Evolution

Compare and contrast different views of natural philosophers prior to Darwin

Essentialism (Plato)
o Two coexisting worlds
Perfect (ideal) we conceive
Illusionary (imperfect) we perceive
o Variation in organisms were imperfect representations of their essential (ideal)
forms
Teleology
o Aristotle
Believe in the divine order or purpose of life
Order species according to Scala Naturae (Great Chain of Being)
Species are fixed forms
Scale from simple to complex
All characteristics are purposeful
o Carolus Linnaeus
Bring order to the diversity of life in the book Systema Naturae
dedicated to the greater glory of God
Develop binomial nomenclature
Hierarchical not evolutionary
Based on morphological similarity
Assume fixity of species
Copernicus, Galileo, Newton and Descartes replaced the will of God or Aristotelian
teleology (purpose of events) with concept of efficient cause (mechanisms that cause
events)
Explain how Cuvier used catastrophism to oppose evolution
Georges Cuvier
o Lifes history is recorded in fossil-containing strata
o Each stratum has unique fossils
o Older strata has more fossil organisms that differed from modern organisms
o Extinction is common is Earths history
o Doctrine of Special Creation
o Fixity of species
o Boundaries in strata and extinction events corresponded to catastrophes (E.g.
Biblical flood)
o Catastrophism; Theory that major changes in Earths crust are result of
catastrophic events rather than gradual process of change
How principles of gradualism and uniformitarianism by Charles Lyell influenced
Darwins idea of evolution

 Gradualism; theory that profound change is the cumulative result of slow continuous
processes
o James Hutton
Proposed major landforms can be explained by mechanisms currently
operating in the world
o Charles Lyell wrote Principles of Geology
Darwin concluded from Hutton and Lyell theory of gradualism
o Earth must be ancient
o Very slow and subtle processes persisting over long time can result in dramatic
change
Jean Baptiste Lamarks model for adaptation evolution
Transformism
o Organisms arise from inanimate matter by spontaneous generation and
progress
o Progress along Scala Naturae towards greater complexity
o Path of organisms evolution is guided by changing environment
o Inheritance of acquired characteristics
Differential use or disuse of organisms during individuals life
Transmitted to subsequent generation
How Thomas Malthus essay influenced Darwin
Essay on the Principles of Population
o Intrinsic rate of increase in human population
o Lead to food shortage and mass starvation
Inferences Darwin made from his observations the lead him to propose natural selection
as a mechanism for evolutionary change
Darwin saw the gradation and diversity of structure in one intimately related groups
of birds
o Original paucity of birds in Galapagos
o One species had been taken and modified for different ends
o Darwins finches
o Preservation of favourable variation and rejection of injurious variations
Principles of Darwins theory
o Organisms beget like organisms; Lamarck
o Number of individuals in each generation that survive and reproduce is
smaller than number initially produced, (struggle for existence) ; Malthus
o Chance of variation among individual organisms in any given population and
some variation is heritable
o Favourable variation is inherited and becomes more common form one
generation to next by differential reproduction (Natural selection)
o Natural selection leads to slow accumulation of adaptive changes that
differentiate groups of organisms from one another (speciation); Lyell

 How was Darwins theory revolutionary

o Dynamic biological change and variation challenge essentialism
o Undirected, purposeless variation made teleological/ metaphysical
explanations unnecessary
o Challenged Doctrine of Special Creation and Fixity of Species
How Darwin used his observations from the voyage to formulate his evolution theory
Darwins claims
o Modern species evolved from ancestral forms
o The mechanism for evolution is natural selection
*Theories; conceptual frameworks with great explanatory and predictive power
supported by many lines of evidence
*Colloquial use of theory is referred to as a hypothesis
Hypthetico-deductive method
o Hypothesis is tested by determining whether the deductions drawn conform to
further observation
Evidence that show species change over time
Natural selection is too slow to be observed directly
Microevolution
o Artificial selection in plants and animal breeding experiments
o Importance of natural variation
o Role of selection in evolution below level of species
Macroevolution
o Above level of species or speciation
o Biogeography
Geographical distribution of species
First evidence for common descent
Biogeographical patterns only made sense in light of evolution
Island of endemic species closely related to species in nearest
mainland/ neighbouring island
Oceanic currents
Continental drift
o Diversity of species
Biodiversity itself challenge special creation
So many species!!
o Fossil record
Fossils are evidence of continuity and descent with modification
Not repeated catastrophes and multiple special creation events
Absence of transitional fossils; the missing links
o Homologous structures
Structures that are the result of common ancestry
Can be used for different functions
Might or might not appear similar

