Evolution Flashcards
Two views of evolution
1) change in character traits of a population (Darwin, phenotypic change)
2) change in population allele frequencies (genotypic change)
Gene
1) region of DNA that codes for a specific polypeptide
2) can be regulatory, act as switches for gene expression, of express given trait
Genome
large recipe book in the form of amino acids (composed of genes)
Locus
Physical location of specific gene on a chromosome
Allele
Version of a specific gene at a given locus
Causes of change in allele frequencies
1) Natural selection
2) Genetic drift
3) Gene flow
4) Mutation
All are microevolutionary processes
Neutral theory
Kimura (1983)
1) Much of genetic variation is neutral, do not affect phenotype (occur by chance and not selection)
2) They either do not alter the product of genes OR affect non-coding regions of DNA
I.e. Traits may change, but they do not influence fitness
Evolutionary Biology pg. 17, 354
Is neutral theory subject to natural selection?
No.
Importance of neutral theory
Can look into ancestry through neutral mutations
Hardy-Weinberg Equilibrium
In the absence of disturbance, gene frequencies will remain constant
If you know allele frequencies in one generation, you can predict genotype frequency in next generation
Hardy-Weinberg Equilibrium assumptions
1) No evolution (no genetic drift, no gene flow, no natural selection, no mutation)
2) Random mating
Genetic diversity
Number and relative frequency of alleles in a population
What are the benefits of high genetic diversity
Ability to respond to change
Natural Selection
1) Evaluates alleles on basis of fitness
2) Occurs when one phenotype has higher fitness
3) There are three types of natural selection, directional, stabilizing, disruptive
Evolutionary Biology p. 60
Directional Selection
1) Allele frequencies change in one direction
2) Favors one extreme of trait distribution
3) Decreases genetic diversity
Stabilizing Selection
1) Alleles associated with mean trait values increase
2) No change in average trait value
3) Decrease in genetic diversity over time
4) Increase in trait and gene frequency over time
Disruptive Selection
1) Alleles associated with both extremes of a trait are favored
2) Distribution bifurcates
Sexual Selection
When there is variation in mating success based on different characters
Evolutionary Biology p. 60
Genetic Drift
1) Independent of fitness
2) Due to random chance
2) More likely in small populations
3) Leads to fixation and loss of alleles
4) Two causes: founder effect and genetic bottleneck
Founder effect
1) Small number of individuals found a new population, and new population does not reflect the allele frequency of source population
2) Also called peripatric speciation
Evolutionary Biology p. 234
Genetic bottleneck
Drastic and/or random reduction in population size (not due to differences in fitness)
Gene Flow
1) Movement of alleles from one population to another (immigration/emigration-functionally)
2) Equalizes allele frequencies between populations
3) Can restore lost alleles
Mutation
1) Production of new allele via damage or replication errors to DNA
2) Increases genetic diversity
3) Variable effect on fitness
Inbreeding
1) NOT a microevolutionary process
2) Changes genotype frequency and phenotypic expression
2) Does NOT change allele frequency
3) Homozygous recessive genotypes often decrease fitness
Speciation
1) enough microevolutionary processes occur to create a new species
2) formation of a new species from an ancestral species
3) two components: reproductive isolation and genetic divergence
Species
1) Evolutionary independent population
2) Distinguished by common characteristics
3) Three functional distinct definitions: biological, morphological, phylogenetic
Biological Species Concept
1) Reproductively isolated populations
2) Member of a species can interbreed and produce viable offspring
3) Can only be applied to organisms that we can observe breeding (can’t apply to fossils and extinct species or geographically isolated species, e.g. blue whales)
4) Hybridization is still allowed (not all or nothing)
Evolution p. 215
Morphological Species Concept
1) Based on differences in morphology
2) Widely applicable to fossils and both sexual and asexual organisms
3) Criteria is subjective and intra-specific variation can sometimes be greater than that of inter-specific (e.g. sexual dimorphism in buffleheads and zebras look different but can successfully interbreed)
Phylogenetic Species Concept
1) Based on ancestral analysis
2) Smallest identifiable group assigned species status (monophyletic group)
3) Can be applied widely
4) Unfortunately, there are only a few thorough phylogenies available
What is the relationship between reproductive isolation and genetic divergence
As isolation increases, so does divergence
Reproductively Isolating Mechanisms
1) How populations reproductively isolate
2) pre- and post-zygotic isolation
Pre-zygotic isolation
Pre-mating
1) Ecological Isolation (temporal isolation (Gryllus crickets reproducing at different times) or habitat isolation (soapberry bugs feeding on different plants))
2) Behavioral isolation (sexual isolation (female mate choice, song of finches) or pollinator isolation (plants are specialized to specific pollinator and there is no overlap))
Post-mating
1) Mechanical isolation (reproductive structures do not fit)
2) Copulatory isolation (female not stimulated by male-occurs in some fly species)
3) Gametic isolation (failure to fertilize, e.g. in larval spawning by abalone or corals).
