stereotyped response

▪ biology

Introduction

      unlearned behavioral reaction of an organism to some environmental stimulus. It is an adaptive mechanism and may be expressed in a variety of ways. All living organisms exhibit one or more types of stereotyped response.

General considerations

      The capacity for unlearned behaviour is genetically determined in much the same sense as are the position, size, shape, and function of organs. Like structural features, stereotyped responses are the result of a continuing process of evolutionary modification and refinement. Those actions that most successfully aid the animal or plant in its basic drives (e.g., reproduction, search for nourishment, escape from predators) are the ones most likely to be retained in succeeding generations. As environmental conditions change, inherently determined responses also become modified in order to ensure continuation of the species.

      The problems that arise in the study of stereotyped responses are many and varied. Particular responses in animals do not readily lend themselves to identification in highly evolved forms because learned behaviour patterns obscure the underlying unlearned behaviour; in addition, stereotyped responses provide the building blocks of instinctive behaviour, the complexity of which may obscure the integral parts (see instinct). In lower animals, just as in plants, in which learned behaviour is absent or nil, the analysis of behavioral mechanisms is limited by the fact that many of the most fundamental cell processes are not well understood.

      Animal behaviour, as a branch of psychology, represents a confluence between the disciplines of ethology and comparative psychology. Most of the pioneer work in stereotyped responses of animals was done by ethologists. During the first half of the 20th century, when much of the groundwork in experimental psychology was laid, ethologists (who were for the most part European) concerned themselves with behaviour in insects, fishes, and birds and were particularly interested in the evolution of instinct. The comparative psychologists during this formative period were mostly Americans. They studied primarily behaviour in common laboratory animals such as guinea pigs, mice, rats, and monkeys, and their interest tended to focus on environmental influences on behaviour as opposed to genetic influences. Since the 1950s, psychologists in general have recognized that both environmental and genetic factors play essential roles in any biological phenomenon. As a consequence of the separate development of ethology and comparative psychology, however, some difficulties have arisen in the use of terminology. The German-American biologist Jacques Loeb (Loeb, Jacques) applied the term tropism to all oriented movements of organisms, and he proposed that all behaviour is composed of tropisms. Subsequently, to avoid confusion, the terms taxes (singular: taxis) and kineses were introduced by other investigators to refer to animal responses other than those of sedentary, plantlike forms. The terms also have been applied to certain plant movements. Although a variety of discrete stereotyped response movements occur in plants, particularly in higher forms such as flowering plants, these autonomous movements usually occur too slowly to permit detection by casual observation. That movements of plants or plant organs actually take place can be strikingly demonstrated by time-lapse photography (photography, history of), in which single photographs are taken at regular intervals as brief as seconds or as long as several days or more. The photographs are then compared or viewed in rapid sequence as a motion picture.

Types of stereotyped responses

      Stereotyped response in animals may be separated into the following four categories: unorganized or poorly organized response, reflex movements of a particular part of an organism, reflex-like activity of an entire organism, and instinct.

      Unorganized or poorly organized responses are given by early embryos or by animals (such as sponges) that lack nervous systems.

reflex

      Reflexes proper, or reflex-arc movements, include responses such as the immediate withdrawal of the hand on touching a hot surface. The basic components of the reflex arc are the receptor, or sensory-nerve cell, which senses the stimulus, and the affector, the nerve cell that directly activates the muscle. These are a theoretical minimum rather than an observed functional arrangement of cells in the body of an animal (see instinct: Varieties of instinctive behaviour (instinct)).

Reflex-like activities

      Reflex-like activities of entire organisms may be unoriented or oriented. Unoriented responses include kineses—undirected speeding or slowing of the rate of locomotion or frequency of change from rest to movement (orthokinesis) or of frequency or amount of turning of the whole animal (klinokinesis), the speed of frequency depending on the intensity of stimulation. Examples of orthokinesis are seen in lampreys, which are more active in high intensities of light, and in cockroaches, which are more active in low intensities; flatworms and many kinds of fly larvae, among other invertebrates, exhibit orthokinesis. Klinokinesis is well demonstrated by the movements of the wood louse (Porcellio scaber). When wood lice are placed in dry air, they crawl about actively but without direction until they become gradually dehydrated. When the wood lice are placed in humid air, they move at first, but any activity they exhibited soon ceases and they become quiet. Wood lice placed in a container with dry air at one end and humid air at the other gradually congregate at the humid end. This transfer is achieved through what appear to be random rather than directed movements.

