Mechanisms

of Evolution
Bui Hong Thuy, Ph.D.
School of Biotechnology,
International University
Email: [email protected]

1
 OUTLINES
1. Descent with Modification:
A Darwinian View of life

2. The Evolution of
Populations

3. The Origin of Species

2
 OUTLINES
1. Descent with Modification:
A Darwinian View of life

2. The Evolution of
Populations

3. The Origin of Species

3
 Overview: Endless Forms
Most Beautiful

 A new era of biology began in 1859 when

Charles Darwin published The Origin of
Species which focused biologists’ attention on
the great diversity of organisms.
 Darwin noted that current species are
descendants of ancestral species.
 Evolution can be defined by Darwin’s phrase:
descent with modification.

4
 Intellectuals / Ideas in Darwin’s Era

Linnaeus (classification)
Hutton (gradual geologic change)
Lamarck (species can change)
Malthus (population limits)
Cuvier (fossils, extinction)
Lyell (modern geology)
Darwin (evolution, natural selection)
Wallace (evolution, natural selection)
American Revolution French Revolution U.S. Civil War
1750 1800 1850 1900
1795 Hutton proposes his theory of gradualism.
1798 Malthus publishes “Essay on the Principle of Population.”
1809 Lamarck publishes his hypothesis of evolution.
1830 Lyell publishes Principles of Geology.
1831–1836 Darwin travels around the world on HMS Beagle.
1837 Darwin begins his notebooks.
1844 Darwin writes essay on descent with modification.
1858 Wallace sends his hypothesis to Darwin.
1859 The Origin of Species is published.

5
Scientists …

 Carolus Linnaeus was the founder of

taxonomy, the branch of biology concerned
with classifying organisms.
 Paleontology, the study of fossils, was largely
developed by French scientist Georges Cuvier.
 The study of fossils helped to lay the
groundwork for Darwin’s ideas.
 Fossils are remains or traces of organisms from
the past, usually found in sedimentary rock,
which appears in layers or strata.

6
 Formation of sedimentary
strata with fossils

Layers of deposited
sediment

Younger stratum
with more recent
fossils

Older stratum
with older fossils

7
 Lamarck: Use & Disuse …
Darwin: Natural Selection ….

 Lamarck hypothesized that species evolve

through use and disuse of body parts and the
inheritance of acquired characteristics.
 During his travels on the Beagle, Darwin
collected specimens of South American plants
and animals. He observed adaptations of
plants and animals that inhabited many
diverse environments.

8
 Darwin’s Voyage on the Beagle

GREAT
BRITAIN EUROPE
NORTH
AMERICA
ATLANTIC
OCEAN
The AFRICA
Galápagos Pinta
Islands
Genovesa
Marchena Equator
SOUTH
Santiago AMERICA
Daphne
Islands
Pinzón AUSTRALIA
Fernandina PACIFIC
Isabela Santa OCEAN Cape of
Santa
Cruz
Fe San Good Hope
Cristobal
Florenza Tasmania
Española
Cape Horn New
Zealand
Tierra del Fuego

9
 Darwin’s Focus on Adaptation

 Darwin perceived adaptation to the

environment and the origin of new species
as closely related processes.
 Recent biologists have concluded that
speciation is indeed what happened to the
Galápagos finches.

10
 Speciation of Galapagos Finches

(a) Cactus-eater (c) Seed-eater

(b) Insect-eater
11
 In 1844, Darwin wrote an essay on
the origin of species and natural
selection but did not introduce his
theory publicly, anticipating an
uproar
 In June 1858, Darwin received a
manuscript from Alfred Wallace, who
had developed a theory of natural
selection similar to Darwin’s

 Darwin published his first

version of the book On the
Origin of Species by Means
of Natural Selection in 1859
Charles Darwin
(1809-1882) 12
 The Origin of Species
 Darwin developed two main ideas:
– Descent with modification explains
life’s unity and diversity.
– Natural selection is a cause of adaptive
evolution.