Evolutionary Biology p. 221
Post-zygotic isolation
1) Extrinsic: hybrids are formed but have low fitness for environmental reasons
a) ecological inviability
b) behavioral sterility
2) Intrinsic (low hybrid fitness is independent of environmental context
a) hybrid inviability (reduced survival due to genetic incompatibility)
b) hybrid sterility (reduced production of viable gametes)
Mechanisms of speciation (5)
1) Allopatric speciation
2) Dispersal/colonization
3) Vicariance
4) Sympatric speciation
5) Polyploidy
Allopatric Speciation
1) Populations become geographically isolated
2) Gene flow ceases
3) Populations diverge genetically (via drift, selection, and mutations) making them separate species
4) Results from dispersal/colonization and vicariance
Dispersal/Colonization
1) Small number of individuals disperse to new habitat
2) Founder effect increases genetic drift
3) If new environment differs, selection may occur
4) Populations diverge genetically (via drift, selection, and mutations) making them separate species
4) Common on islands
5) Type of allopatric speciation
Vicariance
1) Large populations split into two or more sub-populations
2) Usually occurs due to emerging geographic barriers
3) New populations are genetically isolated (no gene flow)
4) Populations diverge genetically (via drift, selection, and mutations) making them separate species
5) Type of allopatric speciation
Evolutionary Biology p. 236
Sympatric Speciation
1) Speciation without geographic isolation
2) Occurs when natural selection overwhelms gene flow (via temporal and behavioral isolation and polyploidy; e.g. soapberry bugs, beak length correlates with fruit size, they mate where they eat)
3) Example: linkage disequilibrium between beak size and color, strong mate choice for same color, reinforces mating of same color and beak size, speciation occurs
4) Example, band-rumped storm petrel has split in sympatric populations with separate breeding seasons
Evolutionary Biology p. 238-40
Example of behavioral disruption of mating
1) Soapberry bugs specialize on different types of food (based on mouth morphology). As a result they occupy different plants and do not mate with bugs on other plants.
2) Apple maggot exploits two different types of apples that have different peak fruiting times. Therefore the timing of emergence of adults differs resulting in reproductive isolation.
Example of allopatric speciation
1) Galapagos finches colonizing new islands and then undergoing speciation.
2) Turtles and the Isthmus of Panama.
Polyploidy
1) Special case of sympatric speciation
2) Usually caused by mutation that creates extra chromosome copy
3) Two types: autoploidy and alloploidy
Example of polyploidy
Allium (garlic and onion) species have evolved via polyploidy.
Autoploidy
Mutation doubles chromosome number, resulting in individuals that can only self-fertilize to produce viable offspring
Alloploidy
Two different species mate, there is a mutation in the offspring that doubles the chromosome number, allows for self-fertilization
Hybrids
1) Formed when isolated populations reconnect if sufficient genetic divergence has occurred through pre- and post-zygotic isolation
2) If viable hybrids form, they may have differential fitness than both parent species
Hybrid Zones
1) Areas of overlap where interbreeding of separate species occurs
2) If hybrid fitness is higher than either parent species, homogenization occurs
3) If hybrid fitness is is lower than either parent species the hybrid zone narrows and reinforcement of separate traits in parent species emerges
Hybrid zone with high hybrid fitness, description and example
1) Fusion occurs reproductive barriers are weakened, there is substantial gene flow between species, this can cause the parent species to become a single species over time.
2) Cichlid fish in Lake Victoria, as turbidity increases, less selection for male coloration from females so hybrid fitness
3) Hypothetical example: Two species of fish where females select for a blue and a red color morph. Waters become more turbid so color is more difficult to see. Hybridization occurs and the hybrids are better equipped to exploit resources in turbid environment and mate choice disintegrates due to turbidity. This would cause fusion of the two species.
Hybrid zone with low hybrid fitness, description and example
1) Reinforcement of barriers between species occurs and rate of hybridization decreases.
