Natural selection acts on individuals (survival and reproductive success). Evolution acts on populations. Concept 23.

1 Population genetics provides a foundation for studying evolution Evolution is studied through population genetics Modern evolutionary theory integrates Darwinian selection with Mendelian inheritance Continuum (qualitative) traits are influenced by genes at multiple loci that each follows Mendelian inheritance. This Mendel-Darwin reconciliation gave rise to population genetics. Themes (Modern Synthesis) Populations are units of evolution Natural selection leads to adaptive evolution Gradualism allows minute changes to accumulate over time A populations gene pool is defined by its allele frequencies. Population = group of individuals living in same area who can mate and produce fertile offspring Individuals near populations center are more closely related to one another Gene Pool = total aggregate of all alleles of all genes at all loci in a population Fixed gene = gene for which only a single allele exists in a population Allele frequency = copies of allele in population / total number of alleles in population p-frequency = 2(homozygous-p) + heterozygous-p q-frequency = 1 - (p-frequency) The Hardy-Weinberg Theorem describes a non-evolving population. Hardy-Weinberg Theorem Frequencies of alleles and genotypes in gene pool remain constant over generations Given random mating: Probability of pp (homozygous p) is (p-frequency)^2 Probability of qq (homozygous q) is (q-frequency)^2 Probability of pq (or qp) (heterozygous) is 2pq This must also equal 1. Thus: p^2 + 2pq + q^2 = 1 This formula allows conversion from allele frequency to genotype frequency Five Conditions Extremely large population size (to avoid chance fluctuations genetic drift) Because rules of probability might not correspond to reality on a small scale No gene flow (AKA altering allele frequency due to transfer of genes due to migration) No mutations (altering allele frequency through creation of new alleles) Random mating (altering allele frequency because of reproductive fitness) No natural selection (altering allele frequency because environmental fitness) Evolution results when any of these five conditions is not met. Evolution Evolution = change in allele frequency Natural Selection = due to non-chance events o Some organisms naturally more fit o Leads non-random mating Genetic Drift = due to chance events o Bottleneck Effect (nondiscriminatory event wipes out random portion of population) o Founder Effect (nonrepresentative set of individuals get separated and start new population with different allele frequency from original) Gene flow = migration in and out of population Mutation = forms new alleles and allows adaptation

Concept 23.2 Mutation and sexual recombination produce the variation that makes evolution possible New alleles of genes originate only by mutation. Mutation = change in nucleotide sequence Only mutations in gametes (not somatic cells) passed on Point mutations usually harmless (noncoding DNA, redundancy leads to same amino acid, etc.) Can be harmful (altering gene expression) Can be beneficial by altering gene expression as well, especially when environment changing Chromosomal mutations (delete or rearrange multiple loci) almost always harmful Cases were beneficial: Translocation causes good gene linkage Shuffled up exons (coding DNA) Gene duplication adding to genome Transposons (shifting sequences of DNA) can add to genome Activation of previously inactive helpful gene Mutation rates low for animals and plants; high for most microorganisms Sexual recombination is more prevalent for genetic variation, without changing allele frequency Concept 23.3 Natural selection, genetic drift, and gene flow can alter a populations genetic composition (See outline of Evolution above.) Concept 23.4 Natural selection is the primary mechanism of adaptive evolution Quantitative characters = vary along continuum (tall to short) and usually due to polygenic inheritance Discrete characters = vary either/or and are usually determined by a single locus o Phenotypic polymorphism = two discrete phenotypes (morphs) Genetic variation is measured by: o Gene variability/average heterozygosity = the average percent of gene loci that are heterozygous o Nucleotide variability = average level of difference between nucleotides in individuals o Typically, average heterozygosity (gene variability) is greater than nucleotide variability because even a single different nucleotide makes the entire gene different. Since there are fewer genes than nucleotides, the percentage of heterozygous genes works out be a lot higher. Geographic variation = difference in phenotype or genotype between populations or subgroups o Caused by slightly different local environmental factors o Also caused by genetic drift (randomness) o Cline = graded change along a geographic axis (I.E. proportional to geographic gradient) May be because of gradually increasing/decreasing resource May also be intergrade zones between populations Evolutionary Fitness Reproductive success depends on more than survival of the fittest although competition for mating is important for most animals Relative fitness = contribution of a genotype to the next generation relative to alternative genotypes at same locus o Most successful is arbitrarily set at 1.0 o No offspring gives 0, even if long, healthy life o 0.8 would mean 80% as successful as most successful o However, phenotype, not genotype is subject of natural selection There are three modes of selection: directional, disruptive, and stabilizing. Directional selection = shifts frequency curve for a phenotypic character in one direction Disruptive selection = individuals at both extremes favored; gives m-shaped curve Stabilizing selection = favors intermediate variants; gives taller (less wide) curve Diploidy and Balancing Selection preserve genetic variation.