13
 Descent
Hyracoidea
(Hyraxes)

Sirenia
(Manatees With
Modification
and relatives)
Moeritherium

Barytherium
This evolutionary tree of
Deinotherium
elephants and their relatives is
based mainly on fossils-their
Mammut
anatomy. order of appearance
in strata. and geographic
Platybelodon distribution Note that most
branches of descent ended in
Stegodon extinction. {Time line not to
scale.}
Mammuthus

Based on the tree shown

Elephas maximus
(Asia) here. approximately when did
the most recent ancestor
Loxodonta
africana shared by .. Mammuthus
(Africa)
Loxodonta cyclotis
(woolly mammoths), Asian
(Africa) elephants, and African
34 24 5.5 2 104 0
elephants live.
Millions of years ago Years ago
14
 Artificial Selection, Natural
Selection, and Adaptation
 Darwin noted that humans have modified
other species by selecting and breeding
individuals with desired traits, a process
called artificial selection.
 Darwin then described four observations of
nature and from these drew two inferences.

15
Observation #1: Members of a population
often vary greatly in their traits.

Variation in a population
16
 Observation #2: Traits are inherited from
parents to offspring.
 Observation #3: All species are capable
of producing more offspring than the
environment can support.
 Observation #4: Overproduction leads to
competition for food or other resources.
The individuals best adapted to their
environment will survive and reproduce.

17
 Descent With Modification:
 Inference #1: Individuals whose inherited
traits give them a higher probability of
surviving and reproducing in a given
environment tend to leave more offspring than
other individuals.
 Inference #2: This unequal ability of individuals
to survive and reproduce will lead to the
accumulation of favorable traits in the
population over generations.

18
 Darwin’s theory: evolution occurs
by natural selection
Darwin’s theory of evolution was based on four general
observations
– Overproduction – each species produces more offspring than
will survive to maturity
– Variation – individuals in a population vary, and some of the
variation is heritable (this was expanded by others later, as
genetics came to be understood)
– Competition – there is competition among the individuals of a
population for limited resources (struggle for existence)
– Differential reproductive success – individuals that possess
more favorable characteristics (in the pool of variation) are
more likely to survive and reproduce; those with less
favorable characteristics are less likely to survive and
reproduce
Natural selection will produce a population of
individuals more suited to their environment through time
19
 Natural Selection: A Summary

 Survival of the Fittest: Individuals with

certain heritable adaptive characteristics
survive and reproduce at a higher rate than
other individuals.
 Natural selection increases the adaptation
of organisms to their environment over time.
 Speciation: If an environment changes over
time, natural selection may result in adaptation
to these new conditions and may give rise to
new species.

20
Predation and Coloration in Guppies :
Scientific Inquiry in Natural Selection

 John Endler has studied the effects of predators

on wild guppy populations.
 Brightly colored males are more attractive to
females.
 However, brightly colored males are more
vulnerable to predation.
 Guppy populations in pools with fewer predators
had more brightly colored males.

21
 EXPERIMENT
Natural
Selection Predator: Killifish; preys
mainly on juvenile
Experimental
guppies (which do not
transplant of Pools with
express the color genes) guppies killifish,
but no
Guppies: Adult males have guppies prior
brighter colors than those to transplant
in “pike-cichlid pools”

Can predation result

in natural selection
Predator: Pike-cichlid; preys mainly on adult guppies
for color patterns in
Guppies: Adult males are more drab in color
guppies? than those in “killifish pools”

RESULTS

12 12

colored spots
10 10

Number of
8 8
6 6
4 4
2 2
0 0
Source Transplanted Source Transplanted
population population population population
22
 Fossil Evidence 0
of Change Over Time 2
4
4 6 4 Bristolia insolens
8 3 Bristolia bristolensis
Fossil evidence of evolution
10
in a group of trilobites.
These fossils are just a few in 3 12
2 Bristolia harringtoni
a series discovered in the 14
Latham Shale bed, which was 16
deposited between 513 and 18 1 Bristolia mohavensis
512 million years ago. The 3
sequence shows change over
time in the location and 2

angle of the spines of the head

shield (the area marked by red
dots).

