Irrelevant behaviour,information processing and arousal homeostasis

Summary This paper expands a new hypothesis on the causal mechanisms underlying irrelevant behaviour. It begins with a critical summary of earlier theories which attempted to explain displacement activities, but failed to predict the consistency with which certain types of behaviour are shown in stressful situations by a variety of species. Behavioural and physiological studies suggest that these behaviour patterns are closely associated with the incipient activation of sleep. The functional significance of this link and some of the causal processes which may be responsible for it are discussed. Paradoxically, however, displacement activities occur when animals are in a state of high arousal. The concept of arousal is reconsidered in the light of information theory and assumed to be closely correlated with the information processing rate in the nervous system. The relationships between neural and autonomic arousal are considered in this context. It is argued that over-arousal may occur when information handling exceeds the limited channel capacity of the system, with a consequent loss of efficiency. It is pointed out that there are mechanisms capable of controlling the information influx into the brain, and it is hypothesized that they are tied up in a feedback mechanism which regulates arousal and which involves the activation of a de-arousal system, corresponding to the neurological sleep mechanism. Displacement activities are viewed as consequences of this regulatory activation of the sleep system. This hypothesis is then compared with existing theories of displacement and its relationship with them is discussed.

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Zusammenfassung Die Arbeit erweitert eine neue Hypothese über die kausalen Mechanismen, die dem übersprungverhalten zugrunde liegen. Sie beginnt mit einer kritischen Übersicht der früher vorgeschlagenen Theorien und zeigt auf, daß diese nicht die Beständigkeit, mit der gewisse Verhaltensweisen von einer Anzahl von Tierarten in Stress-Situationen gezeigt werden, erklären. Verhaltens- und physiologische Studien deuten an, daß diese Verhaltensweisen in einem engen Zusammenhang mit einer partiellen Schlafaktivierung stehen. Die funktionelle Bedeutung dieses Zusammenhanges und die möglichen physiologischen Mechanismen, die dafür verantwortlich sein könnten, werden diskutiert. Paradoxerweise werden aber Übersprungsverhalten dann gezeigt, wenn sich Tiere in einem gesteigerten Wachzustand befinden. In diesem Zusammenhang wird das Arousal-Konzept im Lichte der Informationstheorie neu beleuchtet und es wird angenommen, daß der Arousal-Zustand eines Tieres im engen Zusammenhang mit der jeweiligen Informationsverarbeitungsrate im Nervensystem steht. Die Verhältnisse zwischen neuralem und autonomischen Arousal werden diskutiert. Es wird vorgeschlagen, daß sich Überarousal dann einstellt, wenn die Informationsverarbeitungsrate die Kanalkapazität des Systems übersteigt mit einem sich daraus ergebenen Wirkungsgradverlust. Es wird darauf hingewiesen, daß es Mechanismen gibt, die den Informationsfluß in das Gehirn regeln und angenommen, daß diese innerhalb eines Rückkopplungssystems funktionieren, welches Arousal reguliert. Diese Regelung benötigt die Aktivierung eines Arousal herabsetzenden Systems, das in den neurologischen Schlafmechanismen besteht. Das Auftreten von Übersprungsverhalten wird als eine Konsequenz dieser regulatorischen Aktivierung des Schlafsystems angesehen. Die Hypothese wird dann mit den vorhandenen Übersprungsverhaltenstheorien verglichen.

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The research underlying this paper has been supported by grants from the U.S. Air Force Office of Scientific Research through the European Office of Aerospace Research, the Science Research Council and the Royal Society to Professor N. Tinbergen, F. R. S. and myself. It was partly carried out at the Department of Zoology, Oxford. I am grateful to Dr. S. A. Hillyard who by revising an earlier version contributed much to whatever merits this paper has.     

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