When did life begin?
The Origin and Evolution of
Life on Earth
When did life begin?
• Stromatolites (3.5 bill. Yr)
– Rocks with distinctive layer
structure
• Look identical to living
mats of microbes
– Layers of microbes and
sediment
– Top layer uses
photosynthesis
– Lower layers use top layer’s
byproducts
When did life begin?
• Evidence in metamorphic
rocks that life existed
3.85 billions years ago
• Low C12/C13 fraction in
rock layers suggests life
– Biological processes prefer
C12 to C13
• Find lower fraction of C13
– Non-biological processes
have no preference, so find
equal amounts
Living Fossils
• DNA used as living fossil
• The more alike the DNA sequence
between species, the more recent their
divergence and extinction of their common
ancestor
• Quite early in Earth’s history
• Cannot pinpoint time, but can narrow
down a time period with 3 lines of
evidence
When did life begin?
• Microfossils dating to
3.5 billion years ago
• Difficult to distinguish
from mineral
structures
• Analysis shows that
some structures
contain organic
carbon
-found in at least 3 sites
When did life begin?
• Rocks before ~4 billion years old are rare and
hard to find
• Time of heavy bombardment ended about 3.8-
4.0 billion years ago
– Last devastating impact between 4.2-3.9 bill. Yr ago
• Evidence suggests life as long as 3.85 billion
years ago and definitely at 3.5 billion years ago
• Life rose and dominated the planet between
100-500 million years
Living Fossils
• Bacteria and Archaea: genetic material NOT separated
from rest of cell
• Eukarya: DNA separated from rest of cell by membrane
• Extremophiles (live near deep-sea vents or in hot
springs) closest to root of tree of life
Where did life begin?
• Land is unlikely
– No O2, no ozone: UV destroys molecular bonds
• Shallow ponds
– Once favored, full of organic material
– When evaporated, organic chemical concentration
increases making it easier to combine complex
molecules leading to life
– Current experiments indicate lack of chemical energy
sufficient to support life
• Deep-sea vents/hot springs
– DNA evidence suggests that early organisms survived
in conditions similar to deep-sea vents
– Plenty of chemical energy available
Organic Chemistry on Early Earth
• In 1920’s, scientists hypothesized that the
chemicals in the early atmosphere, fueled
by sunlight, would spontaneously create
organic molecules
• Tested by Miller-Urey experiment 1950’s
Variations of Miller-Urey Experiment
• Different mixes of gases to represent
atmosphere
• Different energy sources, like UV
(sunlight)
• Results: ALL PRODUCE AMINO ACIDS AND
COMPLEX ORGANIC MOLECULES
– Not as much as original experiment
– MUST be more sources of organic material
How did life begin?
• Simplest organisms today and those dated
3.5 billion years ago are remarkable
advanced
• What are the natural chemical processes
that could have led to life?
• Assumptions
– Life began under chemical conditions of early Earth
– Life did not migrate to Earth
Miller-Urey Experiment
• First flask partially filled with water and heated
to produce water vapor (sea)
• Water vapor was moved to a second flask where
methane and ammonia vapor was added
(atmosphere)
• Electric sparks (lightening) in second flask was
energy source for chemical reactions
• Below second flask, water vapor cooled (rain)
and recycled to first flask (sea)
• Result: turned brown with amino acids and other
complex organic molecules
Time to think……
We have discussed the formation of the
solar system and the formation of the
terrestrial planets. Now, what is wrong
with the Miller-Urey experiment?
Sources of Organic Molecules
• Chemical reactions in atmosphere
– Lab experiments show this is likely
• Organic material brought by impacts
– Chemical analysis of comets and
carbonaceous chondrites show that they have
organic molecules
• Chemical reactions near deep-sea vents
– Heat from undersea volcano can fuel chemical
reactions between water and minerals
Transition from chemistry to biology
• Organic molecules are building blocks of
life.
• Low probability of forming life even if
repeated several times.
• Intermediate steps of high probability are
necessary
Search for Self-Replicating Molecule
• Work backward from organisms that live
today
• DNA is double-stranded = complicated
• RNA obvious candidate, more simple than
DNA
– Hereditary information
– Can serve as template for replication
– Fewer steps to produce backbone structure

Search for Self-Replicating Molecule Search for Replicating Molecule

• Problem: RNA and DNA require enzymes
to replicate
• In 1980’s determined that RNA might
catalyze their own replication instead of
other enzymes
• Early Earth was an RNA-world
• On Early Earth, short strands of RNA-like
molecules were produced spontaneously
partially or completely
• RNA-like molecules that could replicate faster
with less errors soon dominated population
• Copying errors introduced mutations, ensuring
the production of many variations of successful
•
molecules
Allowed molecular evolution to continue
• RNA-world gave way to DNA-world
– DNA less prone to copying errors
– DNA more flexible hereditary material
– RNA kept some of its original functions

