Evolution

The Historical context of Darwin's life and ideas

Picture (slide 13)

Carl von Linne (Linnaeus) – Swedish Botanist

 Founder of taxonomy (Linnean)

 Developed the binominal system of naming species, and a system by
which similar species were placed in a genus, similar genera in a
family, similar families in order.
 Based off characteristics
 1st part = genus & 2nd part = species

George Cuvier – French Paleontology

 Developed the study of fossils (paleontology)

 Recognized that fossils were evidence of earlier lie forms. Observed
changes in fossils in different rock strata
 Catastrophism: Believed that there must have been successive
catastrophes (flood/drought) that wiped out most forms of life

Picture (slide 16)

Charles Lyell – British Geologist

 Promoted idea of uniformitarianism i.e., most geological features are

caused by slow, gradual changes – not catastrophic events
 Earth’s surface sculpted by earthquakes, volcanoes …
 Laws of nature have been constant over time – so that we can use
modern, observable phenomena to explain past events

Jean Baptiste de Lamark – French Scientist

 Recognized that species change through time in response to their

environment
 Suggested that spp. Are plastic and can be modified in response to the
environment
 Characteristics are passed from generation to generation
 Proposed a mechanism for the process of evolution
 Promoted idea of gradualism
 His ideas that organisms strive for perfection and that complexity is
achieved by “will" is not “accepted" today
 E.g. Use &Disuse – giraffes’ long necks
  Acquired characteristics are NOT passed on

Thomas Malthus – British Political Economist

 Malthus was concerned about the decline of living conditions in 19h

century England
 In 19798he wrote “Essays on the Principle of Population"
 He observed that in nature plants and animals produce far more
offspring than can survive – there is a struggle for survival

By 19th century …

 Accepted that earth is in the range of millions of years old

 Acceptance of fossil record
 Similarities among different species noted
 Descended from previously existing organisms
 Questions??
1. What natural mechanisms could explain why organisms change?
= Evolution by Natural Selection (Darwin)
2. What hereditary mechanism could enable organisms to change?

Charles Darwin

 1831: post as naturalist on the H.M.S Beagle

 Voyage around the world lasted about 5 years
 Collected a wide range of plants, insects, and animals, including fossils.
 Also studied geological formations
 Beagle voyage seeded his ideas about evolution -> Principle of
Natural selection

Beagle Voyage:

o Argentina
o Darwin found fossils and realized that they were similar too
modern forms
o Saw common characteristics shared among organisms on
different continents
o Fossil record indicates that there are some fossils that are closely
related to present organisms
o Time period = environmental changes which would have caused
some species to evolve to be better suited to the environment
o Living 3 toed sloths in South America (tree-living) are similar too
giant fossil sloths (ground living)
 - Small body size: escape predators, easy to find shelter and
food
o Andes
o Darwin found invertebrates high in the Andes; he reasoned that
geological processes must have caused an uplift
- Indicating possible high sea levels and/or new mountain
formation
o Galapagos Islands
o 950km off NW coast of South America
o Volcanic; no contact with the mainland
o Islands are fairly close to each other – potential movement of
some animals between them
o Far from other land masses
 Darwin’s Finches
 Birds on Galapagos very similar to mainland forms
 One species on mainland, but 14 on island; more
dusky-coloured (to suite the environment)
 Diverse use of habitat
 Ancestor was probably seed-crushing ground dweller
– Geographic speciation could have occurred

Picture (slide 33)

 Cormorant
 Flightless: has no natural predators therefore no
reason to fly
 Mocking bird
 Different species on different islands -> environment
dictated differences
 Galapagos tortoises
 Each island was “inhabited by a different set of
being"
 Two main types based on shell shape:
1. ‘Saddle-backed’ – feeds on taller cactus
2. ‘Dome-shaped’ – feeds on vegetation close to the
ground
 Plants
 Darwin found plants related to daises and
sunflowers, but they grew to tree-size
o Sea point
 o Looked at the rocks in the Cape, noticed the contact zone
between Malmesbury & Granite – concluded that the Granite
intruded into the Malmesbury
o Rocks gave clues as to the periods of changes in the
environment

Formulating Darwin's hypothesis on evolution

 Darwin's observations:
o Diversity of life
o Adaptations – organisms well suited to particular environments
o Fossils and extant forms e.g. Glyptodonts & armadillos
o Galapagos islands – flora and fauna
o Widely distributed species often in different regions
o Geology
 Darwin’s approach was scientific:
1. Induction (fact finding & synthesis – general statement/ hypothesis)
2. Deduction (certain things should follow if general statement is true)
3. Hypothesis testing – if hypotheses are to be useful, there must be
ways of testing their validity

Picture (Slide 39)

Central ideas of Darwin’s theory of Evolution

1. More offspring are born than survive to reproduce (influenced by

Malthus)
 Due to Natural Selection: individuals that where better suited to the
environment survived to reproductive age
2. Variety exists in all species
3. Those varieties that are “better adapted” to the environment are more
likely to survive and reproduce ‘survival of the fittest'. Less ‘fit'
individuals are weeded out by Natural Selection
4. If the characteristics of different varieties are inherited by their
offspring, then the nature of a species will slowly change -> DESCENT
WITH MODIFICATION
 D.W.M: descended from a common ancestor BUT
modified(adaptations) suited to time and
environment

Significance of Darwin's hypothesis

 1. Provides a volume of evidence that evolution occur
2. Provides a mechanism for how evolution occurs i.e. through natural
selection
3. Unifying theory for biology – explains why related organisms are
similar; explains relationship between structure and function; leads
to testable hypotheses.