[] Sexual selection may lead to pronounced secondary differences between the sexes. [] Sex is an evolutionary enigma. Previously, the answer to why is sexual reproduction so common was answered by because it allows future adaptations. (Variation over sheer number of offspring.) However, natural selection only takes into account what is going on now. Therefore, the current answer is the Red Queen hypothesis. Because parasites are constantly evolving, sexual reproduction allows variation and thus resistance to more disease. Natural selection cannot fashion perfect organisms. Evolution limited by historical constraints (adapts existing features for new situations; birds cant have wings AND legs; only two pairs of limbs) Adaptations are often compromises (seals must walk and swim) Chance and natural selection interact (genetic drift vs. environmental adaptability) Selection can only edit existing variations (only variants, no new alleles on demand)

Microevolution = study of adaptive change in a population Macroevolution = study of evolutionary changes above species level Speciation = the origin of new species; source of biological diversity; lack of gene flow o Anagenesis (phyletic evolution) = changing of one species into another o Cladogenesis (branching evolution) = division of one species into multiple

Concept 24.1 The biological species concept emphasizes reproductive isolation Species traditionally defined through morphology; now defined through biochemistry, etc. Biological species concept defied through reproduction Species = group of populations whose members have potential to breed with each other to produce fertile offspring; largest set of populations in which genetic exchange is possible; interfertility, not similarity Reproductive barriers isolate the gene pools of biological species. Prezygotic Barriers = before fertilization o Habitat isolation (different habitats, even if same geographic area; i.e. land vs. water) o Behavioral isolation (different courtship behaviors) o Temporal isolation (breed during different times of day or seasons) o Mechanical isolation (parts dont fit) o Gametic isolation (fertilization cannot occur; no zygote formed) Postzygotic Barriers = sperm does fertilize ovum, but other challenges o Reduced hybrid viability = fetus aborted at some embryonic stage o Reduced hybrid fertility = hybrids are infertile o Hybrid breakdown = next generation after hybrids feeble or sterile The biological species concept has some major limitations. Fossils cannot be tested based on ability to reproduce Entirely asexually reproducing species typically classified based on morphology Bacterial conjugation is different from sexual combination and does not distinguish species Evolutionary biologists have proposed several alternative concepts of species. Ecological species = in terms of niche (job in environment), accommodates asexual and sexual species Paleontological species = focuses on morphologically discrete species only known from fossil record Phylogenic species = set of organisms with a unique genetic history o Physical characteristics of molecular sequences o Sibling species = appear identical morphologically but different genetically o Confirms if phylogenic distinction agrees with reproductive incompatibility Morphological species = unique set of structural features (most practical) o Can be applied to sexual or asexual, even without knowing anything about gene flow o Problem: subjective criteria Concept 24.2 Speciation can take place with or without geographic separation Allopatric speciation = geographic barriers lead to origin of species o Significance of barrier depends on ease of movement (fly vs. walk) o Gene pools diverge through mutations, sexual selection, selective pressures, genetic drift o Most new species quickly perish; few survive to become new species Sympatric speciation = occurs in overlapping populations when biological factors restrict gene flow o Reproductive barriers must evolve Plants Autopolyploid mutant = individual that has more than two sets of chromosomes (from same species) due to error in mitosis/meiosis; can sometimes mate with other to produce fertile offspring but cannot mate with normal diploid species

Allopolyploid mutant = mating of two different species; chromosomes from different species o Hybrids are usually sterile but future generations can propagate asexually o Cannot breed with parent species; thus, they are a new species o Many plants important for agriculture are like this (wheat, oats, cotton, etc.)

Animals Mate preference can restrict gene flow (sexual selection in polymorphic population) Individuals exploit resources (often food) not used by parents Adaptive radiation = explosive formation of new species (many diversely adapted species from common ancestor when new environmental opportunities arise) o Example: Lake Victoria, 600 species of very similar fishes; nonrandom mating (females will only mate with certain color) o Occurs when organisms enter new area or when extinction opens up ecological niches

Researchers study the genetics of speciation. The tempo of speciation is important. Common trend: species suddenly appears, doesnt change for millions of years (stasis), and just as suddenly disappears Explanation: appearing suddenly could actually take tens of thousands of years but leave no fossil record o During equilibrium, changes in behavior, internal anatomy, and physiology may not leave a fossil record Concept 24.3 Macroevolutionary changes can accumulate through many speciation events