BYU11102 Organisms to Ecosystems.

Section B: Evolution
Selection, adaptation and the modern synthesis
Trevor Hodkinson

Darwin and son Erasmus Wallace’s flying frog

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A new era of biology began in 1859 when Charles Darwin
published The Origin of Species

The Origin of Species focused biologists’ attention on the

great diversity of organisms and their origins

Darwin never used the word

‘evolution’ in the first edition of The
Origin of Species

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 Ideas from The Origin of Species

§ Darwin explained three broad

observations:
§ The unity of life (life has
common/shared
characteristics; descent with
modification; tree of life)
§ The diversity of life (natural
selection and speciation)
§ Adaptation (the ways
organisms become suited to
life in their environments)

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 Unity of life
§ Darwin noted that current species
are descendants of ancestral
species
All organisms are related through
descent from an ancestor that lived in
the remote past

Evolution can be viewed as both a Darwin’s sketch 1837

pattern and a process

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 Unity of life and diversity of life
In the Darwinian view, the history of life is like a tree with branches
representing life’s diversity
Diversity of life can be explained byReptiles Darwin’s phrase descent with
(including
modification andOTHERthe
TETRA-
tree of life dinosaurs and birds)
Dimetrodon
Articular
Quadrate
Dentary
Squamosal
†
PODS

Synapsids
Early cynodont (260 mya)
Tree of life

Therapsids
Cynodonts
† Very
late (non-
mammalian) Temporal
(phylogeny) cynodonts
Mammals
fenestra
(partial view)

Hinge
Synapsid (300 mya)

Later cynodont (220 mya)

Temporal
fenestra

Hinge Original hinge

New hinge
Therapsid (280 mya)

Very late cynodont (195 mya)

Temporal
fenestra

Hinge Hinge
Canine tooth

http://tolweb.org/tree/phylogeny.html Descent with modification

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 Darwin explained adaptation (adaptive evolution)

Dead leaf moth

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 Juvenile stage (caterpillar)
of the dead-leaf moth

Natural selection is the

only mechanism that
consistently causes
adaptive evolution

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Education Ltd. Dublin, The University of Dublin
College
Leafy seadragon (Phycodurus) & pygmy seahorse (Hippocampus)

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The Darwinian revolution challenged traditional views
of a young Earth inhabited by unchanging species

§ The Greek philosopher Aristotle

viewed species as fixed and
arranged them on a scala
naturae
§ The Old Testament holds that
species were individually
designed by God and therefore
perfect and unchanging
§ God is a powerful enduring
meme
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 Darwin’s ideas had1809
deep historical roots
Lamarck publishes his
hypothesis of evolution.

1798 Sketch of a flying

Malthus publishes 1812 frog by Wallace
‘‘Essay on the Principle Cuvier publishes his
of Population.” extensive studies of
1858
vertebrate fossils.
While studying species in the
1795 Malay Archipelago, Wallace
Hutton proposes 1830 (shown above in 1848) sends
his principle of Lyell publishes Darwin his hypothesis of
gradualism. Principles of Geology. natural selection.

1790 1870
1809 1831–1836 1859
Charles Darwin Darwin travels The Origin of
is born. around the world Species is published.
on HMS Beagle. 1844
Darwin writes his
essay on descent
Darwin saw with modification.
marine iguanas
in the
Galápagos
Islands.

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 Lamarck’s (1809) hypothesis of
evolution

French biologist Jean-Baptiste

de Lamarck hypothesized that
species evolve through use
and disuse of body parts and
the inheritance of acquired
characteristics

Mechanisms are unsupported by

evidence but he was one of the
earliest to seek explanation for
biological evolution

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College
 Descent with modification by natural selection
explains the adaptations of organisms and the unity
and diversity of life

The Voyage of the Beagle

§ During his travels on the Beagle, Darwin collected
specimens of South American plants and animals
§ He observed that fossils resembled living species from
the same region, and living species resembled other
species from nearby regions
§ He experienced an earthquake in Chile and observed the
uplift of rocks

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 The Voyage of the Beagle
December 1831–October 1836

Darwin in HMS Beagle in port

1840, after
his return
from the Great
voyage Britain EUROPE
NORTH
AMERICA
ATLANTIC
OCEAN
The PACIFIC AFRICA PACIFIC
Galápagos Pinta OCEAN
Islands OCEAN
Genovesa SOUTH Equator Malay
Marchena Equator AMERICA Archipelago
Santiago
Brazil
Andes Mtns.

