Evolution and Speciation - Modern Evidence: A2 Edexcel Biology Xavier DANIEL, PH.D

Session 4.
Unit 4: Development, Plants and the Environment

Topic 5: On the wild side
Evolution and Speciation – Modern evidence

Neo-Darwinism
Population Genetics
- Populations and gene pools
- The Hardy-Weinberg principle and disturbing factors
- How changing gene pools may lead to speciation
- Natural selection and speciation
- New evidence for evolution
- The role of the scientific community in validating evidence
A2 Edexcel Biology Xavier DANIEL, Ph.D.

Session 4.5


Neo-Darwinism
Population Genetics

Neo-Darwinism
Evolution = development of new species from pre-existing ones by accumulation

of genetic differences over long periods of time
 Idea of evolution closely associated with Charles Darwin (1809-1882)

See session 13 of Unit 2
 In C. Darwin’s time, no knowledge of - Genetics (G. Mendel)

- Chromosomes, Genes and DNA
People believed that offspring was the result of “blending” the characteristics of the parents
If that was true, genetic variation would decrease, not increase
 Modern genetics showed blending does not occur
 Neo-Darwinism = Evolution by natural selection restated through Modern genetics

Neo-Darwinism
Ideas in Neo-Darwinism 1/2
 Genetic variation in a population arises through:
1. Mutations
Chromosome mutations and gene mutations
2. Random assortment of homologous chromosomes

During Anaphase I of meiosis (process that forms gametes)
3. Recombination of segments between homologous chromosomes

a.k.a. Crossing-over during Prophase I of meiosis
4. Random fusion of male and female gametes

During fertilisation
Was understood in Darwin’s time
Asexual reproduction
Sexual reproduction

Neo-Darwinism
Ideas in Neo-Darwinism 2/2
 After genetic variation has arisen in a population,
Genetic variation is expressed in the phenotypes of individuals in the population

Different alleles are present at the same time in different individuals
 Some phenotypes are better suited for the environment

Phenotypes are dictated by the alleles in the genotypes
Better chances to survive / reach age of reproduction
Better chances to reproduce Survival of the fittest
Better suited alleles are passed on to offspring
Population evolves
Better suited alleles increase in frequency in the population
More and more individuals possess and express the better suited alleles
New species may arise with time

Session 4.5


Neo-Darwinism
Population Genetics

Population Genetics
Population Genetics = Study of genes in populations
Population = Group of individuals of the same species living in the same place at the same time
Can interbreed

Population Genetics
Gene pool = All the genes and all their different alleles present in
the gametes of the individuals in an interbreeding population
= all the alleles of all genes that can be passed on to offspring
 After breeding has occurred

A subset of the alleles of the gene pool make up the gene pool of the next generation
 Process repeated again and again between succeeding generations
Allele frequency = Frequency of a particular allele in a given population
 May be stable If all alleles’ frequency are stable, population is static = not evolving
 May be changing If allele frequencies are changing, evolution is going on
Evolution = change in allele frequency

e.g. alleles giving an advantage to the individuals that carry them
Will produce more offspring
Evolution can be detected even before a new species arises

Session 4.5


Neo-Darwinism
Population Genetics

Population Genetics
-The Hardy-Weinberg principle and disturbing factors
The Hardy-Weinberg formula is used to detect changes (or no changes) in gene pools
Originally to try and explain

WHY dominant
characteristics do not
eventually take over in
populations

Population Genetics
 Main issue in finding allele frequencies was

Impossible to distinguish Aa from AA based on phenotypes
Because both are A in phenotype
 Hardy-Weinberg equation makes it possible

Starting from numbers of aa = q2
q = square root of q2
 Since p + q = 1
p calculated with p = 1 - q

Population Genetics

Population Genetics
The Hardy-Weinberg formula predicts gene pool of a population
does not change in succeeding populations
 True... only if
1. Breeding population is large
So that mating is random
2. Mating is random
Any genotype can mate with any genotype
Not one genotype is favorised
"chosen", more/less predated...
3. No new alleles appear

No mutation
No immigrant bringing new alleles
 But composition of gene pool does change