Assembling Complex Organic Assembling Complex Organic

Molecules
• Organic soup was too dilute to favor the creation
of complex organic molecules
• Lab experiment with possible solution: When
hot sand, clay or rock is placed in dilute organic
solution, complex molecules self-assemble
– Organic molecules stick to surface of clay
– Increases density and likelihood of reactions
– Strands of RNA up to 100 bases have been produced
this way
Molecules
• Other inorganic minerals may have also
had a similar role
• Iron pyrite (fool’s gold)
– Positive charges on surface which allows
organic molecules to adhere
– Formation of pyrite releases energy which
could be used as fuel for chemical reactions

Early Cell-like Structures Early Cell-like Structures

• Advantages to • Lab experiments suggest that membrane
enclosing enzymes structures existed on early Earth
with RNA molecules
• Close proximity
• Form spontaneously
increases rate of – Cool down warm-water solution of amino
reactions between acids
them – Mix lipids (fats) with water
• Isolate contents from
outside world
Nonliving Pre-Cells have Lifelike
Handedness
Behavior
• Grow in size until • Organic molecules come in left- and right-
unstable then split to
form a ‘daughter’ cell
handed forms
• Selectively allow other • Non-biological processes have not
types of molecules to preference
pass in/out of
membrane • Biological processes DO have a preference
• Store energy in the • If both left- and right-handed RNA
form of electric developed, why did one die out?
voltage

Quick Summary Panspermia?

• Panspermia = life originated elsewhere
and migrated to Earth
• Life began in rock, then kicked off the
planet by an impact
• Support: organic material is everywhere,
and some bacteria can withstand large
amounts of radiation and go dormant
under low atmospheric conditions

Panspermia
• 2 schools of thought
• School 1: life did not evolve as easily as
imagined on early Earth in timescales
we’ve determined
• Problem: entire solar system was under
heavy bombardment at the same time
• Other possibility: interstellar migration
• Problem: rock to be ejected out of its own
system, then fall into ours and hit the tiny
planet of Earth
Panspermia
• School 2: life evolved easily and was
everywhere with suitable conditions
• Earth was not first planet with suitable
conditions
• Migration of life from another planet (say
Mars) dominated before early life on Earth
could
– We’re Martians!!!!

Panspermia
• Martian meteorites
• Both have possible
fossil evidence of life
on Mars

Living cyanobacteria Microfossils in carbonaceous chondrites

Time to think…… Early Evolution and Rise of O2

Work out Exercise I. When, Where and
How? in the class activity ‘Origins of Life
on Earth’.
• First organisms had simple metabolism
• Atmosphere was O2 free, must have been
anaerobic
• Probably chemoheterotrophs
– Obtained nutrients from organic material
– Obtained nutrients from inorganic material
• Modern archaea appear to be close to the root of
the tree of life
• Obtaining energy from chemical reactions involving
hydrogen, sulfur and iron compounds (all
abundant on early Earth)

Early Evolution Photosynthesis

• Natural selection probably resulted in rapid • Most important new metabolic process evolved
diversification gradually
• Modern DNA has enzymes that reduce the • Organisms that lived close to ocean surface
probably developed means of absorbing sunlight
rate of mutations (UV in particular)
• RNA is not so lucky, more likely to have • Once absorbed, developed method of turning it
copying errors into energy
• Higher mutation rate in early evolution – Modern organisms of purple sulfur bacteria and green
sulfur bacteria much like early photosynthetic
than now microbes, use H2S instead of H2O for photosynthesis

Photosynthesis
• Using water for photosynthesis developed later, perhaps
3.5 billion years ago
• First appearing in cyanobacteria (blue-green algae)
• By product of O2, released into atmosphere
• Changed the world!
Rise of O2
• O2 is highly reactive
• All initial O2 would react with rock and minerals
in water
• O2 could not accumulate in atmosphere until
surface rock was saturated
• Rocks 2-3 bill. Yr old called banded iron
formations, show atmosphere had <1% of
current amount of O2
• Rock evidence suggests that O2 amounts in
atmosphere began to rise about 2.0 bill. Yr ago
• Clear evidence of O2 near current levels appears
only 200 million yr ago
– Find charcoal (fossil fuel)
– Indicates enough O2 in atmosphere for fires to burn