Chile AUSTRALIA
Fernandina
Isabela Santa Santa PACIFIC Cape of
Cruz Fe San
OCEAN Good Hope
Cristóbal
0 20 40 Argentina Tasmania
Floreana Española
Kilometers
Cape Horn New
Zealand

He hypothesized that species from South America had

colonized the Galápagos and speciated on the islands
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 Darwin’s focus on adaptation

How did this flightless

bird come to live on the
isolated Galápagos
Islands?

§ Darwin perceived adaptation to the environment and the

origin of new species as closely related processes

§ From studies made years after Darwin’s voyage,

biologists have concluded that this is what happened to
the Galápagos finches
 Cactus-eater Insect-eater

Beak variation

Seed-eater
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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

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 Artificial selection generated all these
vegetables from the same common ancestor

Cabbage

Selection for
apical (tip) bud

Brussels Selection for Selection Broccoli

sprouts axillary (side) for flowers
buds and stems

Selection
Selection for stems
for leaves
Kale Wild mustard Kohlrabi

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 Mechanism of natural selection
§ Darwin drew two inferences from two observations

§ Observation 1: Members of a population often vary in

their inherited traits (genes)

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 Puffball fungus
• Observation 2: All species
can produce more offspring Spore
than the environment can cloud
support, and many of these
offspring fail to survive and
reproduce

Darwin was influenced by Thomas

Malthus, who noted the potential for
the human population to increase
faster than food supplies and other
resources leading to famine

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§ Inference 1 Some individuals in a given environment
leave more offspring than other individuals - because of
their inherited traits (that give them a higher probability
of surviving and reproducing)

§ Inference 2 This unequal ability of individuals to survive

and reproduce will lead to the accumulation of favorable
traits (genes) in the population over generations
(adaptation)

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 Therefore natural selection changes populations over time
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 Examples

Wikipedia
Peppered moth in
the industrial
revolution evolved
dark (melanic) form
for camouflage

https://evolution.berkeley.edu/evolibrary/art
icle/bergstrom_02
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Natural selection - explains close fit of organisms
to environment (adaptation)
nothing else can explain such a match

Echinocactus

E.g.
Cacti are succulent
CAM photosynthesis
Lost leaves, gained spines
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§ Note that individuals do not evolve; populations evolve
over time
§ Natural selection can only increase or decrease heritable
traits that vary in a population
§ Adaptations vary with different environments

But Darwin did not know what was inherited, so it

was an incomplete theory – until ‘the modern
synthesis’ (the update since then with new genetic
knowledge)
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 Mendel and the modern synthesis

1822-84
§ Darwin was not aware of Mendel’s work and did not
know that the unit of inheritance was the gene
§ Evolution acts primarily on the gene
Gregor Mendel worked out the units of inheritance ‘factors’ now
called genes
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 BB bb

Mendel’s law of
segregating genes
(alleles)

Bb Purple dominant over

recessive white pea flowers

Bb x Bb

because

3:1 ratio

Mendel’s discovery led to recognition of genes

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 Modern synthesis incorporated genetic knowledge into
evolutionary theory – evolution acts on variants of genes
(they are the heritable unit of selection)
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Plant breeders use this knowledge to manipulate useful traits
via artificial selection

If you want a true

breeding white you
can just take white
types from a
population and it will
only have white allele
type (bb).