Principles of evolution

 Natural Selection leads to adaptation (if the environment doesn’t

change and there if variety in the offspring produced)
 Evolution (change) driven by changing environment. (Populations may
change very little when conditions are constant)
 Natural Selection NOT necessarily life or death: improved fitness may
simply improve reproductive success
 Accumulation of changes may lead to speciation ((i.e. development of
new species)
 Extinction is common (evident in the fossils records)
 Evolution never yields a predictable outcome, although natural
selection and past evolutionary history may “steer" the evolution
species in a particular direction. Mutations arise by chance and are
then acted upon by natural selection

Testing the Hypothesis

1. Age of the earth should be “old" (Why? Evolution occurs over a long
period of time)
2. Fossil Evidence: change in composition; simple to complex; increasing
diversity
3. Morphology, physiology, biochemistry and genetics should all reflect an
evolutionary tree: closely related species share most characteristics
 Apes & Humans: shared common ancestor – both have opposable
thumbs
4. Existence of intermediate forms (transitional species: shared features
between two different species)
5. Populations of species should change (adapt) as conditions change
6. Change from one species to another over a period of time
7. Biogeographical distribution should reflect evolutionary history
(PANGAE)

1. Age of the Earth should be old

4.6 billion years ago

 Solar nebula (swirling cloud of dust and gas) contracts due to gravity
and forms the sun
 Remaining material forms planetesimals through-accretion the sun
 Planetesimals collide and merge to form planets including Earth

How could organic molecules be generated in an abiotic environment?

 Stanley Miller Experiment 1950s

 Methane, ammonia, hydrogen, and water vapour used to form organic
molecules (amino acids)
o E.g. Alanine and glutamic acid (common in protein)
 Amino acids identified on meteorites
o Panspermia hypothesis?
 The theory that the building blocks of life exist throughout the
universe, and are distributed by meteorites, asteroids and
space dust

Earth – 4.5 billion years

 Origin of heredity (replicators/ self-replicating entity like DNA

or RNA) -> within the DNA and RNA are the essential genes
that lead to evolution
 Once life evolved, Evolutionary Processes led to the
development of amazing diversity and complexity

Stratigraphy

 The branch of geology that studies rock layers (strata) and layering
(stratification) – primarily used in the study of sedimentary and layered
volcanic (igneous) rocks
 Fossil record is set in a framework for geological time
 Rocks are “dated” by radiometric dating i.e. half- life decay radioactive
elements

Picture (Slide 51)

Dating rocks

 Radioactive elements decay into other elements at a fixed rate

 Results in 50% loss of the original element over a fixed period of time
 If the first 50% is lot in 5600 years – refer to as “half-life” of the
element
[Link] Evidence

What are our sources of information about biodiversity?

1. Living forms
 Extant: still in existence; surviving
2. Fossil record
 What are FOSSILS?
 “Fossil” refers to the preserved traces or remains of ancient
organisms

Picture (Slide 57)

Examining the fossil record

 Fossil record provides vital info about:

1. Origination
2. Diversification
3. Extinction

Major episodes in the history of life

 Supported by the fossil records

Picture (Slide 60)

Oldest traces of life

 Barberton Greenstone Belt, South Africa, 3.5 billion years old

 SW Greenland – 3,75 billion years old
 Microbially-generated tubular structures containing carbon-support a
biogenic origin

Stromatolites – mounds formed by colonies of cyanobacteria

 Mounds grow 1mm per year

 Sediment trapped with calcium carbonate – forms layers

Cambrian Explosion ~ 545 mya

 Explosion of diversity over 5 – 10 million years

 All extant phyla occur
 Relatives of crustaceans, starfish, sponges, molluscs, chordates, algae
etc.

Picture (Slide 71)

[Link], physiology, biochemistry and genetics

Three domains of life

Picture (slide 72)

Cladograms

 Show how different animals are related to each other

 Oldest groups at the base and at the top = the youngest group
 Branch = common ancestor between organisms

Picture (side 74)

Picture (Slide 77)

Homologies

 Shared derived characteristics = HOMOLOGOUS (in this case the

presence of 4 limbs in a homology)
 Criteria for Homologies:
o Same basic structure
o Same relationship to other features
o Same development

Picture (slide 3)

Vertebrate diversity

 Diverse body plans

 Each group with their own SYNAPOMORPHIES

Analogies

 Bird and bat wings = ANALOGOUS

 They have separate evolutionary origins but are superficially similar
because they evolved to serve the same function
 Analogous Morphology = CONVERGENCE EVOLUTION

Picture (slide 5)

Vestigial structure

 Ancestral features no longer used become vestigial

o Vestigial limb bones in whale and python
o Wisdom teeth - helped human ancestors grind down plant tissue
o Coccyx – remnant of lost tail

Genetic similarity
  Genetic code is similar across entire range of living organisms –
suggest common ancestry
 Genetic info is encoded in a set of 3 bases (a codon) that specify
specific amino acids during protein synthesis

Picture (slide 11)

What makes us human?