Rise of O2 Early Eukaryotes

• Rise of O2 would have created a crisis for
life
• O2 reacts with bonds of organic materials
• Surviving species avoided effects of O2
because they lived or migrated to
underground locations
– Many anaerobic microbes found in such
locales today
• Fossil evidence dates to 2.1 bill. Yr ago
• Dates to when O2 rising in atmosphere
• DNA evidence suggests that prokaryotes and
eukaryotes separated from common ancestor
much earlier
• O2 played a key role in eukaryote evolution
– Cells can produce energy more efficiently using
aerobic metabolism than anaerobic metabolism
– Adaptations of aerobic organisms could develop
adaptations that required more energy that would be
available for anaerobic organisms
The Cambrian Explosion Colonization of Land

• Animal branch of the tree of life
• Different classifications based on body
plan
• All known body plans made appearance in
fossil record in a time span of 40 million
years
– <1% of Earth’s age
– Animal diversity began 545 mill. Yr ago
• Life flourished where liquid water exist
• Life on land was more complicated
– Had to develop means of collecting solar
energy above ground and nutrients below
• Life in shallow ponds or edges of lakes
– Water evaporates
– Natural selection favored that which could
withstand periods of drought

Colonization of Land Mass Extinctions

• DNA evidence suggests that plants evolved from
an algae
• It took only 75 mill. Yrs for animals to follow
plants out of water

Mass Extinctions Primate Evolution

• Possible Causes • Monkeys, apes, lemurs and humans have

– Impacts common ancestor that lived in trees
• Impact sites found for K-T boundary • Tree life
• Suspected for Permian extinction 245 mill yr ago
– Limber arms for swinging between branches
– Active volcanism
– Eyes in front of head for depth perception
• Climate change
– Offspring would be born more helpless than
– External influence for copying errors
other animals
• Increase in solar particles or radiation hitting
surface
• Local supernova

Emergence of Humans Emergence of Humans

• Did NOT evolve from gorillas or monkeys

• Share a common ancestor that lived just a
few million years ago
• 98% of human genome is identical to
genome of the chimpanzee
• 2% difference in genome separates the
success of humans verses chimps
– Also indicates evolution of intelligence is
complex
Emergence of Humans Emergence of Humans
• After hominids diverged from chimps and
gorillas, evolution has followed a complex path
• Numerous hominids species existed, some
during the same time period
– All humans are the same species
• First skull fossils that are identical to modern
human skulls dates to 100,000 yr old
• Our ancestors shared the Earth with
Neanderthals
– Went extinct 35,000 years ago

Cultural and Technological

Time to think……
Evolution
• Have not undergone biological evolution in
40,000 years Work out Exercise II. From Microbes to
– Mutation rates are slow Complex Organisms in class activity
• Dramatic cultural changes ‘Origins of Life on Earth’.
– Transmission of knowledge between generations
• Spoken to written word, thousands of years
• agriculture
• Technological evolution
– Result of coupling between science and technology
– About 100 years between industrial revolution to
landing on the Moon and generating weapons of
mass destruction

Historical Context for

evolutionary Theory
■ Plato (427-347 B.C) – believed in two worlds: a real world (ideal and
eternal) and an illusory world (imperfect and perceived through the
senses)

■ Aristotle (384-322 B.C) – believed that living things could be arranged

on a scale, or ladder of increasing complexity later called the ‘scale of
NATURAL SELECTION nature’.
BRINGS EVOLUTION
Mrs. Guerra ■ Carolus Linnaeus (1707-1778) – founder of taxonomy. He came up
BIO 231 Evolution & Biodiversity with the binomial system

Georges Cuvier

■ French scientists
■ Founder of study of fossils
■ Theory of Catastrophism
■ Provided evidence in the extinction of organisms on earth
 James Hutton
■ Theory of Gradualism
Jean-Baptiste Lamarck
Theory of evolution
1. Use and Disuse
2. Inheritance of acquired
traits
•Darwin made tremendously
important observations in many
places including the
Galapagos Islands
DARWIN
•Darwin wrote for a long time,
and was finally motivated to
publish his "Great Work" only
after almost being "scooped"
by Alfred Russell Wallace
(1823-1913).
Charles Lyell
■ Theory of Uniformitarianism
■ Wrote the book called
‘Principles of Geology’
Charles Darwin (1809-1882)
Voyage of the Beagle;
Origin of Species
Career in science really began with
his voyage on HMS Beagle (1831-
1836).
•After the voyage is apparently when
Darwin’s ideas came together.
•He had come to doubt the
fixity of species, and he struggled to
provide an explanation.
•He also read Malthus's "Essay
on the Principle of Population"
(1798) during this period
After a joint paper that
Wallace presented to the Royal
Society, Origin of Species was
published in 1859.
 Two main parts to Origin of Species
There are really five ideas or
"theories" that were present in Origin
of Species
■ 1. Common Descent with Modification
1. The biological world is changing.
■ 2. Natural Selection 2. Concept of Common Descent
3. Gradualism.
4. Multiplication of Species
5. Natural Selection

