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Unifying principles of biology
evolutionary theory according to Theodosius Dobzhansky
Evolution
a change in genetic composition of biological populations over time
Scientist are able to demonstrate and observe evolution by?
fossil records
laboratory experiments
natural populations
Evolutionary theory
includes the entire set of ideas about HOW evolution takes place
Why should we care about evolution and evolutionary theory?
diseases
agriculture and industry
conservation
small populations evolve more rapidly
Characteristics of a scientific theory
well-supported
parsimonious
simplest explanation
falsifiable
leads to predictions that can be tested
Predictions of Evolutionary Theory
1. Spatial and temporal patterns in fossil records
2. Genetic Variation in most traits
3. A hierarchy of relatedness
4. Demonstrate evolutionary processes
Early Theories of Evolution
Eramus Darwin
Agreed that the population and species
change overtime- he had no evidence
Jean-Baptist Lamark
Looked at fossil evidence and noted that
organisms do change overtime. He concluded that offsprings inherit traits that the parents acquired which allowed them to become better adapted. (wrong) First to propose a mechanism of evolution.
Darwin was influenced by two academic works which where....
Charles Lyell's, Principles of Geology (1830)
Geologic uniformitarianism- natural process have been at work for years and explain geography changes.
Thomas Malthus, An Essay on the Principle of Population 1789
Populations may grow quickly at first but their growth will eventually decrease due to limited resources
Evolution through natural selection
Species change overtime and are different from common ancestors because of ns. Individuals with desirable traits are more reproductively fit and thus pass their genes on to the future generations. Less favorable traits are less like to reproduce...
Three key concepts of natural selection
- population must exhibit some variation in traits
aka genetic mutation
- the trait that exhibits variation must be heritable
- trait must impact an individual's reproductive success
Evolution is CHANGE IN POPULATIONS not individuals.
Natural selections is ONE MECHANISM by which evolution occurs.
Alred Wallace and Charles Darwin
Charles was developing his idea from 1830s but would publish it until he gathered enough information. Alred Wallace, independently, created a similar theory of evolution.
They both co-published the theory in 1858. But, the break though came from Darwin's published book- On the Origin of Species.
Charles Darwin discovered natural selection by observing artificial selection
What is the basis of theory of evolution by natural selection?
He noted that more individuals are born that can survive to reproduce and that individuals tend to resemble parents. So, individual differences can help increase or decrease the chance... Therefore, beneficial traits will increase in frequency in future generations
Gene pool
sum of all the alleles for a particular locus found in the population
Adaptions
favored by natural selection; process by which the trait emerges in the population
The populations must exhibit some variation in a trait
genetic mutation; change in nucleotide sequence- bad, neutral and good mutations
natural selection can only act on existing variation
The trait that exhibits variation must be heritable
phenotypic variation sources are genetic variation and environmental variation
Vp= Vg + Ve
H^2= Vg/Vp
It is the fraction of VARIABILITY that is genetic and not a fraction of the trait that id determined.
IF ALL HAVE THE SAME GENOTYPE= then Vg is 0 and heritability is 0
IF ALL HAVE THE SAME ENVIRONMENT= the Vg would equal Vp --> heritability would be 1
Parent offspring regression line
which relates the values for the traits of an offspring with the values for the traits of its biological parent
Natural selection has the greatest potential to shape traits in a population when traits have ____
high heritability
Breeder's equation
response to natural selection in terms of heritability of the trait and selection differential
R= H^2 x S
S= measure of intensity
Allows people to predict the strength of the response towards artificial selection.
Three Modes of Selection
Directional
Stabilizing
Disruptive
Directional Selection
When extreme phenotype is the fittest and that trait moves in the direction of phenotype. There is an evolutionary trend for entire frequency distribution to shift toward extreme direction.
Positive: favors trait on the high end of phenotypic spectrum (right)
Negative: low end (left)
The means changes, but variance (spread) doesn't.
Stabilizing Selection
When genetic variation decreases as the population stabilizes on an intermediate trait.
Frequency of each phenotypes changes and fitness varies... variance distribution becomes slimmer.
mean stays the same
EXAMPLE: human birth is stabilizing around 7-8
Disruptive Selection
When an intermediate trait is selected against and that traits tend toward both low and high extremes in phenotypes.
