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Samuel Hank
RSE Paper 2
10/26/2017
Mathematical Models Describing the Inheritance of Traits via Natural Selection:

When a scientific theory is first proposed, it is almost always the case that the theory

comes first and then a mathematical model is later created that is found to be in congruence with

both the theory and observed behavior of the subject of the theory. Such is the case with Darwin

and the theory of evolution by natural selection. When Darwin first wrote The Origin of Species,

he made no attempt at mathematical formulation; rather, he proposed the theory itself. Centuries

later biologists proposed mathematical models that could begin to approximate aspects of

Darwin’s theory. Those mathematical models all had the goal of creating a strong model for

evolution; that is, they sought to be able to predict the paths natural selection would take, and

especially the traits it would favor. These mathematical formulas strengthen Darwin’s model. A

scientific study can only be used to demonstrate the evolution of small micro-evolved features

within the span of a few centuries at most. Mathematical models strengthen the theory of

evolution by explaining why certain features evolved as they did. In addition, it provides new

areas for exploration and explanation of organismal behaviors.

The first mathematical models of the mechanisms of natural selection appeared in the

twentieth century. The pioneer of this field was W.D. Hamilton in the 1960s. Although

rudimentary mathematical models of genetic inheritance had been created years earlier, few had

attempted to model the processes that select for or against a trait based on its ability to aid or

harm an organism and its genetic progeny. Hamilton pioneered the study of a specific trait’s

ability to aid survival when he created the concept of inclusive fitness. Previously, most studies

of natural selection had focused on the obvious necessity of personal fitness. Personal fitness is

how likely an organism’s genes are to be passed on based on the number of progeny the
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individual produces. This is the obvious mechanism of natural selection because it is the simplest

to understand - if an organism has traits that benefits its survival it will live long enough to pass

on those traits to its children. However, in human society as well as in the natural world there are

a myriad of traits and behaviors that organisms possess which directly harm their ability to

survive. So, in contrast to the concept of personal fitness, inclusive fitness refers to the number

of “offspring equivalents” an individual affects.

Like all mathematical formulations attempting to describe natural processes, Hamilton

first had to base his equations on a set of assumptions. The main assumption Hamilton made was

that animals must have some ability to determine who their kin were, either intentionally or

unintentionally, and they must be able to know who is most closely related to them. He proposed

two mechanisms by which this could occur. The first mechanism is straightforward in that the

organism has some way to intentionally distinguish kin from non-kin, such as in humans

whereby one can distinguish one’s family by the physical features they possess. The other

mechanism is in sedentary species that do not migrate. Often one’s kin are most likely those in

close proximity to oneself (Hamilton). Using these assumptions, Hamilton created a general

equation describing inclusive fitness and likelihood of specific acts an organism might undertake

to the benefit of another organism in its kin group. This equation is called Hamilton’s rule:

Equation 1:

In this equation, c represents the relative cost to the individual committing the act or the actor, r

represents the genetic relatedness of the organisms (one’s own offspring would have a value of

one), and b is the relative benefit gained by the recipient of the act. An important part of the
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equation to note is that it does not describe the cost to the actor’s ability to survive or even the

cost to their kin’s ability to survive, rather it describes merely how much an action aids the

organism’s genes chances of survival. Because the equation measures strictly the benefit gained

by the actor’s own genes in their fight to be passed on (de Catanzaro). the benefit to the recipient

must be modified by the relatedness of actor and recipient. Thus, the equation shows that in order

for an action to be favored by natural selection, the cost of the action to the actor’s genes

(meaning the fecundity of their genes) must be less than the benefit gained by those genes as

they are present to varying degrees within the recipients of the action.

How does one arrive at values for c and b? This question illustrates both the elegance as

well as the shortcomings of this equation. In simple examples where the cost of an action is

immediate and its effects obvious, such as in a case where one organism sacrifices itself to save

its children, the cost is equal to the potential of that organism to reproduce while the benefit is

the potential of that organism’s offspring to reproduce. In a situation where both the cost and

benefit are much less than life and death for an organism, however, the arithmetic becomes much

more complicated as the values of b and c become much harder to define. It becomes clear then

that Hamilton’s rule provides a framework for judging an action and the underlying genes

causing it as likely to be favored via natural selection or not, despite the fact that exact arithmetic

and calculation is likely to be difficult. Thus, Hamilton’s rule was the first step toward creating a

strong model describing traits as either favorable or not.

Hamilton’s rule, however, was only the beginning. In the 1980s de Catanzaro expanded

the Hamilton's mathematical models of inclusive fitness. The equation he created was termed

“the residual capacity of an organism to promote inclusive fitness.” Rather than focusing on

individual traits an organism possessed, de Catanzaro sought to create a model that could
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describe the continued impact an organism’s existence would have on the likelihood of its own

genes being passed on. The is Equation 2. It incorporated and expanded upon aspects of

Hamilton’s rule.

