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Gene
A segment of DNA that carries the instructions for a trait or protein
Allele
One of the alternative versions of a gene (e.g. a flower-color gene may have a purple allele and a white allele)
Locus
The physical location of a gene on a chromosome
Chromosome
A long coiled molecule of DNA that carries many genes
Genotype
The combination of alleles an individual carries for a gene (such as AA Aa or aa)
Phenotype
The observable trait produced by an organism's genotype interacting with its environment
Dominant allele
An allele that shows up in the phenotype even when only one copy is present (written as a capital letter)
Recessive allele
An allele expressed only when two copies are present; it is masked by a dominant allele (written lowercase)
Homozygous
Having two identical alleles for a gene (AA or aa)
Heterozygous
Having two different alleles for a gene (Aa)
Trait
A measurable characteristic of an organism (like height or color) that can differ among individuals
Heritable
Able to be passed from parent to offspring through genes; only heritable traits can evolve by natural selection
Population
A group of individuals of the same species living in the same area and interbreeding
Species
A group of organisms that can interbreed and produce fertile viable offspring and are reproductively isolated from other groups
Evolution
A change in the heritable characteristics (allele frequencies) of a population across generations
Descent with modification
Darwin's phrase for evolution — all species share common ancestors and accumulate changes over time
Common ancestor
An ancestral species from which two or more later species descended
Natural selection
The process by which individuals with heritable traits better suited to the environment survive and reproduce more; the main mechanism of adaptive evolution
Requirement 1 for natural selection
There must be variation among individuals in the population
Requirement 2 for natural selection
Some of that variation must be heritable (genetically passed on)
Requirement 3 for natural selection
More offspring are produced than can survive (a struggle for existence)
Requirement 4 for natural selection
Survival and reproduction are non-random — certain traits give a reproductive advantage
Fitness
An individual's reproductive success — how many surviving fertile offspring it leaves (not strength or speed for its own sake)
Adaptation
A heritable trait shaped by natural selection that improves survival or reproduction in a particular environment
Artificial selection
Humans choosing which organisms reproduce to promote desired traits (like breeding low-fat corn or dog breeds)
Variation
Differences in traits/alleles among individuals; the raw material that evolution acts on
Misconception - individuals evolve
False — individuals do not evolve during their lifetime; only populations evolve across generations
Misconception - evolution has a goal
False — evolution is not goal-directed or progressive and does not strive toward perfection
Misconception - traits arise because needed
False — mutations occur randomly and the environment then selects among the variation that already exists
Evidence for evolution
Fossils and transitional forms; homologous structures; shared genes and development; vestigial structures; biogeography
Transitional fossil
A fossil showing intermediate features between an ancestral group and its descendants
Tiktaalik
A famous transitional fossil (a fish-to-tetrapod intermediate) with fins containing wrist-like bones and a neck; documents the move onto land
Homologous structures
Body parts inherited from a common ancestor with the same developmental origin even if they now have different functions (like the tetrapod limb bones in a human arm bat wing and whale flipper)
Analogous structures
Body parts that look or function similarly but did NOT come from a common ancestor; they arose independently (like a bird wing vs an insect wing)
Vestigial structure
A reduced nonfunctional remnant of a structure that was useful in an ancestor (like the pelvic and leg bones inside a boa constrictor or the human tailbone)
Homology
Similarity between species due to shared ancestry; the key evidence of evolutionary relatedness
Homoplasy
Similarity between species that is NOT from common ancestry but from convergent evolution (an analogous trait)
Convergent evolution
When distantly related species independently evolve similar traits because they face similar environments (produces analogous structures)
Divergent evolution
When related lineages become increasingly different over time and a shared structure takes on new functions
Biological species concept
Defines a species as populations whose members can interbreed and produce fertile viable offspring but cannot breed with other groups
Reproductive isolation
The inability of members of different species to successfully interbreed; maintained by reproductive barriers
Prezygotic barrier
A reproductive barrier that blocks mating or fertilization before a zygote can form
Habitat isolation
A prezygotic barrier in which two species live in different habitats and rarely encounter each other
Temporal isolation
A prezygotic barrier in which two species breed at different times of day season or year
Behavioral isolation
A prezygotic barrier in which different courtship rituals or mating signals keep species from attracting each other
Mechanical isolation
A prezygotic barrier in which reproductive structures are physically incompatible
Gametic isolation
A prezygotic barrier in which sperm and egg are chemically incompatible and cannot fuse
Postzygotic barrier
A reproductive barrier that acts after fertilization by making hybrids unfit
Reduced hybrid viability
A postzygotic barrier in which hybrid offspring die early or are weak
Reduced hybrid fertility
A postzygotic barrier in which hybrids survive but are sterile (such as a mule)
Hybrid breakdown