1 Latham Shale dig site, San

Bernardino County, California
23
 Paleontologists
study fossils of
possible
transitional forms:

Whale (a) Pakicetus (terrestrial)

Ancestors

The transition to life in the sea

(b) Rhodocetus (predominantly aquatic)
The hypothesis that whales and
other cetaceans evolved from
terrestrial organisms predicts that
cetacean ancestors were four- Pelvis and
hind limb
legged. Paleontologists have (c) Dorudon (fully aquatic)
unearthed fossils of elctinct
cetaceans that had hind limbs.
This similarity strongly suggests Pelvis and
that cetaceans are most closely hind limb
(d) Balaena
related to this group of land (recent whale ancestor)
24
mammals.
 Anatomical and
Molecular Homologies

 Homology is similarity resulting from

common ancestry.
 Homologous structures are anatomical
resemblances that represent variations on a
structural theme present in a common
ancestor.

25
 Comparative anatomy of related species
Organs or structures that have similar form due to a
common evolutionary origin are called homologous
features
– Example: the similarity between the human arm, the
dolphin's flipper, the bat's wing, and the bird's wing

Humerus

Radius

Ulna

Carpals

Metacarpals

Phalanges
Human Cat Whale Bat
26
Related species have similar patterns of development
Very young embryos of reptiles, birds, mammals, and
humans are indistinguishable
Studies of developmental biology are revealing the
common genetic basis for such similarities.
“Evo/devo study is one of the hottest fields in biology
today (evolutionary developmental biology--examines
the relationships between embryonic development and
evolutionary changes)

Pharyngeal
pouches

Post-anal
tail

Chick embryo (LM) Human embryo 27

 Homologies and “Tree Thinking”

 The Darwinian concept of an evolutionary

tree of life can explain homologies.
 Evolutionary trees are hypotheses about the
relationships among different groups.
 Evolutionary trees can be made using
different types of data, for example,
anatomical and DNA sequence data.

28
Homologies and “Evolutionary Tree”
Branch point
(common ancestor)
Lungfishes

1 Amphibians

2 Mammals
Tetrapod limbs

3 Lizards
Amnion and snakes

4 Crocodiles
Homologous
characteristic 5
Ostriches

Birds
6
Feathers
Hawks and
other birds

29
 Convergent Evolution
 Convergent evolution is the evolution of
similar, or analogous, features in distantly
related groups.

NORTH
Sugar AMERICA
glider
• Analogous traits
arise when groups
independently adapt
AUSTRALIA
to similar
environments in
similar ways.
Flying
squirrel
30
 Darwin’s observations of biogeography, the
geographic distribution of species, formed an
important part of his theory of evolution.
Darwin : Natural Selection
Observations
Individuals in a population Organisms produce more
vary in their heritable offspring than the
characteristics. environment can support.

Inferences
Individuals that are well suited
to their environment tend to leave
more offspring than other individuals
and
Over time, favorable traits
accumulate in the population. 31
The central role of evolution
in modern biology
 The modern synthetic theory of evolution is
accepted today by most biologists as:
– a robust and well-supported model
– the central framework for the study of life

 Studies of evolution itself today focus largely on

– the causal processes of evolution, such as:
• the speed of evolutionary change
• the role of chance in evolution
– molecular comparisons between and within
species by
• comparing DNA sequences
• comparing genomes
• comparing proteins and proteomes
32
 OUTLINES
1. Descent with Modification:
A Darwinian View of life

2. The Evolution of
Populations

3. The Origin of Species

33
 Overview: The Smallest Unit
of Evolution
 Natural selection acts on individuals, but
only populations evolve.
 Genetic variations in populations contribute
to evolution.
 Microevolution is a change in allele
frequencies in a population over
generations.
 Two processes, mutation and sexual
reproduction, produce the variation in
gene pools that contributes to differences
among individuals.

34
 Nonheritable (a) (b)
variation
The moth Nemoria
arizonaria owe their
different appearances
to chemicals in their
diets, not to their
genotypes. Caterpillars raised on a diet of oak flowers resembled
the flowers (a), whereas their siblings raised on oak
leaves resembled oak twigs (b).

