Population regulation: Historical context and contemporary challenges of open vs. closed systems

By definition, a population is regulated if it persists for many generations with fluctuations bounded above zero with high probability. Regulation thus requires density-dependent negative feedback whereby the population has a propensity to increase when small and decrease when large. Ultimately, extinction occurs due to regulating mechanisms becoming weaker than various disruptive events and stochastic variation. Population regulation is one of the foundational concepts of ecology, yet this paradigm has often been challenged, during the first half of the 20th century when the concept was not clearly defined, and more recently by some who study demographically open populations. The history of ecology reveals that earlier manifestations of the concept focused mostly on competition as the mechanism of population regulation. Because competition is often not evident in nature, it was sometimes concluded that population regulation was therefore also absent. However, predation in the broadest sense can also cause density dependence. By the 1950s, the idea that demographic density dependence was essential (but not sufficient) for population regulation was well established, and since then, challenges to the general concept have been short lived. However, some now believe that metapopulations composed of demographically open local populations can persist without density dependence. In particular, some recent manifestations of the Recruitment Limitation Hypothesis all but preclude the possibility of regulation. The theory of locally open populations indicates that persistence always relies on direct demographic density dependence at some spatial and temporal scale, even in models reportedly demonstrating the contrary. There is also increasing empirical evidence, especially in marine, systems where competition for space is not self evident, that local density dependence is more pervasive than previously assumed and is often caused by predation. However, there are currently insufficient data to test unequivocally whether or not any persistent metapopulation is regulated. The challenge for more complete understanding of regulation of metapopulations lies in combined empirical and theoretical studies that bridge the gap between smaller scale field experiments and larger scale phenomena that can presently be explored solely by theory.