Evolution and Natural Selection

Nature encourages no looseness, pardons no errors

- Ralph Waldo Emerson

I have called this principle, by which each slight variation, if useful, is preserved,
by the term Natural Selection.
- Charles Darwin, The Origin of Species

In this lesson, we wish to ask:

 How did observations in nature lead to the formulation of the theory of evolution?
 What are the main points of Darwin's theory of evolution?
 How does the process of natural selection work?
 What evidence do we have for local adaptation?
 How can natural selection affect the frequency of traits over successive generations?

The (R)Evolution of Theory

The theory of evolution is one of the great intellectual revolutions of human history,
drastically changing our perception of the world and of our place in it. Charles Darwin put
forth a coherent theory of evolution and amassed a great body of evidence in support of this
theory. In Darwin's time, most scientists fully believed that each organism and each
adaptation was the work of the creator. Linneaus established the system of biological
classification that we use today, and did so in the spirit of cataloguing God's creations.

In other words, all of the similarities and dissimilarities among groups of organisms that are
the result of the branching process creating the great tree of life (see Figure 1), were
viewed by early 19th century philosophers and scientists as a consequence of omnipotent
design.

Figure 1: A phylogenetic "tree of life" constructed by computer analysis of cyochrome c molecules in the organisms
shown; there are as many different trees of life as there are methods of analysis for constructing them.

However, by the 19th Century, a number of natural historians were beginning to think of
evolutionary change as an explanation for patterns observed in nature. The following ideas
were part of the intellectual climate of Darwin's time.

 No one knew how old the earth was, but geologists were beginning to make
estimates that the earth was considerably older than explained by biblical creation.
Geologists were learning more aboutstrata, or layers formed by successive periods
of the deposition of sediments. This suggested a time sequence, with younger strata
overlying older strata.
  A concept called uniformitarianism, due largely to the influential geologist Charles
Lyell, undertook to decipher earth history under the working hypothesis that present
conditions and processes are the key to the past, by investigating ongoing,
observable processes such as erosion and the deposition of sediments.
 Discoveries of fossils were accumulating during the 18th and 19th centuries. At first
naturalists thought they were finding remains of unknown but still living species. As
fossil finds continued, however, it became apparent that nothing like giant dinosaurs
was known from anywhere on the planet. Furthermore, as early as 1800, Cuvier
pointed out that the deeper the strata, the less similar fossils were to existing
species.
 Similarities among groups of organisms were considered evidence of relatedness,
which in turn suggested evolutionary change. Darwin's intellectual predecessors
accepted the idea of evolutionary relationships among organisms, but they could not
provide a satisfactory explanation for how evolution occurred.
 Lamarck is the most famous of these. In 1801, he proposed organic evolution as the
explanation for the physical similarity among groups of organisms, and proposed a
mechanism for adaptive change based on the inheritance of acquired characteristics.
He wrote of the giraffe:
"We know that this animal, the tallest of mammals, dwells in the interior of Africa, in places
where the soil, almost always arid and without herbage, obliges it to browse on trees and to
strain itself continuously to reach them. This habit sustained for long, has had the result in
all members of its race that the forelegs have grown longer than the hind legs and that its
neck has become so stretched, that the giraffe, without standing on its hind legs, lifts its
head to a height of six meters."

In essence, this says that the necks of Giraffes became long as a result of continually
stretching to reach high foliage. Larmarck was incorrect in the hypothesized mechanism, of
course, but his example makes clear that naturalists were thinking about the possibility of
evolutionary change in the early 1800's.

 Darwin was influenced by observations made during his youthful voyage as naturalist
on the survey ship Beagle. On the Galapagos Islands he noticed the slight variations
that made tortoises from different islands recognizably distinct. He also observed a
whole array of unique finches, the famous "Darwin's finches," that exhibited slight
differences from island to island. In addition, they all appeared to resemble, but
differ from, the common finch on the mainland of Ecuador, 600 miles to the east.
Patterns in the distribution and similarity of organisms had an important influence of
Darwin's thinking. The picture at the top of this page is of Darwin's own sketches of
finches in his Journal of Researches.
 In 1859, Darwin published his famous On the Origin of Species by Means of Natural
Selection, a tome of over 500 pages that marshalled extensive evidence for his
theory. Publication of the book caused a furor - every copy of the book was sold the
day that it was released. Members of the religious community, as well as some
scientific peers, were outraged by Darwin's ideas and protested. Most scientists,
however, recognized the power of Darwin's arguments. Today, school boards still
debate the validity and suitability of Darwin's theory in science curricula, and a whole
body of debate has grown up around the controversy (see the WWW
site Talk.Origins for an ongoing dialogue). We do not have time to cover all of
Darwin's evidence and arguments, but we can examine the core ideas. What does
this theory of evolution say?
Darwin's Theory
Darwin’s theory of evolution entails the following fundamental ideas. The first three ideas
were already under discussion among earlier and contemporaneous naturalists working on
the “species problem” as Darwin began his research. Darwin’s original contributions were
the mechanism of natural selection and copious amounts of evidence for evolutionary
change from many sources. He also provided thoughtful explanations of the consequences
of evolution for our understanding of the history of life and modern biological diversity.