1.0

The frequency of the Ldh-Bb 0.8

allele for the enzyme lactate
dehydrogenase-B decreases 0.6
in fish from Maine to
Georgia. Individuals with the 0.4
Ldh-Bb allele can swim faster
0.2
in cold water than can
individuals with other alleles. 0
46 44 42 40 38 36 34 32 30
Maine Latitude (°N) Georgia
Cold (6°C) Warm (21°C)
35
 Mutation
 Mutations are changes in the nucleotide
sequence of DNA.
 Mutations cause new genes and alleles
to arise.
 Only mutations in cells that produce
gametes can be passed to offspring.
 A point mutation is a change in one
base in a gene.

36
 The effects of point mutations can vary
– Mutations in noncoding regions of DNA are
often harmless.
– Mutations in a gene might not affect protein
production because of redundancy in the
genetic code.
– Mutations that result in a change in protein
production are often harmful.
– Mutations that result in a change in protein
production can sometimes increase the
fitness of the organism in its environment.

37
 Mutations That Alter Gene /
Chromosome Number or Sequence

 Chromosomal mutations that delete,

disrupt, or rearrange many loci are typically
harmful.
 Mutation rates are low in animals and
plants.
 Mutations rates are often lower in
prokaryotes and higher in viruses.

38
 Sexual Reproduction
 Sexual reproduction can shuffle existing
alleles into new combinations.
 In organisms that reproduce sexually,
recombination of alleles is more important
than mutation in producing the genetic
differences that make adaptation possible.

39
 2.2. Hardy-Weinberg equation tests
whether a sexually reproducing
population is evolving

 A population is a localized group of

individuals (a species in an area) capable of
interbreeding and producing fertile offspring.
 A gene pool consists of all the alleles for all
loci in a population.
 A locus is fixed if all individuals in a
population are homozygous for the same
allele.

40
 The Hardy-Weinberg Theorem
The Hardy-Weinberg theorem describes the
frequencies of genotypes in a population based
on the frequency of occurrence of alleles in the
population that is in a state of genetic
equilibrium (that is, not evolving)
– the usual case for calculations: if allele “A” is
dominant to “a”, and they are the only two alleles
possible at the A-locus, then
• p = freq[A] = the frequency of occurrence of the
A-allele in the population
• q = freq[a] = the frequency of occurrence of the
a-allele in the population
– Then p + q = 1 (following the sum rule for
probability) 41
 Allele associations follow the product rule
for probability, so you multiply to predict the
genotype frequencies
– ( p + q ) x ( p + q ) = p2 + 2 pq + q2
– p2 = frequency of homozygous dominant
genotypes
– 2 pq = frequency of heterozygous genotypes
– q2 = frequency of homozygous recessive
genotypes
– note that ( p + q ) x ( p + q ) = 1 x 1 = 1, so
p2 + 2 pq + q2 = 1

42
43
Hardy-Weinberg Ideal Conditions
 The five conditions for nonevolving
populations are rarely met in nature:
– No mutations
– Random mating
– No natural selection
– Extremely large population
– No gene flow (no migration – no exchange of
alleles with other populations)

44
 Applying the Hardy-Weinberg Principle

• We can assume the locus that causes phenylketonuria

(PKU) is in Hardy-Weinberg equilibrium given that:
– The PKU gene mutation rate is low
– Mate selection is random with respect to whether or not
an individual is a carrier for the PKU allele
– Natural selection can only act on rare homozygous
individuals who do not follow dietary restrictions
– The population is large
– Migration has no effect as many other populations have
similar allele frequencies

45
• The occurrence of PKU is 1 per 10,000
births
– q2 = 0.0001
– q = 0.01
• The frequency of normal alleles is
– p = 1 – q = 1 – 0.01 = 0.99
• The frequency of heterozygotes / carriers is
– 2pq = 2 x 0.99 x 0.01 = 0.0198
– or approximately 2% of the U.S. population.

46
2.3. Natural selection, genetic drift, and gene
flow can alter allele frequencies in a population

• Three major factors alter allele

frequencies and bring about most
evolutionary change:
– Natural selection - nonrandom
– Genetic drift - random
– Gene flow - random

47
 Natural Selection and Genetic Drift

• Natural Selection: Differential success in

reproduction results in certain alleles being passed
to the next generation in greater proportions by the
more fit individuals.
• Genetic drift: describes how allele frequencies
fluctuate randomly from one generation to the next.
• The smaller a sample, the greater the chance of
deviation from a predicted result.
• Genetic drift tends to reduce genetic variation
through losses of alleles.