Common descent with Modification Natural Selection

Ernst Mayr has described
the logic of Darwinian natural
selection. He identifies 5 facts and 3
inferences.
■ It holds that particular groups or species of living things can
undergo modification in successive generations with such
change sometimes resulting in formation of new separate
species.
Observation 1:
All species have potential fertility which could
allow exponential increase in population size.
■ Eg. like a tree with multiple branches
Observation 2:
But most populations are stable in size.

Observation 3:
Natural resources are limited.

■ Inference 1: ■ Observation 4:
■ Potential fertility produces more offspring than the limited There is variation in many traits among individuals in a population.
natural resources can support;

■ Observation 5:
■ This leads to a “struggle for existence” among individuals -- only
Much of the variation among individuals is heritable.
a fraction of offspring and adults survive.

■ Inference 2:
- Survival in the struggle for existence is not random, but
depends on the inherited traits of the surviving individuals.

- Those whose inherited traits make them most “fit” will

survive and leave more descendents than the less “fit”
individuals.

■ Inference 3:
- The differences in survival and reproduction will lead, over
time, to an increase in the frequency of those individuals
with the favorable traits, and this trend will accumulate over
generations.
 Evidence for Evolution
•Idea of evolution occurring was
controversial

•Evolution is defined as change -- but

change in what?
•“Evolution” implies change
large enough to lead to separate
species.
Species Concepts! -- topic for
discussion

Three lines of evidence usually cited: 1.Observed change:

Examples
1. Observed Change: data from small-scale examples of evolution, or short
term experiments A. Change in the beak length of soapberry bugs
2. Observations on homologous structures (comparative anatomy)
3. Observation on embryos (comparative embryology)
4. The fossil record
5. Biogeography
6. Molecular Biology
7. Radiometric data B. Resistance to Pesticides
e.g., rapid appearance of DDT resistance in flies,
mosquitos.
In less than 40 years, over 200 insect species are know
to have evolved resistance to DDT.

2. Observations on Homologous
structures
C. Industrial Melanism Homologies are the subject of comparative anatomy.
e.g., pepper moth Biston betularia Similarities can be homologous or analogous.

Analogous structures are

similar and related functionally.
E.g., Wings of insects and bats

•Homologous structures are

similar, but not necessarily
functionally related.

•(Superficially they may be quite

different.)
 3. Observations on embryos
■ Also called comparative embryology
■ Here closely related organism go through similar stages in their
embryonic development.

■ Eg. all vertebrate embryo go through a stage in which they have gill
pouches on the sides of their throats.

Pharyngeal
pouches

Post-anal
tail
WHAT CAUSES THESE
Chick embryo Human embryo
SIMILARITIES???

NON HOMOLOGOUS Other examples of homologies:

STRUCTURES - universality of the genetic
code

- vestigial structures

VESTIGIAL STRUCTURES

Pakicetus (terrestrial)

Rhodocetus (predominantly aquatic)

Pelvis and Dorudon (fully aquatic)

hind limb

Pelvis and
hind limb
Balaena (recent whale ancestor)
 WHICH OF THE FOLLOWING 4. The Fossil Record
•Formerly, the fossil record was very spotty. Today it is
PAIRS ARE HOMOLOGOUS? Fairly complete for some lineages, and clearly shows the
ANALOGOUS? scope of evolutionary change.

■ The dorsal fins of porpoise and a salmon? •Fossils -- mainly in sedimentary rocks. Not all types of
organisms fossilize.

■ The jointed leg of a ladybird bettle and a robin

■ Rhesus monkey’s tail and a human coccyx

TRANSITIONAL FORMS

e.g., Archeopteryx - has both

reptile-like and bird-like
features. True “missing link”
discovered in 1861.

A Skull of Homo erectus

C Fossilized organic matter of a leaf 40 MYO

Horses -- very complete fossil record.

Earliest known horse-like

animal
Eohippus = Hyracotherium
(ca. 20” high, ca. 60 MY bp.)