o Adaptations
Organisms influenced by changing environments over evolutionary
time
Convergence; evidence of unrelated organisms occupying similar
niches adapting in similar ways
Imperfect adaptation
Vestigial organs
o Rudimentary structures of little or no use to organism
o Why an intelligent designer retain vestigial organs
Why variation is important in Darwins theory
Distinguish between artificial selection and natural selection
Artificial selection in dogs
o Likely origin of domestic dogs for Middle Eastern gray wolf
o Modern breeds appearing through intensive artificial selection
Drug resistant HIV due to the use of drugs
How population genetics was the turning point for evolutionary theory
Atavistic traits
o E.g. Dolphin with 4 fin is evidence for ancestral cetacean with modified hind
limbs, but the gene is now vestigial
Modern synthesis
Mendel
o Inheritance of a trait is determined by genes which are passed on
o Individual inherits 1 gene from each parent
o Recessive traits can be passed on to the next generation
Mutation
o Was regarded as the only significant force to bring about evolutionary change
o Origin of new species/forms
o Sports of nature
Then Mendelian genetics and Darwinian evolution was reconciled
o Synthetic theory
o Neo-Darwinian theory
Hardy-Weinberg equilibrium
o Stability of frequency of alleles in idealised populations of diploid organisms
o Assume
1.No mutation
2.No gene flow
3.No genetic drift; population is large enough
4.Panmictic (random mating)
5.No selection; no difference in reproductive success conferred by
alleles

o Provides a null hypothesis about an idealised population in which allele

frequencies does not change from generation to the next
Modern synthesis
o Emphasized
Importance of populations as unit of evolution
Central role of natural selection as the primary mechanism of
Adaptive evolutionary change
Random genetic drift, gene flow
Gradualism as the explanation of how large changes can result from
the accumulation of small changes over long period of time
Hardy-Weinberg theorem, use it to calculate allele, genotype frequencies
Hardy-Weinberg equilibrium
o Stability of frequency of alleles in idealised populations of diploid organisms
o Assume
1.No mutation
2.No gene flow
3.No genetic drift; population is large enough
4.Panmictic (random mating)
5.No selection; no difference in reproductive success conferred by
alleles
o Provides a null hypothesis about an idealised population in which allele
frequencies does not change from generation to the next
o Calculate expected frequencies of genotypes
Given observed frequencies of alleles
Single locus with 2 alleles
Allele frequencies p+q=1
Genotype frequencies p2+2pq + q2=1
E.g. 64 pink, 36 white
o Pink A is dominant to a which is recessive
o q2=36/100 q=6/10
o 1-0.6=0.4
o p2= 0.42= 0.16
o Homozygous dominant= 16
o Heterozygous= 64-16=48
How genetic variation is created and preserved in populations
Microevolution process gives rise to macroevolution pattern
Adaptive radiation is the occasional strong selection of heritable characteristic in a
variable environment (geographic isolation)
E.g. Beak evolution in Darwins finches
o Beak depth influences feeding efficiency of medium ground finch
o Has considerable variation in population
o Heritable

o Fulfil requirement for natural selection

o Periodic episodes of drought provide opportunity to measure impact of
environmental events on differential survival
Wet; birds have small seeds-average beak depth decreases
Dry; small seeds are rare and big seed must be cracked-average beak
depth increases
Adaptations of population
o Environmental change
o Natural variation in heritable trait
o Natural selection (differential survival and reproduction)
Phenotypes are influenced
o Heritability
o Environmental variation
Polygenic variation
o Influenced by multiple genes
o Assume
Additive polygenic
No environmental variation
Genes are codominant
o Influence with polygenes make continuous variation
How genetic drift, gene flow, mutation, non-random mating and natural selection can
cause microevolution
o Evolution: change in allele frequencies in a population
o Biological species concept: a group of organisms that actually (potentially)
interbreed in nature producing reproductively viable offsprings, and are
reproductively isolated form other such groups
o Gene pool: Most inclusive reproductive community- defined as the totality of
genotypes of all individuals in a reproductive community
o Population: any group of individuals of one species that occupy a given area at
the same time
o Sexual reproduction
Most important in promoting genetic variability in populations of
diploid organisms
Recessive refuge
Diploidy preserves variation by sheltering rare, recessive alleles
from selection
o Inbreeding
Breeding between close relative
Loss of variation
Promotes homozygosity
*Incest is mating between first-order relatives
o Outbreeding
Promote variability

o Heterozygote advantage
Recessive alleles that are fatal in homozygous state is selective for in
heterozygous state
E.g. Sickle-cell anaemia protects individuals from malaria in
homozygous state
Role of population size in genetic drift

Genetic drift: phenomenon in which certain alleles increase or decrease in frequency

