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ORGANIC EVOLUTION
Chapter 6

Evolution - Defined
• Evolution – a change in the genetic composition of a
population over time.
• A change in the frequency of certain alleles.

• On a larger scale, evolution can be used to refer to

the gradual appearance of all biological diversity.

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Darwin’s Revolutionary Theory

• The Origin of
Species focused
attention on the
diversity of life,
similarities as well
as differences,
and the
adaptations
organisms have
for particular
environments.

Darwin’s Revolutionary Theory

• Charles Darwin
presented evidence
that many modern
organisms are
descended from
ancestral species
that were different.

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Darwin’s Revolutionary Theory

• Prevailing view of the world was that the Earth was
only a few thousand years old and that all life had
been created at the beginning and remained
unchanged.

Pre-Darwinian Evolutionary Ideas

• Several ancient Greek philosophers thought
life changed through time.
• Aristotle recognized fossils as forms of
ancient life.
• He developed the scala naturae (scale of
nature).
• Each form of life had a rung on the ladder.
• Organisms were arranged in order of complexity.
• The ancient Greeks didn’t propose an
evolutionary mechanism.

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Scala naturae
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Pre-Darwinian Evolutionary Ideas

• Lamarck was the

first to suggest an
explanation for
evolution.
• Inheritance of
acquired
characteristics
• Didn’t hold up to
testing

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A Mechanism for Evolution

• Darwin presented a mechanism for evolution –
natural selection.
• Organisms that are in some way more successful at
reproduction will pass on more of their genes.

• Over time the traits responsible for that success will

become widespread in the population.

• This theory holds up very well!

Alfred Russell Wallace

• Wallace independently
developed a theory of
natural selection.
• He sent his manuscript to
Darwin, spurring him to
finally publish his ideas.
• Both ideas were
presented to the Linnean
Society in 1858.
• Darwin finished On the
Origin of Species and
published it in 1859.

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Uniformitarianism

• Charles Lyell’s
principle of
uniformitarianism:
• Laws of physics &
chemistry present
throughout history of
Earth.
• Past geological
events similar to
today’s events.
• Principles of Geology

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Uniformitarianism
• Natural forces could
explain the
formation of fossil-
bearing rocks.
• Lyell concluded the
age of the earth
must be millions of
years.
• He stressed the
gradual nature of
geological
changes.

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Uniformitarianism and the Age of Earth

• Darwin studied the work of Lyell closely. He took the
first volume of Lyell’s Principles of Geology on the
Beagle. He received the second volume while on the
voyage.
• He concluded that Earth must be much older than 6000
years.
• Perhaps these slow changes could work on living things as
well.

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Evolution in Need of a Mechanism

• Darwin was not the first to have the thought that
organisms change through time.
• His grandfather, Erasmus Darwin, and others suggested that
life evolves as environments change.
• But a mechanism for that change was needed.

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Darwin (1809 – 1882)

• Darwin had a
lifelong love of
nature.
• His father wanted
him to study
medicine.
• This was not what
Darwin wanted and
he didn’t finish.

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Darwin
• After leaving medical school he attended
Cambridge University with the intention of entering
the clergy.
• His mentor and botany professor, John Henslow,
recommended him for a position as ship’s naturalist
aboard the Beagle.

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The Voyage of the Beagle

• Darwin started out on a five year trip around
the world aboard the Beagle in 1831. He was
22.
• As ship’s naturalist he spent his time on shore
collecting thousands of plant and animal
specimens and making important observations.

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The Voyage of the Beagle

• Darwin saw that the plants and animals that he
found in temperate areas of South America were
more similar to tropical South American species than
they were to temperate European species.

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The Voyage of the Beagle

• The fossils he found in South America were more like
modern South American species than European
species.

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The Voyage of the Beagle

• During the voyage he read Lyell’s Principles of
Geology.
• He had Lyell’s ideas in mind as he traveled and
observed the geology of South America.

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The Voyage of the Beagle

• He experienced an earthquake in Chile and
observed that the coastline had risen several feet.
• He also found marine fossils high in the Andes
Mountains.
• Darwin concluded that the mountains were formed
by a series of such earthquakes.