E Insect in amber
35 myo

5. Biogeography 6. Molecular Biology

■ Geographical distribution of spp. ■ Supports Darwin’s boldest speculation that all forms of life
are related to some extent through branching descent from
the earliest organism.
 7. Radiometric dating
■ Is a technique used to determine the age of the earth and its
organisms by measuring the decay of the radioactive elements they
contain.
■ Elements can be transformed into another at a fixed rate
■ Eg. U-238 to Pb-206 it takes 4.5 billion yrs. for half a given amount of
(Microevolution)
U-238 to decay to pb-206
■ Half-life
■ 40 different techniques today being used

 Genetic variations in populations

 Overview: The Smallest Unit of Evolution  Contribute to evolution
 One common misconception about evolution is
that individual organisms evolve, in the Darwinian ฀
sense, during their lifetimes
 Natural selection acts on individuals, but
populations evolve

Figure 23.1

 Concept 23.1: Population genetics provides a foundation

for studying evolution The Modern Synthesis
 Microevolution
฀  Population genetics
฀
 Is change in the genetic makeup of a population from  Is the study of how populations change genetically over time
generation to generation
 It is the study of the distributions and changes in allele
frequency in a population, as it is subject to four main
evolutionary processes: natural selection, genetic drift,
mutation and recombination
 Reconciled Darwin’s and Mendel’s ideas

Figure 23.2

Gene Pools and Allele

Frequencies
฀ MAP

 A population AREA

 Is a localized group
of individuals that
are capable of
 The modern synthesis interbreeding and
•

 Integrates Mendelian genetics with the Darwinian theory of producing fertile Fairbanks

evolution by natural selection offspring Fortymile

herd range •

 Focuses on populations as units of evolution Whitehorse

Figure 23.3
 The Hardy-Weinberg
Gene Pool AA aa Aa
Theorem
 The gene pool ฀  The Hardy-Weinberg theorem ฀
 Is the total aggregate of genes in a population at any  Describes a population that is not evolving
one time  States that the frequencies of alleles and genotypes in a
 Consists of all gene loci in all individuals of the population’s gene pool remain constant from generation to
population generation provided that only Mendelian segregation and
recombination of alleles are at work

Generation
1
Preservation of Allele
Frequencies
CRCR CW CW
genotype genotype

฀ ฀
Plants mate

Generation

 Mendelian inheritance
2
All CRCW
(all pink flowers)

Preserves genetic 50% CR 50% CW

 In a given population where gametes contribute to the next
gametes

variation in a population
gametes

Come together at random generation randomly, allele frequencies will not change

Generation
3
25% CRCR 50% CRCW 25% CWCW
For eg. every time a gamete is drawn from the
Pool at random, the chance the gamete will bear
50% CR 50% CW
an A allele is 0.8 & a allele is 0.2.
gametes gametes

Come together at random

Generation
4
25% CRCR 50% CRCW 25% CWCW

Alleles segregate, and subsequent

generations also have three types

Figure 23.4 of flowers in the same proportions

Hardy-Weinberg  A population in Hardy-Weinberg equilibrium

Gametes for each generation are drawn at random from

Equilibrium
the gene pool of the previous generation:

฀ ฀
80% CR (p = 0.8) 20% CW (q = 0.2)

Sperm
CR CW
(80%) (20%)

 Hardy-Weinberg equilibrium p2 pq

 Describes a population in which random mating occurs

(80%)
CR
Eggs

 Describes a population where allele frequencies do not p2

64%
CRCR
16%
CRCW

change
(20%)

16% 4%
CW

qp
CRCW CW CW

q2
If the gametes come together at random, the genotype
frequencies of this generation are in Hardy-Weinberg equilibrium:

64% CRCR, 32% CRCW, and 4% CWCW

Gametes of the next generation:

64% CR from 16% CR from 80% CR = 0.8 = p
+ CRCW homozygotes
=
CRCR homozygotes
4% CW from 16% CW from 20% CW = 0.2 = q
+ CRCW heterozygotes
=
CWCW homozygotes
With random mating, these gametes will result in the same
mix of plants in the next generation:

