The biopsychology of maternal behavior in nonhuman mammals

The term "maternal behavior," when applied to nonhuman mammals, includes the behaviors exhibited in preparation for the arrival of newborn, in the care and protection of the newly arrived young, and in the weaning of those young, and represents a complex predictable pattern that is often regarded as a single, comprehensive, species-specific phenomenon. Although the delivering first-time mammalian mother is immediately and appropriately maternal, a "virgin" with no prior exposure to young does not show immediate and appropriate behavior toward foster young. Nevertheless, the virgin female, and indeed the male, possess the neural circuitry that underlies the pattern referred to as maternal behavior, despite not exhibiting the pattern under normal circumstances. At parturition, or after extensive exposure to young, what emerges appears to be a single stereotyped maternal behavior pattern. However, it is actually a smoothly coordinated constellation of simpler actions with proximate causes that, when sequenced properly, have the appearance of a motivated, purposive, adaptive pattern of caretaking. Over the past 50 years, much research has focused on finding the principal external and internal factors that convert the nonmaternal behavior patterns of the nonpregnant nullipara, the virgin, to the almost immediate and intense maternal behavior characteristic of the puerpera, the mother. This review is an attempt to summarize the many comprehensive, even encyclopedic, reviews of these factors, with an emphasis on brain mechanisms, and to highlight the gaps that remain in understanding the processes involved in the almost immediate onset of maternal caretaking behaviors observed in mammals at delivery. Where possible, the reader is directed to some of those excellent reviews.