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trivial situation from the standpoint of the H-W equilibrium at this point
once you get to fixation, population wont evolve
ex: consider a single locus that determines flower color with two alleles with incomplete dominance: C R
(red) and CW (white); heterozygote has pink flowers
assume conditions for H-W equilibrium apply
if p represents the relative frequency of CR in the parental population, and q represents the relative
frequency of CW, then p and q remain unchanged in the next generation
if only 2 alleles in a population, one of the relevant equations: p + q = 1, in both the parental and
offspring generations
p = frequency of A (dominant allele)
p2 = AA homozygotes
q = frequency of a (recessive allele)
q2 = aa homozygotes
2pq = Aa heterozygotes
if HW conditions apply:
1) p and q will remain constant
2) regardless of starting genotype frequencies after one generation of mating, will remain constant
thereafter no evolution occurs
Genotype
Frequency
CRCR
p2
CRCW
2pq
CWCW
q2
^ as long as HW conditions apply regardless of genotype frequency, after one generation of mating
imagine the populations producing gametes in this proportion that reflects the gametes they produce
gametes combined at random
a violation to any one of the rules will lead to a change in allele frequency natural selection isnt the
only change that starts evolution: non-adaptive evolution
in real populations, allele and genotype frequencies usually do change over time
the neutral theory: most evolution occurs without the involvement of natural selection
natural selection producing changes in organisms
changes happening regardless of natural selection
both producing evolution simultaneously, but natural selection is still the primary mechanism by which
adaptive changes occur
Finite Population Size: effect of finite population size on evolution
3 contexts in nature in which this can occur:
1) genetic drift: statistical phenomenon; the smaller the population the greater the effect; effect of
small sample size leading to a higher statistical likelihood of changes occurring
gene pool with a certain frequency of alleles when gene pool reproduces, high statistical likelihood
that the next generation will not have the same allele frequency as the previous generation (evolution!)
^ex: repeated instances might cause one allele to be lost, or the population = fixed
cant predict if a population will be fixed or what allele may be fixed
once a population is fixed, the lost allele will not come back
2) The Bottleneck Effect: effect of small population size, but in a particular context: involves a large
population of organisms with a certain amount of genetic diversity then, population size gets reduced
to a small number of individuals then, population is allowed to re-expand but as a result of the
restriction in population size, the population experiences a loss of genetic diversity, rare alleles are lost:
more likely to disappear
ex: sea otters, seals due to hunting
evolutionary effect of a loss of variation: increased likelihood of an extinction event because in the face
of a new selective challenge (ex: disease, environmental change), lower likelihood that there will be
variants that will be able to survive and reproduce well in these circumstances
3) The Founder Effect: occurs when a few individuals become isolated from a larger population
ex: birds blown off the mainland to an island
essentially an isolation bottleneck with similar results
Migration/Gene Flow:
causes a population to gain or lose alleles
certain alleles passed to the next generation with greater success than others; over time natural
selection will maintain favorable genotypes that lead to effective survival, but more importantly
reproduction
allele frequencies stay at 0.5; heterozygotes stay at 0.5 in the population and the two alternative
homozygotes are at the same place at 0.25 means that selection for the heterozygote maintains both
alleles in the population and tends to keep them around at roughly equal frequencies would preserve
the sickle cell allele in the population at a higher frequency: another example in which genetic variation
is preserved
natural selection tends to reduce genetic variation
selection for heterozygotes means higher frequency for sickle cell anemia staying around
modeling discrete characters: one or the other phenotype
^top graph: how a typical quantitative character is normally distributed; in the normal distribution:
illustrated by the top graph bell curve
frequency of individuals that have a particular phenotype on y-axis; phenotypic range on x-axis
the majority of the kangaroo rats have an intermediate fur color: medium brown is most frequent
most frequent because the rats live in a desert environment so most surroundings are medium brown
the rats of this color as least visible to predators