 HAR1 – (Human Accelerated Region 1) 118 letter sequence that change

the most (with 18 differences) since humans and chimps; not much
change between chimps and chickens

Physiological & Biochemical similarities: metabolic pathways e.g. myoglobin,

rhodopsin

 Globin – common in bacteria, plants, animals, fungi, it binds and

carries oxygen; through duplication and divergence difference copies
of globin molecule adapted for different roles
o Myoglobin – functions in muscles
o Haemoglobin – functions in blood, carries oxygen to body
tissues

Molecular Data and the Evolutionary Relationships of Vertebrates

 Differences between human haemoglobin and molecules and other

vertebrates

Picture (slide 16)

 Closely related species share greater proportion of bases than distantly

related species

Picture (slide 17)

“Ontogeny recapitulates phylogeny”

 In simple terms, an organism’s embryonic development mirrors its

evolutionary history
 An old idea – not as simple as originally described; but essentially true
  Even human embryos have a stage with a tail (monkey) and
pharyngeal pouches (fish) – descent from a common ancestor explains
these similarities

[Link] of intermediate forms

Intermediate/ transitional forms

1. Archaeopteryx – intermediate between dinosaurs and birds

 Dated to 150 mya (full skeleton)
 Earliest bird – intermediate form between reptiles and birds
 Shared features:

TETRAPODS(meat-eating BIRDS
dinosaurs)
Skull anatomy Feathers
Forelimb structure Length of arms
Tail Fused clavicles (powered
flight)
Teeth Pelvis structure
No sternum Structure of hindlimb
 Feathered dinosaurs – not all feathers are for flight

Picture (slide 24)

2. Tiktaalik – intermediate between sarcopterygian (lobe-finned fishes)

and tetrapods
 About 375 mya; Canadian Artic
 Pectoral girdle is intermediate between a fin and a limb
3. Diarthrognathus – intermediate between mammal-like reptile
(cynodont)

CYNODONTS(mammal- DIARTHROGNATHUS MAMAMALS

like reptile
Quadrate-articular jaw Skull almost entirely Squamosal-dentary
squamosal and joint
dentary
Unfused lower jar at
front
Shoulder and hip
girdles

4. Pakicetus – intermediate between hoofed mammals (artiodactyls) and

whales
  500 mya

HOOFED MAMMALS PAKICETUS WHALES

(artiodactyls)
Ungulate-like features Semi-aquatic Inner ear (auditory
bulla with sigmoid
process)
Skull anatomy Small brain (as in
early whales)
Overall small size Relatively large skull
(1.8m)
 Other transitional whales:
 Basilosaurus, 40 mya; 16m fully aquatic; has pelvis and hind
limbs detached from spine
 Dorudon, 40 mya; 5m in length; movement of nostrils to top
of skull (blowhole); eyes move to side of head; tail powered
swimming (vestigial limbs)
 Molecular evidence shows that hippos are closer living relatives
of whales

[Link]

 An adaption is a feature that is common in a population because it

provides some improved function e.g., a behaviour that allows better
evasion of predator; a protein that functions better etc.
 Bat echolocation: improved predation
 Creosote bush: produces toxins that prevent other plants from growing
nearby, therefore reduced competition for nutrients and water

Artificial vs Natural selection

 Artificial Selection: Farmers and breeders have long been selecting

particular traits in organisms e.g. GMOs, dogs
 Natural Selection: Traits and characteristics selected by nature
 Natural Selection = survival of the fittest
o What does fittest mean?
 “Fitness” relates to characteristics that affect an organism’s
chances of reproducing and/or surviving
 Examples: speed at which antelope runs to evade predators;
cryptic colouration; ability to sense prey; ability to attract
pollinators

Natural selection -> differential reproductive success

  Organisms have unequal ability to survive and reproduce
 Organisms with traits better suited for the environment, producing
more offspring
 Differential reproduction means that heritable variants are
preferentially passed on to future generations

Natural selection can sometimes be directly observed

1. Peppered moths, Biston Betularia

 In 1844 (northern hemisphere) there were very few black
peppered moths (Biston betularia). By 1900 there was an
increase. WHY?
 Trees were sooted because of coal burning – dark moths were
less visible to predators and thus became dominant
 Later when “clean air” act was introduced, many more lightly
coloured moths were seen
 Different varieties may have different chances of survival
and reproduction
 The fitter variety will produce more offspring with its genes
which have a chance to spread in population
2. Ground finches from Galapagos Islands
 Rosemary and Peter Grant (with student, Peter Boag)
 Ground finches (Geospiza fortis) – beak correlates with size and
hardness of seeds it ate. Beak size was heritable
 Appears to track weather patterns: during drought – fewer small
seeds – birds had to eat larger harder seeds; those with strong
beaks survived and reproduced
 Following droughts, finches have deeper, stronger beaks that
enables them to eat tougher seeds
3. Insecticide resistance

Picture (slide 46)

 NOTE: Insecticide does not create resistant individuals. Resistant

insects that were already present in the population are selected for
4. HIV resistance
 3TC drug interferes with the reverse transcriptase enzyme which the
virus uses to copy its RNA into host DNA
 By chance some of the HIV have a slightly different reverse
transcriptase enzyme
 These variants soon increase and become 100% of the population
Picture (slide 47)

