Yawning in the Greenfinch
JO Harrison
1968; 55; 511

mise à jour du
7 août 2003
Displacement activities and arousal
Juan D. Delius
Department of Zoology, University of Oxford
Pandiculation: the comparative phenomenon of systematic stretching
Les comportements instinctifs G.Richard


Irrelevant behaviour, information processing and arousal homeostasis
Juan D. Delius
Yawning : a displacement activity ?
Le bâillement: une activité substitutive ?
Self-injurious and stereotypic behavior
In 1940 Tinbergen and Kortlandt independently drew attention te a behavioural phenomenon which has since been called displacement activity and has received a good deal of attention. Although no binding rules exist by which displacement behaviour can be recognized, the term is applied to behaviour patterns which appear to be out of context with the behaviour which closely precedes or follows them either in the sense that they do not seem functionally integrated with the preceding or following behaviour or that they occur in situations in which causal factors usually responsible for them appear to be absent or at least weak compared with those determining the behavioural envelope.
Displacement activities occur in three situations: motivational. conflict, frustration of consummatory acts and physical thwarting of performance. Several theories have been put forwayd to explain the causal mechanism involved. A variety of behaviour patterns have been reported as displacement activities, even in a single species, but this variety needs revision.
Monographie treatments of the behaviour of any one species usually indicate only two or three activities which according te the judgment of the observer occur commonly as displacement. None of the theories on displacement activities gives cogent reasons why particular behaviour patterns should be more common than others as displacement activities, apart from stating that the causal agents which usually elicit them in non-displacement situations can also be presumed te be present, if only weakly, in the displacement context, or remarking that those patterns are prepotent in the repertoire of the animal.
In the course of a systematic exploration of the forebrain and brainstem of herring and lesser black-backed gulls (Larus argentatus and L. fuscus) with electrical stimulation, information was obtained which may bear on this particular issue. The gulls with chronically implanted monopolar electrodes (conical active ares, -0-25 mm2) were stimulated with a sine current at 50 c/s ranging between 10 and 150 µamp root mean square current, in repeated trains lasting 30 sec-5 min for several testing sessions, over 2 or more months. The electrode tip locations were checked histologically.
A great variety of responses have been obtained, but here we will only consider a behavioural syndrome which is characterized by preening and staring down, and more rarely by pecking, yawning, squatting, relaxation (fluffing of plumage, shortening of the neck, general diminution of activity, intermittent closure of eyes) and occasional sleep. We find that several, and sometimes all, these component patterns can often be elicited from single loci with the same stimulation strength usually less than 50 µamp, either as a result of a single stimulation train or more frequently in the course of several consecutive trains.
In Table 1 all 202 loci so far explored have been classified into those which gave preening and those which did not. Within each class of loci the percentage which yielded the different other components is shown. All the component patterns wore associated with electrodes eliciting preening rather than with those which did not, and the association is significant. A similar relationship may also hold for mandibulation, shaking the body and head, wagging the tail and shaking the foot, but because those patterns are also frequent during control periods without stimulation, a decision is difficult. No such association could be dotected for some twentyfive other various behaviour patterns examined.

It is significant that the preening positive points clustered in several discrete anatomical areas of the telencephalon and diencephalon and that ten electrodes responsible for more than half the entries in the non-preening class also lay within or close to these areas. There is net sufficient information to decide whether the associations of components are stronger in some areas than in others, although some evidence points in this direction.

The conclusion that these diverse behaviour patterns reflect the activation of a more or less unitary system leading to dearousal, and are not a result of the simultaneous stimulation of contiguous but otherwise unrelated neural systems, is supported by observations on unstimulated normal gulls which suggest a high temporal and sequential association between the component patterns including sloop. Furthermore, preliminary experiments indicate that at least two hypnotic drugs, pentobarbital sodium and tribromoethanol, given systemically, reliably elicit the syndrome just described. Preening, staring down and pecking the ground are also the patterns most often involved in displacement behaviour in gulls. A similar relationship seems te hold in some other species.

In the cat Parmeggiani stimulated several different and separate areas of the forebrain and brainstem and obtained a behavioural complex consisting of sniffing, grooming, yawning, lying down, curling up, dozing and sleeping. He emphasizes that this behaviour is normal in unstimulated cats. Rowland and Gluck presont some evidence that in a certain conditioning procedure grooming replaced the synchronization of the electroencephalogram shown by sleeping cats when those were tested awake. Again, Leyhausen lists grooming, sniffing and lying down as displacement behaviour for the cat.