      Oriented reflex activities of entire organisms include tropisms, taxes, and orientations at an angle. Tropisms in animals are those directed growth-curvature movements of sessile (i.e., sedentary) forms that lead to equal intensities of stimulation of symmetrically placed body parts. These movements are demonstrated by hydroid animals such as Eudendrium.

Taxes

      Taxes may be described as oriented locomotory reactions of motile organisms. They exist in purest form as oriented, forced movements; that is, as reflex actions of entire organisms. When exposed to a single source of stimulation, the body is oriented in line with the source. Movement toward the source is said to be positive; that away from it is negative.

      Klinotaxis is the achievement of orientation by alternate lateral movements of part or all of a body; there appears to occur a comparison of intensities of stimulation between one position and another and a “choice” between them. Klinotaxis is shown by animals with a single intensity receptor such as the protozoan Euglena, earthworms, and fly larvae. For several days before going into the pupal (or resting) state, the blowfly (blow fly) maggot tends to move away from a light source. As it crawls, it swings its head alternately left and right. Presumably a light receptor is located on the maggot's head, and differences in intensity between successive light stimuli as it moves its head determine the direction in which the maggot travels. This type of response is given more commonly to chemical stimuli than to light.

      In tropotaxis, attainment of orientation is direct, resulting from turning toward the less stimulated (negative) or more stimulated (positive) side as simultaneous, automatic comparisons of intensities on two sides of the body are made. No deviations (trial movements) are required. Tropotaxis is shown by animals with paired intensity receptors. If exposed to stimulation from two sources, orientation is to some intermediate point and is determined by the relative intensity of the sources. If one receptor is effectively covered, the animal moves in spirals (circus movement). Tropotaxis is shown by many arthropods, especially insects.

      In telotaxis, known only for responses to light, attainment of orientation is direct and without trial movements. When between lights from two sources, the animal orients to one light, rather than to some intermediate point. The animal switches orientation from one source to the other at unpredictable intervals and consequently follows a zigzag course. The response is given to the source as though it were a goal. Bilateral balance is not necessary, and circus movements, if they occur, demonstrate that the animal is reacting tropotactically rather than telotactically. Honeybees (Apis) and hermit crabs (Eupagurus), among others, show telotaxis.

      Orientations at an angle (transverse orientations) may or may not be accompanied by locomotion. They include the light-compass reaction (menotaxis) and dorsal (or ventral) transverse reaction. Menotaxis is shown by foraging insects such as ants (ant) and bees that return to a fixed nest. It has been demonstrated experimentally by covering for 2 ¹/₂ hours an ant returning to its nest. After being uncovered, the ant proceeds not toward the nest but at the same angle to the Sun that it had been moving at the time it was hidden from the light.

      In another demonstration of menotaxis, the sea slug Elysia viridis has been shown to move at angles of from 45° to 135° in relation to a steady source of light. No satisfactory explanation for this type of response in the sea slug is known.

      Dorsal (or ventral) transverse reaction is demonstrated when the impact of the stimulus is kept at right angles to both longitudinal and transverse axes of the body. Locomotion need not occur. This reaction is given to light by various aquatic crustaceans—Argulus, the fish louse, and Artemia, the brine shrimp—and is given to gravity by crayfish.

Ed.

Additional Reading

Studies include Jacques Loeb, Forced Movements: Tropisms and Animal Conduct (1918, reprinted 1973); J.D. Carthy, Animal Navigation: How Animals Find Their Way About (1956, reissued 1963); Gottfried S. Fraenkel and Donald L. Gunn, The Orientation of Animals: Kinesis, Taxes, and Compass Reactions, new ed. (1961); and Hermann Schöne, Spatial Orientation: The Spatial Control of Behavior in Animals and Man (1984; originally published in German, 1980), written as an overview of the field of orientation for advanced readers in animal behaviour.Stereotypic behaviour, a repetitive behaviour with no obvious goal, is an important behaviour of caged or intensive animals; it may be an indication of poor welfare and is thought to be the animal's attempt to cope with a sterile environment. Alistair B. Lawrence and Jeffrey Rushen (eds.), Stereotypic Animal Behaviour: Fundamentals and Applications to Welfare (1993), covers all the theories of the importance of stereotypic behaviour, the ways in which they are carried out, and the implications for animal welfare.

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