Figure 23.5 64% CRCR, 32% CRCW and 4% CWCW plants

Conditions for Hardy-

Weinberg Equilibrium
฀ ฀
 If p and q represent the relative frequencies of the only two  The Hardy-Weinberg theorem
possible alleles in a population at a particular locus, then  Describes a hypothetical population
 p2 + 2pq + q2 = 1  In real populations
 And p2 and q2 represent the frequencies of the homozygous  Allele and genotype frequencies do change over time
genotypes and 2pq represents the frequency of the
heterozygous genotype

p+q=1
 Population Genetics and
Human Health
฀ ฀
 We can use the Hardy-Weinberg equation
 To estimate the percentage of the human population carrying
the allele for an inherited disease
Activity:
฀ ฀
Problem # 2: Within a population of butterflies, the color brown (B) is dominant
over the color white (b). And, 40% of all butterflies are white. Given this simple
information, which is something that is very likely to be on an exam, calculate the
following:
a. The percentage of butterflies in the population that are heterozygous.
b. The frequency of homozygous dominant individuals.
Problem # 3: A very large population of randomly-mating laboratory mice
contains 35% white mice. White coloring is caused by the double recessive
genotype, "aa". Calculate allelic and genotypic frequencies for this population.
Practice on your own
฀ ฀
 2. a. Walking through the forest, you find a large population of
toadstools. From your extensive knowledge of the kingdom
fungi, you know that the allele for being spotted (S) is
dominant over the allele for being plain (s). In this population
of 1007, you find 14 toadstools that are not spotted. What are
the allele frequencies?
 b. In a different forest, you find a somewhat smaller population
of 548. Through genetic testing, you determine that there are
308 homozygous spotted, 206 heterozygous, and 34
homozygous plain toadstools. (i) Is this what you expected? (ii)
If not, what are the allele frequencies of this population?
CAUSES OF
MICROEVOLUTION
฀ ฀
 Concept 23.2: Mutation and sexual recombination
produce the variation that makes evolution possible
 Two processes, mutation and sexual recombination
 Produce the variation in gene pools that contributes to
differences among individuals
Class Activity:
 PROBLEM #1. You have sampled a population in which you know that the percentage
of the homozygous recessive genotype (aa) is 36%. Using that 36%, calculate the
following:
A. The frequency of the "aa" genotype.
B. The frequency of the "a" allele.
C. frequency of the "A" allele.
D. The frequencies of the genotypes "AA" and "Aa."
E. The frequencies of the two possible phenotypes if "A" is completely dominant over "a."
Hardy Weinberg Practice
using Chi square
 1. a. In a certain population of newts, being poisonous (P) is
dominant over not being poisonous (p). You count 200 newts,
and 8 are not poisonous. What are the allele frequencies of the
parent population?
 b. Fifty newts are washed downstream after a big storm and
colonize a new pond. What do you expect the frequency and
number of each genotype to be?
 c. You count the new population of newts and find 21
homozygous poisonous newts, 23 heterozygous poisonous
newts, and 6 homozygous non-poisonous newts. (i) Is this what
you expect? (Test using Chi square). (ii) If it is not, what are the
new allele frequencies?
Conditions of
Microevolution
 The five conditions for non-evolving populations are
rarely met in nature
 A locus with 2 or more alleles will be in Hardy-Weinberg
Equilibrium if these 5 conditions are met:
 Extremely large population size (many individuals in a pop)
 No gene flow ( no movement of individuals from pop. To
pop.)
 No mutations (no biochemical change in DNA that produces
new alleles)
 Random mating ( individuals mate at random)
 No natural selection ( differential genotypes have equal
fitness)
Mutation
 Are changes in the nucleotide sequence of DNA
 Cause new genes and alleles to arise
Figure 23.6
 Mutations That Alter
Point Mutations
Gene Number or Sequence
฀ ฀
 A point mutation  Chromosomal mutations that affect many loci
 Is a change in one base in a gene  Are almost certain to be harmful
 Can have a significant impact on phenotype  May be neutral and even beneficial
 Is usually harmless, but may have an adaptive impact

Mutation Rates
฀ ฀
 Gene duplication  Mutation rates
 Duplicates chromosome segments  Tend to be low in animals and plants
 Average about one mutation in every 100,000 genes per
generation
 Are more rapid in microorganisms

Concept 23.3: Natural selection, genetic drift, and gene flow can alter a

Sexual Recombination population’s genetic composition

฀ ฀
 Three major factors alter allele frequencies and bring about
 In sexually reproducing populations, sexual recombination
most evolutionary change
 Is far more important than mutation in producing the genetic
differences that make adaptation possible  Natural selection
 Genetic drift
 Gene flow

Natural Selection Genetic Drift

฀ ฀
 Differential success in reproduction  Statistically, the smaller a sample
 The greater the chance of deviation from a predicted result
 Results in certain alleles being passed to the next generation
in greater proportions
 The disproportion of results in a small sample is known as
sampling error. It is an important factor in the genetics of
small populations.
 Genetic drift
 Describes how allele frequencies can fluctuate unpredictably The Bottleneck Effect
฀  In the bottleneck effect
฀
from one generation to the next