as you move to either extreme, the number of individuals that possess that particular character state
drops off; very few
darker and lighter colors stand out more against the medium brown background easier for a predator
to detect
bottom right graph: stabilizing selection extremes selected against so the shape of the distribution
narrows around the peak; narrower bell curve
bottom middle: disruptive selection creates a bimodal distribution: two peaks with a valley in the
middle
ex: dark colors would be favored if habitat changed to be more dark and light, selection against the
medium brown
bottom left: directional selection curve remains the same, but curve shifts in the favored direction
ex: soil background changes to darker brown darker rats blend in better compared to anything lighter
^effect of genetic drift, not infinite population, over time allele frequencies are changing
^all became fixed, in a small population size, the degree of genetic drift is such that theres a higher
likelihood one or other alleles will be lost, fixation
along same plane with minimal variation: genetic drift reduce population 10-fold: greater genetic drift
50 individuals: fixation
ever-decreasing size of population: the smaller the population, the greater amount/degree of genetic
drift: a random process that does not involve evolution
^Relative fitness: infinite population size, lethal dominant allele allele wont reproduce, will be fixed
and go away in one generation
allele that is lethal dominant goes to zero, other/alternative allele goes to one; both happen in just one
generation
genotype frequencies: in one generation, beginning in HW expectations: go from homozygotes with A1
allele goes to 1; alternative homozygotes and heterozygotes go to 0
^recessive deleterious allele is not eliminated in nearly as quick a fashion, able to persist at really low
frequencies over a long time frame: as A1 allele frequency increases, number of individuals increase;
because alternative allele going down, heterozygotes and the recessive allele go down but not
eliminated in one generation, instead the homozygotes asymptotically approach 0; heterozygotes not
affected by allele because danger is held in the recessive allele
^allele frequencies stay at 0.5; genotypes: heterozygotes stay at 0.5 and 0.25 respectively
selection for heterozygotes maintains both alleles, keeping them around and at equal frequencies
preserves sickle cell allele at higher frequency
normal hemoglobin allele also maintained at high frequency; both at 0.5
another way genetic variation is preserved
natural selection tends to reduce genetic variation
selection for heterozygotes means higher frequency for sickle cell anemia staying around
Speciation:
speciation: process/mechanism by which new species arise/evolve into more than one species
involves the evolution of reproductive isolating mechanisms between populations of organisms
life started with a prokaryote and the millions of species today = result of that one species undergoing
speciation all life is related at some level
to have species form, have to evolve reproductive isolating mechanisms
species names are Latin binomials (two words/names): first name = genus, second = species epithet:
never used alone because a lot of species share this term
first letter of genus is capitalized, others are lower case
ex: American Alligator = Alligator mississippiensis; Chinese Alligator = Alligator sinensis; humans =
homo sapiens
on computers, latin words in english are designated by italics but when writing, underlined
domain kingdom phylum class order family genus species: taxonomic system that
expresses evolutionary relationships
several different definitions/concepts of what a species is, all have advantages and disadvantages:
morphological species concept: used in species descriptions, especially for larger organisms;
members of a species are those who possess a unique combination of characteristics that is possessed
by no other collections of organisms
ex: turtles
phylogenetic species concept: defines a species as the smallest group of individuals that share a
common ancestor, forming one branch on the tree of life
ecological species concept: emphasizes a species as occupying a niche
evolutionary species concept: species is a single lineage of ancestor-descendant populations which
maintains its identity from other such lineages and which has it own evolutionary tendencies and
historical fate
biological species concept: standard theory; a species is one or more populations of individuals that
interbreed under natural conditions and produce fertile offspring and are reproductively isolated from
other such populations
barrier that prevents two populations from mating: reproductive isolating mechanisms:
Prezygotic Isolation: comes into play before formation of the zygote and prevent the formation of the
zygote something prevents fertilization
habitat isolation: potential mates occupy different habitats within the same area