 NOTE: a drug does not ‘create’ a resistant pathogen; it ‘selects for’

resistant individuals in the population
5. Mussle shell thickness in response to invasive Asian shore crabs
 1998
 Asian shore crab found on New Jersey coast (USA)
 Feeds on mussels
 Bred profusely
 Mussles with thicker shells not eaten; survive and reproduce;
population develop thicker shells; counters them being preyed on

Natural selection can affect level of variation

Picture (slide 48)

*Selection favours heritable traits through differential reproductive success

Natural Selection does not produce perfect traits

1. Natural selection
 Galapagos bird variation
2. Constraints due to evolutionary history
 Evolution of bird flight from tetrapod ancestor
3. Trade-offs
 Seal flippers on land and in water
 Natural selection can affect levels of variation by stabilizing or
diversifying, or directional selection

[Link] from one species to another

Picture (slide 51)

*Living horse is the only surviving branch of an evolutionary tree with many
divergent trends (“dead ends”)

Evolution of horses

 Trend from browsers (leaf-eating) to gazers (grass-eating)

Hyracotherium to Equus – exposed; need to be able to eat grass
(physical and dentition change) an evade predators through running
(foot became a single toe)
 Changes linked to environmental changes and spread of grasslands
 Changes in skull + teeth in response to shift from browsing to grazing
  Reduction in numbers of toes (4 – 1) and legs become longer ->
increased running ability

[Link] distribution should reflect evolutionary history

 Distribution of fossil plants and animals on the southern continents in

the Permian and Triassic (about 286 -200 mya)

Picture (slide 55)

 Marsupials – pouched mammals

 Fossil record – late Cretaceous (+- 70 mya)
 Oldest fossils are from North America
 By Miocene/ Early Pliocene (+- 5 mya) most marsupials become
extinct in South America, however Australia and North American
forms still existed (e.g. possum, Didelphis)
 Marsupials are not as diverse as placentals, but have a range of
different forms
 Isolation of Australia -> convergent evolution between
marsupials and placentals; marsupials and placentals evolved to
fill the same ecological niches
 Galapagos Islands
 Variety based on conditions of each island

Picture (slide 58)

Summing up

1. None of the lines of evidence that can be used to test the hypothesis of
evolution has allowed rejection of the concept
2. Evolution (descent with modification) is real
3. Darwin’s four central ideas/postulates about mechanism of
evolution are strongly supported by evidence
4. Darwin’s views on the mechanisms of evolution pre-date genetics so
are insufficient to explain all aspects of evolution especially with regard
to (a) how new varieties arise and are inherited, and (b) the origin of
new species

Species and Speciation

What is a species?

 A group of individuals that can interbreed and produce fertile offspring

in nature
 The biological species concept is based on inter-fertility rather than
physical similarity.

Causes of Speciation

1. Geographic Isolation
 One population can be divided into two or more smaller
populations by mountains, rivers changing course, continental
drift
 Populations thus prevented from interbreeding

Picture (slide 65)

2. Reduction in gene flow

 No specific extrinsic barrier
 Consider species that inhabit broad geographical ranges –
individuals in the far west don’t mate with individuals in the far
east
 Zero gene flow, but not total isolation – could lead to speciation

Picture (slide 66)

A summary of reproductive barriers between closely related species

 Prezygotic Barriers
 Impede mating or hinders fertilization if a mating attempt
occurs
 Postzygotic Barriers
 Prevents a hybrid zygote from developing into a viable, fertile
adult

Picture (slide 67)

Modes of Speciation

1. Allopatric (allo=other)
 Geographically isolated
 Examples:

Picture (slide 69)

Picture (slide 70)

 2. Sympatric (sym = same)
 Within range of ancestral population
 Example: Apple maggot flies
 200yrs ago – only laid eggs on hawthorns; today they lay eggs
on hawthorns and apples
 Females tend to lay eggs in fruit they grew up in; and males
tend to find mates in fruit they grew up in
 Gene flow is reduced between population on different fruit
 Sympatric speciation often involves chromosomal changes and
non-random mating that reduces gene flow
 What about Polyploidy in sympatry?
 Example: doubling of chromosome number from diploid (2n)
to tetraploid (4n). Tetraploid may not be able to breed with
parent species but can reproduce with each other or through
self-pollination. This can lead to speciation
 Many agricultural crops are polyploid e.g. cotton, oats,
tobacco, wheat
3. Parapatric (para = beside)
 A continuously distributed population
 Example:
 Plants near mines have selected genotypes tolerant of heavy
metals; and have evolved different flowering times
 Restriction of gene flow between two species
4. Peripatric (peri = near)
 Small population isolated at the edge of a large population
 Example: (Founder Effect)
1. Geographic isolation of small population of Hawaiian fruit flies
2. Small group have “rare” genes
3. Rare genes become common in subsequent generations
4. Island flies become reproductively isolated from mainland
ones

Picture (slide 75)

Is there a barrier to gene flow?