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The Voyage of the Beagle

• Darwin became interested in the

geographic distribution of organisms after
visiting the Galapagos Islands.

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After the Voyage

• After returning, Darwin

realized that
adaptation to the
environment and the
origin of new species
were closely linked
processes.
• Galapagos finch
species have evolved
by adapting to
specific conditions on
each island.

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Natural Selection

• After reading a paper by

Thomas Malthus
concerning the fact that
human populations
increase faster than
limited food resources,
Darwin noticed the
connection between
natural selection and this
ability of populations to
overreproduce.

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Natural Selection
• Only a small fraction of all offspring produced by any
species actually reach maturity and reproduce.

• Natural populations normally remain at a constant

size.

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Natural Selection
• Those that survive may have heritable traits
that increased their chances of survival.

• They will pass those traits on.

• The frequency of those traits will increase.

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Artificial Selection

• Artificial selection –
people selectively
breed organisms
with desired traits.
• Darwin noticed that
considerable
change can be
achieved in a short
period of time.

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Natural Selection
• Natural selection occurs when organisms with
particular heritable traits have more offspring
that survive and reproduce.

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Natural Selection

• Natural selection
can increase the
adaptation of an
organism to its
environment.

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Natural Selection
• When an environment changes, or when individuals
move to a new environment, natural selection may
result in adaptation to the new conditions.
• Sometimes this results in a new species.

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Natural Selection
• Individuals do not evolve; populations evolve.
• Evolution is measured as changes in relative
proportions of heritable variations in a population
over several generations.

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Natural Selection
• Natural selection can only work on heritable traits.
• Acquired traits are not heritable and are not subject to
natural selection.

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Natural Selection
• Environmental factors are variable.
• A trait that is beneficial in one place or time may be
detrimental in another place or time.

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Darwinian Evolutionary Theory: Evidence

• The main premise underlying evolutionary theory is
that the living world is always changing.
• Perpetual change in form & diversity of organisms
over the last 700 million years can be clearly seen in
the fossil record.

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Fossils

• Fossils are remnants of

past life preserved in the
earth.
• Complete remains –
insects in amber.
• Petrified skeletal parts
infiltrated with silica or
other minerals.
• Or traces of organisms
such as molds, casts,
impressions, trackways, or
fossilized excrement.

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The Fossil Record

• Fossils provide
support for the idea
that life changes
through time.
• Fossil intermediates
• Whales descended
from land mammals.
• Birds descended from
one branch of
dinosaurs.
• The oldest fossils are
of prokaryotes.

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Dating Fossils
• Geological time can be
measured in sedimentary rock
layers.
• The Law of Stratigraphy

• Dates oldest layers at the bottom and youngest at the top.

• Time is divided into eons, eras, periods and epochs.

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Dating Fossils
• Radiometric dating methods are based on the
decay of naturally occurring elements into other
elements.
• Different methods used for different time periods.

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Dating Fossils - example

• 40K
has a half life of 1.3 billion years – meaning half of
the 40K will have decayed to 40Ar and 40Ca. Half of
what remains will decay in the next 1.3 billion years.
• Measure ratio of remaining 40K to the amount of 40K
originally there (remaining 40K plus 40Ar and 40Ca).

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Fossil Record

• The fossil record of

macroscopic
organisms begins in the
Cambrian period: 505–
570 MYA.
• Fossil bacteria and
algae, casts of
jellyfishes, sponges
spicules, soft corals,
and flatworms are
found in Precambrian
rocks.
• Mostly microscopic

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Evolutionary Trends
• The fossil record shows that species arise and go
extinct repeatedly throughout geological history.
• Trends appear in the fossil record – directional
changes in features or patterns of diversity.

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Evolutionary Trends

• The evolution of horses

from the Eocene
epoch (57.8 MYA) to
the present is a well
studied trend.
• Body size – increasing
• Foot structure – fewer
toes
• Tooth structure –
larger grinding surface

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Common Descent

• Darwin proposed that all organisms have

descended from a single ancestral form.
• Life history is shown as a branching tree
called a phylogeny.