^dont interbreed because though they live in the same location, they live in two different habitats
temporal isolation: different timing of breeding/mating; organisms that have different breeding
seasons
ex: locusts: have breeding years, not seasons; egg that hatches makes larvae in soil and eats, etc. stays
there underground for years when it is time to mate, undergo metamorphosis and mate for weeks
species of locusts have different mating years: some 5 year, some 4 year
behavioral isolation: species (not plants) with unique behaviors regarding mating that are not shared
with other species
ex: 3-spined sticklebacks have own unique mating dance as well as blue footed boobies
mechanical isolation: physical/mechanical barrier exists that prevents sperm from meeting the egg,
e.g.parts dont fit
ex: elephant and mouse
gametic isolation: egg puts up chemical barrier so only a sperm of the same species with proper
chemical signature can gain entry into the egg
important for organisms who release sperm in water (ex: sea urchins)
Postzygotic Isolation: will get zygote and might get a fully intact and functioning organism (though
not sexually functioning)
reduced hybrid viability: the fertilized egg fails to develop into a viable organism; zygote formed, but
fails developmentally, ability to survive is severely compromised
reduced hybrid fertility: get a viable organism that may be quite good at surviving but is
sterile/infertile
ex: mules: result of mating of horse and donkey; very strong, desirable for work but sterile
heterozygous at many loci
hybrid breakdown:
hybrid = result of two species interbreeding
members of two species interbreed and produce fertile offspring but either:
1) mate back into one or the other species, dont maintain genetic identity; dilution effect
rate of exchange of alleles is not high enough to compromise the identity of each species
2) hybrid formed from two species interbreeding are fertile and are able to interbreed with one another
but after several generations, start to see certain genetic defects in the offspring that ultimately
compromises their ability to survive/become inviable and die off
two views of evolutionary change:
anagenic change: change within a lineage over time
question: Can you then recognize two or more different species within one lineage over time?
cladogenic change: speciation; lineage splitting
historical events and relationships represented by cladograms or phylogenetic trees
^earlier at bottom, most recent at top: over time the species has changed
left: species has evolved, hasnt split into two species not speciation: anagenic change
right: has evolved into two species speciation: cladogenic event
this cladogenic event involves anagenic change because it requires some evolution for the two species
to become different enough to officially become species
above is called a cladogram/evolutionary tree
^ex: shrimp and other species closely related on either side after the closure of the isthmus
^founder effect combined with natural selection new species evolving fairly rapidly to reproductive
isolation
underwent numerous cladogenic events
if part of archipelago: island hopping migration events where locally adapts to conditions
local adaption + founder effect leads to relatively rapid speciation of many species starting from a single
ancestor adaptive radiation
^ex: Darwins finches, underwent adaptive radiation: burst of divergence from a single lineage that
gave rise to many new species
1) DNA
although a bacterium that has possibly replaced phosphorus with arsenic in its DNA and other molecules
has been found wasnt an independent organism of life
2) universal genetic code: turning codons into amino acid sequences = universal process
3) ribosomes: all cells have ribosomes
4) cell membrane: universal boundary between the inside of the cell and the outside environment
taxonomy = the science of naming things
time goes from left right or bottom top: either way, base tips
taxa = entities we are trying to determine the relationships of
^taxa that have a related ancestor = sister taxa
branching points = node: each one represents a hypothesis of cladogenesis/speciation events
very base = common ancestor of all taxa
polytomy: reflects an inability to distinguish a relationship; no basis to further establish a relationship
^The branch leading to Node 1 represents the common ancestor of ABCDE, the branch leading to Node 2
is the common ancestor of BCDE (but excluding A)
B, C, D, and E share a more recent common ancestor not shared with A, they are more closely related to
one another
^All three of these trees are equivalent: trees can be swiveled/rotated about their nodes without
changing the expressed relationships; the important thing to look at is the pattern of common ancestry
Phylogenetic Reconstruction and Subsequent Taxonomic Considerations:
to determine relationships:
1) collect data: if trying to compare organisms to establish a relationship between/tree, not confined to
living things can refer to paleontological record (fossils) but also not confined to physical
characteristics of the body (morphology/anatomy) can collect behavioral data, molecular data (DNA
sequences)
2) analyze data to produce trees; many methods of analysis
3) taxonomic classification: based on those trees, can rearrange taxonomy if necessary
Cant you reconstruct phylogeny by just grouping organisms by overall similarity? first glance, it would
seem you dont need to worry about all the different methods of analysis/that organisms that share a
more recent common ancestor would always be more similar to one another (compared to other
organisms that dont share the common ancestor)
Sources of error: if using overall similarity to determine relationship
1) Symplesiomorphies: shared ancestral characteristic, arose earlier than in the stage where you are
establishing relationships didnt arise late enough to be used in the analysis
ex: mouths: showed up early, doesnt help to see mouths of monkeys and humans these similarities
can swamp out the similarity trying to be made
can detect a symplesiomorphies by using a proper outgroup in your analysis
2) unequal evolutionary rates among lineages
^hypothesis: things like alligators and birds are more closely related than crocodiles to a lizard or snake
crocodilians and birds share a common ancestor and therefore are more closely related than crocodiles
to lizards and snakes unequal evolutionary rates produced this: in the time since crocodilians and birds
shared a common ancestor, crocodiles didnt change much but birds evolved very rapidly look very
different misleading
3) homoplasy = any phenomenon where two different lineages change in the same way independently
and wind up changing the same character state, not because they inherited it from a common ancestor,
but happened independently; 3 subtypes that cause this:
a) convergent evolution (analogous characters)
b) parallel evolution
c) reversals
many credible methods of phylogenetic analysis attempt to identify synapomorphies that reveal
relationship/common ancestry
dont just take any tree building method available on the internet or elsewhere and apply it to your data
without knowing more about the assumptions of the method!
the method of analysis matters a lot: your tree(s) will be garbage if you dont use a correct method of
analysis
a given data set is informative for investigating relationships only within a certain range; no data set is
applicable to all levels of relationship
*overall similarity does not = relationship trees that are only based on overall symmetry are less
reliable: evolutionary rates assumed, parallel evolution
Cladistics:
cladistic analysis = best way to analyze data
many different methods of analysis, but cladistics favors methods that try to establish a tree based on
shared derived characteristics
once tree is made, cladistics favor a classification scheme that directly reflects it: accomplished by
constructing monophyletic groups
Mono-, Para-, and Polyphyletic Groups:
monophyletic group = a group that contains all of the descendants of a given ancestor
all taxonomic classification based on monophyletic groups is the only one that is completely equivalent
with the phylogenetic tree of relationships
paraphyletic group = a group that has a shared common ancestor, but omits one or more of the
descendants
polyphyletic group = a group that has no common ancestor not shared with groups
even less favored by cladistics than the paraphyletic group
^para: includes some descendents but excludes G, poly: C isnt even closely related to highlighted
organisms
^monophyletic groups allow you to make a tree from taxonomic categories: 1:1 relationship makes this
desirable
ancestor at base, includes all descendants: kingdom; first branch point: 2 phyla: 1) includes A, 2)
includes B, C, D, E, F, G phylum split into 2 classes: 1) B, 2) CDEFG; order: C, D and other = E,F,G; 2
families: one that includes E, other includes F,G
in this approach, directed correspondence between tree and taxonomy
if made tree of life: would have to have many taxonomic categories, but whatever groups we do have
should be monophyletic (express relationship)
ex: reptiles = paraphyletic group
importance of knowing the assumptions involved in a given method of analysis when constructing
hypotheses of relationship (phylogenetic trees)
especially the reasons that trees constructed using overall similarity can be problematic I went into
detail regarding the specific reasons that overall similarity does not necessarily indicate relationship (and
what we mean by relationship in the first place), so maybe I think this is important
other important topics: HW equilibrium conditions and how violation of the conditions leads to evolution,
speciation ppt