YES

 Parapatric speciation occurs when new species evolve from a

geographically continuous population (geographic gradient) with
limited gene flow between them, often due to differences in habitat
preferences
NO

 Sympatric speciation, on the other hand, happens when new species

arise within the same geographic area, without any physical barrier to
gene flow. Reproductive barriers can limit breeding

Co-speciation

One species speciation is dependent on another

Picture (slide 78)

Mechanisms of Evolution

1. Natural Selection
2. Descent and Heredity
3. Genetic Variation:
 Mutations = i.e changes in the DNA
 Gene flow = movement of genes from one population to
another
 Sexual Reproduction = results in new gene combinations
4. Genetic Drift
5. Co-evolution
6. Sexual selection

Mutations

 Changes in DNA (hereditary material)

 Causes of mutations
o DNA not copied accurately during cell division
o Point mutation – change in one base
e.g. substitution of one amino acid (valine) by abnormal one
(Glutamic acid) in haemoglobin protein -> sickle cell disease
o External influences (radiation, chemicals, etc.) cause DNA to
break down
o Mutations can alter gene number and sequence e.g.
chromosome duplication/ deletion
o Examples:
 Down’s syndrome – extra chromosome 21 or part of it
(variable effects – physical and mental development; features)
 Wolf-Hirschhorn syndrome – deletion of part of chromosome 4
(craniofacial changes; intellectual disability; microcephaly)
o Mutations are random – Mutations can be beneficial, neutral
or harmful
 o Not all mutations matter to evolution – Mutations in non-
reproductive cells will not be passed on to offspring
o Germ-line mutations (mutations in the reproductive
cells/gametes) are passed on to offspring

Gene Flow

 Gene flow varies depending on organism e.g. sedentary vs mobile

organisms
o Corn: Wind pollinated – unlikely to reach other plants more than
15m away
 Low rate of gene flow
o Fruit flies:
 High rate of gene flow
 Effect of gene flow on a population
o Within a population: Introduce or re-introduce genes into a
population – thereby increasing genetic variation in the
population
o Across populations: By moving genes across population –
makes populations similar to one another (reduces chances of
speciation)

Sexual Reproduction

 Sexual Reproduction results in genetic shuffling – introduces new gene

combinations in a population and results in genetic variation

Picture (slide 87)

 During meiosis matched pairs exchange DNA (recombination) resulting

in 4 gametes that are different from each other

Picture (slide 88)

DISADVANTAGES ADVANTAGES
Need to find a mate Gives variation in descendants
(through meiosis – crossing over and
recombination)
Only females bear young (as Combining of ‘useful’
opposed to asexual reproduction characteristics/genes
where all individuals breed)
 Most eukaryotes are diploid – large amount of variation is ‘hidden’ from
selection in the form of recessive alleles (a)
  Thus, potentially ‘harmful’ alleles can persist in populations in
heterozygous individuals (Aa). The recessive allele is expressed only
when two heterozygous individuals produce offspring with both
recessive alleles (aa)
e.g. Cystic fibrosis (defective chloride transport across membranes)
Picture (slide 90)
e.g. Huntington’s disease in humans (a degenerative disease of the
nervous system; late onset). The allele that causes the disease is
dominant. Allele is located on chromosome 4
Picture (slide 91)
 Heterozygote advantage
o Homozygous recessive – causes sickle cell disease
o Homozygous dominant – more susceptible to Malaria
o Heterozygous individuals are protected from severe effects of
Malaria
o Heterozygotes favoured
o Frequency of sickle cell allele is highest in areas where
Malaria is highest.

Picture (slide 93 *whole slide)

Genetic Drift

 In each generation, some individuals may just by chance have more

offspring than others
 Genes/ alleles of these individuals will have higher frequencies (this
doesn’t mean that they are healthier or better)
 Allele frequencies can drift/ fluctuate randomly from generation to
generation
> GENETIC DRIFT

Picture (slide 94)

 Effect of genetic flow

1. Can reduce genetic variation
2. More drastic in smaller populations (especially rare and endangered
species)
 Two circumstances where genetic drift can have a significant impact on
a population:
1. Founder Effect:
  Few individuals become isolated from larger populations,
small group may establish a new population whose gene pool
differs from source population

Picture (slide 97)

 Example:
1814:
o 15 people founded British colony on Tristan da Cunha
o 1 individual had recessive allele for retinitis pigmentosa
(a progressive form of blindness – affects homozygous
individuals)
Late 1960s:
o 4 out of 240 people had the disease, and 9 were
heterozygous
o Frequency of the allele is 10x higher on the island than
from the original population from which the founders
came
2. Bottleneck effect:
 Size of population is greatly reduced by sudden event (e.g.
fire)
 Remaining population no longer represents original
 By chance – some alleles can be overrepresented,
underrepresented, or even absent from the surviving
population
 Even if the population size recovers, genetic variation may
remain low due to legacy of genetic drift on small
populations

Picture (slide 99)

 Example:
o Northern elephant seal population drastically reduced
(to about 20 individuals) in 1890s because of hunting
o Today there are about 30 000 – but genetic variation is
limited

Co-evolution

 Species reciprocally affect each other’s evolution

 Happens among species that have ecological relationships:
o Predator/ prey
o Plant/ pollinator
 o Parasite/ host
o Competitive species and mutualistic species
 Example:
o Bats and Moths
 Bats
 Evolved echolocation
 Moths
 Remain stationary
 Fly in opposite direction
 Complex flight manoeuvres
 Emit ultrasound at bat
o Orchid (Angraecum sesquipedale) from Madagascar has a long
nectary (30cm long tube)
 Darwin predicted in 1862 that there must be an insect that
had a long feeding proboscis
 Only in 1922 was such an insect described
 The giant hawk moth has a long whip-like proboscis that
can suck the nectar from the long tube of the orchid)