Emergence of two sep. species....
looks like "m"
Population genetics
study of how evolutionary forces cause allele frequencies to change over time
How to calculate Genotype frequency
# of individuals with the genotype of interest / # of individuals in the population
(or) fAA= NAA/N
FREQUENCY is between 0 to 1
SUM is equal to 1
How to calculate Allele Frequency
# of copies of a particular allele in pop./ # of copies of all alleles in pop.
(or) f(A)= 2NAA +1NAa / 2N
Genetic structure
describes the frequency of different alleles and genotypes in that population
p=1 or q=1 then its is a fixed allele- monomorphic
p<1 or q<1 then they have more than one allele on a locus- polymorphic
Basics of Hardy-Weinberg equation
p^2 + 2pq + p^2 = 1
states that allele frequencies should stay the same between one generation to anotherr... if it isn't then evolution is occuring
Assumptions of Hardy-Weinberg equillibrium
- no mutations
- no selection
- random mating
- population is infinitely large
- no gene migration between populations
if one isn't followed --> evolution
Genetic drift
unpredictable, random fluctuations in allele frequencies from one generation to the next because of a population's small size
random and unpredictable
larger impact when population size is smaller
( part of genetic drift) Fixation
occurs when one allele is lost and the other becomes the only one present in the population
basically gone forever
speed depends on side
The bottleneck effect
reduction of a population, usually natural disaster, such that the surviving population is no longer genetically representative of the original population
loss of genetic variation/diversity
example: cheetahs and elephant
The founder effect
when a few individuals become isolated from a larger population, with the result that the new population's gene pool is not reflect of original population
incorporates gene flow**
examples: lady bugs and tristan da cunha
Gene flow (migration)
genetic ADDITIONS or SUBTRACTIONS from a population, resulting from the MOVEMENT of fertile individuals or gametes
exchange of genetic information by migration
usually leads to reduction of differences by introduction of new alleles ( increases diversity )
Two factors:
- number of migrants
- size of the population
Comparing Gene flow to Genetic drift
Gene flow= can increase genetic diversity
migration and reproduction
exchange of information
Genetic drift= tends to decrease genetic diversity
random events leading to small size
Selection
acts on heritable variation- introduced by gene flow or mutation- favors the trait that lead to reproductive fitness and promotes adaptive evolution
higher fitness if it passes more alleles
Processes that create/maintain variation
Mutation- "creator"
Migration ( gene flow )- introduce
Disruptive selection- maintains multiple phenotypes
Neutral Evolution
Two kinds of mutations
- synonymous substitutions
- nonnsynonymous substitutions
Synonymous substitutions
nucleotide substitution has NO AFFECT on amino acid and phenotypes
silent mutations
more than one codon can code for the same amino acid UUA/UUG
Nonsynonymous substitutions
substitutions DOES change amino acid and affects phenotype
inserting different codes... missense
can be under selection
Nonsynonymous substitutions TYPES
Frameshift: adding or subtracting nucleotides
Nonsense: premature stop
In general
rates of syn. are higher than nonsyn.
( rates of silent mutation are greater than alterations in amino acid sequence )
substitutions in PSEUDOGENES are higher- no longer functional (syn. are more likely...)
Neutral Evolution
most molecular variation in most populations are selectively NEUTRL, meaning they don't convey a selective adv. or dis.
so genetic drift explains accumulation (not selection)
***Rate of fixation equation
2Nu x 1/2N = u
***Molecular clock
approach to calculating the amount of time since two species diverged
How to determine whether a particular portion of the genome is under....
- positive
- neutral selection
- purifying
Positive selection
more nonsyn. than syn.
directional selection!!
Neutral selection
nonsyn. and syn. are equal
genetic drift changes don't provide adv. or dis.
Purifying selection
nonsyn. is less than syn.
more resistant to change (silent) allowing stabilization
Genome Size, Organismal Complexity and population size
there is a strong correlation between organismal complexity and the size of the coding portion of an organism's genome
noncoding proteins STILL maintain chromosome structure etc... some have parasitic transposable elements
population size influences have effective selection is at weeding out
Genome evolution includes
- sexual recombination
- lateral gene transfer
- gene duplication
Sexual recombination
during meiosis-> genetic variability- selection can act
Asexual vs. Sexual
sexual evolved from asexual...
Sexual disadvantages;:
- reduces rate of reproduction
- reduces rate of gene transfer from mother (50%)
Sexual advantages:
- can separate malicious gene combinations
- homologous chromosomes can be used to repair DNA
- increase genetic variation ("shuffling") ( new combinations for selection to act on )
Asexual:
- no way to eliminate mutations
- every offspring is similar
Lateral Gene transfer
process by which genes, organelles or fragments of entire genomes move from one lineage to another
aka horizontal gene transfer
transfer genes from other species like viruses and bacteria
Gene duplication
when an entirely new copy of a gene appears in a genome over evolutionary time
- increase in production of protein
- genes can be expressed at different times
- one original, other accumulates deleterious (pseudogenes)
- one original, accumulates advantages (new function)
What is more likely to occur with gene duplication...?
