Altruism

Altruism is selfish.

It is to your advantage to be advantageous to others. It is never advantageous to debase someone who is not malicious.

"The work of Kruuk (1964) has been reinforced by further
studies, summarized by Lack (1968). The latter seem to have shown that all
marginal nests failed to rear young, mainly due to predation. Perhaps,
nevertheless, the gulls that could not get places in the centre of the colony
were right in nesting on the edge rather than in isolation where, for a conspicuous
bird like a gull, the chances would have been even worse. "
Hamilton, William D. "Geometry for the selfish herd." Journal of theoretical Biology 31.2 (1971): 295-311.

## Example: Evolution of altruism

To study the evolution of a genetic predisposition to altruism, altruism will be defined as the genetic predisposition to behavior which decreases individual fitness while increasing the average fitness of the group to which the individual belongs. First specifying a simple model, which will only require the simple Price equation. Specify a fitness *w**i* by a model equation:

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{\displaystyle w\_i = \frac{n'\_i}{n\_i} = k - a z\_i + b z}

where *z**i* is a measure of altruism, the *az**i* term is the decrease in fitness of an individual due to altruism towards the group and *bz* is the increase in fitness of an individual due to the altruism of the group towards an individual.
Assume that *a* and *b* are both greater than zero. From the Price equation:

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{\displaystyle w\Delta z = -a\operatorname{var}\left(z\_i\right)}

where var(*z**i*) is the variance of *z**i* which is just the covariance of *z**i* with itself:

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{\displaystyle \operatorname{var}(z\_i) \;\stackrel{\mathrm{def}}{=}\; \operatorname{E}(z\_i^2) - \operatorname{E}(z\_i)^2}

It can be seen that, by this model, in order for altruism to persist it must be uniform throughout the group. If there are two altruist types the average altruism of the group will decrease, the more altruistic will lose out to the less altruistic.

Now assuming a hierarchy of groups which will require the full Price equation. The population will be divided into groups, labelled with index *i* and then each group will have a set of subgroups labelled by index *j*. Individuals will thus be identified by two indices,
*i* and *j*, specifying which group and subgroup they belong to. *n**ij* will specify the number of individuals of type *ij*. Let *z**ij* be the degree of altruism expressed by individual *j* of group *i* towards the members of group *i*. Let's specify the fitness *w**ij* by a model equation:

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{\displaystyle w\_{ij} = \frac{n'\_{ij}}{n\_{ij}} = k - a z\_{ij} + b z\_i}

The *a z**ij* term is the fitness the organism loses by being altruistic and is
proportional to the degree of altruism *z**ij* that it expresses towards members
of its own group. The *b z**i* term is the fitness that the organism gains from the altruism of the members of its group, and is proportional to the average altruism *z**i* expressed by the group towards its members. Again, in studying altruistic (rather than spiteful) behavior, it is expected that *a* and *b* are positive numbers. Note that the above behavior is altruistic only when *az**ij* >*bz**i*. Defining the group averages:

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{\displaystyle \begin{align}
 n\_i &= \sum\_j n\_{ij} \\
 z\_i &= \frac{1}{n\_i}\sum\_j z\_{ij}n\_{ij} \\
 w\_i &= \frac{1}{n\_i}\sum\_j w\_{ij}n\_{ij} = k + (b - a)z\_i \\
 n\_i'&= \sum\_j n\_{ij}' = n\_i[k + (b - a)z\_i] \\
 z\_i'&= \frac{1}{n\_i'}\sum\_j z\_{ij}n\_{ij}'
\end{align}}

and global averages:

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{\displaystyle \begin{align}
 n &= \sum\_{ij} n\_{ij} = \sum\_i n\_i \\
 z &= \frac{1}{n}\sum\_{ij} z\_{ij}n\_{ij} = \frac{1}{n}\sum\_i z\_in\_i \\
 w &= \frac{1}{n}\sum\_{ij} w\_{ij}n\_{ij} = \frac{1}{n}\sum\_i w\_in\_i \\
 n'&= \sum\_{ij} n\_{ij}' = \sum\_i n\_i' \\
 z'&= \frac{1}{n'}\sum\_{ij} z\_{ij}n\_{ij}' = \frac{1}{n'}\sum\_i z\_i'n\_i'
\end{align}}

It can be seen that since the *z**i* and *z**i* are now averages over a particular group, and since these groups are subject to selection, the value of Δ*z**i* = *z*′*i*−*z**i* will not necessarily be zero, and the full Price equation will be needed.

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{\displaystyle \Delta z = \operatorname{cov}\left(\frac{w\_i}{w}, z\_i\right) + \operatorname{E}\left(w\_i\,\Delta \frac{z\_i}{w}\right)}

In this case, the first term isolates the advantage to each group conferred by having altruistic members. The second term isolates the loss of altruistic members from their group due to their altruistic behavior. The second term will be negative. In other words, there will be an average loss of altruism due to the in-group loss of altruists, assuming that the altruism is not uniform across the group. The first term is:

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{\displaystyle \operatorname{cov}\left(\frac{w\_i}{w}, z\_i\right) = \left(b - a\right)\operatorname{var}(z\_i)}

In other words, for *b*>*a* there may be a positive contribution to the average altruism as a result of a group growing due to its high number of altruists and this growth can offset in-group losses, especially if the variance of the in-group altruism is low. In order for this effect to be significant, there must be a spread in the average altruism of the groups.