Sexual Selection

 Refers to natural selection that acts on mate-finding and reproductive

behaviour
 Sexual Selection acts on an organism’s ability to obtain a mate and
pass on one’s genes to the next generation
 Extreme lengths for sex: sexual dimorphism (size, colour,
ornamentation)
 Individuals with certain inherited characteristics are more likely to
obtain a mate and pass on their genes to the next generation
 Can result in sexual dimorphism and trade-offs
 Sexual selection acts in two ways:
1. Intrasexual selection
o Individuals of one sex compete directly for mates of opposite sex
o Example: male-male competition
 Males can compete for territories, access to females, amount
of time spent mating and even whose sperm fertilises her
eggs (sperm competition)
2. Intersexual selection (mate choice)
o One sex (usually female) chooses which individual of the
opposite sex to mate with
 o Why the female?
 Female ‘investment’ in reproduction is higher than male
investment
 Cost of reproduction:

FEMALES MALES
Eggs = expensive Sperm = cheap
Lay eggs
Incubation/ gestation Sometimes (seahorses)
Care of young Sometimes (seahorse
Lactation (milk production)
 Female’s choice often depends on ‘showiness’ of males – appearance,
dances, structures etc.
 Leks
 During the mating seasons, males may even assemble at a
traditional place (‘lek’) and engage in competitive displays
that attract females
 ‘Showiness’ may pose some risk e.g. increased predation
 Females choose good genes:
 If the trait preferred by female is indicative of the male’s
overall genetic quality, male trait and female preference for it
should increase in frequency

Macroevolution

 All the changes, diversification, and extinctions that occurred over the
3.8 b.y. of history of life on earth are the patterns of macroevolution
 Mutations + Gene Flow + Genetic Drift + Natural Selection + 3.8 b.y. =
MACROEVOLUTION
 i.e. grandest trends and transformations in evolution; patterns we
observe at the large-scale history of life; reconstructed via geology,
fossils and living organisms
 Contrast: Microevolution
 Generational change in a population’s frequency of alleles

Four Main Themes of Macroevolution

1. Stasis
 Characteristics in populations that don’t change much other time
 Many lineages don’t change much over time, e.g. cockroaches,
coelacanth, horseshoe crab
Picture (slide 3)

2. Character change
 Characteristic in populations that rapidly change over time
 E.g. Ammonite, Trilobite, Horses

Picture (slide 4)

3. Lineage-splitting
 Anagenesis: The accumulation of changes in a species that
gradually transforms it into a new species
 Cladogenesis (or branching evolution): The splitting of a
gene pool into 2 or more branches resulting in new species
 This results in BIOLOGICAL DIVERSITY

Picture (slide 6)

Tempo of Speciation

 Gradualism model: some species evolve gradually over long periods

 Punctuated equilibrium model: some species evolve rapidly, potentially
in response to environmental changes

Picture (slide 7)

Hominid Evolution

 Punctuated/Gradual?
 Stasis/change?
 Anagenesis/Cladogenesis?
 Rapid/slow tempo? (in geological time)

4. Extinction
 Five mass extinctions over the 3.8 b.y. of life on Earth

Picture (slide 12)

 End of Permian extinction (252 mya)

o Biggest mass extinction event:
 70% of land species – insects, plants, vertebrates
 Only 3 out 41 therapsid genera survive
 50% marine invertebrates’ families
 96% of all marine species
 What caused the End Permian mass Extinction?
 1. Merging of continents to form a single land mass, PANGAEA
2. Extreme volcanism in Siberia (currently)
3. Asteroid impact – Wilke’s land, Eastern Antarctica
 After the End Permian extinction, the Triassic begins with a much-
depleted fauna
 Triassic sees the origin of mammals, origin and radiation of
dinosaurs and pterosaurs
 K-T Extinction (65 mya)
o Survivors:
 In water -> Gastropods, bivalves, Fishes, other
invertebrates
 In the air -> some birds
 On land –> Mammals, Crocodiles, Plants, Snakes,
Amphibians, Tortoises, Lizards
 No Evidence for Selectivity
 K-T Extinction Processes
o Terrestrial
1. Biological
 Competition
 Egg predation
 Angiosperms origins
 Sex ratios skewed
2. Physical
 Climate cooling
 Volcanism
 Sea level changes
 Continental drift
3. Cosmic
 Viruses from space
 Asteroid impact
 Orbital eccentricity

Iridium Anomaly or Alvarez Hypothesis

 Clay layer at Cretaceous-Tertiary boundary

 High concentration of iridium
 Indicates that there must have been an asteroid impact
 At the time, no large crater was known – 180km Chicxulub impact
crater (plus 3 others, similar carte sites: Ukraine, North Sea and Shiva
Crater)
Evidence for an impact at the end of Cretaceous

1. 180km diameter crater in Mexico

2. Iridium anomaly
3. Glass spherules (Tektites)
4. Shocked quartz

What about volcanism?