Equation 2:

In this equation, p sub i represent the expected reproduction of the individual in question past the

age that is being considered; delta p sub k is the increment or decrement in expected

reproduction of each kinship member; and r sub k is the same r from Hamilton’s rule

representing the degree of relatedness from the organism in question to the kin member being

considered (de Catanzaro). As in the original equation this second equation examines the impact

of an organism’s continued existence on the chance of its genes being passed on. It does so by

adding the chance of the organism reproducing to the effect of that organism’s existence on those

who share genes with the organism in question. An important note regarding delta p sub k is that

it can be either positive or negative. Thus, this model provides a more comprehensive view of

inclusive fitness. Overall de Catanzaro’s equation for inclusive fitness expands upon the

foundation laid by Hamilton to produce an equation that, if effective, could be used to model the

ideal age of death for an organism in terms of increasing its genes’ fitness.

These equations can be used then to examine behaviors that personal fitness fails to

account for in organism, such as suicide and other self-sacrificial behaviors. In the study of

suicide in humans it is useful to examine archaic societies in order to attempt a study of human

behavior unmodified by institutions and compounded centuries of culture. Thus, for this example

we will examine patterns of suicide among the Eskimo. In a study on Eskimo suicide, Alexander

and Hughes found that the elderly in Eskimo society often committed suicide once they could no
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longer sustain themselves via hunting and became an overall burden to their tribe (Leighton and

Hughes). We can examine this behavior pattern using de Catanzaro’s equation. If we take an

elderly Eskimo man of roughly age 60 and assume that his chances of reproducing are small

although still present, thus we could assign a value of .1 for p sub i (representing a 10% chance

that he could father another child). For the second half of the equation, we first need to

determine how many kin members he effects by his existence. For simplicity we shall assume

one child and two grandchildren, each having r values of 1 and .5, respectively. Last is the most

difficult part and that is assigning a value for delta p sub k. It is difficult to determine the

negative or positive effect a being’s continued existence will have on its kin. However, that does

not mean we cannot get a close approximation. In this case, we would weigh all relevant factors.

For instance, if the man continued to live, we could assume that he could have a small positive

impact by way of sharing his wisdom and experience; however, in a survival society such as the

Eskimo where food is scarce a man unable to gather his own food would represent a large burden

to those caring for him. Taking all this into consideration we could estimate a delta p sub k value

of -.25. Putting all these variables together and solving the equation provide an answer of -.4

units of genetic equivalency, meaning that the elderly man continuing to exist would exert a

negative pressure on his genes and their ability to propagate. Thus, in this case, it makes sense

that the Eskimo man would commit suicide. In doing so he eliminates the negative factor his

continued existence would have on his genes' ability to survive. We can assume that, over the

course of human evolution, natural selection favored the behavior of the elderly to commit

suicide when they felt worthless to their kin members.

Similarly, we can apply Hamilton’s rule to the suicide in the case of the elderly Eskimo

man. In this case, the cost to the man is that he would give up any chance of having more
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children; earlier we ascribed the value of this happening to be .1. If we take the recipient of the

act to be his child, the value of r is 1 while the benefit to that child is the absence of the negative

impact the burden of caring for the elderly man would have had on the child; thus, b is +.25.

Plugging these values into the equation gives: .1<.25 and since this is true we can say once again

that the behavior of suicide in an elderly Eskimo man will be favored by natural selection since it

increases the chances of his genes being passed on.

Through these examples it becomes clear that the mathematical models created by the

two researchers provide a lens not only for examining why certain behaviors seem to be favored

by natural selection but also a model for predicting if certain behaviors and traits will be

preserved when certain environmental conditions change. For instance, one can use these

equations again with regard to human suicide in the modern era. As shown above, it is clear that

suicide was beneficial in early human society. However, in the developed world where food is

abundant this equation and its variables must be reexamined as the impact of an elderly man’s

existence on his children and their ability to reproduce will certainly not be negative to the extent

it is in a survival-oriented society, and it may even be positive. In this way, the equations can be

used as the basis to create a strong model to predict which behaviors are conserved or discarded,

in humans, looking forward into the deep future. These models, as such, show the value of

mathematical analysis of verbalized theories even when perfect values for many variables are

nearly impossible to ascertain. The creation of the equation itself provides valuable frameworks

for examining the original theory.

Mathematical models thus strengthen the theory of evolution by explaining why certain

features evolved as they did, as well as provide new areas for exploration in and explanation of

organismal behavior. In all the social sciences and even in the natural sciences, the theory always
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comes before the model and equation and oftentimes when examining something as complex as

an organism’s behavior, the equation may be severely lacking. Nevertheless, mathematical

models provide an invaluable lens to examining theories, especially those as complex as evolved

behavior.
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Works Cited

Hamilton, W. (1964). "The genetical evolution of social behaviour. I". Journal of

Theoretical Biology. 7 (1): 1–16. PMID 5875341. doi:10.1016/0022-5193(64)90038-4.

Evolutionary limits to self- preservation, de Catanzaro, Denys, Ethology and

Sociobiology , Volume 12 , Issue 1 , 13 – 28

Leighton, Alexander H., and Charles C. Hughes. "Notes on Eskimo Patterns of Suicde."
The University of Chicago Press Journals, vol. 11, no. 4, Winter 1955, pp. 327-38.

Lester, David. Why People Kill Themselves: A 1990s Summary of Research Findings on
Suicidal Behavior. 3rd ed., Springfield, Thomas, 1992.