Due to factors that will change the proportions of some alleles
Called "disturbing factors"

Population Genetics
1. Mutations
Random and rare
Have to happen in cells that are/will become gametes to be passed on to offspring
2. Selective predation
The "selected for prey" individuals will decrease in number
So the frequency of the allele responsible for the characteristic "target" will decrease
Vice-versa for the "non-selected for predation" individuals and their allele
3. Emigration and immigration

Emigration causes decrease in the frequency of alleles carried by the individuals that leave the population
Immigration can bring new alleles carried by the individuals that join the population
Immigration may increase the frequency of alleles carried by the individuals that join the population
4. Random genetic drift

Sudden hostile physical conditions can sharply reduce population to a few survivors
Favourable environment again: population goes back to normal
BUT from a lot of inbreeding: some alleles are lost in the process
5. Founder effect (another form of genetic drift)

Small sample of population suddenly isolated from the rest
Small sample grows into a new population
Based on small sample, so gene pool not representative
So new population will have a different gene pool
Session 4.5


Neo-Darwinism
Population Genetics

Population Genetics
-How changing gene pools may lead to speciation
Speciation = Appearance of a new species

= Process where one species evolve from another
1. Speciation by isolation
2. Reproductive isolation mechanisms
3. Speciation by polyploidy

Population Genetics

1. Speciation by isolation
3 Divergence of species
after migration
Galapagos Islands
Tortoise, Finches, Iguanas
Allopatric speciation
Speciation due to separation in space
Population Genetics

2. Reproductive isolation mechanisms

Population Genetics

3. Speciation by polyploidy
Polyploidy = a form of chromosome mutation

Ploidy = number of sets of chromosomes in nucleus
Euploidy = normal ploidy = 2 sets of chromosomes
Polyploidy = more than 2 sets of chromosomes
Polyploidy comes about when additional sets of chromosomes come from

- A member of the same species
Problem in Anaphase in meiosis: gametes diploid instead of haploid
- A member of a different species
Usually offspring sterile because old and new sets of chromosomes likely not homologous
So no pairing in meiosis
So no gametes formed
BUT if total number of chromosomes doubled (duplication of DNA occurred, but not mitosis)
So pairing in meiosis possible
Gametes can be formed, new species is polyploid

Population Genetics


Session 4.5


Neo-Darwinism
Population Genetics

Population Genetics
-Natural selection and speciation
Natural selection = pressure of environment on population to make it evolve
1. Stabilising selection
OR
2. Directional selection

Population Genetics
Natural selection = pressure of environment on population to make it adapt to it
 Environment conditions are stable and mostly not changing
 Stabilising selection does NOT lead to evolution
It - maintains favourable characteristics

- eliminates unfavourable characteristics
Population will be - more and more homogenous

- more and more fit for the environment

Population Genetics

Population Genetics
 Environment conditions are unstable and changing
 Most individuals of the population may not be fit for the changing/new environment
A subset of the population may have a selective advantage
Will be selected for
The frequency of the "better-fitted" allele will increase
Selected individuals form a new population
Evolution took place

Population Genetics
Antibiotic resistance
in bacteria
Melanism in
peppered moths

Session 4.5


Neo-Darwinism
Population Genetics

Population Genetics
-New evidence for evolution
Before "new evidence"...
 Evidence from fossils

Palaeontology
But fossilisation rare
Most fossils not found
 Fossils that were recovered
- Petrified remains: organic matter replaced by mineral ions
- Moulds: organic matter gone, vacated space becomes a mould
- Traces: impression of leaves, footprint...
- Intact, preserved whole organisms (in amber or acidic peat)

Population Genetics
Before "new evidence"...
 Dating fossils with isotopes
1. Known rate of decay of Carbon = 14C

During its life, an organism incorporates 14C
When it dies, it stops incorporating 14C
14C gradually disappears after death
Rate of decay of 14C is known
Can calculate how long ago the organism died

Gives age of fossils in the last 60 000 years
2. Ratio of Potassium to Argon = 40K / 40Ar