 Massive volcanic eruptions in west-central India

o Deccan traps (lava beds) cover 500 000km
o Up to 2km thick in some places
o Largest volcanic features on Earth
o Formed 63-69 mya (episodically)
o Coincides with mass extinction …
 K-T event may have been the result of multiple causes i.e. combination
of volcanism, asteroid impact and marine regression

‘Age of The Mammals’ = Cenozoic Era (65 mya – now)

 The Messel Pit provides the single best fossil site which contributes to
the understanding of evolution and past environments during the
Palaeogene, a period which saw the emergence of the first modern
mammals
 Palaeogene- a period which saw the emergence of the first modern
mammals
 Age of placental mammals
 Palaeocene Epoch (66-58 mya)
 Messel, near Frankfurt
o Middle Eocene (50 mya)
o Best preserved fossil mammals
o Deposit contains also plants, invertebrates, fishes and
birds in an Eocene Lake

Evolutionary Developmental Biology

Evo-deco: Interface between evolutionary biology and how organisms

develop

 Genes that control development play a major role in evolution

 Large morphological differences can result from genes that alter the
rate, timing and spatial patterns of developmental change
Patterns in development

 Large morphological differences can result from genes that alter the
rate, timing and spatial pattern of developmental change*
o Changes in rate and timing of development (Heterochrony e.g.
paedomorphism, allometric growth)
 Change in Allometric Growth (i.e. relative growth rate of
different parts of an organism during development)
Paedomorphosis (retention of juvenile characteristics as
adults)
o Changes in spatial pattern or organisation (Homeotic genes)

Heterochrony: Allometric growth

 Refers to a change in the rate of growth of a dimension or feature

relative to other features
 Limbs that grew larger had those genes switched on for a longer period
 Allometric growth: Timing of foot development in salamanders
o Ground dwelling salamander
 Longer period of foot growth
 Longer digits and less webbing
o Tree – dwelling salamander
 Foot growth stops sooner
 Shorter digits, more webbing (enables efficient vertical
climbing)

Heterochrony: Paedomorphosis

 A slowing down of development of body, but not of maturation of

reproductive system can result in an adult that is similar to the
ancestral juvenile form
o Larval stages of salamanders have external gills (1), which they
lose as adults (2). Because of heterochrony, axolotls retain gills
as fully reproductive adults (3). This is called paedomorphosis
o Paedomorphosis is humans: Retention of juvenile cranial morph
as adults

Picture (slide 50)

Changes in spatial patterning

 Changes in homeotic genes -> Macroevolution

o Homeotic genes control the placement and spatial organisation
of body parts e.g. they determine where a pair of legs and a pair
 of wings should develop on a bird; or how a plant’s flower parts
should be arranged
-> MASTER REGULATING GENES
 Plants: MADS-box genes
 Animals: Homeobox/ Hox genes

Hox vs Homeotic genes

 Homeotic genes are master regulator genes that control the

development of body segments and structures. Hox genes are a
particular group/ subset of homeotic genes that share a common DNA
sequence called the Homeobox/ MADS-box and are often arranged on
the chromosome in the same order as they are expressed along the
body axis (regulate the expression of other genes) *

Maternal effect genes

Picture (slide 55)

 Maternal effect genes refer to products (RNA or protein) present in the

egg prior to fertilization. Influences early embryonic development
 Egg-polarity genes:
o During embryonic development-positional information (anterior,
posterior, dorsal-ventral etc.) is determined by molecular cues
(morphogens produced by mother) present in the cytoplasm of
the egg
o Egg polarity genes are a type of maternal effect gene
 Morphogen= molecules produced by mother genes - cytoplasmic
determinant
 The anterior – posterior axis of the developing embryo is set
early by gradients of morphogens (e.g. Bicoid protein formed by
Bicoid genes)
 If the mother lacks/ mutant Bicoid gene – lethal for embryo
 E.g. A mutant mother has two mutant alleles for Bicoid gene –
larva has two tails and lacks anterior structure

Picture (slide 56)

How is mRNA involved in creating the gradient of morphogens

 mRNA plays a crucial role in establishing morphogen gradients by

acting as the blueprint for the morphogen protein. Specifically, the
 localized expression and translation of mRNA determine the source and
distribution of the morphogen, which then diffuses to create a
concentration gradient. This gradient, in turn dictates cell fate and
development

Gradients of morphogens also affect regional expression of segmentation

genes

Picture (slide 60)

 Anatomical identity of the segments is determined by the homeotic

genes
 i.e. Homeotic genes specify the type of appendages and other
structures that each segment will form
 Segment duplication and loss probably occurred several times among
the arthropods

Picture (slide 61)

Modification for a function

 Homeotic genes are also responsible for modified appendages e.g. to

form pincers

Hox genes
 A type of homeotic gene – they provide positional info in an embryo;
this positional info prompts cells to develop into particular structures
appropriate for that location.

How does this relate to macroevolution?