 Species (populations of interbreeding organisms) change over time and space. The
representatives of species living today differ from those that lived in the recent past,
and populations in different geographic regions today differ slightly in form or
behavior. These differences extend into the fossil record, which provides ample
support for this claim.

 All organisms share common ancestors with other organisms. Over time,
populations may divide into different species, which share a common ancestral
population. Far enough back in time, any pair of organisms shares a common
ancestor. For example, humans shared a common ancestor with chimpanzees about
eight million years ago, with whales about 60 million years ago, and with kangaroos
over 100 million years ago. Shared ancestry explains the similarities of organisms
that are classified together: their similarities reflect the inheritance of traits from a
common ancestor.

 Evolutionary change is gradual and slow in Darwin’s view. This claim was supported
by the long episodes of gradual change in organisms in the fossil record and the fact
that no naturalist had observed the sudden appearance of a new species in Darwin’s
time. Since then, biologists and paleontologists have documented a broad spectrum
of slow to rapid rates of evolutionary change within lineages.

The primary mechanism of change over time is natural selection, elaborated below. This
mechanism causes changes in the properties (traits) of organisms within lineages from
generation to generation.

The Process of Natural Selection

Darwin’s process of natural selection has four components.

1. Variation. Organisms (within populations) exhibit individual variation in appearance

and behavior. These variations may involve body size, hair color, facial markings,
voice properties, or number of offspring. On the other hand, some traits show little
to no variation among individuals—for example, number of eyes in vertebrates.
2. Inheritance. Some traits are consistently passed on from parent to offspring. Such
traits are heritable, whereas other traits are strongly influenced by environmental
conditions and show weak heritability.
3. High rate of population growth. Most populations have more offspring each year
than local resources can support leading to a struggle for resources. Each
generation experiences substantial mortality.
 4. Differential survival and reproduction. Individuals possessing traits well suited for
the struggle for local resources will contribute more offspring to the next generation.

From one generation to the next, the struggle for resources (what Darwin called the
“struggle for existence”) will favor individuals with some variations over others and thereby
change the frequency of traits within the population. This process is natural selection. The
traits that confer an advantage to those individuals who leave more offspring are called
adaptations.

In order for natural selection to operate on a trait, the trait must possess heritable variation
and must confer an advantage in the competition for resources. If one of these
requirements does not occur, then the trait does not experience natural selection. (We now
know that such traits may change by other evolutionary mechanisms that have been
discovered since Darwin’s time.)

Natural selection operates by comparative advantage, not an absolute standard of

design. “…as natural selection acts by competition for resources, it adapts the inhabitants
of each country only in relation to the degree of perfection of their associates” (Charles
Darwin, On the Origin of Species, 1859).

During the twentieth century, genetics was integrated with Darwin’s mechanism, allowing us
to evaluate natural selection as the differential survival and reproduction of genotypes,
corresponding to particular phenotypes. Natural selection can only work on existing
variation within a population. Such variations arise by mutation, a change in some part of
the genetic code for a trait. Mutations arise by chance and without foresight for the
potential advantage or disadvantage of the mutation. In other words, variations do not
arise because they are needed.

Evidence of Natural Selection

Let's look at an example to help make natural selection clear.

Industrial melanism is a phenomenon that affected over 70 species of moths in

England. It has been best studied in the peppered moth, Biston betularia. Prior to
1800, the typical moth of the species had a light pattern (see Figure 2). Dark colored
or melanic moths were rare and were therefore collectors' items.

During the Industrial Revolution, soot and other industrial wastes darkened tree trunks and
killed off lichens. The light-colored morph of the moth became rare and the dark morph
became abundant. In 1819, the first melanic morph was seen; by 1886, it was far more
common -- illustrating rapid evolutionary change.

Eventually light morphs were common in only a few locales, far from industrial areas.
The cause of this change was thought to be selective predation by birds, which
favored camouflage coloration in the moth.