2) example European flycatchers. In sympatric species, females will never mate with other species, so female choice selects for divergence in male secondary sexual characteristics, so hybrids with intermediate characteristics have low fitness.
Macroevolution
dramatic and rapid evolution
Two views on the pace of evolution
1) gradualism
2) punctuated equilibrium
Gradualism
1) genetic change is continuous through time
2) these genetic changes lead to changes in phenotype and eventually to new species
3) predicts transitional forms (e.g. cetacean evolution)
Punctuated equilibrium
1) change occurs in short bursts
2) long periods of minimal change
3) few or no transitional forms
4) rapid appearance of new species
Orthologs
1) Genes that diverge from a common ancestral gene by phylogenetic splitting at the organismal level
\_\_\_\_\_\_\_\_\_ A_Species 2 A | \_\_\_\_\_\_\_\_X | |\_\_\_\_\_\_\_\_\_ B_Species 2 | ---( ) | \_\_\_\_\_\_\_\_\_\_\_\_\_ B_Species 1 | | | \_\_\_\_X B |\_\_\_\_\_\_\_\_\_\_\_\_\_ A_Species 1
Evolutionary Biology p. 40
Paralogs
1) Genes that originate from an ancestral gene duplication
Evolutionary Biology p. 40
Cambrian Explosion
1) Major radiation of multicellular animals, beginning of Paleozoic era (540 mya)
2) Within 40 million years almost all major animal phyla appear
Homeotic Genes
1) found in all multicellular organisms
2) control development (transcription)
3) can be specific to regions of the body
4) are turned on or off by regulatory genes
5) highly conserved (extreme selective pressure to maintain them, similar sequences in very different organisms)
Hox genes and complexity
1) More hox genes typically yields more complex organisms
2) Gene duplication mutations produce more Hox genes (these hox genes are paralogs)
3) influenced by heterochrony and heterotropy
Hox genes
Control development (growth and body region)
Heterochrony
difference in duration of expression, e.g. skull formation in chimps and humans (chimps have longer duration development of jaw bones)
Heterotropy
difference in location of expression
Example of heterochrony and heterotropy
1) Fin versus limb bud growth in vertebrates
2) both mouse limbs and fish fins form from limb buds in embryo
3) hoxd-11 and shh are genes that regulate direction of limb growth
4) hoxd-11 is expressed early in development, but there is differential expression of this gene later in development, leading to different shapes of limbs and fins
Phylogenetic trees
1) Series of branches (population), nodes (forks where ancestor splits), and tips (extinct and extant species) that trace phylogenetic relationships
2) Sister taxa occupy linked by nodes
Clade
Ancestor and all descendants (starts at node)
Ingroup
Member of clade
Evolutionary Biology p. 36
Outgroup
1) Taxon with common ancestor that does not fit into clade
Importance: Give nearest reference point to understand clade
Evolutionary Biology p. 36
Monophyletic Group
Complete clade
Paraphyletic Group
Clade with erroneous omissions (common ancestor without all descendants)
Polyphyletic Group
Clade with erroneous inclusion (different common ancestors)
What can be used to create phylogenies
1) Fossil record
2) Comparative molecular biology (molecular clocks and orthologs)
3) Comparative anatomy and embryology
4) Biogeography
Molecular Clock
1) When differences in DNA sequences can be used as a marker for how much time has passed
2) Must be specific to character traits and taxa
Evolutionary Biology p. 42
Sympleisomorphy
Ancestral trait in cladistics
Snyapomorphy
1) Shared derived trait in cladistics (where nodes form)
2) Ingroups share synapomorphy
Evolutionary Biology p. 402
Problems with cladistics
1) Must select good, homologous traits for character matrix
2) Homoplasies can occur: when analagous traits are included in cladistics, as opposed to homologous traits
3) Many possible tree formations (but select tree with maximum parsimony)
Homoplasies
1) The independent evolution of a character or character state in different taxa
2) Trait is not inherited from a common ancestor
3) Convergent evolution, parallel evolution, and evolutionary reversal
4) E.g. wings are present in birds and bats but they are analogous, not homologous
Evolutionary Biology p. 47
Adaptive Radiation
1) Appear in phylogenies as polytomies
2) Rapid speciation events
3) Occurs as a result of colonization events, mass extinctions, morphological innovations
4) E.g., Galapagos finches
Evolutionary Biology p. 50
Polytomy
1) Node with multiple branches
2) Star phylogeny (indicative of adaptive radiation)
Colonization event
1) Habitat unoccupied by a competitor is colonized by a species
2) Colonizing species quickly radiates to exploit new resources
Mass extinction
1) Removes major competitors from the system
2) Allows surviving species to exploit resources that were previously occupied
Morphological innovation
Allows new resources to be exploited
Whale evolution
Order Cetartiodactyla, closest relative is hippo. All Cetartiodactyls have astralagus but cetaceans. Why? Most likely explanation is that the astralagus was lost in cetaceans (rather than lost for hippos and cetaceans, adn then regained in hippos) based on idea of maximum parsimony. Supported by discovery of ambulocetus (early cetacean without astralagus)
Timeline for life on earth
1) Eons: Hadean, Archaean, Proterozoic, Phanerozoic (includes Paleozoic, Mesozoic, Cenozoic eras)
2) PreCambrian includes Hadean, Archaean and Proterozoic.