(or disappear) as a result of chance events (not selection)
Loss of genetic variation
3 Forms
o Generational
Generational variance in populations of constant size
o Bottleneck
Population decline
Only few of original alleles survive
o Founder effect
Saddleback population
Allele frequencies change following each trans-location

Mutation without selection has little quantitative effect on a large population

o Mutation
Change in organisms DNA
Mutation in gametes immediately changes gene pool of population by
creating new allele
*Back mutation extremely rare
Random: happen in any gene in any cell without regard for benefit of
cell
Only sources of true genetic novelty
Raw material for natural selection
Contribution of mutation is substantial on a population-wide basis
Most are silent (neutral)
*Pseudo genes; dysfunctional genes that lost ability to code for protein
or no longer expressed
Effect of gene flow between populations
Gene flow
o Movement of alleles into or out of local gene pool
o Introduce new alleles
o Alter proportion of alleles already present
o Maintain similarity of allele frequencies among population
o Maintain variation within population
Why natural selection is the only agent of microevolution that result in adaptation

 Natural selection: differential reproduction of genotypes resulting from interactions

between individual organisms and their environment
o Operates whenever genotypes differ in fitness
Genetic contribution of an individual to succeeding generations
relative to the contribution of thither individuals in the population
o The phenotype (complex expression of the interaction between genes and the
environment that constitute an organism) is selected
Heritability of phenotypic character set the limits of natural selection
Character can be single gene (genetic diseases) or polygenic
(morphological character)
o The population evolves by change in allele frequencies
Evidence of adaptation
o Clines: gradual variation that follow a geographic distribution
o Ecotypes: distinct groups of phenotypes of the same species occupying
different habitats
Result in reproductive isolation
Incipient speciation
o Convergent evolution
Unrelated species come to resemble each other in some attribute
Adaptation to similar selection pressures
Not common ancestry
E.g.
fully aquatic mammals and reptiles
Adaptive radiation in placental and marsupials resulting in
similar forms (Wolf and Tasmanian wolf)
o Coevolution: result of selective forces exerted by interaction between species
Mimicry (pattern)
Mullerian: two or more harmful species, not closely related
mimic each others warning signals (cooperation)
Batesian: harmless species evolved to mimic warning of
harmful species directed at a common predator (cheating)
*Test coevolution: forms similar in allopatry?
Relative fitness and the role in adaptive evolution

How selection acts and what factors contribute to the overall fitness of a genotype
Stabilising selection, directional selection, diversifying selection
Stabilising selection: extreme phenotypes eliminated
Disruptive selection: extreme phenotypes selected at the expense of intermediate
forms
Directional: One of the extremes is favoured, driving population in a particular
direction

Rate of decline for a deleterious allele depends upon whether the allele is dominant or
recessive
Examples of organisms phenotype can be influenced by the environment
Why natural selection cannot result in perfect organisms

Organisms are often locked into historical constraints (developmental limits to form)
Adaptations often compromised by other forces (sexual vs natural selection)
Not all evolutionary change is adaptive (genetic drift is not adaptive)
Selection only edit variation that exists in gene pool

Define sexual dimorphism and how it can influence evolutionary change

Sexual selection
o Result of competition for mates
o Responsible for the elaborate mating displays and secondary sexual
characteristics
o Operates within sexes (intra-sexual: same sex) and between sexes (intersexual)
o Sexual selection on secondary sexual characteristics are strongest in
polygamous mating systems
Polygamy
Polygyny: many females
Polyandry: many males
Distinguish allopatric and sympatric speciation
Biological species concept
o Proposed by modern synthesis
o A species is a group of natural populations whose members can interbreed
with one another but cannot interbreed with members of other populations
o Hard to do for rare or widely distributed species
Modes of speciation
o Allopatric speciation
Geographical isolation of populations
Most common mode of speciation
Dispersal: move to new habitat
o E.g. Hectors dolphin vs related dolphins
Founder effect
Genetic drift and mutations
Adaptations
Vicariance: geographic disruption of distribution
o Sympatric speciation
No geographic isolation
By genetic or social isolating mechanisms

 Polyploidy: increase in genome number often coupled with

hybridisation
Disruptive selection (assortative/ preferential mating) creating
genetic discontinuities
Distinguish prezygotic and postzygotic isolating mechanisms
Prezygotic
o Habitat: live in different areas
o Ecological
o Temporal: mating at different times
o Behavioural (cultural):
o Mechanical: structural differences in genitalia
o Gametic: gametes not fertilise each other
Postzygotic
o Zygotic mortality
o Reduced reproductive success: hybrid fertile
o Hybrid breakdown: hybrids have lower viability/ fertility
Allopatric speciation model and the role of intraspecific variation and geographical
isolation
Sympatric speciation and how polyploidy can cause reproductive isolation
Distinguish anagenesis and cladogenesis