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Homology
• The phrase “descent with modification” summarizes
Darwin’s view of how Evolution works.
• All organisms descended from common ancestor.
• Similar species have diverged more recently.
• Homology – when similar structures result from shared
ancestry.

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Anatomical Homologies

• Homologous structures – variations on a

structural theme that was present in a common
ancestor.
• Example – vertebrate forelimbs have different
functions, but share the same underlying
structure.

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Anatomical Homologies
• Vertebrate embryos have a tail and pharyngeal
pouches.
• These structures develop into different but
homologous structures in adults.
• Gills in fishes
• Part of ears & throat in humans.

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Ontogeny & Phylogeny

• There are many parallels between ontogeny (an
individual’s development) and phylogeny
(evolutionary descent).
• Embryological similarities
• Features of an ancestors ontogeny can be shifted earlier or
later in a descendant's ontogeny.

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Ontogeny & Phylogeny

• Heterochrony – evolutionary
change in timing of
development.
• Characteristics can be
added late in development
and features are then
moved to an earlier stage.
• Ontogeny can be shortened
in evolution.
• Terminal stages may be
deleted causing adults of
descendants to resemble
youthful ancestors.
• Paedomorphosis
• Retention of ancestral
juvenile characters by
descendant adults.

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Developmental Modularity and

Evolvability
• Heterotopy – a change in the physical location of a
developmental process in an organism’s body.
• Process must be compartmentalized into semi-autonomous
modules to be expressed in new location
• Ex: Location of toepad development in geckos.

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Developmental Modularity and

Evolvability
• Evolvability – denotes the great evolutionary
opportunities created by semi-autonomous
developmental modules whose expression can be
moved from one part of the body to another.
• Allows for “experimentation” with the construction of many new
structures.

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Vestigial Organs
• Vestigial organs – remnants of structures that served
important functions in an ancestor.
• Remnants of pelvis and leg bones in snakes
• Appendix in humans

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Molecular Homologies
• Similarities can be found at the molecular level, too.
• The genetic code is universal - it is likely that all organisms
descended from a common ancestor.
• Different organisms share genes that have been inherited
from a common ancestor.
• Often, these genes have different functions, like the mammalian
forelimbs.

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Homologies & the Tree of Life

• Darwin’s evolutionary tree of life can explain
homologies.
• The genetic code is shared by all species because it goes
back deep into the ancestral past.
• More recent homologies are shared by only a smaller
branches of the tree.

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Homologies & the Tree of Life

• Homologies result in
a nested pattern.
• All life shares the
deepest layer.
• Each smaller group
adds homologies to
those they share
with larger groups.

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Speciation
• Speciation refers to the formation of new
species.
• Defining a species is difficult…
• Descent from common ancestral population.
• Ability to interbreed.
• Maintenance of genotypic & phenotypic
cohesion.
• Reproductive barriers prevent species from
interbreeding.
• Where do they come from?

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Allopatric Speciation
• Allopatric (another land) populations occupy
separate geographic areas.
• Separated geographically, but able to interbreed if brought
together.
• Over time, reproductive barriers may evolve so that
they could not interbreed.
• Allopatric speciation

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Allopatric Speciation
• The geographical separation can arise in two ways:
• Vicariant speciation is initiated when climatic or geological
changes fragment a species’ habitat, forming impenetrable
barriers.
• Founder events occur when a small number of individuals
disperse to a distant place where no other members of their
species exist.

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Hybrids

• Much can be
learned by studying
what happens when
previously isolated
populations come
into contact again.
• Hybrids are offspring
of members of two
closely related
species.

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Hybrids
• Species eventually become different enough that
they can’t produce hybrids.
• Premating barriers prevent mating from occurring in the first
place.
• Postmating barriers impair growth, survival, or reproduction of
the offspring.

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Sympatric Speciation

• Sympatric (same land) speciation occurs when

speciation occurs in one geographic area – a
lake for example.
• Individuals within the species become specialized
on a food type, shelter, part of the lake etc.
• Eventually reproductive barriers evolve.

