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Lotka–Volterra equations

math Maturity 9-11 Next article →

The Lotka–Volterra equations are a pair of first-order nonlinear differential equations. They are used to describe the dynamics of biological systems where two species interact. In these systems, one species acts as the predator and the other acts as the prey. These equations help scientists understand how population densities change over time. They show that the populations of predators and prey are not static. Instead, they follow a continuous and deterministic path where generations constantly overlap.

Lotka Volterra dynamics
Lotka Volterra dynamics

To understand the mechanism, we must look at how the variables interact. The model uses the variable $x$ for prey population density and $y$ for predator population density. The rate of change for the prey population depends on two main factors. First, the prey grows at a maximum per capita rate. Second, the prey population decreases based on the rate of predation. The rate of predation is proportional to how often predators and prey meet. For the predators, the rate of change depends on their consumption of prey. However, this is balanced against their own intrinsic death rate, which causes exponential decay if no prey is available.

Lotka-Volterra model (1.1, 0.4, 0.4, 0.1)
Lotka-Volterra model (1.1, 0.4, 0.4, 0.1)

This model relies on several specific biological assumptions. It assumes the prey has an unlimited food supply and reproduces exponentially. It also assumes the predator's food supply depends entirely on the size of the prey population. Furthermore, the model assumes that predators have a limitless appetite. It treats populations as single variables, meaning it ignores spatial distribution or different age groups. While these assumptions are rarely perfectly true in nature, the model reveals fundamental truths about ecological oscillations. These oscillations are the rising and falling cycles of population numbers seen in the wild.

Lotka Volterra dynamics
Lotka Volterra dynamics

The history of these equations involves two different researchers working independently. Alfred J. Lotka first proposed these ideas in 1910 regarding chemical reactions. He later extended the model to organic systems in 1920 and published work on biomathematics in 1925. Around the same time, the mathematician and physicist Vito Volterra developed the same equations. Volterra was inspired by the marine biologist Umberto D'Ancona. D'Ancona observed a strange trend in the Adriatic Sea during World War I, from 1914 to 1918. Even though fishing effort decreased during the war, the percentage of predatory fish caught actually increased. Volterra created his model to explain this specific observation.

Milliers fourrures vendues en environ 90 ans odum 1953 en
Milliers fourrures vendues en environ 90 ans odum 1953 en

One significant finding of the model is the nature of population equilibrium. The equilibrium density of the prey depends on the parameters of the predator. Conversely, the predator's equilibrium density depends on the prey's parameters. This leads to a surprising result: making the environment better for the prey often benefits the predator more than the prey. For example, adding iron to the ocean can cause a bloom of phytoplankton. While this increases the prey, it often simply leads to a higher density of predators, which limits the overall effect. This phenomenon is sometimes called the paradox of enrichment.

Lotka Volterra dynamics
Lotka Volterra dynamics

Mathematical analysis shows that these equations have periodic solutions. These solutions can be visualized in a phase-space plot. In a phase-space plot, one axis represents the number of prey and the other represents the predators. The resulting paths are closed curves. However, the model has a limitation known as the "atto-fox problem." In some mathematical cycles, the prey population drops to an extremely low number. In a real ecosystem, such a small number might lead to accidental extinction due to chance fluctuations.

Beyond biology, these equations apply to many other fields. In economics and marketing, the model describes markets with multiple competitors. It can show how one firm might drive others out or how firms reach a stable market share. In 1965, Richard Goodwin applied these ideas to economics. He used the predator-prey interaction to reinterpret the relationship between unemployment and wage changes. This connection shows that the mathematical patterns found in nature can also describe the complex movements of human systems.

Lotka Volterra dynamics
Lotka Volterra dynamics
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File:Milliers fourrures vendues en environ 90 ans odum 1953 en.jpg
Milliers fourrures vendues en environ 90...
File:Lotka Volterra dynamics.svg
Lotka Volterra dynamics.svg
File:Lotka-Volterra model (1.1, 0.4, 0.4, 0.1).svg
Lotka-Volterra model (1.1, 0.4, 0.4, 0.1).svg
File:Predator prey dynamics.svg
Predator prey dynamics.svg
File:Lotka-Volterra orbits 01.svg
Lotka-Volterra orbits 01.svg
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