For the rat, Caspers has shown that grooming and some other unfortunately unspecified "motorautomatisms" are associated with shifts in the cortical d.c. potential making the surface positive, which otherwise are typical of sloop, while shifts towards a negative surface are characteristic of the waking animal. Grant mentions grooming, digging and sniffing as typical displacement activities of rats.

Theee facts suggest that grooming or preening and certain other movements are largely controlled by neurophysiological mechanisms which are also responsible, for de-arousal and sleep. On the other hand they are often involved in displacement bohaviour. There is little doubt, however, that the behavioural situations loading to displacement, that is, conflict, frustration and thwarting, are effective in increasing arousal. I suggest that the occurrence of at least some displacement activities is the reflexion of a homeostatic process operating towards cancelling the arousal increment so generated, through the activation of an arousal inhibiting system. The existence of arousal homeotasis has been suggested by Borlyne, who also marshals supporting empirical evidence. Such regulation appears logically necessary if arousal is correlated with the rate at which information is handled and if the nervous systera is considered as a communication channel of restricted and specific capacity whore for maximum efficiency the information handling rate must be hold within certain limits.

The striking occurrence of slep as displacement in several species may be regarded in this context as a regulatory overshoot. Why grooming and other patterns should be activated by a neural de-arousal system is functionally not obvious, but perhaps those types of activities are conducive to de-arousal in a similar way to other activities, possibly through stimulus reduction (cut-off), switch of attention to stimuli of little novelty or generation of repetitive stimulation.