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Parapatric Speciation
• Parapatric Speciation – geographically
intermediate between allopatric and sympatric
speciation.
• Two species are parapatric if their geographic ranges are
primarily allopatric but make contact along a borderline that
neither species successfully crosses.

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Adaptive Radiation
• Adaptive radiation – the production of ecologically
diverse species from a common ancestral stock.
• Common in lakes & islands – sources of new evolutionary
opportunities.

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Adaptive Radiation

• Archipelagoes increase opportunities for

both founder events and ecological
diversification.
• Entire archipelago isolated from the continent.
• Each island is geographically isolated from the
others.
• Ex: Galápagos Islands

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Gradualism

• Darwin’s theory of
gradualism proposes
that small differences
accumulate over time
producing the larger
changes we see over
geologic time.
• Certainly, this process is
always at work, but
probably does not
account for all changes.

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Punctuated Equilibrium

• Punctuated
equilibrium states
that phenotypic
evolution is
concentrated in
relatively brief
events of branching
speciation followed
by periods of stasis.

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Populations Evolve

• Variation exists within

any population.
• When natural
selection acts to
favor one trait over
another that trait will
increase in the
population.
• The population has
evolved, not any
one individual.

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The Modern Synthesis

• Population Genetics – the study of how populations
change over time.
• Dependent on both Darwin’s theory of natural selection and
Mendel’s laws of inheritance.
• All heritable traits have a genetic basis, some are controlled
by multiple genes – not as simple as in Mendel’s studies.

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The Modern Synthesis

• The modern synthesis is a comprehensive
theory of evolution that brings in ideas from
many fields.
• R. A. Fisher (statistician)
• J. B. S. Haldane (biologist)
• Theodosius Dobzhansky (geneticist)
• Sewall Wright (geneticist)
• Ernst Mayr (biogeographer)
• George Gaylord Simpson (paleontologist)
• G. Ledyard Stebbins (botanist)

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Populations
• Population – a localized, interbreeding group of
individuals of a particular species.
• Separate populations of a species may be isolated from
each other.

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Populations

• Sometimes the
populations
overlap, but little
interbreeding
occurs.

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Microevolution
• Microevolution – evolutionary changes
in the frequency of alleles in a
population.
• Polymorphism – occurrence of different
allelic forms of a gene in a population.
• If there is only one allele for a gene in the
population – every individual is homozygous
for the trait – it is fixed in the population.
• All alleles of all genes possessed by all
members of a population form a gene
pool.

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Microevolution
• Population geneticists measure the
relative frequencies of alleles in the
population.
• Allelic frequency

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Genetic Equilibrium
• According to Hardy-Weinberg equilibrium, the
hereditary process alone does not produce
evolutionary change.
• Allelic frequency will remain constant generation to
generation unless disturbed by mutation, natural selection,
migration, nonrandom mating, or genetic drift.
• Sources of microevolutionary change.

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Frequency of Alleles
• Each allele has a frequency
(proportion) in the population.
• Example population of 500 wildflowers.
• CRCR = red; CRCW = pink; CWCW = white
• 320 red, 160 pink, 20 white
• Frequency of CR =
(320 x 2) + 160 / 1000 = 800/1000 =.8 = 80%

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Frequency of Alleles
• p is the frequency of the most common allele (CR in
this case).
• p = 0.8 or 80%
• q is the frequency of the less common allele (CW in
this case).
•p+q=1
• q = 1- p = 1 – 0.8 = 0.2 or 20%

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Hardy-Weinberg Theorem

• Populations that are

not evolving are
said to be in Hardy-
Weinberg
equilibrium.
• As long as Mendel’s
laws are at work,
the frequency of
alleles will remain
unchanged.

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Hardy-Weinberg Theorem
• The Hardy-Weinberg theorem assumes random
mating.
• Generation after generation allele frequencies are
the same.

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Hardy-Weinberg Theorem
• At a locus with two alleles, the three genotypes will
appear in the following proportions:
• (p + q) x (p + q) = p2 + 2pq + q2 = 1

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Hardy-Weinberg Theorem
• Conditions:
• Very large population
• No gene flow into or out of the population
• No mutations
• Random mating
• No natural selection
• Departure from these conditions results in evolution.