 Molecular analysis of Drosophila’s homeotic genes showed that it
included a 180-nucleotide sequence called a hox/homeobox
sequence. An identical sequence has been found in many
invertebrates and vertebrates
 Hox genes have been relatively well conserved during evolution.
Similar sequences of hox genes found among wide variety of
organisms e.g. insects, nematodes, fish, frogs, mammals
Similarities reflect common ancestry of life on Earth
 Animals have homeobox/ hox genes but plants have MADS-box
genes, suggesting that plants and animals diverged earlier and ten
developed different developmental regimes

Evolution of vertebrates from invertebrates: major evolutionary step

Picture (slide 65 – whole slide)

Picture (slide 66 – whole slide)

 Homeotic genes that control anterior and posterior structure

 Changes in expression of hox genes results in different body plans
 How the gene is expressed will determine how the organism looks

Picture (slide 68 – whole slide)

One of the major transitions in vertebrate evolution of tetrapods from fishes

(sarcoterygians):

 Change from fins to walking/ supporting limbs

Picture (slide 69)

Picture (slide 70)

Cambrian explosion – 545 mya

 Explosion of diversity
 Developmental genes may have played an especially important role
(intrinsic theory)
 Changes in developmental genes can result in new morphological
forms
 Adaptive evolution then occurs by natural selection
 Extrinsic theory – oxygen levels in the atmosphere triggered the
explosion in diversity

Evolution of complex structures

 “Irreducible complexity” used by anti-evolutionists

 Term has been applied to any system of interacting parts in which the
removal of any one part destroys the function of the entire system. An
irreducible complex system requires that every component needs to be
in place before it will function

Evolution of complex structures: EYES

 Rhodopsin (visual purple)

 Photosensitive pigment (photoreceptors) sensitive to light
 Found in all animals that are sensitive to light (even bacteria)
 Detection of light
 When light strikes rhodopsin – it splits into opsin and retinene
– leads to nerve impulses that relay info to brain
 In the dark, opsin and retinene are recombined
 Eye Evolutionary Stages
  Limpets: simple eyecup with photoreceptors, no lens
 Slit-shell mollusc: slightly more advanced, slit allows light
direction sensing
 Nautilus: fluid- filled pinhole eye, helps focus light (still no
lens)
 Independent Evolution of Eyes
 Eyes with lenses evolved independently in:
o Insects
o Molluscs
o Vertebrates
 Convergent evolution -> similar solutions evolved separately
 Different Eye Types
 Insects (compound eyes):
o Thousands of small units (ommatidia) each with lens
and few receptors
o Produce a mosaic image
 Molluscs (e.g. squid, octopus, cuttlefish):
o Complex camera-like eyes, with lens and retina
o Can form sharp images
 Vertebrates (e.g. birds, humans):
o Highly sophisticated eyes
o Birds: can adjust both cornea and lens -> sharp vision
at different distances
 The Cambrian explosion saw the evolution of the eye from simple
structures to complex structures within a short period of time
(evolved from simple light detectors to complex image
forming organs)
 Rapid diversification of eye types across animal groups

Relevance of understanding Evolution

1. Agriculture
 Importance of genetic variation:
 Mass production of genetically similar food makes them
vulnerable to disease and pests
 Low genetic variation = greater vulnerability to change in
environmental conditions
 Example: Irish potato famine
  Planted “lumper” potato variety. Vegetatively reproduce so
they were “clones”. Potato crops were devastated when a
potato disease swept through Ireland.
 Planting a single genetically uniform crop might increase yield
in short-term but could be disastrous if conditions change
 Evolution of insecticide resistance in insect populations

Picture (slide 88)

Picture (slide 89)

Can slow the spread of resistant genes by having refugia i.e.

fields without pesticides near fields planted with pesticide
GMO crops
2. Conservation
 Why small populations struggle
 Bottleneck effect -> drastic reduction in population size
 Leads to low genetic variation, making populations:
o Susceptible to disease
o At risk of abnormalities from inbreeding
o More vulnerable to extinction
 Threatened species: African Wild Dogs, Cape Vultures, Right Whales
 Cheetahs:
 10 000 left in the wild
 Threats:
o Human encroachment (lost 80% of range)
o Habitat loss & fragmentation
o Conflict with framers
o Genetic inbreeding -> abnormalities, low fertility
o High cub mortality
o Poaching for skins
 Population Recovery vs Genetic Diversity
 Even if population size increases -> genetic variation
does not necessarily increase
 This keeps populations vulnerable in the long run

Picture (slide 94)

 Genetic Rescue
 Introducing new individuals (outbreeding) can restore genetic
variation and fitness
 Examples:
 o Vipera Berus (adder in Sweden)
 Small isolated population -> inbreeding ->
deformities & stillborn young
 Outbreeding -> recovery, more viable offspring
o Scandinavian Wolves
 Only ~430 left, highly inbred (all closely related)
 Problems: dental defects, skeletal problems, male
sterility
 Genetic rescue: immigrants from Finland & Russia
improve breeding success
 Population had suffered a serve bottleneck in
1980s
3. Medicine
 Hereditary diseases
- Understanding the evolutionary histories of the disease-
causing genes may help to control the disease, as well as limit
inheritance and predisposition for disease
- Genetic counselling
 Heterozygote advantage
- Homozygous recessive -> causes sickle cell disease.
- Homozygous dominant-> more susceptible to Malaria.
- Heterozygous individuals are protected from severe effects of
Malaria
-> Heterozygotes favoured
 Pathogenic disease – understanding the evolutionary patterns of
disease-causing organism (or hosts) is important

Picture (slide 99)

 Emerging disease
- Zoonoses
o Diseases transmitted from animals to humans e.g.
Rabies, Malaria, bird flu, Covid 19
o Understanding pathogen evolution assists in the
development of new drugs and vaccines