

Part 2: The LotkaVolterra Model*
Vito Volterra (18601940) was a famous Italian mathematician who retired from a distinguished career in pure mathematics in the early 1920s. His soninlaw, Humberto D'Ancona, was a biologist who studied the populations of various species of fish in the Adriatic Sea. In 1926 D'Ancona completed a statistical study of the numbers of each species sold on the fish markets of three ports: Fiume, Trieste, and Venice. The percentages of predator species (sharks, skates, rays, etc.) in the Fiume catch are shown in the following table:
1914  1915  1916  1917  1918  1919  1920  1921  1922  1923 
12  21  22  21  36  27  16  16  15  11 
As we did with Canadian furs, we may assume that proportions within the "harvested" population reflect those in the total population. D'Ancona observed that the highest percentages of predators occurred during and just after World War I (as we now call it), when fishing was drastically curtailed. He concluded that the predatorprey balance was at its natural state during the war, and that intense fishing before and after the war disturbed this natural balance  to the detriment of predators. Having no biological or ecological explanation for this phenomenon, D'Ancona asked Volterra if he could come up with a mathematical model that might explain what was going on. In a matter of months, Volterra developed a series of models for interactions of two or more species. The first and simplest of these models is the subject of this module.
Alfred J. Lotka (18801949) was an American mathematical biologist (and later actuary) who formulated many of the same models as Volterra, independently and at about the same time. His primary example of a predatorprey system comprised a plant population and an herbivorous animal dependent on that plant for food.
We repeat our (admittedly simplistic) assumptions from Part 1:
If there were no predators, the second assumption would imply that the prey species grows exponentially, i.e., if x = x(t) is the size of the prey population at time t, then we would have
dx/dt = ax.
But there are predators, which must account for a negative component in the prey growth rate. Suppose we write y = y(t) for the size of the predator population at time t. Here are the crucial assumptions for completing the model:
These assumptions lead to the conclusion that the negative component of the prey growth rate is proportional to the product xy of the population sizes, i.e.,
dx/dt = ax  bxy.
Now we consider the predator population. If there were no food supply, the population would die out at a rate proportional to its size, i.e. we would find
dy/dt = cy.
(Keep in mind that the "natural growth rate" is a composite of birth and death rates, both presumably proportional to population size. In the absence of food, there is no energy supply to support the birth rate.) But there is a food supply: the prey. And what's bad for hares is good for lynx. That is, the energy to support growth of the predator population is proportional to deaths of prey, so
dy/dt = cy + pxy.
This discussion leads to the LotkaVolterra PredatorPrey Model:
dx/dt = ax  bxy,
dy/dt = cy + pxy,
where a, b, c, and p are positive constants.
The LotkaVolterra model consists of a system of linked differential equations that cannot be separated from each other and that cannot be solved in closed form. Nevertheless, there are a few things we can learn from their symbolic form.
Here is a link for a biological perspective on the LotkaVolterra model that includes discussion of the four quadrants and the lag of predators behind prey.
Additional links are provided in Part 6 for various extensions of the model.
* This material is based in part on section 6.4 of Differential Equations: A Modeling Perspective, by R. L. Borrelli and C. S. Coleman, Wiley, 1996.


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