The definition of biological fitness of an individual

Gustavo Gollo

Abstract

A precise definition of biological fitness of an individual is presented.

Keywords

Fitness, Fitness Definition.

Potential fitness 

The word “fitness” is often losely defined, meaning sometimes “potential fitness”, and other
times, “real fitness”. I will clarify the differences between both. 

When comparing two individuals from the same species, if we notice one of them is robust
and healthy, while the other is skinny and sick, we are tempted to say that the robust
individual is fitter than the other. Such a conclusion derives naturally from our expectation
that a healthy and robust individual survives longer, and also reproduces easier than
other. 

Such belief produces the expectation that the most robust individual generates more
offsprings than others. Moreover, as its descendants inherit the same potentials that made
it robust, we also expect a tendency of its offsprings to generate more descendants than
the skinny one. Thus, if we compare such individuals, we expect that, after some
generations, the robust individual has produced more offsprings than the other. When we
have such an expectation we can say the individual is potentially fitter than the other. 

However, many events may frustrate such expectation. Every living being is subject to a
large number of uncertainties that may drastically change its life. An animal, for example,
may eventually be victimized by an avalanche, a lightning, or many other natural factors
that would annihilate it. 

One of many difficulties in establishing the individual potential fitness can be clarified by
the following example: be an island inhabited by two hypothetical mammal populations,
one of them predating the other. The expected dynamics derived from such a simple
situation is that both populations strongly oscillate out of phase.

An increase or the prey number consists in a raise of food to predators, what induces its
population raise. A large number of predators than tend to eat a lot, reducing drastically the
prey number, and, consequently its own food. Food scarcity then reduces the predator
population what permits prey population to grow, returning to its initial number and closing
the cycle. This well known predator-prey dynamics provides the result of the two out-of-
phase sinusoidal-like population curves.  

Let us imagine that on such a hypothetical island inhabited only by these two hypothetical
species, a new mutation arises turning a predator faster than others, with no additional
cost. We can assume that such individual, faster than other, becomes an expert hunter,
having, thus, a propensity to feed more than others and also to provide a larger amount of
food to its progeny. Thus, we expect such an individual to be fittest than others, and
probably more prolific than the population mean. We can also expect its descendants to
inherit its fast speed, such that we can predict, along generations, the increase of fast
hunters until all predators inherit the advantageous character. The former argument seems
to clearly establish the advantage of mutant character acquired by species and attest
mutation fitness raise. 

Nevertheless, predators speed increment would induce an excessively strong predating

pressure leading to a reduction of the minimum number achieved by the prey population.
Such a reduction would cause food scarcity thus reducing the predator population. At
extreme, predators raise of speed could lead them to extinction.

Real fitness 

a) Asexual populations 

For exclusively asexual populations, the real, or effective fitness of an individual can be
described by the function φ(t) that equals the number of its adult living descendants at
each time. (The individual, itself, must be included at computation if it is alive at the time
considered).

Thus, for exclusively asexual species, the real fitness of an individual is: 

φ(t)=i. 

Where i corresponds to the (integer) number of its living fertile descendants at instant t
(plus 1, if itself is still alive). 

b) sexual populations 
More sofisticated than that, is the evaluation of real fitness of sexual reproducing
individuals. Before presenting it, however, some considerings must be done. 

In order to an asexually reproducing population keeps its population number constant,

each individual must let, in average, one descendant when itself dies. If each individual at
such population let one descendant and then die, its population number remains constant
over time. The same occurs if, for each individual that lets two descendants before dying,
there is another one that lets no descendant, what matter is the mean descendant
number. 

On the other hand, the same doesn't occur for sexually reproducing populations. 

Let's imagine a group of eight individuals from an exclusively sexually reproducing

species; if each individual lets a descendant and dies, the following generation will be
composed by only four individuals. This happens because, in a sense, we can say that, at
a sexually reproduction population, the generated individual is only half descendant of
each of its parents. Thus, for a sexually reproducing population to remain in equilibrium
(keeping its population number constant), each individual have to generate two
descendants before dying. In this case, each couple generates a pair of individuals, in
such a way that the couple number remains constant along generations (what suggests
that the couple, and not the individual, should be considered the replicative unit).

The above considerations suggest the following measurement for the real fitness of
sexually reproducing individuals: the real fitness “φ” of an individual “I”, at the time of
generation g, is the number of its living offsprings, divided by 2 g, where “g” is the number
of generations separating the individual and its offspring: 

This simplified calculation is only applicable to species subjected to seasonal generation or

other processes that successively result at simultaneous death to all the species adults.

For general populations, in which different generations individuals coexist, the fitness of an
individual at an instant t is expressed by:

where d is a descent line connecting the individual to its living offsprings; m is the number
of descent lines; and g(d) is the length of descent line, i.e. the number of reproductions
encompassed by the line.

This calculation is similar to that of inclusive fitness as defined by Hamilton (1964). The
above proposal can be clarified by the following example.

The figure below shows an individual and its descendants. Arrows indicate offspring paths.
All individuals were generated by sexual reproduction. Ancestors that were not
descendants of the original individual were all omitted in such a way that, when only one
arrow points the individual, only one of its parents is presented in the figure. Generations
are not cleared defined, as can be seen for I 2,3 individual, which belongs simultaneously to
generations 2 and 3. For simplifying purposes, only, let us consider that all individuals are
born, and die, at the time stipulated by the generation at which it occurs (in spite of
individual I3,1 from generation 1 generates individual I2,3 from generation 3).