A postzygotic barrier in which first-generation hybrids are healthy but their offspring are weak or sterile
Speciation
The evolutionary process by which one species splits into two or more new species
Allopatric speciation
New species forming when a population is split by a geographic barrier (such as squirrels separated by the Grand Canyon)
Sympatric speciation
New species forming without geographic separation while living in the same area (often by polyploidy or a host shift)
Polyploidy
Having one or more extra full sets of chromosomes; can instantly create a new plant species in the same area (as in the potato which doubled its chromosomes)
Host shift speciation
Sympatric speciation when a population specializes on a new host and stops interbreeding (as in Rhagoletis flies on apple vs hawthorn trees)
Adaptive radiation
The rapid diversification of one ancestral species into many new species that exploit different niches (like Galapagos finches or Anolis lizards)
Hybrid zone
A region where two species meet and interbreed producing some hybrid offspring
Reinforcement
A hybrid-zone outcome where selection strengthens prezygotic barriers because hybrids are unfit so the species stay separate
Fusion
A hybrid-zone outcome where weak barriers let two species merge back into one
Stability (hybrid zone)
A hybrid-zone outcome where hybrids keep being produced over time without fusing or fully separating
Gradualism
The model that species evolve slowly and steadily through small changes over long periods
Punctuated equilibrium
The model that species stay mostly unchanged for long periods then change rapidly in short bursts often during speciation
Population genetics
The study of allele and genotype frequencies in populations and how they change over time
Gene pool
All of the alleles of all genes in every individual of a population
Allele frequency
The proportion of a given allele among all copies of that gene in a population
Genotype frequency
The proportion of individuals in a population that have a particular genotype
Microevolution
A change in allele frequencies within a population from one generation to the next
Hardy-Weinberg principle
The idea that in a non-evolving population allele and genotype frequencies stay constant across generations; used as the baseline null model
Hardy-Weinberg allele equation
p + q = 1 where p is the dominant allele frequency and q is the recessive allele frequency
Hardy-Weinberg genotype equation
p squared + 2pq + q squared = 1 for the three genotype frequencies
p squared
The frequency of the homozygous dominant genotype
2pq
The frequency of the heterozygous genotype
q squared
The frequency of the homozygous recessive genotype (the only genotype directly visible under full dominance)
Hardy-Weinberg condition 1
No mutation (no new alleles are being created)
Hardy-Weinberg condition 2
Random mating (no mate choice or inbreeding)
Hardy-Weinberg condition 3
No natural selection (all genotypes survive and reproduce equally)
Hardy-Weinberg condition 4
Very large population size (so chance/drift is negligible)
Hardy-Weinberg condition 5
No gene flow (no alleles entering or leaving the population)
Solving Hardy-Weinberg problems
The visible recessive phenotype equals q squared; take its square root to get q; then p = 1 minus q; then compute 2pq and p squared
HW example - heterozygotes
If the recessive allele frequency q = 0.3 then p = 0.7 and heterozygotes = 2pq = 0.42
HW example - homozygous dominant
If the recessive allele frequency is 0.8 then the dominant allele is 0.2 and homozygous dominant = p squared = 0.04
HW example - disease frequency
If 1 in 2500 people are homozygous recessive then q squared = 0.0004 so q = 0.02 and p = 0.98
Why Hardy-Weinberg matters
It predicts a non-evolving population so any deviation from it means the population is evolving at that gene
Deviation from Hardy-Weinberg
A sign that one of the five conditions is violated and the population is evolving at that locus
Sources of genetic variation
Mutation (makes new alleles); recombination and independent assortment (reshuffle alleles); gene flow (brings in alleles)
Why heritable variation is required
Natural selection can only change a population across generations if the favored trait can be inherited
Mutation
A random change in DNA sequence; the ultimate source of all new alleles
Genetic drift
Random change in allele frequencies due to chance events; strongest in small populations and not adaptive
Bottleneck effect
A form of genetic drift where a disaster sharply reduces population size leaving survivors with a random unrepresentative set of alleles
Founder effect
A form of genetic drift where a few individuals start a new isolated population carrying only part of the original variation
Gene flow
The movement of alleles between populations through migration and interbreeding (such as a baboon leaving to join another troop)
Effect of gene flow
It increases genetic variation within a population and makes different populations more similar to each other
Effect of genetic drift on variation
It decreases variation within a population and increases genetic differences between populations
Non-random mating
Choosing mates based on traits or inbreeding; it changes genotype frequencies (for example inbreeding raises the number of homozygotes)
The five evolutionary forces
Natural selection; genetic drift; gene flow; mutation; non-random mating
Only adaptive force
Natural selection is the only evolutionary force that consistently produces adaptation (a better fit to the environment)
Directional selection
Selection that favors one extreme of a trait and shifts the whole population toward it (like lizards evolving shorter legs on treeless islands)
Stabilizing selection
Selection that favors the intermediate phenotype and removes both extremes (like a population of butterflies all becoming medium gray or average human birth weight)
Disruptive selection
Selection that favors both extremes of a trait against the intermediate and can split a population into two