 Tends to reduce genetic variation  A sudden change in the environment may drastically reduce
the size of a population
 The gene pool may no longer be reflective of the original
CRCR CRCR CWCW CRCR CRCR population’s gene pool
CRCW Only 5 of CRCW Only 2 of CRCR CRCR
10 plants 10 plants
leave leave
CWCW CRCR offspring CWCW offspring CRCR CRCR
CRCR

CRCW CRCW CRCR CRCR

CRCR CRCW CWCW CRCR CRCR (a) Shaking just a few marbles through the
narrow neck of a bottle is analogous to a
CRCR CRCW CRCW CRCW CRCR CRCR drastic reduction in the size of a population
after some environmental disaster. By chance,
Generation 1 Generation 2 Generation 3 Bottlenecking
blue marbles are over-represented in the new Original Surviving
p (frequency of CR) = 0.7 p = 0.5 p = 1.0
q (frequency of CW) = 0.3 q = 0.5 q = 0.0 population and gold marbles are absent. population event population

Figure 23.7 Figure 23.8 A

 Understanding the bottleneck effect The Founder Effect

฀ ฀
 Can increase understanding of how human activity affects  The founder effect
other species  Occurs when a few individuals become isolated from a
larger population
 Can affect allele frequencies in a population

(b) Similarly, bottlenecking a population

of organisms tends to reduce genetic
variation, as in these northern
elephant seals in California that were
once hunted nearly to extinction.

Figure 23.8 B

Gene Flow
฀ ฀
 Concept 23.4: Natural selection is the primary mechanism
 Gene flow
of adaptive evolution
 Causes a population to gain or lose alleles
 Natural selection
 Results from the movement of fertile individuals or gametes
 Accumulates and maintains favorable genotypes in a
 Tends to reduce differences between populations over time
population

Variation Within a
Genetic Variation
Population
 Genetic variation
฀
 Occurs in individuals in populations of all species
฀
 Both discrete and quantitative characters
 Is not always heritable  Contribute to variation within a population

(a) Map butterflies that (b) Map butterflies that

emerge in spring: emerge in late summer:
orange and brown black and white
Figure 23.9 A, B
 POLYMORPHOSM
฀  Phenotypic polymorphism
฀
 Discrete characters
 Can be classified on an either-or basis  Describes a population in which two or more distinct
morphs for a character are each represented in high enough
 Quantitative characters
frequencies to be readily noticeable
 Vary along a continuum within a population
 Genetic polymorphisms
 Are the heritable components of characters that occur along a
continuum in a population

MEASURING GENETIC Variation Between

VARIATION Populations
 Population geneticists ฀  Most species exhibit
฀
 Measure the number of polymorphisms in a population by geographic variation
determining the amount of heterozygosity at the gene level 1 2.4 3.14 5.18 6 7.15

 Differences between gene

and the molecular level
pools of separate 8.11 9.12 10.16 13.17 19 XX

 Average heterozygosity populations or population

 Measures the average percent of loci that are heterozygous in subgroups
a population

1 2.19 3.8 4.16 5.14 6.7

9.10 11.12 13.17 15.18 XX

Figure 23.10

 Some examples of geographic variation occur as a cline, A Closer Look at

which is a graded change in a trait along a geographic axis
Natural Selection
฀ Heights of yarrow plants grown in common garden ฀
 From the range of variations available in a population
 Natural selection increases the frequencies of certain
Mean height (cm)

EXPERIMENT Researchers observed that the average size

of yarrow plants (Achillea) growing on the slopes of the Sierra
genotypes, fitting organisms to their environment over
Nevada mountains gradually decreases with increasing
elevation. To eliminate the effect of environmental differences
generations
at different elevations, researchers collected seeds
from various altitudes and planted them in a common
garden. They then measured the heights of the
resulting plants.

RESULTS The average plant sizes in the common

garden were inversely correlated with the altitudes at
Atitude (m)

which the seeds were collected, although the height

differences were less than in the plants’ natural
environments.
Sierra Nevada Great Basin
Range Plateau

CONCLUSION The lesser but still measurable clinal variation

in yarrow plants grown at a common elevation demonstrates the Seed collection sites
role of genetic as well as environmental differences.