48
Genetic Drift

CR CR CR CR CW CW CR CR CR CR

CR CW CR CW CR CR CR CR

CW CW CR CR CR CR CW CW CR CR CR CR

CR CW CR CW CR CR CR CR

CR CR CR CW CW CW CR CR CR CR

CR CR CR CW CR CW CR CW CR CR CR CR

Generation 1 Generation 2 Generation 3

p (frequency of CR) = 0.7 p = 0.5 p = 1.0
q (frequency of CW ) = 0.3 q = 0.5 q = 0.0
Only 5 of 10 Only 2 of 10
plants leave plants leave
offspring offspring 49
 Genetic Drift: The Founder Effect

 The founder effect occurs when a few

individuals become isolated from a larger
population.
 Allele frequencies in the small founder
population can be different from those in the
larger parent population.

50
 Genetic Drift: The Bottleneck Effect
 The bottleneck effect is a sudden reduction in
population size due to a change in the
environment, such as a natural disaster.
 The resulting gene pool may no longer be
reflective of the original population’s gene pool.

Original Bottlenecking Surviving

population event population
51
 Effects of Genetic Drift: A Summary

1. Genetic drift is significant in small

populations.
2. Genetic drift causes allele frequencies to
change at random.
3. Genetic drift can lead to a loss of genetic
variation within populations.
4. Genetic drift can cause harmful alleles to
become fixed.

52
 Gene Flow:
Immigration & Emmigration
 Gene flow consists of the movement of alleles
among populations.
 Alleles can be transferred through the
movement of fertile individuals or gametes (for
example, pollen).
 Gene flow tends to reduce differences
between populations over time.
 Gene flow is more likely than mutation to alter
allele frequencies directly.

53
 Gene flow and human evolution

The migration of people throughout the world has increased gene flow
between populations that once were isolated from one another. The
computer-generated image on this magazine cover illustrates how
gene flow can homogenize the gene pools of such populations.
54
 2.4. Natural selection is the only
mechanism that consistently causes
adaptive evolution

 Only natural selection consistently results

in adaptive evolution.
 Natural selection brings about adaptive
evolution by acting on an organism’s
phenotype.

55
 Natural Selection: Relative Fitness
 The natural selection phrases “struggle for
existence” and “survival of the fittest” are
misleading as they imply direct competition
among individuals.
 Reproductive success is generally more subtle
and depends on many factors.
 Relative fitness is the contribution an individual
makes to the gene pool of the next generation,
relative to the contributions of other individuals.
 Selection favors certain genotypes by acting on
the phenotypes of certain organisms.

56
 Natural Selection
Original
population

Original Evolved
Phenotypes (fur color)
population population

• Directional • Disruptive • Stabilizing

selection favors selection favors selection favors
individuals at one individuals at intermediate
end of the both extremes of variants and acts
phenotypic the phenotypic against extreme
range. range. phenotypes. 57
 The Key Role of Natural
Selection in Adaptive Evolution
• Natural selection increases the frequencies of
alleles that enhance survival and reproduction.
• Adaptive evolution = the match between an
organism and its environment.
Movable bones

(a) Color-changing ability in

cuttlefish (b) Movable jaw
bones in
snakes 58
 Sexual Selection
 Sexual selection is natural selection for mating
success.
 There is intrasexual selection between competing
males, followed by intersexual selection when the
females choose among the showiest males.

59
 OUTLINES
1. Descent with Modification:
A Darwinian View of life

2. The Evolution of
Populations

3. The Origin of Species

60
 Speciation, the origin of new species, is at the
focal point of Darwin’s evolutionary theory.
 Microevolution consists of adaptations that
evolve within a population, confined to one
gene pool.
 Macroevolution refers to evolutionary change
above the species level.

In the Galápagos
Islands Darwin
discovered plants
and animals found
nowhere else on
Earth

61
 3.1. The Biological Species Concept
emphasizes reproductive isolation

 The basis of macroevolution in sexual species

is microevolution coupled with reproductive
isolation
 Biologists compare morphology, physiology,
biochemistry, and DNA sequences when
grouping organisms.