3) Cambrian (in Paleozoic era) yields large multicellular organisms
Hadean eon
1) Formation of earth
2) Cooling of earth, formation of oceans from rain
3) Idea of Primordial Soup (Miller), life formed from rich organic compounds
Archaean eon
1) Origin and radiation of prokaryotes
2) Photosynthesis begins and oxygen increases in atmosphere along with aerobic respiration
Proterozoic eon
1) Origin and radiation of eukaryotes
2) Larger cells with distinct nucleus and mitochondria (mitochondria came from a eukaryote that ingested bacteria and adopted aerobic respiration)
3) Chloroplasts appear
4) Simple multicellular organisms
Phanerozoic eon
1) Diversification of multicellular organisms
2) Broken up into 3 eras: Paleozoic (ancient animals), Mesozoic (dinosaurs), and Cenozoic (mammals)
Paleozoic era
1) era in the Phanerozoic eon
2) Ancient animals
3) Cambrian explosion
4) First vertebrates
5) Diversification of fish
6) Invasion of land
7) Extensive fern forests
8) First gymnosperms
9) Permian extinction
Proximate causes
Immediate, mechanical causes of biological phenomena, e.g. what causes a male bird to sing? Testosterone and other hormones, the syrinx, and the operation of certain centers in the brain.
Ultimate causes
Historical causes, especially via natural selection, e.g. what events led to birds singing? Past individuals who were inclined to sing were more successful in attracting mates.
Horizontal gene transfer
Transfer and change of genes from one taxon to another through non-reproductive means.
E.g. the evolution of mitochondria in cells when eukaryote consumed bacterium, aphids acquired gene for synthesizing carotenoids from fungi, viruses to humans
E.g., HGT between mitochondria and nucleus
Evolutionary Biology p. 38, 350
Vertical gene transfer
Transfer of genes through reproductive means, parent to offspring
Anagenesis
Evolutionary change of features within a single lineage (species)
Evolutionary Biology p. 33
Cladogenesis
1) Branching of a lineage into two or more descendant lineages.
2) Following cladogenesis, anagenesis occurs between each lineage causing divergence
Evolutionary Biology p. 33
Sister groups (phylogeny)
Two clades that originate from a common ancestor
Evolutionary Biology p. 33
Homolog
Shared traits among taxa that share a common ancestor, e.g. forelimb skeletons of tetrapod vertebrates
Evolutionary Biology p. 35
Analog
Shared trait among non-related taxa, typically related to exploitation of similar niche or resource, e.g. wings in bats and birds
Evolutionary Biology p. 35
Hybrid speciation
When various phenotypic and DNA markers throughout the genome reveal two ancestral sources
Evolutionary Biology p. 38
Time of divergence between species
D=2rt, r=rate at which these genes evolve in given taxa
the factor 2 appears since this gene has diverged in two lineages
Evolutionary Biology p. 42
Evidence for evolution
1) hierarchical organization of life
2) homology
3) Embryologic similarities
4) Vestigial characteristics
5) Convergence
6) Suboptimal design
7) Geographic distributions
8) Intermediate forms
Evolutionary Biology p. 44
Mosaic evolution
1) Evolution of different characters at different rates within a lineage
2) It tells us that a species evolves not as a whole but piecemeal
3) Many of its features evolve more or less independently
4) Every species is a mosaic of plesiomorphic and apomorphic characters
Evolutionary Biology p. 44
Plesiomorphic
Ancestral