Biol 117 F 2016

Lecture 9/2
The imperfection of life
I.

Introduction
The beauty in evolution is not in how organisms have adaptations that are perfectly
created to allow survival in their environment, but in that they are well-suited for
survival at all. Though natural selection results in adaptation, nature is full of
examples where organisms are not ideally suited for survival in their current
environment. There are several reasons why this is so.
A. Adaptations are compromises.
The adaptive traits of most organisms are compromises produced by competing
selection pressures. To successfully exist in any environment, organisms must
have several adaptations (whether behavioral or morphological) to survive.
Often, these adaptations exist in direct conflict with each other, one providing
success in a situation, while another acting as a detriment. For example, sea
turtles must lay their eggs on beaches because their embryos cannot acquire
enough oxygen in water to fulfill developmental needs. Females are excellent
swimmers due to the presence of flippers. However, these flippers are not wellsuited for movement on land. If the sea turtle had legs, allowing it to move more
freely on land, it would not be nearly as efficient at swimming in the water.
B. The environment changes over time.
When selection occurs in a population, it preserves alleles that are successful
under the prevailing environmental conditions. That is, each generation is
adapted for survival in the environment that existed during their parents
generation. When the environment changes, adaptations lag behind. If the
environment changes with every generation, adaptations will always lag behind,
and, as a result, may never look appropriate given the current environment. In
this specific case of constant change, a species may evolve a more generalistic
trait that allows some success in any environment, which may look imperfect in
any one environmental condition.
C. Adaptations are limited by historical constraints.
Natural selection is not an engineer that designs new organisms from scratch.
Instead, it acts on new mutations and existing genetic variation. Because new
mutations are fairly rare, natural selection works primarily with alleles that have
been present for many generations. Thus, adaptive changes in the morphology of
a species are often based on small modifications of existing structures. For
example, the bipedal posture of humans evolved from the quadrupedal posture of
our ancestors. Natural selection did not produce an entirely new skeletal design to

accompany this radical behavioral shift. Instead, existing characteristics of the

spinal column, and corresponding musculature of the legs and back, were
modified, albeit imperfectly, for an upright stance. The result is a species that has
the benefits of an upright posture and increased agility, but is more likely to suffer
with chronic back and leg problems.
One may ask, therefore, how do you determine if an adaptation is truly novel (arising
from a mutation in the genetic line) or if it is only a modification of previously
existing behavioral or morphological phenomena? Generally speaking, almost all
adaptations are built as slight modifications of ancestral traits and the variation that
existed concurrently with them. Additionally, many adaptations are seen as imperfect
solutions to current environmental problems, resulting from the reasons stated in A
through C above. While examining a population through time, if you see a novel
adaptation that in no way resembles the behaviors or mechanisms of the ancestor,
then you may have a truly novel genotype and resulting phenotype.
Below are a few terms that are frequently associated with this sort of discussion.
Students are responsible for knowing these terms, and how they are used with the
discussion above.
1) Adaptation: a phenotype that is selected for within a population. Heritable trait
that increases the fitness of the individual with that trait.
2) Contrivance: an adaptation in an organism, where the adaptation exists as a result
of the modification of the original (or previous) adaptation in an ancestor.
3) Vestigial trait: a trait the entire species has that is no longer used or has no use in
the present environment, but was likely an adaptation for an ancestor.
4) Atavism: a trait possessed by only a few members of the population that is no
longer used or has no use in the present environment, but was likely an adaptation
for an ancestor. (Although technically not correct, think of it as a vestigial trait
that only a few possess.
5) Exaptation (pre-adaptation): A trait that was adaptive under a prior set of
conditions and later provides the initial stage for evolution of a new adaptation
(contrivance) under a new set of conditions (in descendants).
6) Homology vs. Convergence: As covered in previous lectures, homology is the
similarity in traits between individuals as a result of the inheritance of those traits
from a common ancestor. Convergence, on the other hand, is when organisms
have traits in common, but do not have a common ancestor that provided them
with those traits. Typically, the traits in question arose from the independent
evolution of the traits by the different species or genetic lines. For example, the

glass snake has several traits in common with regular snakes, including being
limbless. However, the glass snake is really a lizard, having, among other lizardly
things, moveable eyelids and external ear openings. The most recent common
ancestor between the true snakes and glass snake had legs. The genetic line that
eventually gave rise to the true snakes evolved the loss of limbs for ease of
movement through burrows. Similarly, the genetic line that gave rise to the glass
snake also evolved to lose its limbs so that it too could move through burrows
more easily. However, since their most recent common ancestor had limbs, we
say that the two groups (true snakes and glass snakes) have adaptations that
converged to be similar as a result of similar environmental pressures.