The young of many species are born relatively helpless: in songbirds, rats, cats, dogs, and primates, the hatchling or newborn infant is wholly dependent on its parents. These are altricial species. In other species, such as domestic fowl, ducks, geese, ungulates, and guinea pigs, the hatchling or newborn is at a more advanced stage of development. These are precocial species, and their young are capable, among other things, of walking independently within a few minutes or hours of birth, and therefore of wandering away from their parents. Since mammals are dependent on their mothers for nourishment, and even birds are still dependent on parental guidance and protection, it is important that the precocial infant not get lost in this way. The phenomenon of filial imprinting ensures that, in normal circumstances, the precocial infant forms an attachment to its mother and never moves too far away.

Although imprinting was first studied by the Englishman Douglas Spalding in the 19th century, Konrad Lorenz is usually, and rightly, credited with having been the first not only to experiment on the phenomenon but also to study its wider implications. Lorenz found that a young duckling or gosling learns to follow the first conspicuous, moving object it sees within the first few days after hatching. In natural circumstances, this object would be the mother bird; but Lorenz discovered that he himself could serve as an adequate substitute, and that a young bird is apparently equally ready to follow a model of another species or a bright red ball. Lorenz also found that such imprinting affected not only the following response of the infant but also many aspects of the young bird’s later behaviour, including its sexual preferences as an adult.

Imprinting, like song learning, involves a sensitive period during which the young animal must be exposed to a model, and the learning that occurs at this time may not affect behaviour until some later date. In other words, one can distinguish between a process of perceptual or observational learning, when the young animal is learning to identify the defining characteristics of the other animal or object to which it is exposed, and the way in which this observational learning later affects behaviour. In the case of song learning, observation establishes a template that the bird then learns to match. In the case of imprinting, observation establishes, in Lorenz’ phrase, a model of a companion, to which the animal subsequently directs a variety of patterns of social behaviour.

With imprinting, as with song acquisition, one can ask why learning should be necessary at all. Would it not be safer to ensure that the young chick or lamb innately recognized its mother? There are, in fact, genetic constraints on the range of stimuli to which most precocial animals will imprint. A model of a Burmese jungle fowl (the species whose domestication produced domestic chickens) serves as a more effective imprinting object for a young chick than does a red ball; there is even evidence that imprinting in the latter case involves different neural circuits from those involved in imprinting to more natural stimuli. Nonetheless, it is clear that the innate constraints are not very tight and that a great deal of learning normally occurs. The most plausible explanation, as in the case of song learning, is that imprinting involves some measure of individual identification. Lorenz argued that one of the unique characteristics of imprinting was that it involved learning the characteristics of an entire species. It is true that imprinting results in the animal directing its social and mating behaviour toward other members of its own species, and not necessarily toward the particular individuals to which it was exposed when imprinting occurred. But learning usually involves some generalization to other instances, and there does not seem to be anything peculiar to imprinting here. The primary function of imprinting, however, is to enable the young animal to recognize its own mother from among the other adults of its species. This no doubt is particularly important in the case of such animals as sheep, which live in large flocks. Only learning could produce this result.

There is also an important element of individual recognition in at least some cases of imprinting’s effects on sexual behaviour. Experiments with Japanese quail have shown that their sexual preferences as adults are influenced by the precise individuals to whom they are exposed at an earlier age. Their preferred mate is one like, but not too like, the individuals on whom they imprinted. The preference for some similarity presumably ensures that they attempt to mate with members of their own species. The preference for some difference is almost certainly a mechanism for reducing inbreeding, since young birds will normally imprint on their own immediate relatives.

The difference between imprinting and song learning lies in the consequences of observational learning. The effect of imprinting is the formation of various forms of social attachment. But what mechanism causes the young chick or duckling to follow its mother? Lorenz thought that imprinting was unrewarded, yet the tendency of a young bird to follow an object on which it has been imprinted in the laboratory can be enhanced by rewarding the bird with food. Rewards also occur outside the laboratory: the mother hen not only scratches up food for her young chicks, she also provides a source of warmth and comfort. Moreover, following is also rewarded by a reduction in anxiety. As chicks develop over the first few days of life, they show increasing fear of unfamiliar objects; they allay this anxiety by avoiding novel objects and approaching a familiar one. This latter object must be one to which they have already been exposed—in other words, one on which they have imprinted. Imprinting works because newly hatched birds do not show any fear of unfamiliar objects, perhaps because something can be unfamiliar only by contrast with something else that is familiar. On the contrary, the newly hatched birds are attracted toward salient objects, particularly ones that move. Once, however, a particular object has been established as familiar and its features identified, different objects will be discriminated from it. These will be perceived as relatively unfamiliar, and hence they will provoke anxiety and the attempt to get as close as possible to the more familiar object. The imprinting of the young bird on one object necessarily closes down the possibility of its imprinting on others, as these will always be relatively less familiar. Thus, there is normally a relatively restricted period in the first few hours or days of life during which imprinting can occur. The only way to prolong this period is to confine the newly hatched bird to a dark box where it is exposed to no stimuli; prevented from imprinting during this period of confinement, the bird imprints on the first salient object it sees after emerging.