, that a man may explicity support monism, and in the same breath use a dualistic terminology. The obvious example is the persistence of the division of illness into " mental " and physical." We can see-this very clearly with the " psychoneuroses " and the " psychosomatic disorders." As their names imply, the mind or psyche isheld to be involved in both : the first, however, are commonly regarded as strictly mental " disorders, while the second are, by definition, diseases in which the manifestation is somatic." The distinction, however, disappears when we find that the two kinds of condition may alternate or co-exist in the same individual : a man with persistent pulmonary tuberculosis, given a phrenic crush which improves the condition of the lung, develops a conversion hysteria instead (Day 1946) ; similarly, anxiety may be without any manifestation in talk or other behaviour when there are " somatic " indications of stress, although the somatic state disappears when anxiety is removed. .
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"
From this point of view the common practice of using the term " neurosis," instead of psi/cAoneurosis," is appropriate ; but " neurosis " should include, not only conditions such as anxiety states or obsessional behaviour, but also " adaptive bodily changes with repression or denial of anxiety and other disturbances in feeling, with no verbalisation of conflicts except after considerable enquiry " (Wolff 1953) ; in other words, it should include the psychosomatic disorders, and all these disorders might more reasonably be called neurogenic." The has been suggested, not very term " somato-somatic for this seriously, approach (Schorstein 1954). Russian physiologists use- the term " cortico -visceral " to refer to the effects of emotion on smooth muscle or glandular secretion. (It would be wrong, no doubt. to pursue etymology But it is remarkable that the terms much further. anxiety " and " angina " both have the same origin, which they share with the Latin verb angere, to choke. There are other instances of monism in etymology.) A difficulty in the approach suggested here is that one can palpate a spastic colon, while the neural changes of an anxiety state can be inferred only from general behaviour or from speech. A psychologist or physician, accustomed either to the somatic or the " psychic " frame of reference, finds difficulty in switching to the other. This example further underlines the desirability of psychiatrists, psychologists, and ethologists learning each other’s languages. Another, related, difficulty is that the way of speaking adopted here is, to some people, distasteful or even odious : to them it seems inhuman. This attitude deserves respect, since it often reflects a real and necessary concern with the needs of the whole human personality. The question remains, however, whether the formulations suggested in this paper help towards the successful application of knowledge to human ends. The ends to which knowledge is put depend on the motivation of scientists, physicians, and others, not on their prose style. It is not suggested for a moment that only doctors use imprecise or obscurantist terminologies. The example Drive " of instinct has already been mentioned. remains an instance of a mendicant word-one which goes around begging for a meaning (Ogden and Richards 1931). The whole of learning terminology, despite the efforts of Thorpe (1950) and others, is in a most unsatisfactory state (cf. Haldane 1954). This state of affairs will not be remedied by logical meditation or semantic analysis alone. Greater precision must depend on improved understanding, and this will develop only from further experiment. The proposals put forward here for future research and for more rigorous interpretation are therefore interdependent. "
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-
"
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"
Summary and Conclusions Disturbed behaviour, in which one reaction pattern replaces another, more " appropriate " one, is widespread in vertebrates. It
occurs when an activity is interrupted prevented (i.e., in conflict or in frustrating circumstances) or when a particular sensory situation is not achieved (e.g., when the number of eggs in a nest is reduced, or when the skin-temperature is outside a certain, optimum, range). In the study of animal behaviour (ethology), responses of this sort are called displacement behaviour. These responses have been mainly described as substitutes for other activities (usually for consummatory acts such as breathing, eating, or copulating), but it is suggested that the immediate cause of such behaviour must always be some deficiency in the afferent input to the central nervous system. All activity is accompanied by -a (and proprioceptive interoceptive) input, and this is presumably responsible for the, normal cessation of "
or
"
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1208
activity which occurs when a consummatory performed.
act has been
In human behaviour there is an almost complete absence of innate " behaviour in the sense of unlearnt stereotyped reaction patterns. Human behaviour of the kind which is said to arise from " instinct " is infinitely variable (apart from reflexes) on the motor side ; but it tends to achieve particular sensory inputs (consummatory situations). If such ends are not attained, behaviour may be disturbed: psychoneurotic behaviour-e.g., in hysteria or obsessional statesis analogous in certain respects to displacement behaviour in birds or fishes. Psychoneurotic behaviour is, however, not innate, but in a sense learnt. In this it resembles the behaviour fixations developed by, for instance, rats in frustrating situations. In these examples one motor output is substituted for another. The same applies in the " psychosomatic disorders of man, in which an autonomic discharge, resulting from some form of stress," produces a condition such as asthma. It is suggested that the distinction between psychoneurotic and psychosomatic disorder is unjustified ; that both might properly be called " neurotic or neurogenic " ; and that the distinction arises from a conventional mind-body dualism which is now a handicap in the interpretation of behaviour. It is further suggested that displacement behaviour and neurotic responses of all kinds may represent a general feature of the functioning of nervous systems-namely, that if there is an excitatory state which would ordinarily produce a motor discharge, and if the motor output cannot take its " normal " form, then it will follow some other pattern. This proposal may be regarded as an hypothesis. It requires investigation and clarification
BIOPSY OF THE RECTUM IN ULCERATIVE COLITIS
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"
GEORGE LUMB M.D. READER
IN
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"
WESTMINSTER
MEDICAL
SCHOOL;
CONSULTANT PATHOLOGIST
R. H. B. PROTHEROE B.A., M.B. Camb.
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"
Lond., M.R.C.P.
