From Freud to a modern understanding of behavioral, physiological, and brain development.

Seymour Levine's first "early experience" experiments were inspired by Freud. Yet, Levine's lifetime of work, and the work of his colleagues and scientists who followed, unveiled a myriad of early experience effects that even Freud himself could not have imagined. Related to and extending beyond his work on early experience, Levine also made important, often seminal, contributions to overlapping and related areas, such as early maternal separation and deprivation, maternal behavior and physiology, sexual differentiation, perinatal malnutrition, attachment in non-human primates, hypothalamic-pituitary-adrenal (HPA) stress reactivity and its adaptive significance, and the development of the HPA system. Moreover, his work spawned new lines of research by investigators active today. The papers contained in this special issue provide a sampling of research demonstrating some of the important directions in which those earliest experiments have led, many with clinical applications.

Dorsolateral columns of the spinal cord are necessary for both suckling-induced neuroendocrine reflexes and the kyphotic nursing posture in lactating rats.

Maternal behavior in rats consists of active behaviors, such as retrieval and licking of pups, and quiescent nursing, including the suckling-induced kyphotic (upright, dorsally-arched) posture. Because lesions of the dorsolateral, but not of the dorsal, columns are known to prevent the suckling-induced milk-ejection reflex, we asked whether the same is true for kyphosis as well. Bilateral lesions of the dorsolateral funiculus (DLF) or dorsal columns (DC) at spinal segments C(4-6) were made on day 5-8 postpartum; controls (CON) were subjected to a sham procedure. All aspects of maternal behavior and lactation were present in CON and DC dams soon after treatment. Among DLF dams, two had poor postural, ambulatory, and ingestive recovery that was associated with large lesions extending to the ventrolateral columns, while one with very small lesions continued to lactate. Of the remaining eight DLF dams, milk ejection was lost while recovery of retrieval and licking of pups occurred in all (between 1 and 4 days after surgery). All eight were quiescent for long periods in response to suckling but they did not display sustained kyphosis; rather, they nursed while prone or hunched over the pups, with little or no leg support, or while supine. Ventral trunk cutaneous sensitivity was present in all subjects. These data suggest that the dorsolateral funiculus relays both suckling-induced neuroendocrine and postural nursing reflexes that are mediated by separate supraspinal regions, hypothalamus and the ventrolateral sectors of the caudal periaqueductal gray, respectively.

Thermal control of mother-young contact revisited: hyperthermic rats nurse normally.

Morphine (MOR) is known to inhibit maternal behavior and induce hyperthermia; at appropriate doses, concurrent administration of naloxone (NAL) counteracts its disruption of maternal behavior but not the hyperthermia. We used these findings to evaluate the view that lactating rats terminate nursing due to intolerable hyperthermia. After a dam-litter separation of 4 h on Day 7 postpartum (PP), mother-litter interactions were observed continuously for 1 h. One hour before reunion, the dams received two injections (1 ml/kg ip each) of saline (SAL), MOR (20 mg/kg) and/or NAL (1 mg/kg) in the following combinations (n = 7 each): SAL + SAL, SAL + NAL, MOR + SAL or MOR + NAL. MOR profoundly disrupted maternal behavior, thereby preventing litter weight gains; these effects were completely counteracted by NAL, which alone had no discernible effects. In contrast, MOR-induced hyperthermia (approximately 0.7 degrees C increase in each hour, before and after reunion with pups) was not antagonized by NAL at the doses used. Thus, an additional 0.7-1.4 degrees C of body temperature (T) did not delay the onset or reduce the duration of nursing compared with SAL-treated controls. Further, there were no group differences in behaviors displayed both shortly before and after a nursing bout that included milk ejections or in the resumption of nursing. Together with earlier methodological and empirical criticisms of the thermal control theory, as well as knowledge about the somatosensory determinants of nursing, the present results suggest that nursing bouts in lactating rats are not limited by the mother's T.

Acute hunger of rat pups elicits increased kyphotic nursing and shorter intervals between nursing bouts: implications for changes in nursing with time postpartum.

Earlier findings, based on limited behavioral observations, indicate that nursing behavior in rats declines dramatically in duration over time postpartum-despite increasing ingestion of milk by rat pups to meet their growth and metabolic needs-although hungry pups elicit more nursing than do well-nourished pups. The authors compared the nursing pattern in detail for 6 hr on Days 7 and 14 and induced hunger in pups acutely with mammary-duct-ligated dams unable to provide milk. Compared with Day 7, on Day 14, supine nursing and the interval between nursing bouts increased, whereas hovering over pups and kyphotic nursing decreased. When pups were increasingly hungry, these age-related changes were counteracted. Thus, the ingestive motivation of pups largely regulates the nursing pattem over time.