Types of learning

Habituation

A classic example of habituation is the following observation on the snail Helix albolabris. If the snail is moving along a wooden surface, it will immediately withdraw into its shell if the experimenter taps on the surface. It emerges after a pause, only to withdraw again if the tap is repeated. But continued repetition of the same tapping at regular intervals elicits a briefer and more perfunctory withdrawal response. Eventually, the stimulus, which initially elicited a clear-cut, immediate response, has no detectable effect on the snail’s behaviour. Habituation has occurred.

Habituation can be defined in behavioral terms as a decline in responding to a repeatedly presented stimulus. As such, it is a very widespread phenomenon, one that can be observed in animals ranging from single-celled protozoans to humans. Most animals behave differently to novel and familiar stimuli: the former sometimes elicit startle responses, sometimes investigatory or exploratory responses; the latter often apparently are ignored. The suggestion that habituation is a simple form of learning, however, implies that it can be distinguished from some even simpler potential causes of this sort of change in behaviour. One reason why an animal might stop responding to a stimulus is that it no longer detects the stimulus; i.e., some form of sensory adaptation might have occurred. Another potential cause is fatigue: perhaps some temporary refractory state is produced by repeated elicitation of the same response, making it impossible to perform that response again. Whether or not one would want to call either of these processes a form of learning is doubtful. But both behavioral and physiological evidence establishes that habituation cannot be explained in these terms.

The critical behavioral evidence is that habituation can be disrupted by almost any change in the experimental conditions. If repeated presentation of one stimulus leads to habituation of a response, the same response can still be elicited by a different stimulus. Even if the experimenter presents a novel stimulus that does not itself elicit the response in question, its presentation may restore the response on the next trial in which the originally habituated stimulus is presented. This latter observation, usually referred to as an instance of dishabituation, seems to rule out any simple sensory adaptation; both observations rule out simple effector fatigue.

Neurophysiological analysis of habituation in various mollusks—for example, in the sea snail Aplysia—has confirmed that habituation need not depend on changes in the activity of sensory or motor neurons. In the case of Aplysia, researchers have studied the gill withdrawal reflex, a response that rapidly habituates to repeated stimulation of the snail’s siphon or mantle shelf. But habituation still occurs even if it is elicited by direct, electrical stimulation of the motor nerve, bypassing the sensory receptors completely; and recording from the sensory nerve during normal habituation reveals no decline in its level of activity. These observations eliminate sensory adaptation as a possible cause of the animal’s having ceased to respond to the stimulus. Effector fatigue can be ruled out by showing that direct stimulation of the motor neurons controlling the withdrawal response can still elicit a perfectly normal reaction even after the response has completely habituated. Research shows that habituation in Aplysia depends on changes in the activity of more central neurons. Repeated tactile stimulation of the siphon, leading to habituation of the withdrawal response, causes changes in the activity of the motor neurons innervating the response. Specifically, these motor neurons show a decline in excitatory postsynaptic potential, which is the electrical change that enables the nerve impulse to cross the gap (synaptic cleft) that separates one neuron in the pathway from the next. The decline in excitatory postsynaptic potential short-circuits the response. Moreover, the presentation of a novel stimulus, sufficient to dishabituate the behavioral response, restores the postsynaptic potential.

Habituation occurs even in animals without a central nervous system—probably in single-celled protozoans; certainly in animals such as the coelenterate Hydra, which have a diffuse nerve net and do not appear to be capable of associative learning. Among mammals, habituation of certain reflex responses can be observed even in “spinal” subjects, that is, those whose spinal cord has been severed from the brain. There can be little doubt, then, that habituation is not only widespread, but that it also can be a relatively simple phenomenon. There is, however, no guarantee that it is the same phenomenon wherever it appears. The waning response to a repeatedly presented stimulus admits of a number of different explanations. In principle, as we have already seen, it might be due to sensory adaptation, effector fatigue, or a more central neural change. These distinctions make rather little sense in the case of a single-celled animal. And one should not necessarily expect the habituation observed in a spinal mammal to involve precisely the same mechanisms as those responsible for comparable behavioral effects in an intact animal. Some psychologists have proposed theories of habituation that appeal to processes of classical conditioning. Such a theory is not likely to apply to the habituation observed in an animal that shows no capacity for classical conditioning.

Habituation is usually, as here, classified as an instance of simple, nonassociative learning. It is supposedly nonassociative because all that happens in the course of habituation is that a stimulus is repeatedly presented and the animal’s behaviour changes; there is, on the face of it, no other event with which the stimulus can be associated. Habituation must therefore, it appears, be understood by reference to some change in the pathway between stimulus and response, and the work with Aplysia and other mollusks shows how this analysis may proceed at the physiological level. But if habituation is not always the same phenomenon, it is possible that different processes may underlie the habituation of the startle response to a loud noise in an intact mammal. And despite appearances to the contrary, those processes may involve some associative learning. One suggestion is that novel stimuli elicit a biphasic response: an initial increase in startle responses, which include components of emotion or anxiety, followed by a rebound in the opposite direction. Habituation occurs when the latter, rebound response becomes conditioned to the stimulus, occurring sooner and sooner with each repetition of the stimulus and thereby damping down and eventually canceling out the initial reaction. An alternative possibility is that long-term habituation depends on associating the repeatedly presented stimulus with the context in which it occurs, a suggestion that would explain why presentation of the stimulus in a different context sometimes leads to dishabituation.

The generality of habituation implies that this behavioral phenomenon has considerable adaptive significance; if true, it would be quite reasonable to expect that a number of different mechanisms might have evolved to produce the behavioral result. The adaptive value of habituation is not difficult to see. A novel stimulus may signify danger, and an animal should react to this stimulus either by withdrawing or at least by orienting toward it to see what will happen next. But if the same stimulus occurs again with no further consequence, it is probably safe: regular repetition of the same stimulus implies that it is part of the background, such as the waving of a branch in the wind or the shadow caused by a piece of seaweed floating with the waves. If the stimulus is not dangerous, time should not be wasted on it. Withdrawal, especially in the case of a snail into its shell, is a time-consuming effort, incompatible with such vital activities as searching for food. If it is important, therefore, for animals to be wary of novel stimuli, it is equally important that they should discriminate the novel and potentially dangerous from the familiar and probably safe.