PATHOLOGY,
SENIOR
IN
REGISTRAR
PATHOLOGY,
AND
RESEARCH FELLOW
IN ULCERATIVE COLITIS
WESTMINSTER
HOSPITAL,
LONDON
RAPID autolysis has hitherto greatly impeded study of the morbid histology of the gastro-intestinal tract. For this reason post-mortem material is of very limited value for detailed study, and examination of surgical specimens also has its limitations because of the degenerative changes that occur before fixation is complete. The stomach and the rectum, however, are accessible for biopsy and the gastric mucosa has been carefully studied with the biopsy gastroscope. Palmer (1954) has used this method extensively and reviewed the published reports. Rectal
biopsy
has been used almost
entirely
for
diagnosing
by neurophysiologists. In general, there is need for more effective communication between ethologists, psychologists, physiologists, and psychiatrists, and for the development of research in which - physiologists and students of behaviour
collaborate. Given such combination, it should be to devise possible experimental programmes which would lead to a rapid advance in the understanding of behaviour. I am grateful to Prof. T. Ferguson Rodger and his colleagues of the Department of Psychological Medicine, Glasgow University, and to Prof. J. B. S. Haldane, F.R.S., and Dr. H. Spurway, for illuminating discussions on the subject-matter of this paper. Especial acknowledgment is due tq Dr. A. B. Sclare for reading and criticising the manuscript. ADDENDUM
After this paper was written Kortlandt (1955) published a long and valuable discussion of the concept of instinct, with many references to the relationships betweenethological and psychiatric theories. I have also been allowed to see, in advance of publication, a paper by Morris in which an interesting aspect of the connection between autonomic function and behaviour is presented. Both papers can be recommended to anyone interested in the topics I have discussed. REFERENCES
Alexander, (1952a) Psychosomatic Medicine. — (1952b) In Alexander and Ross’s Dynamic Psychiatry. Chicago. Armstrong, E. A. (1950) Symp. Soc. exp. Biol. 4, 361. Barnett, S. A., Spencer, M.M. (1951) Behaviour, 3, 229. Bastock, M., Morris, D., Moynihan, M. (1953) Ibid, 6, 66. Darwin, C. (1872) The Expression of the Emotions in Man and
Fig. I-Age-distribution
—
cases
of ulcerative colitis.
Hardy, J. D. (1954) Harvey Lect. 49, 242. Hebb, D. O. (1949) The Organization of Behaviour. New York. Jones, E. (1953) Sigmund Freud, Life and Work. London ; vol. i: Kennedy, J. S. (1954) Brit. J. anim. Behav. 2. 12. Kortlandt, A. (1955) Arch. Néerl. Zool. 11, 155. Lissman, H. W. (1950) Symp. Soc. exp. Biol. 4, 34. Lorenz, K. Z. (1950) Ibid, p. 211. Maier, N. R. F. (1949) Frustration. New York. Miller, N. E. (1948) J. abn. soc. Psychol. 43, 155. Morris, D. Behaviour (in the press). Moynihan, M. (1953) Ibid, 5, 58. Ogden, C. K., Richards, I. A. (1931) The Meaning of Meaning London.
F.
Animals. London. Day, G. (1946) Lancet, ii, 703. Fessard, A. E. (1954) In Brain Mechanisms and Consciousness (edited by J. F. Delafresnaye). London. Freud, S. (1949) An Outline of Psychoanalysis. London. Fulton, J. F. (1949) Functional Localisation in the Frontal Lobes and Cerebellum. Oxford. Haldane, J. B. S. (1949) In Genetics, Paleontology, and Evolution. Princeton and Oxford. (1954) Behaviour, 6, 256. Halstead, W. C. (1951) In Cerebral Mechanisms in Behaviour. New York.
in 150
Russell, R. W. (1950) Brit. J. Psychol. 41, 95. (1953) Int. J. Psycho-Anal. 34, 1. Pretty, R. G. F. (1951) Quart. J. exp. Psychol. 3, 151. Schorstein, J. (1954) Personal communication. Selye, H. (1950) The Physiology and Pathology of Exposure to -
-
Stress. Montreal.
Spitz, R. A., Wolf, K. M. (1946) Genet. Psychol Monogr. 34, 57. Spurway, H., Haldane, J. B. S. (1953) Behaviour, 6, 8. Stellar, E. (1954) Psychol. Rev. 61, 5. Thorpe, W. H. (1950) Symp. Soc. exp. Biol. 4, 387. (1951) Bull. anim. Behav. 9, 34. (1954) Nature, Lond. 174, 101. Tinbergen, N. (1951) The Study of Instinct. Oxford. (1952) Quart. Rev. Biol. 27, 1. (1953) The Herring Gull’s World. London. van Iersel, J. J. A. (1947) Behaviour, 1, 56. Whitehorn, J. C. (1952) In Alexander and Ross’s Dynamic —
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Psychiatry. Chicago. Wolff, H. G. (1953) Stress and Disease. Springfield, Ill. Young, J. Z. (1950) Doubt and Certainty in Science. Oxford.