Article,

The Ecological and Genetic Consequences of Density-Dependent Regulation in Plants

, and .
Annual Review of Ecology and Systematics, 11 (1): 411-452 (1980)
DOI: 10.1146/annurev.es.11.110180.002211

Abstract

Understanding the mechanisms of population size regulation is of vital importance to both the pure and applied ecologist. Throughout the history of ecology one school of thought has maintained that populations of plants and animals are in some way regulated by "density-dependent factors"­ i.e. processes that either increase mortality or decrease fecundity as the density of the population increases (133). Thus there would be a density­ dependent feedback that holds the population within certain limits. A sec­ ond school of thought has maintained that "density-independent factors" (e.g. weather conditions or disturbance) are more important in determining population size. Whether populations are regulated by density-dependent or density­ independent factors has been the subject ofintensive debate and controversy over the past twenty-five years, especially in the zoological literature ( l0,4 1 , 146). In part, the basis for this controversy may lie in different spatial and temporal scales of reference, as well as in differing expectations about the presence of equilibrium vs nonequilibrium states in natural populations (37). Yet much of the controversy stems from a lack of adequate informa­ tion about density-dependent regulation, particularly with regard to its frequency of occurrence, intensity, spatial pattern, and interactions with density-independent factors. Technical problems in censusing populations, particularly those of animals, and in studying the operation of the phenome­ non itself contribute to the paucity of information. Paradoxically, density­ dependent factors may operate during seasons when population numbers are low, as when the regulating force is the number of refuges or "safe-sites" available to tide the population over the harsh season. The operation of density-dependent factors may be sporadic, will interact with density­ independent factors, or (in populations undergoing unpredictable fluctua­ tions in time and space) may only be one component necessary for the continuing long-term persistence of those populations (37). Indeed, such fluctuating populations have been cited as paradigms of density-independ­ ent regulation (e.g. 9). The tractability of plants for population studies contrasts markedly with that of animals. As a consequence, future understanding of the nature of density-dependent regulation, and of its mode of operation with regard to predictability, spatial and temporal scale of action, severity, and biological basis, is likely to come from studies on plants. A clearer understanding of the ecological, genetic, and evolutionary consequences of different kinds of density dependence is also likely to be forthcoming. This review describes the manifold causes and consequences of density­ dependent regulation, formulates new conceptual frameworks for their study, and offers a predictive synthesis of ecological genetics as related to plant density. This synthesis will, we hope, stimulate further research on the process of density regulation. We review density-dependent processes in natural populations; studies on experimental and agricultural plant popula­ tions have recently been reviewed extensively by Harper (76). We focus on natural plant populations, but many of the concepts developed here are equally relevant to animal populations, though perhaps harder to apply in practice.

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