- directional selection; because the original gene is free to serve it original function
- when an entire genome is copied then opportunities for evolution of a new function may occur
Gene family
a group of homologous genes that have related functions- from gene duplications and subsequent evolution
Evolutionary developmental biology
field of biology that compares developmental process of different organisms in an attempts to determining the ancestral relationships between them and how the developmental process evolved
Fibroblast growth factor (Fgf8)
outgrowth of the limb- expressed along the outside of the growing limb bud (apical ectodermal ridge)
Sonic hedgehog (Shh)
determining which digit is which- poster side of growing limb (zone of polarizing activity)
Animals inherit a basic genetic framework for embryonic development from a common ancestor....
snakes, mice and chickens alike
Speciation
The process in which biological lineages diverge due to reproductive isolation, resulting in new species
Three species concepts
- Morphological
- Biological
- Lineage
Morphological species concepts
based on OBSERVABLE morphological characteristics- body shape, size, color, structure, etc.
**easy to apply
Morphological species definition was by
Carlos Linnaeus - binomial nomenclature
birdwatchers
Problems of morphological species concept
1. Members of the same species may not look alike
(caterpillars look nothing like butterflies)
2. Members of different species may look similiar
CRYPTIC species look alike but cant interbreed
( male and female ducks have diff. features and age appearance )
( frogs look alike but can't breed- tetra. & diploid )
Biological species concept
by....
Ernest Mayr
"groups of actually or potentially interbreeding natural populations that are reproductively isolated from other such groups"
importance: isolation
horse and donkey can mate but aren't fertile.... (ex)
Problems of biological species concept
1. Doesn't apply to asexual organisms
( prok. wouldn't be a species then...)
2. Definition can't be applied to extinct animals
( fossils can't tell us about isolation- we make inferences )
3. It isn't clear what "potentially interbreed" really means
( ... could in the future- organisms can mate, but can be infertile )
** Ring Species
4. Definition is impractical
( we can't cross each organism to determine if they can interbreed )
5. Infertility is an ancestral characteristics
( when two species arise from one, they often remain infertile for a period of time before they are reproductively isolated )
Lineage species concept
by
GG Simpson
two groups that share a branch on the tree of life
relies on molecular similarity and phylogeny
( Cali. ravens and Canada ravens would be diff. species )
Morphological= reproductive isolated species undergo physical changes
Biological= reproductive isolation results in evolution of distinct lineages over time
Role of reproductive isolation
most important factor in speciation
speciation REQUIRES an interruption in gene flow between two groups... to evolve
How can gene flow be disrupted??
- habitat selection
- assortative mating
- geographical barriers
Dobzhansky-Muller model
explains how a single lineage can split into 2 reproductively isolated species
subdividing a population...
alleles will not be unable to mix-> different species
How isolation of gene flow occurs?
- allopatric speciation
- sympatric specitation
Allopatric speciation
population is divided by a physical or geographic barrier, forming two isolated populations that evolve independently
other homeland
Via dispesal or vicariance
Dispersal Allopatric Speciation
moving to a new area that is separated from the rest of the population and colonize- this is called FOUNDER EFFECT
Galapagos Finches-> migrated and developed different characteristics
Hawaiian honeycreepers
Vicariance allopatric speciation
climate change, glacial advance and retreat, etc.
Freshwater steam fishes-glacial advance divided them
Panama- land arose and separated Caribbean and Pacific populations
Sympatric speciation
species arise from a population that remains connected
Hint-- SS, same
Two types of sympatric speciation
- disruptive speciation
- polyploidy
Disruptive sympatric speciation
favors two extremes in same population from behavioral difference
palm trees that prefer different soil
Polyploidy sympatric speciation
mistake during cell division results in an individual with cells that have more than one copy of the genome
generally viable but genetically isolated, can't reproduce normally with others
Two kinds of polyploidy sympatric speciation
- autopolyploidy
- allopolyploidy
Autopolyploidy sympatric speciation
duplication of a set of chromosomes WITHIN a single species
Allopolyploidy sympatric speciation
when the chromosomes of two DIFFERENT species are combined
combine chromosomes by interbreeding- forming a hybrid with abnormal chromosomal number
Incipient species
species that are in the process of diverging into separate species but they can currently interbreed
Two possibilities of incipient species...
1. reinforcement
hybrids might be less reproductive- NS will favor parents who mate with their own kinds that parents who hybridize
2. merger
hybrids are as fit as non-hybridizing
can become a single species again
Prezygotic isolation mechanisms
mechanisms
temporal
behavioral
habitat
gametic
Mechanism Isolation
anatomical differences
the right "fit"
"match" between a plant and pollinator
Temporal Isolation
isolated that they never have the opportunity to reproduce
breeding time
Behavioral Isolation
animals have mating rituals that must occur
acceptance
Habitat Isolation
animals are isolated by space and never have the opportunity to come into contact with one another
relates to pre.