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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