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Practice with Hardy Weinberg

• Hardy & Weinberg studied the frequencies of alleles
in populations.
• Frequency – the proportion of individuals in a category in
relation to the total number of individuals. 100 cats, 84 black,
16 white – frequency of black = 84/100 = 0.84, white =0.16.
• Two alleles – p is common, q is less common.
• p+q = 1

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Practice with Hardy Weinberg

⚫ (p + q)2 = p2 + 2pq + q2
Individuals Individuals Individuals
homozygous heterozygous homozygous
for allele B for alleles B for allele b
&b

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Practice with Hardy Weinberg

• Used to calculate allele frequencies (p & q) in a
simple way.
• 100 cats, 16 white (bb) so q2 = 0.16
• q = square root of 0.16 = 0.40.
• Since p + q = 1; p = 1 – q = 0.60.
• p2 = 0.36; so 36 homozygous dominant (BB)
• 2pq = 0.48; 48 heterozygous (Bb)

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Where Does Variation Come From?

• Natural selection acts on the variation that is already
present in the population.
• But, where did that variation come from?

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Where Does Variation Come From?

• Two processes provide the variation in gene pools.
• Mutation
• Sexual recombination

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Mutation

• New genes or
alleles only result by
mutations.
• Mutations are
changes in the
nucleotide
sequence of DNA.

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Point Mutations
• Point mutation – a change in a single base pair.
• Often harmless
• Much of the DNA does not code for protein products.
• Genetic code is redundant.
• CGU, CGA, CGC, CGG all code for arginine.
• Occasionally significant
• Sickle cell disease.

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Mutations
• Beneficial mutations of any kind are very rare.
• Mutations that alter gene number or sequence are
almost always harmful.

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Gene Duplication
• Gene duplication occasionally provides an
expanded genome with new loci that may take on
new functions as selection continues.
• New genes can also appear when non-coding
introns get shuffled into the coding portion of the
genome.

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Sexual Recombination
• Sexual recombination is a much more common way
of producing variation in populations.
• Reshuffling of allele combinations already present in the
population is how variation is maintained in populations.
• Sexual reproduction rearranges alleles into fresh
combinations every generation.

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Natural Selection
• When natural selection is occurring, some individuals
are having better reproductive success than others.
• Alleles are being passed to the next generation in
frequencies that are different from the current generation.
• Hardy-Weinberg equilibrium is upset.

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Genetic Drift
• The smaller the sample, the greater the chance of
deviation from expected results.
• These random deviations from expected frequencies are
called genetic drift.
• Allele frequencies are more likely to deviate from the
expected in small populations.

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Genetic Drift

• Which allele was lost is due to random

chance.
• Over time, drift tends to reduce genetic
variation through random loss of alleles.

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The Bottleneck Effect

• Sometimes a
catastrophic event
can severely
reduce the size of a
population.
• The random
assortment of
survivors may have
drastically different
allele frequencies.
• Bottleneck effect

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The Bottleneck Effect

• The actions of people
sometimes cause
bottlenecks in other
species.
• N. California elephant seal
population reduced to 20-
100 individuals in the 1890s.
• Current population >
30,000.
• Variation drastically http://www.sealexperience.com/index.html

reduced – 24 genes with 1

allele.

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The Founder Effect

• Founder effect – When a small group of individuals
becomes separated from the population and form a
new population, the allele frequencies in their gene
pool may be different than the original population.

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Gene Flow
• The population can gain or lose new alleles through
gene flow.
• When individuals move into or out of a population,
they may carry the only copy of certain alleles in the
gene pool with them.
• Gene flow usually reduces differences between
populations.

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Natural Selection & Adaptation

• Natural selection is the only one of these ways of
altering the gene pool that results in adaptation.
• Selection depends on variation.

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Genetic Variation
• Variation in a population is always present.
• Heritable variation is the raw material of natural
selection.

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Genetic Variation

• Not all genetic variation is heritable.

• Environmental influences sometimes effect
phenotype.

