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Population dynamics is the type of mathematics used to model and study the size and age composition of populations as dynamical systems. Population dynamics has traditionally been the dominant branch of mathematical biology, which has a history of more than 220 years, although over the last century the scope of mathematical biology has greatly expanded. The beginning of population dynamics is widely regarded as the work of Malthus, formulated as the Malthusian growth model. According to Malthus, assuming that the conditions (the environment) remain constant (ceteris paribus), a population will grow (or decline) exponentially. This principle provided the basis for the subsequent predictive theories, such as the demographic studies such as the work of Benjamin Gompertz and Pierre François Verhulst in the early 19th century, who refined and adjusted the Malthusian demographic model. A more general model formulation was proposed by F. J. Richards in 1959, further expanded by Simon Hopkins, in which the models of Gompertz, Verhulst and also Ludwig von Bertalanffy are covered as special cases of the general formulation. The Lotka–Volterra predator-prey equations are another famous example, as well as the alternative Arditi–Ginzburg equations. Simplified population models usually start with four key variables (four demographic processes) including death, birth, immigration, and emigration. Mathematical models used to calculate changes in population demographics and evolution hold the assumption ('null hypothesis') of no external influence. Models can be more mathematically complex where "...several competing hypotheses are simultaneously confronted with the data." For example, in a closed system where immigration and emigration does not take place, the rate of change in the number of individuals in a population can be described as: where N is the total number of individuals in the specific experimental population being studied, B is the number of births and D is the number of deaths per individual in a particular experiment or model.
Stewart Cole, Charlotte Avanzi, Philippe Busso