Displacement Behavior

Maestripieri, D., Schino, G., Aureli, F. & Troisi, A. A modest proposal: displacement activities as an indicator of emotions in primates. Anim. Behav. 1992; 44; 967-979.
Displacement behavior is usually thought of as self-grooming, touching, or scratching, which is displayed when an animal has a conflict between two drives, such as the desire to approach an object, while at the same time being fearful of that object. With the fall of drive theory into disfavor, animal behaviorists paid little attention to displacement behavior until Maestripieri et al. (1992) pointed out that displacement behavior might be a good measure of anxiety levels.
Since then a sizeable literature evaluating the effectiveness of displacement behavior measures (also called self directed behavior, or SDB) as indicators of anxiety has grown. Particularly attention has been paid to primates, including humans. Measures of displacement behavior, for example, have been applied in psychiatric studies of anxiety (Troisi et al. 2000).
Castles et al (1999) found that SDB's increased in wild olive baboons between depending on whether the nearest animal (to the animal being watched) was dominant or not. Dominant animals caused in increase in self directed behaviors by 40 %, indicating a higher level of social anxiety caused by the proximity of a dominant animal.
Chimpanzees also display higher levels of SDB's in anxiety inducing situations. Baker and Aureli (1997) found, in captive chimps, that vocalizations from animals in neighboring cages stimulated more SDB's when the chimps were housed in groups. Socially isolated chimps, however, did not respond with SDB's to vocalizations coming from neighboring cages. The vocalizations may suggest to the chimps that an attack is imminent.
Baker and Aureli (1997) suggest that the isolated animals realize that no other chimps are in their cage, and consequently they feel safe even when hearing the vocalizations. Leavens et al. (2001) gave captive chimps problems of varying difficulty to solve. If their chimps started with an easy problem and then progressed to more difficult problems they displayed more SDB's when confronted with the difficult problems. Chimps who only received difficult problems did not display more SDB's. Positive auditory reinforcement during the problem reduced SDB's.
Research on animal welfare has used displacement behavior as a measure of anxiety induced by captivity or other manipulations. The effectiveness of nterventions intended to ameliorate the effects of captivity on animals may be measured by reductions in self directed behaviors.
Baker K. C., Aureli F. 1997. Behavioural indicators of anxiety: An empirical test in chimpanzees. Behaviour 134: 1031-1050.
Castles D. L., Whiten A., Aureli F. 1999. Social anxiety, relationships and self-directed behaviour among wild female olive baboons Animal Behaviour 58: 1207-1215.
Hughes B. O., Duncan I. J. H. 1988 The notion of ethological need, models of motivation and animal-welfare. Animal Behaviour 36: 1696-1707.
Leavens DA, Aureli F, Hopkins WD, Hyatt CW 2001 Effects of cognitive challenge on self-directed behaviors by chimpanzees (Pan troglodytes) American Journal of Primatology 55 (1): 1-14
Maestripieri D., Schino G., Aureli F., Troisi A. 1992. A modest proposal - displacement activities as an indicator of emotions in primates. Animal Behaviour 44: (5) 967-979
Manson JH, Perry S 2000 Correlates of self-directed behaviour in wild white faced capuchins Ethology 106 (4): 301-317
Troisi A., Belsanti S., Bucci A. R., Mosco C., Sinti F., Verucci M. 2000 Affect regulation in alexithymia - An ethological study of displacement behavior during psychiatric interviews. Journal Of Nervous And Mental Disease 188: (1) 13-18
Tous les travaux de B. Deputte
Tinbergen N Quart Rev Biol 1952;27:1-32
An interpretation of the "displacement phenomenon"
Bindra D British J Psychology 1959:32:236-268
Displacement activities and arousal
Delius J Nature-1967;214:1259-1260
Displacement activities as a behavioral measure of stress in nonhuman primates and human subjects
Troisi A Stress 2002;5(1):47-54
A modest proposal: displacement activities as an indicator of emotions in primates
Maestripieri D, Schino G, Aureli F, Troisi P Anim Behav 1992;44:967-979
The effects of fluoxetine and buspirone on self-injurious and stereotypic behavior in adult male rhesus macaques
Fontenot MB, Padgett EE et al Comp Med 2005;55(1):67-74
Effects of outdoor housing on self-Injurious and stereotypic behavior in adult Male Rhesus Macaques (Macaca mulatta)
Fontenot MB, Wilkes MN, Lynch CS J Am Ass Laboratory Animal Science 2006; 45(5):35-43
Extinction deficits in male rhesus macaques with a history of self-injurious behavior
Lutz C, Tiefenbacher S, Meyer J, Novak M. Am J Primatol 2004;63(2):41-48
Inhibition of social behavior in chimpanzees under high-density conditions
Aureli F, de Waal FB Am J Primatol 1997;41(3):213-28
Frequencies and contexts of gape yawn displays of free-ranging Patas Monkeys
Zucker EL, Gerald MS, Kaplan JR Am J Primatol 1998;45(2):215
Pandiculation: the comparative phenomenon of systematic stretching
Fraser AF Appl Anim Behav Sci 1989;23:263-268
 An ethological interpretation of stereotypy induced by environmental stimulus
Beckmann H, Zimmer R Arch Psychiatr Nervenkr 1981;230(1):81-89
Revue sur le comportement de bâillement chez les vertébrés.
Deputte BL Bull interne société française pour l'étude du comportement animal.1974;1:26-35.
Uber das Gähnen bei Vögeln
Bergmann H Die Vögelwelt 1966;87(5):134-138
Zur Frage des Gähnens bei der Vögel
Löhrl H Die Vögelwelt 1967;88(3):85-86
Maintenance activities
Dilger W Zeitsch. Tierpsychologie 1960;17:649-685
Yawning in the Greenfinch
Harrison JO AUK 1968;55:511
Yawning and other maintenance activities in the South African Ostrich
Sauer EG, Sauer EM The Auk 1967;84:571-587
Zum geruchlichen Beutefinden und Gähnen der Kreuzkröte
Heuser H Zeitschrift für TierPsychologie 1958;15:94-98
New evidence for a locus coeruleus norepinephrine connection with anxiety.
Redmond DE, Wang Y Life Sciences 1979;25(26):2149-2162
Limbic-midbrain lesions and acth-induced excessive grooming
Colbern D et al. Life Sciences. 1977;21:393-402
Aggression does not increase friendly contacts among bystanders in geladas (Theropithecus gelada) Leone A, Mignini M, Mancini G, Palagi E. Primates. 2010;51(4):299-305.

Herrelko ES, Buchanan-Smith HM, Vick SJ .Perception of Available Space During Chimpanzee Introductions: Number of Accessible Areas Is More Important Than Enclosure Size. Zoo Biology 2015;xx:1-9