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Polymorphism

• Different versions of
discrete characters
are called morphs.
• When a population
has two or more
morphs that are
common in the
population, it is
called polymorphic.
• This is phenotypic
polymorphism

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Protein Polymorphism
• Different allelic forms of a gene code for
slightly different proteins – protein
polymorphism.
• If the difference affects the protein’s net
electric charge, the different forms can be
separated using protein electrophoresis.

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Quantitative Variation
• Quantitative traits are those that show continuous
variation.
• Influenced by many genes.
• Height in humans, tail length in mice
• When trait values for a population are graphed, they follow a
bell shaped curve.

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Modes of Selection

• Directional –
variants at one of
the extremes are
favored.
• Disruptive –
variants at both
extremes are
favored.
• Stabilizing –
removes the
extremes.

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Evolutionary Fitness
• Fitness – the contribution an individual makes to the
gene pool of the next generation.
• Relative fitness – the contribution of one genotype
relative to the contribution of other genotypes at the
same locus.
• Natural selection acts on phenotypes.

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Preserving Variation
• Some variation is hidden from the natural selection
process in the form of recessive alleles in
heterozygotes.
• Less favorable recessive alleles can be maintained in
the population because they do not harm
heterozygous individuals.

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Sexual Selection

• Sexual selection –
natural selection for
mating success.
• May result in sexual
dimorphism –
differences
between the sexes.
• Secondary sexual
characteristics – not
directly involved in
reproduction.

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Intrasexual Selection
• Intrasexual selection – selection within the same sex
– results when individuals of one sex are competing
with each other for members of the other sex.
• Features that make the male a better fighter or more
intimidating to other males would be favored.

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Intersexual Selection

• Intersexual
selection – mate
choice – individuals
of one sex are
choosy in selecting
a mate.
• Features that make
an individual more
attractive to the
opposite sex would
be favored.

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Intersexual Selection
• Showiness that results from mate choice can be risky.
• Flashy tails of guppies make them more visible to predators.
• Benefits of finding a mate outweigh potential costs.
• Showiness may reflect overall health.

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Macroevolution
• Macroevolution refers to grand scale events in
evolution.
• Evolution of new structures
• Major trends in the fossil record

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Gould’s Tiers of Time

• Stephen Jay Gould
recognized three tiers of
time for evolutionary
processes:
• Tens to thousands of
years – population
genetic processes.
• Millions of years –
speciation and
extinction can be
measured and
compared among
different groups of
organisms.
• Tens to hundreds of
millions of years –
marked by episodic
mass extinctions.

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Speciation and Extinction Through

Geological Time
• A species has two possible fates:
• Become extinct or
• Give rise to new species.
• Speciation and extinction rates vary among species.
• Lineages with high speciation and low extinction
produce the greatest diversity.

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Speciation and Extinction Through

Geological Time
• Species Selection
• Differential survival and multiplication of species based on
variation among lineages.
• Species-level properties include mating rituals, social
structuring, migration patterns, geographic distribution, etc.

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Mass Extinctions
• Mass extinctions are episodic events where many
species go extinct at the same time.
• Permian extinction – 225 MYA – half the families of
shallow-water marine invertebrates and 90% of the
marine invertebrate species went extinct over a few
million years.
• Cretaceous extinction – 65 MYA – marks the end of the
dinosaurs as well as many other species.

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Mass Extinctions
• Many possible explanations
for mass extinctions have
been suggested.
• Alvarez hypothesis –
bombardment of the earth by
asteroids would send debris
into the atmosphere, altering
climate.
• Search for evidence
• Craters
• Atypical iridium concentrations

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Mass Extinctions
• Catastrophic species selection would result from
selection by these events.
• Mammals were able to use resources due to dinosaur
extinction.
• Paleontologist Elisabeth Vrba uses term Effect
Macroevolution to describe differential speciation
and extinction rates among lineages caused by
organismal-level properties.

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Endurance of Darwin’s Theory

• The beauty of Darwin’s theory is that it explains so
many different kinds of observations: anatomical
and molecular homologies that match patterns in
space (biogeography) and time (fossil record).

"Nothing in biology makes sense except in the light of evolution." Theodosius Dobzhansky, Geneticist

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