Anagenesis (phyletic): gradual change within a single lineage over time

Cladogeneis: evolutionary change produced by branching of populations to form new
species
o Responsible for the diversity of species

Adaptive radiation
Adaptive radiation: rapid formation of many new species form a single ancestral
group, characteristically to fill a new ecological zone
Extinction: disappearance of a species from the earth
o Fossil records show
Low steady rate of extinction
Interrupted periodically by mass extinctions
5 events
Initiated adaptive radiation
Gradualism vs punctuated equilibrium
Punctuated equilibrium
o Bursts of rapid speciation among small peripheral poulations
o New species replace or displace many previously existing species

o Species persist for long periods with little change then abruptly become
extinct
Understand patterns of descent as a phylogeny
Phylogeny
o Evolutionary systematics: construct hierarchical groups of species that reflect
their evolutionary history (phylogeny)
According to inferred histories of speciation events which take the
form of bifurcating branches of a tree
Taxonomy: discovering, describing, naming organisms
Systematics
o Deals with phylogenetic relationships between taxa (esp. higher levels)
o Influenced by data derived from DNA, mitochondria, chloroplast
Phylogenetic reconstructions:
o Analysis of characters and changes in character states (presence/ absence or
modifications)
o *Characters: morphological, molecular, behavioural
o Ancestral (primitive) characteristic inherited with little or no change from
remote ancestors
o Derived characters: undergone recent change and may be shared only by
closely related species or taxa
Homologous and analogous structures
Homologous:
o Structures share a common ancestry although not necessarily a common
function
Analogous:
o Similar character due to adaption to similar environments
o Misleading in determining phylogenetic relationships
Monophyletic group and polyphyletic group

Grades: groups which have evolved new adaptive features

Clades: groups including only monophyletic branching events
Contributions of phonetics and cladistics to phylogenetic systematics
Traditional taxonomy
o Relied on morphological similarities
o Often rank taxa according to evolutionary grades of change
Cladistics
o Most widely accepted
o Based on branching relationships (clades)
o Determined by shared derived Character states
o Ignore overall similarities (ancestral character)
E.g. holes in head
o Synapsid: mammals
o Anapsid: turtles
o Diapsid: birds, crocodiles, lizards
o Euryapsid: dinosouars
E.g. whales
o All artiodactyls have modified ankle bond
o Whales not have this
o DNA analysis: whales closely related to artiodactyls
o Confirmed by fossil evidence
o Now a new order: Cetartiodactyla
o Whales achieved a new grade but must be included in the new larger order
*This is all a hypothesis: can change anytime
Prosimian, anthropoid, hominoid, primates

Prosimians: lorises, bush babies, lemurs, tarsiers

Anthropoids : New/ Old world monkeys and apes
Hominoids: humans (hominins) and other apes
Hominid: group consisting of all modern and extinct Great apes (modern humans,
chimpanzees, gorillas, orang-utans + immediate ancestors)
Hominin: groups consisting of modern humans, extinct human species and all
immediate ancestors (genera Homo, Austrualopithecus, Paranthropus, Ardipithecus
o Bipedal (walking upright)

Origin of modern humans

Relatives
o Chimpanzee the closest living relative to modern humans
o Homo erectus : tall, similar skeleton to modern humans, heavier skulls, stone
tools, fire
o H. neanderthalis : Europe, fire, inhabit caves, hunted large animals, wear
clothes

o H. sapiens: Cro-Magnons (anatomically modern humans) , replace

Neanderthals, improved tools, developed symbolic representation
Eve hypothesis
o Modern humans originated in Africa
o Spread rapidly throughout Europe and Asia
o Replace other archaic humans
Multiregional model
o Geographically distant population of archaic humans linked by gene flow
o Neanderthal genes in humans
How mass extinctions could occur and affect the evolution of surviving forms
65 million years ago asteroid impact on Earth with speed of 10 km/s
o Enormous energy liberated caused environmental disasters
o Nuclear winter (fire storms)
o Huge reptiles removed
o Iridium layer
o Phase of adaptive radiation for mammals
o Punctuated equilibrium
Bursts of rapid speciation among small peripheral populations
New species replace many of the then-existing species
Species persist for long periods with little change then abruptly extinct
How inbreeding affects a populations allele frequencies and genotype frequencies
Cheetahs
o Caused by historical bottleneck
o Aberrant sperm
o Lower reproductive ability
o High neonate mortaility
o Low genetic variation
o Increase risk of disease