The influence of four types of heterogeneity (heterogeneity in spatial arrangement, initial size heterogeneity, environmental heterogeneity in space and in time) on the development of a plant population was studied using a simple model of plant growth and competition. Parameters were estimated for Scots pine (Pinus sylvestris). The model showed that without heterogeneity the population became very unstable, so that even a small decrease in environmental favourability caused population collapse, because there was not any differentiation and competition was extremely severe. The presence of any type of heterogeneity created a basis for social differentiation of individuals. As a consequence, self-thinning occured and lowered the competition. The population became smaller, but more tolerant against decreases in environmental favourability. However, a glut of heterogeneity is rather deleterious, because it further diminished the population while adding nothing to its tolerance.
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A. B. Yastrebov (1996) studied this question.
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