Potassium is gradually replaced by Argon in rocks
Gives age of sedimentary rocks + fossils
Up to Cambrian period = 580 mya
But unreliable for the last 500 000 years
Population Genetics
 Comparative physiology and biochemistry
 Most living organisms use DNA as their genetic material

Chemical structure of DNA is universal
Transcription and translation processes almost identical all over evolution
Genetic code almost universal
 Processes are extremely conserved

Cellular respiration, Transcription and translation, Photosynthesis in cyanobacteria/Plants
 Many biochemical processes have been inherited during evolution
 BUT some changes appear with time

Sequence of DNA and proteins
Use those differences to measure relatedness between species

The longer ago two species diverged,
The more differences will be found between their DNA/Proteins
 Proteomics = study of all proteins in organisms

Qualitative and quantitative
Population Genetics
Immunological studies
The more related the species,

the more precipitate is formed

Population Genetics
Differences in DNA sequences
The more related the species,

the higher the temperature needed to separate the hybrids

Population Genetics
DNA as a molecular clock
 Nucleus contains 99% of the cell's DNA
 Mitochondria also contain their own DNA

Mitochondrial DNA (mtDNA) = 16 500 base pairs
 Mutations happen in all DNAs Common ancestor

Nucleus DNA has genes encoding for proteins
that repair mutations
mtDNA does not possess these genes
mtDNA evolves 5-10 times faster than nuclear DNA

1-2 base change every 100 nucleotide per million year
Time since separation of two species from

their common ancestor calculated from
their mtDNAs' differences

Population Genetics
DNA as a molecular clock

Population Genetics
Ribosomal RNA studies
 Number of species of micro-organisms has been underestimated
 Especially those living in extreme conditions

Extremophiles
- Survive in very high temperatures: Thermophiles, found in hotsprings

- Require very high pH: Alkalinophiles, found in soda lakes
- Survive in very high salt concentrations: Halophiles, found in salt lakes
- Survive in very high pressure: Barophiles
- Survive very cold temperatures: Psychrophiles
 Numbers of known extremophiles increasing rapidly
 Extremophiles look like prokaryotes as we know them

No nucleus nor organelles
 But metabolism of extremophiles different from known prokaryotes
 Information about their ribosomal RNA was a keystone in their analysis

WHY choose rRNA for comparisons for measuring evolution? A2 Edexcel Biology Xavier DANIEL, Ph.D.
Population Genetics
1. Extraction of Ribosomes from extremophiles
Ribosomes are made of a mix of proteins and RNA
2. Ribosomal RNA separated from proteins in ribosomes
3. Different ribosomal RNAs separated from each other
4. Ribosomal RNAs sequenced: determination of their base sequence

e.g. 5' UACGUUAACGCGAUCGA... 3'
5. Comparison of their sequence with known RNA sequences from prokaryotes and eukaryotes

Population Genetics
6. Conclusions from ribosomal RNA comparisons
- Discovery of new evolutionary relationships between organisms
- Development of a new scheme of classification of organisms
The three domains

Classification based on their similarities/differences in ribosomal RNA
Archaea = Archaeabacteria = Extremophiles prokaryotes
Bacteria = Eubacteria ("true" Bacteria)
Eukarya = Eukaryotes = Protoctists + Fungi + Plants + Animals

Population Genetics
The Three Domains

Session 4.5


Neo-Darwinism
Population Genetics

Population Genetics
-The role of the scientific community in validating evidence
The example of the use of ribosomal RNA as a new evidence for evolution
 First, a small team of scientists mastering techniques of ribosomal RNA analysis
Compared rRNAs from many micro-organisms
Discoveries (by chance?) of many differences between rRNAs
 Confidence in their results: - write papers/articles

- send for publication in top-notch scientific journals
 Editors send copies of articles to scientists skilled in the same domain of research
Peer review
Rejected, need additional data for publication, or immediate publication
 Publication = recognition of the research and researchers

Will more easily get grants/funds for future research
Invited to international conferences to give seminars/talks
There, will present further advanced results
 Slowly ideas/theories get accepted (sometimes after hot debates)
 Anytime, contradictory results may disprove these results


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Session 4.

Unit 4: Development, Plants and the Environment