In the 1950's, the biologist Kettlewell did release-recapture experiments using both
morphs. A brief summary of his results are shown below. By observing bird predation
from blinds, he could confirm that conspicuousness of moth greatly influenced the
chance it would be eaten.
 Recapture Success

light moth dark moth

non-industrial woods 14.6 % 4.7 %
industrial woods 13 % 27.5 %

Local Adaptation - More Examples

So far in today's lecture we have emphasized that natural selection is the
cornerstone of evolutionary theory. It provides the mechanism for adaptive change.
Any change in the environment (such as a change in the background color of the
tree trunk that you roost on) is likely to lead to local adaptation. Any widespread
population is likely to experience different environmental conditions in different parts
of its range. As a consequence it will soon consist of a number of sub-populations
that differ slightly, or even considerably.

The following are examples that illustrate the adaptation of populations to local
conditions.

o The rat snake, Elaphe obsoleta, has recognizably different populations in

different locales of eastern North America (see Figure 3). Whether these
should be called geographic "races" or subspecies is debatable. These
populations all comprise one species, because mating can occur between
adjacent populations, causing the species to share a common gene pool (see
the lecture on speciation).

Figure 3: Subspecies of the rat snake Elaphe obsoleta, which interbreed where their
ranges meet.

o Galapagos finches are the famous example from Darwin's voyage. Each island
of the Galapagos that Darwin visited had its own kind of finch (14 in all),
found nowhere else in the world. Some had beaks adapted for eating large
seeds, others for small seeds, some had parrot-like beaks for feeding on buds
and fruits, and some had slender beaks for feeding on small insects (see
Figure 4). One used a thorn to probe for insect larvae in wood, like some
woodpeckers do. (Six were ground-dwellers, and eight were tree finches.)
(This diversification into different ecological roles, or niches, is thought to be
necessary to permit the coexistence of multiple species, a topic we will
examined in a later lecture.) To Darwin, it appeared that each was slightly
modified from an original colonist, probably the finch on the mainland of
South America, some 600 miles to the east. It is probable that adaptive
radiation led to the formation of so many species because other birds were
few or absent, leaving empty niches to fill; and because the numerous islands
of the Galapagos provided ample opportunity for geographic isolation.

Stabilizing, Directional, and Diversifying Selection

Finally, we will look at a statistical way of thinking about selection. Suppose that
each population can be portrayed as a frequency distribution for some trait -- beak
size, for instance. Note again that variation in a trait is the critical raw material for
evolution to occur.
 What will the frequency distribution look like in the next generation?

First, the proportion of individuals with each value of the trait (size of beak, or body weight)
might be exactly the same. Second, there may be directional change in just one direction.
Third (and with such rarity that its existence is debatable), there might be simultaneous
change in both directions (e.g. both larger and smaller beaks are favored, at the expense of
those of intermediate size). Figures 5a-c capture these three major categories of natural
selection.

Under stabilizing selection, extreme varieties from both ends of the frequency distribution
are eliminated. The frequency distribution looks exactly as it did in the generation before
(see Figure 5a). Probably this is the most common form of natural selection, and we often
mistake it for no selection. A real-life example is that of birth weight of human babies (see
Figure 6).

Under directional selection, individuals at one end of the distribution of beak sizes do
especially well, and so the frequency distribution of the trait in the subsequent generation is
shifted from where it was in the parental generation (see Figure 5b). This is what we usually
think of as natural selection. Industrial melanism was such an example.

The fossil lineage of the horse provides a remarkable demonstration of directional

succession. The full lineage is quite complicated and is not just a simple line from the
tiny dawn horse Hyracotheriumof the early Eocene, to today's familiar Equus.
Overall, though, the horse has evolved from a small-bodied ancestor built for moving
through woodlands and thickets to its long- legged descendent built for speed on the
open grassland. This evolution has involved well- documented changes in teeth, leg
length, and toe structure (see Figure 7).

Under diversifying (disruptive) selection, both extremes are favored at the

expense of intermediate varieties (see Figure 5c). This is uncommon, but of
theoretical interest because it suggests a mechanism for species formation without
geographic isolation (see the lecture on speciation).

Summary
Darwin's theory of evolution fundamentally changed the direction of future scientific
thought, though it was built on a growing body of thought that began to question
prior ideas about the natural world.

The core of Darwin's theory is natural selection, a process that occurs over
successive generations and is defined as the differential reproduction of genotypes.

Natural selection requires heritable variation in a given trait, and differential survival
and reproduction associated with possession of that trait.

Examples of natural selection are well-documented, both by observation and through

the fossil record.

Selection acts on the frequency of traits, and can take the form of stabilizing,
directional, or diversifying selection.