Stimulated by Malthus' "Essay on the Principle of Population", Verhulst (1838) published what he called a "logistique" equation to describe the sigmoidal growth of population density to carrying capacity. The equation was rediscovered by Pearl and Reed (1920). Then, Lotka (1925) derived the same equation mathematically, calling it "the law of population growth", and Gause (1934) demonstrated its validity in laboratory experiments. For an excellent review of these historical events see Kingsland (1985). The discrete form of the logistic equation used in this book was first proposed by Cook (1965) but is, in fact identical to the Ricker (1954) equation (see e.g., Berryman 1978).
Royama (1992) derived the logistic equation by considering a system in which randomly distributed organisms draw resources from a circular area of influence of radius r around themselves, and compete with neighboring organisms when their areas of influence overlap; i.e., when the distance to the neighbor is less than 2r (see figure).
Let Gi be the finite per-capita rate of change of an individual organism competing with i neighbors and Pr(i) be the expected proportion of the population having i overlapping competitors, then the mean finite rate of increase of the population is given by the weighted sum
If the individuals in the consumer population are distributed randomly, then they can be described by a Poisson distribution with mean P, the density of the population (really Pt-1 but we omit the time subscript for convenience). Under these conditions the number of competitors is also Poisson distributed with mean sP, where s = 4πr2. From this the expected proportions are given by the Poisson formula
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which, can be substituted into the previous equation to yield
Now suppose that the addition of a competitor reduces the average rate of change of an individual by the proportion k (0<k<1), so that the finite rate of change in competition with i neighbors is Gi = G0ki, and substituting ki for Gi/G0 in the previous equation, we get
This equation can then be reduced by Taylor's theorem to
which is the well-known Ricker (1954) equation familiar to fisheries ecologists. Continuing after Berryman et al. (1995), we let RP = lnG and AP = lnG0, so that
Noting that the parameter b can be expanded to b = V/H, where V is a measure of the intensity of competition, and H is the density of resources, we obtain the logistic R-function
Notice that V is the same as the demand for prey by a predator, the greater the demand, therefore, the greater the degree of intraspecific competition.
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©1997 Alan A. Berryman