 The biological species concept states that a

species is a group of populations whose
members have the potential to interbreed in
nature and produce viable, fertile offspring;
they do not breed successfully with other
populations.
62
(a) The eastern meadowlark
( left) and the western meadowlark
(right) have similar body shapes
and colorations. They are distinct
biological species because their
songs and other behaviors are (a) Similarity between different species
different enough to prevent
interbreeding

(b) As diverse as we
may be in appearance.
all humans belong to a
single biological species
(Homo sapiens),
defined by our capacity
to interbreed.

(b) Diversity within a species

63
 Macroevolution is essentially the
formation of new species (speciation)
Species “kind of living thing”; the word “species”
is both plural and singular
Relatively easy to define for sexual organisms,
hard for asexual organisms and extinct species
– Biological species concept (for sexual organisms) –
one or more populations whose members are:
• capable of interbreeding
• able to produce fertile offspring
• reproductively isolated from other such groups

64
 Reproductive Isolation =
Barriers to Interbreeding
 Reproductive isolation = means of preventing
gene flow between two species
– for a new species to evolve from an existing
(sexually reproducing) species, there must be a
reproductive isolating mechanism in place

 Reproductive isolating mechanisms can be

classified as either prezygotic or postzygotic
– prezygotic barriers – prevent fertilization (zygote
formation) between gametes from two species
– postzygotic barriers – reproductive isolation after
fertilization has occurred
65
 Reproductive Barriers
Between Species
 Prezygotic barriers block fertilization from
occurring by:
– Impeding different species from attempting to
mate.
– Preventing the successful completion of
mating.
– Hindering fertilization if mating is successful.
Prezygotic barriers maintain reproductive
isolation and include: Temporal, Habitat,
Behavioral, Mechanical, and Gamete
Isolation
 Hybrids are the offspring of crosses between
different species.
66
 Reproductive Barriers
Between Species
Prezygotic Barriers Postzygotic Barriers
Habitat Temporal Behavioral Mechanical Gametic Reduced Reduced Hybrid Hybrid
Isolation Isolation
Isolation Isolation Isolation Hybrid Viability Fertility Breakdown
Individuals Viable,
Mating Fertilization
of attempt fertile
different offspring
species
(a) (c) (e) (f) (g) (h) (i) (l)

(d) (j)

(b)

(k)

67
68
 PreZygotic Reproductive Barriers
 Habitat isolation: Two species encounter each
other rarely, or not at all, because they occupy
different habitats, even though not isolated by
physical barriers.

Water-dwelling Thamnophis Terrestrial Thamnophis

69
PreZygotic Reproductive Barriers

 Temporal Isolation: Species that breed at

different times of the day, different seasons, or
different years cannot mix their gametes.
 Behavioral isolation: Courtship rituals and other
behaviors unique to a species are effective
barriers.
 Mechanical isolation: Morphological differences
can prevent successful mating.
 Gametic isolation: Sperm of one species may not
be able to fertilize eggs of another species.

70
 Mechanical Isolation
Behavioral Isolation The snails’ genital openings
Occurs Without are not aligned, and mating
Appropriate Mating Rituals cannot be completed

Courtship ritual of blue-footed Bradybaena with shells spiraling

boobies in opposite directions

71
 PostZygotic Reproductive Barriers
 Postzygotic barriers prevent the hybrid zygote from
developing into a viable, fertile adult:
– Reduced hybrid viability -- weak offspring
– Reduced hybrid fertility -- sterile offspring
– Hybrid breakdown.

example:
male donkey x female horse
[n=31] [n=32]
generates sterile mule [2n=63]
not an even number, pairing problems
during meiosis

72
 Other Definitions of Species
 The morphological species concept
defines a species by structural features.

 The ecological species concept views a

species in terms of its ecological niche.

 The phylogenetic species concept: defines

a species as the smallest group of individuals
on a phylogenetic tree.