Figure 23.11

Evolutionary Fitness
฀
 The phrases “struggle for existence” and “survival of the  Fitness ฀
fittest”  Is the contribution an individual makes to the gene pool of
the next generation, relative to the contributions of other
 Are commonly used to describe natural selection
individuals
 Can be misleading
 Relative fitness
 Reproductive success
 Is the contribution of a genotype to the next generation as
 Is generally more subtle and depends on many factors compared to the contributions of alternative genotypes for
the same locus
 Directional, Disruptive,
and Stabilizing Selection
฀  Directional selection
฀
 Selection
 Favors certain genotypes by acting on the phenotypes of  Favors individuals at one end of the phenotypic range
certain organisms  Disruptive selection
 Three modes of selection are  Favors individuals at both extremes of the phenotypic range
 Directional  Stabilizing selection
 Disruptive (diversifying)  Favors intermediate variants and acts against extreme
 Stabilizing phenotypes

 The three modes of selection The Preservation of

Genetic Variation
฀ Original population
฀
 Various mechanisms help to preserve genetic variation in a
population
Original Evolved Phenotypes (fur color)
population population

(a) Directional selection shifts the overall
makeup of the population by favoring
variants at one extreme of the
distribution. In this case, darker mice are
favored because they live among dark
rocks and a darker fur color conceals them
from predators.
(b) Disruptive selection favors variants
at both ends of the distribution. These
mice have colonized a patchy habitat
made up of light and dark rocks, with the
result that mice of an intermediate color are
at a disadvantage.
(c) Stabilizing selection removes
extreme variants from the population
and preserves intermediate types. If
the environment consists of rocks of
an intermediate color, both light and
dark mice will be selected against.

Fig 23.12 A–C

Diploidy Balancing Selection

฀ ฀
 Diploidy
 Balancing selection
 Maintains genetic variation in the form of hidden recessive
 Occurs when natural selection maintains stable frequencies
alleles
of two or more phenotypic forms in a population
 Leads to a state called balanced polymorphism

Heterozygote
Advantage
฀  The sickle-cell allele ฀
 Some individuals who are heterozygous at a particular  Causes mutations in hemoglobin but also confers malaria
locus resistance
 Have greater fitness than homozygotes  Exemplifies the heterozygote advantage
 Natural selection
 Will tend to maintain two or more alleles at that locus

Frequencies of the
sickle-cell allele
0–2.5%
2.5–5.0%
Distribution of 5.0–7.5%
malaria caused by 7.5–10.0%
Plasmodium falciparum 10.0–12.5%
(a protozoan)
>12.5%
Figure 23.13
  An example of frequency-dependent selection

฀ ฀ On pecking a moth image

the blue jay receives a
food reward. If the bird
Parental population sample
 Frequency-Dependent Selection does not detect a moth
on either screen, it pecks
the green circle to continue
 In frequency-dependent selection to a new set of images (a
new feeding opportunity).
 The fitness of any morph declines if it becomes too common
in the population

Phenotypic diversity
Experimental group sample 0.06

0.05

0.04

Frequency-
0.03
independent control

0.02
0 20 40 60 80 100
Generation number
Plain background Patterned background
Figure 23.14

Neutral Variation Sexual Selection

 Neutral variation
฀ ฀
 Sexual selection
 Is genetic variation that appears to confer no selective
 Is natural selection for mating success
advantage
 Can result in sexual dimorphism, marked differences
between the sexes in secondary sexual characteristics

Intersexual selection
฀ ฀
 Intrasexual selection  Occurs when individuals of one sex (usually females) are
 Is a direct competition among individuals of one sex for choosy in selecting their mates from individuals of the other
mates of the opposite sex sex
 May depend on the showiness of the male’s appearance

Figure 23.15

The Evolutionary Enigma of

Sexual Reproduction
 Sexual reproduction ฀ ฀
 Produces fewer reproductive offspring than asexual  If sexual reproduction is a handicap, why has it persisted?
reproduction, a so-called reproductive handicap  It produces genetic variation that may aid in disease
resistance
Asexual reproduction Sexual reproduction
Female Generation 1
Female

Generation 2

Male

Generation 3

Generation 4

Figure 23.16
 Why Natural Selection Cannot Fashion Perfect
Organisms Activity
฀ ฀
 Form a group of 3 students. Your group has been asked to study two
recently discovered fish species, both found in waters off a small island.
 Evolution is limited by historical constraints The two species are very similar in general structure, but there are two key
differences between them. In species A, males are brightly patterned with
 Adaptations are often compromises blue, red, and purple scales, whereas the females are drab, and males are
 Chance and natural selection interact much larger than females. In species B, males and females are the same
size, and both are a dull brown color that blends in with the sandy bottom.
 Selection can only edit existing variations
 You’ve decided to do an in-depth study of these two species, but to get your
research grant, you must first develop some predictions about what you’ll
find.
 What differences in behavior do you predict (list as many as you can think
of)?
 How might the evolutionary histories of the two species differ?
 Discuss these questions, and submit a brief report of your answer.