For true breeding

purple need to select
for homozygote BB
(will take longer)
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Sometimes genes show incomplete dominance (pink)

Figure 23.8c

R R
C C

W W
C C

R W
C C

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New knowledge of genetics led to recognition of
other major factors that can alter allele
frequencies in populations (microevolution)
Natural selection – alters allele frequencies over
generations by differences in fitness (reproductive
success; contribution to the genepool of the next
generation)

Genetic drift – alters allele frequencies via

stochastic (random) processes

Geneflow (migration) – alters allele frequencies via

movement of alleles among populations
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Genetic drift can cause evolution via stochastic
processes (random sampling)
5 plants 2 plants
leave leave
offspring offspring
CRCR CRCW CRCR
CRCW CWCW CRCR
CRCR CRCR
CRCR
CRCR CRCR
CRCW
CRCR CRCR
CWCW
CWCW CRCR CRCR
CRCW CWCW CRCR
CRCR
CRCW CRCW CRCR
CRCR CRCR
CRCW CRCW CRCR

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

This is microevolution (population genetic level)

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 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 (like
the selection of red in the previous example)

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 The Bottleneck Effect
§ The related bottleneck effect occurs when there is a
drastic reduction in population size due to a sudden
change in the environment (e.g. overhunting)
§ The resulting gene pool may no longer be reflective of
the original population’s gene pool
§ If the population remains small, it may be further
affected by genetic drift

This is an important consideration

in conservation genetics

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 Bottleneck effect is similar to founder effect but caused by
different factors

Original Bottlenecking Surviving

population event population

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 Gene flow (migration)
§ 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/sperm)
§ Gene flow tends to reduce variation among populations
over time (but can also introduce favourable traits)

Plants can’t walk but they can move seeds or their pollen can move via wind or animal pollination
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§ Gene flow can increase the fitness of a population
Consider, for example, the spread of alleles for resistance to
insecticides (to control pests)
§ E.g. Insecticides have been used to target mosquitoes that
carry West Nile virus and malaria
§ Alleles have evolved in some populations that confer
insecticide resistance to these mosquitoes
§ The flow of insecticide resistance alleles into a population
can cause an increase in fitness (they are no longer killed)

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 Hardy Weinberg theory explains frequencies of
alleles in populations and their genotypes
– Hardy Weinberg can be used to show that
populations are evolving if they don’t fit this

If p (R) =0.5 and q (W) = 0.5.

Then 0.25+0.5+0.25 =1
In absence of selection, mutation, drift and migration populations
fit this equation. If frequencies of alleles don’t fit this model then
the population is evolving in some way.
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 Different modes of natural selection

Selection for extremes Selection against extremes

Selection towards a
particular type

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Balancing selection maintains variation at particular
genetic loci (instead of it being removed by natural selection)
§ Frequency-dependent
selection (fitness depends
on frequency of trait to
other phenotypes)

§ Heterozygote advantage
(heterozygote at given
locus is fitter). E.g. sickle
cell allele (next page)

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Heterozygote has
an advantage in
some populations.
Esp. where malaria
is common. So
populations evolve
to have this allele.

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 Conclusions
§ Darwin’s theory of natural selection (descent with
modification) explains evolution
§ The modern synthesis incorporates genetics and
evolutionary theory
§ Evolution explains unity of life, diversity of life &
adaptation

Next up - species
and speciation
processes
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 Key Features of Natural Selection

§ Individuals with certain heritable traits survive and

reproduce at a higher rate than other individuals
§ Natural selection increases the frequency of adaptations
that are favorable in a given environment
§ If an environment changes over time, natural selection
may result in adaptation to these new conditions and
may give rise to new species

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§ Gene flow can affect adaptation to local environments
§ For example, variation in banding pattern in Lake Erie
water snakes represents adaptation to mainland and
island habitats
§ Most snakes on the mainland are strongly banded; most
island snakes are unbanded or intermediate
§ Unbanded snakes are better camouflaged in island
habitats, but migration of banded snakes from mainland to
island populations maintains alleles for the banded pattern
in island populations

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 Unbanded
N. sipedon
ONTARIO (Pattern D)
Detroit

LAKE ERIE Pelee

Island

OHIO Cleveland Middle

Island
BASS
ISLANDS
LAKE ERIE

Kelleys
Island
OHIO
Banded N. sipedon
(Pattern C) 5 km

100
Percentage of individuals

80

60
40

20
0
A B C D A B C D A B C D
Ohio mainland Islands Ontario mainland
Banding patterns in snake populations
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