73
 3.2 Speciation can take place with or
without geographic separation
 Speciation can occur in two ways:
– Allopatric speciation: geographic barrier separates
populations.
– Sympatric speciation: no geographic barrier

(a) Allopatric (b) Sympatric

Forms a new species Becomes a new species
isolated from its parent without geographic
population separation
74
Allopatric (“Other Country”) Speciation
Continental Drift Caused Allopatric Speciation
Mantellinae
(Madagascar only):
100 species
Rhacophorinae
(India/Southeast
Asia): 310 species

Other Indian/
Southeast Asian
frogs
100 80 60 40 20 0
1 2 3
Millions of years ago (mya)
1 2 3
India

Madagascar
88 mya 65 mya 56 mya

The frog subfamilies Mantellinae and Rhacophormae diverged when

present-day Madagascar separated from India. The maps show the
movement of Madagascar (red) and India (blue) over time. 75
 Sympatric (“Same Country”)
Speciation
 In sympatric speciation, speciation takes place
in geographically overlapping populations.
 Polyploidy is the presence of extra sets of
chromosomes due to accidents during cell
division.
 An autopolyploid is an individual with more than
two chromosome sets, derived from one species.
 An allopolyploid is a species with multiple sets of
chromosomes derived from different species.
 Polyploidy is common in plants. Many important
crops (oats, cotton, potatoes, tobacco, and wheat)
are polyploids.
76
 Sympatric speciation by autopolyploidy in plants

Gametes
2n = 6 4n = 12 produced 2n 4n
Failure of cell division after are diploid.. Offspring with tetraploid
Chromosome duplication gives karyotypes may be viable
rise to tetraploid tissue. and fertile.

One mechanism for allopolyploid speciation in plants

Species B Unreduced
2n = 4 gamete Unreduced
with 4 Hybrid gamete
chromosomes with 7 with 7
chromosomes chromosomes
Meiotic
error

Normal Viable fertile

gamete Normal hybrid
Species A n=3 gamete (allopolyploid)
2n = 6 n=3 2n = 10 77
 Allopatric and Sympatric Speciation
May Cause Reproductive Isolation:
A Review
 In allopatric speciation, geographic separation
restricts gene flow between populations.
 Reproductive isolation may then arise by natural
selection, genetic drift, or sexual selection in the
isolated populations.
 Even if contact is restored between populations,
interbreeding is prevented.
 In sympatric speciation, a reproductive barrier
isolates a subset of a population without
geographic separation from the parent species.
 Sympatric speciation can result from polyploidy,
natural selection, or sexual selection.
78
3.3. Hybrid zones provide opportunities to
study factors that cause reproductive isolation
A hybrid zone is a region in which members of
different species mate and produce hybrids
EUROPE

Fire-bellied
toad range
Hybrid zone
Fire-bellied toad,
Yellow-bellied

Allele frequency (log scale)

Bombina bombina
Yellow-bellied toad, toad range
Bombina variegata 0.99

0.9

0.5

0.1

0.01
40 30 20 10 0 10 20
Distance from hybrid zone center (km)

79
 Hybrid Zones over Time
 When closely related species meet in a
hybrid zone, there are three possible
outcomes:
– Reinforcement -- Strengthening of reproductive
barriers reducing gene flow.
– Fusion -- Weakening of reproductive barriers
with eventual fusion into one species.
– Stabilizing -- Continued formation of hybrid
individuals.

80
Hybrid Zones over Time

81
 Studying the Genetics of
Speciation
 The explosion of genomics is enabling
researchers to identify specific genes
involved in some cases of speciation.
 Depending on the species in question,
speciation might require the change of only a
single allele or many alleles.
From Speciation to Macroevolution:
 Macroevolution is the cumulative effect of
many speciation and extinction events.

82
 Knowledge Testing 1
1. Explain Darwin’s observations and inferences
2. Explain why an individual organism cannot
evolve
3. Describe evidence for evolution by natural
selection

83
 Knowledge Testing 2
1. Explain how sexual recombination
generates genetic variability
2. Explain why natural selection is the only
mechanism that consistently produces
adaptive change
3. Explain the role of population size in genetic
drift
4. Apply the Hardy-Weinberg equation to a
population genetics problem

84
 Knowledge Testing 3
1. Define and discuss the limitations of the four
species concepts
2. Describe and provide examples of prezygotic
and postzygotic reproductive barriers.
3. Explain how polyploidy can cause reproductive
isolation

85