Modes
of Natural Selection
For a
trait influenced by multiple gene loci with approximately
equal, additive effect, the trait
distribution will approximate a normal
distribution (bell curve). Such a trait is
measured over an arbitrary scale of 1-14 units (top
row). The probability
of survival follows a fitness
function that varies over the range of
phenotypes [middle row]. As a result, the
trait distribution after selection is
modified in a (statistically) predictable manner
[bottom row]. The change in trait distribution can be
described in terms of the effect on mean and variance.
Under Directional
Selection (left), the trait mean is shifted
(here, it increases), and the variance is skewed
(unbalanced towards the left or right tail (as shown
here)) over the course of a single generation. Variation in
the offspring generation will be normal around the new mean,
with the same variance. Under Stabilizing
Selection (middle), both tails are truncated.
The trait mean remains constant, but the trait
variance is reduced. Under Disruptive
Selection, both tails are favored at
the expense of the modal phenotypes. The trait mean
remains constant, but the trait variance is increased.
[The technical descriptions of the distributions after stabilizing
and disruptive selection are leptokurtotic
("pointed") and platykurtotic ("flattened"),
respectively'].
The fitness function for directional
selection is shown as a straight line between 0.0 and
1.0: this is unrealistic. Except for medical conditions
involving lethality, fitness differences across trait
distributions tend to be small, < or << 0.01, which
means they are difficult to measure, and will affect evolution
very slowly. As well, the criterion for directional selection
is a fitness function with a positive slope, but no
necessarily a constant positive slope as in the
example. The fitness function for stabilizing selection is
somewhat more realistic, if trait variation around the mean
consistent with viability is small (consider body weight in
newborns). Note that the fitness function is initially
positive, flattens at the optimum, and is negative thereafter.
The criterion for disruptive selection is the reverse of this,
negative, then flat at the minimum, then positive. Disruptive
selection may facilitate local adaptation, if the optimal
trait value varies over the range of a species. In principle,
disruptive selection can accompany speciation, for
example if the trait extremes are for aspects of mating
behavior (Darwin recognized "Sexual Selection").
Selection in each case ensures that the
genetic composition of the next generation is modified.
This is most easily appreciated in the case of directional
selection, where allele(s) at each locus that increase the
trait measure are favored by selection: the frequency of those
alleles will be increased in next generation.