0 I0,0

1 I1,1 I2,1 I3,1

2 I1,2 I2,2 I3,2

 
time
3 I1,3 I2,3 I3,3

The original individual, I0,0 , generated 3 individuals that lived at generation 1. Each of
these individuals derives from the original one through only one descent line. Thus, if I 0,0 is
dead at generation 1, its fitness at this time is:

This means that, at the first generation (t=1), individual I 0,0 has 1.5φ, where φ, the fitness
unit, is read fit. The value arises from the sum of 3 individuals, I 0,0 descendents at
generation 1, each one contributing with a ½ φ. If individual I0,0 was still alive at this time,
the value 1 φ should be added to total (corresponding to its own contribution to it). 

At the second generation, t=2, we can see that the individual I 1,2 is double descendant from
I0,0 ,by I1,1 ,and I2,1. This I0,0 double grandson corresponds, for this reason, to two grandsons,
the I1,1 son, and the I2,1 son. Therefore, there are two descent lines from I 0,0 to I1,2, thus, the
I1,2 contribution for I0,0 fitness calculation is twice the contribution from an usual descendant
from its generation, i.e.

The contribution from I2,2 is the same from I3,2, and corresponds to half this value

. Thus, I0,0(2) real fitness, the fitness of I0,0 at generation two, is their sum:

This calculation presupposes that individuals from former generations exist no more,
otherwise, the contribution from each one should be added to it.

At the third generation, we notice that the individual I 1,3 descends from the double
descendant from I0,0 , I1,2 ; its contribution to I0,0 real fitness corresponds to half the
contribution from its parent I1,2 , it is to say: (¼)φ.

The individual I2,3 presents two intricacies. It corresponds to double descent line of I 0,0 and,
also, belongs to twice distinct generations. Its contribution to the fitness of I 0,0 must be
done summing the contributions from each of these line descents. These contributions
correspond, by its turn, to half the value of each parent's contribution to I 0,0 fitness. Thus,
the contribution of I2,3 to I0,0 fitness from I2,2 is (1/8)φ, whereas the contribution from I 3,1 is
(¼)φ. The total contribution from I0,0 thus adds (3/8)φ to I0,0 fitness at the time of the third
generation.

The contribution of I3,3 to I0,0 fitness is simply calculated: (½3)φ = (1/8)φ.

Hence, the fitness of I0,0 at the time of the third generations is due to all its living
descendants' contribution sum, and corresponds to:
 .

The value of the real fitness of an individual tend to oscillate for a short initial period,
stabilizing after some time, if the population remains stable. 

Relative fitness

Such population equilibrium requirement, however, is not always satisfied. While the value
proposed above describes an absolute evolution of the lineage, it may sometimes be
interesting to evaluate the individual fitness compared to the fitness of others of the same
species or group. This is specially desirable when analising ciclic growth populations, and
can be done introducing the following normalizing factor:

where φrel.(I,t) (relative fitness of individual I at instant t) is the individual real fitness of I
compared to other individual of the same species at time t; φabs.(I,t) (absolute fitness of
individual I) the absolute real fitness of I; P(to) the compared group initial population
(measured at the individual born instant); and P(t) population at an instant t.

For both sexual and asexual species, constant φ rel.(I) means that individuals character are
neutral. Growing φrel.(I) indicates I characters are spreading over species, while reducing
φrel.(I) suggests further extinction of I lineages.

These informations are specially relevant to evaluate new mutations and exotic invasions.

For both sexual and asexual reproduction, a constant value for absolute fitness through
time [φabs(I,t)] shows that the genetic contribution of the focused individual remains
constant along the same period. Reducing fitness values mean that individual genetic
contribution is reducing along time; growing values mean its expansion along time. [φ rel.
(I,t)] shows the same, but analising genetic contribution growth relative to species
population.

A final note

The example above, for didactic reasons, is simple but imprecise. Although correct as put
above, the proposed calculation doesn't derive from reproducing mode, sexual or not, but
from the genetic material parcel that is transmitted from a parent to each of its
descendants. In fact, half the genetic material of each parent is usually transmitted to
descendants through sexual reproduction.

Bees, however, reproduce at an unusual manner; females are diploid, while males are
haploid. Unfertilized eggs provide males, fertilized eggs develop females. Such way,
whenever a male reproduces, it does it sexually. But its fitness calculation must be done
as if it reproduced at assexual manner, as it brings to offsprings its total genetic content.
Females, on the other hand, always bring half its genetic material to offspring, be it by
sexual reproducing, or not. Thus, the calculation of its genetic contribution to each
descendant must reflect this fact, always being divided by 2, independent of being sexual,
or not. The same holds for its fitness calculation, as descendants contribution to it must be
divided by 2 at each generation. 

Real fitness calculations for diploid species that reproduces both sexually or not must take
into account the reproducing way of generating each individual.

Extra-biology

Mainly for artificial life purpose, we can define an n sexual reproduction, it is to say, n
parents generating an individual.

In this context, reproduction, is the production of an individual similar to itself by another.

Sexual reproduction, is the production of an individual that inherit similarities from more
than one parent (what differ to traditional biological definition that asserts sexual
reproduction is the one that includes a meiotic step).

For the n-progenitor sexual reproduction, the contribution from each parent is the parcel
reproduced from it. As a final result, the sum of contributions, can differ from parent to
parent.

Generalizing what was proposed to haplodiploid species, we can define fitness of an

individual by the sum of each “actual living contribution” due to the individual, which is the
sum of the parcel of itself in each living individual at an instant.

On the other hand, the participation of an individual at a reproduction without its own
contribution to reproductive traits must be seen only as a parasitic relation. In this case,
the individual that contributes to the reproduction of another, reproducing none of its own
characters is its host, not its parent.

Reference:

Hamilton WD. The genetical evolution of social behaviour. I. J Theor Biol. 1964 Jul;7(1):1-
16. doi: 10.1016/0022-5193(64)90038-4. PMID: 5875341.