For several decades the spectacular increase and decrease of certain arctic mammals has stimulated research on populations. The crashes of rabbits were dramatized by Seton (1), and the suicidal movements of lemmings were publicized by many authors. However, as is so often the case, the conspicuous features turn out to be merlCy an cxtreme case of a very general phenomenon--namely, the fluctuations of a population. Investigators first sought an explanation for the "crash," but now most of them search for a description and understanding of the interaction and relative importance of the many factors that influence the ups and downs of populations.In this article we describe the current status of our understanding of population fluctuations, emphasizing the regulatory features that prevent populations from destroying the habitat. The research discussed is limited to work with mammals, since the mechanisms are best known for that class. It is assumed that the reader has knowledge of ecological principles such as density dependence and limiting factors.For many years it was assumed that epizootics, famine, and climatic factors terminated the explosive rises in population size and precipitated the often spectacular crashes (2). However, by the early 1940's it had become apparent that none of these mechanisms explained some of the observed declines in population, and it was suggested that factors intrinsic to the population were involved in its regulation Dr. Christian is affiliated with the Research Laboratories of the Albert Einstein Medical Center, Philadelphia, Pa.; Dr. Davis is professor of zoology at Pennsylvania State University, University Park.
(3). The skepticism toward earlier explanations was reflected further in a review by Clarke in 1949 (4), as well as in Elton's classic earlier work (5).Probably the greatest shift in emphasis has occurred since 1949; there has been an upsurge of investigations in which density-dependent changes in the animals themselves have been explored, and of theories in which the observed phenomena of population growth and decline (6-9) are explained in terms of biological mechanisms intrinsic in the populations and not only as results of the action of external factors. It is clear that food, climatic factors, and disease may cause population change. Indeed, it would be foolish to state that these factors do not, under certain circumstances, limit population growth or produce spectacular decline. The early investigations of Emlen, Davis, and their co-workers (8) on populations of Norway rats demonstrated clearly that environmental factors can reduce a population. For example, a drought followed by excessive rain resulted in a notable decline in rats in Baltimore (8). However, as early as 1946 spectacular declines in rat populations were found to be coincident with social disturbances rather than with environmental changes.The suspicion that social phenomena were involved prompted a search for mechanisms that could regulate the growth of populations in a densitydependent manner. No lo...