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Today’s Plan: 2/25/11 Bellwork: Go over test/fly counts (30 mins) Amino Acid Sequence and Evolution Lab (30 mins) Begin Natural Selection Notes (the rest of class) Pack/Wrap-up (last few mins of class) Today’s Plan: 2/26/10 Bellwork: Finish Flies/Compile Class Data (30 mins) Sexual Selection Video Pack/Wrap-up (last few mins of class) Today’s Plan: 3/1/10 Bellwork: Go over lab/Do PTC (15 mins) AP Lab 8: Population Genetics and Evolution-Part B Genetic Drift (20 mins) Finish video Continue Notes-if time Pack/Wrap-up (last few mins of class) Today’s Plan: 3/2/10 Bellwork: Settle in/Grab cards (5 mins) Finish Lab (30 mins) Continue notes (25 mins) Pack/Wrap-up (last few mins of class) Today’s Plan: 3/3/10 Finish Notes (30 mins) Restriction Mapping Exercise (the rest of class) Pack/Wrap-up (last few mins of class) Today’s Plan: 3/4/10 Bellwork: Finish Restriction Mapping (20 mins) Finish Nat. Sel Notes (20 mins) HB fun week (the rest of class) Today’s Plan: 3/5/10 HB Stations-the entire period! Before Natural Selection Recall that Darwin wasn’t the 1st to think about how species have changed over time Aristotle’s Scala Naturae grouped species with similar “affinities” together Linnaeus came up with Binomial Nomenclature and did much classifying based on physical similarity Cuvier noted that fossils of species differed significantly from more modern forms (proposed the idea of Catastrophism-that changes happened b/c of catastrophic events, and not gradually) Lamarck suggested that use/disuse and will could change an organism’s body to fit the environment (he thought that acquired traits were heritable) Malthus also discussed population limits Darwin bred pigeons for various traits (artificial selection) Recall that besides thinking about species change, others before Darwin worked with how the planet changed Hutton proposed that geologic features were the result of gradual changes that are still occurring today Lyell took this a step further and proposed his principle of Uniformitarianism-mechanisms of change are constant over time Figure 24-1 Natural Selection Aka “Descent with Modification” was Darwin’s proposal for how species change over time and was the result of careful ponderance over his Galapagos Island collections Darwin’s main focus was on adaptations that allowed species to survive better in their environments-finches had beaks adapted to their food source Recall that while Darwin came up with the idea first, Alfred Russel Wallace also had the same ideal later, with no knowledge of Darwin’s work The term Natural Selection was coined by Darwin’s friend, T.H. Huxley, who was called “Darwin’s bulldog” because he staunchly defended Darwin’s hypothesis Darwin’s Observations and Inferences Observations: Inferences (Summary of Natural Selection’s Mechanism): Members of a population vary greatly in their traits Traits are inherited from parents to offspring All species are capable of producing more offspring than their environment can support Because of lack of resources, many offspring don’t survive Individuals whose inherited traits give them a higher probability of surviving and reproducing in an environment tend to leave more offspring (have greater reproductive success) This inequality means that favorable traits accumulate in populations over multiple generations Remember: Populations evolve, individuals don’t Traits influenced by natural selection must be heritable Environments are moving targets, so there’s no “perfect” and what’s good in one population isn’t necessarily good in another Current Directly Observable Evidence for Natural Selection Predation and Coloration in Guppies Drug-resistant HIV and other “superbugs” Pools of guppies w/high predation produce more drab colored males There are numerous examples of these “natural experiments” done by scientists It’s natural for some viruses or bacteria to be resistant, but when you treat, what’s all that’s left to reproduce? The Fossil Record We can see trends in the evolution of species Similar patterns of fetal development Homology (forelimb picture) Vestigial structures Looking at what we think happened to the geographic features of the planet to explain distribution of species (ex: how Pangea’s split allowed us to predict where we’d find certain types of fossils) Anatomical Features Biogeography Molecular Similarity Studies of DNA sequence and amino acid sequence can be used to construct “molecular clocks” that give us clues to which organisms diverged from one another, and tells us relatively how long ago the divergance occurred Figure 24-11 1. Large population of 2. Drug therapy begins killing 3. The mutant cells proliferate, 4. A second round of M. tuberculosis bacteria in patient’s lungs makes him sick. most M. tuberculosis. Patient seems cured and drug therapy is ended. However, a few of the original bacteria had a point mutation that made them resistant to the drug treatment. resulting in another major infection of the lungs. The patient becomes sick again. drug therapy begins but is ineffective on the drugresistant bacteria. The patient dies. Figure 24-3 Figure 24-9 Figure 24-8 Figure 24-5 Figure 24-7 Figure 27-7 Tree thinking vs. Convergent evolution In many cases, evolutionary trees are created in order to show how species evolved from common ancestors Sometimes, this happens b/c of adaptive radiation-when organisms evolve in a variety of directions in order to exploit different aspects of the environment Occasionally, organisms resemble each other, not because they’re related, but because some characteristics are advantageous regardless of their ancestry Ex: sugar gliders in Australia look like flying squirrels Figure 27-11 Figure 24-15b Darwinian evolution produces a tree of life. Common ancestor of all species living today The branches on the tree represent populations through time. All of the species have evolved from a common ancestor. None is higher than any other Sexual Selection This is a variation of natural selection where some traits persist, not because they’re advantageous, but because they’re attractive In some cases, the traits that evolve are disadvantageous, but continue to persist Intersexual Selection is based on the “female choice” model-the opposite sex chooses a mate Intrasexual Selection is based on competition within the sex for access to mates or resources that will attract mates Causes sexual dimorphism (variation between sexes) Figure 25-15 Beetle Scarlet tanager Lion Females Males During the breeding season, males of the Male scarlet tanagers use their bright beetle Dynastes granti use their elongated coloration in territorial and courtship horns to fight over females. displays. Male lions are larger than females lions and have an elaborate ruff of fur called a mane. Figure 25-14 Males compete to mate with females. Variation in reproductive success is high in males. Variation in reproductive success is relatively low in females. The survival of the fittest. . . Remember, fitness is relative, and “struggle” isn’t always direct conflict. Depending on which traits are favored, there are 3 ways in which natural selection can influence phenotypic variation Directional selection-one extreme phenotype is favored Disruptive selection-both extreme phenotypes are favored Stabilizing selection-average is favored Figure 25-3 Directional selection changes the average value of a trait. Normal distribution For example, directional selection caused average body size to increase in a cliff swallow population. Original population (N = 2880) Before selection Survivors (N = 1027) Low fitness During selection After selection Change in average value High fitness Change in average value Figure 25-4 Stabilizing selection reduces the amount of variation in a trait. For example, very small and very large babies are the most likely to die, leaving a narrower distribution of birth weights. Normal distribution Mortality Before selection Low High fitness Low fitness fitness During selection After selection Reduction in variation Heavy mortality on extremes Figure 25-5 Disruptive selection increases the amount of variation in a trait. For example, only juvenile black-bellied seedcrackers that had very long or very short beaks survived long enough to breed. Normal distribution Before selection Only the extremes survived High fitness Low fitness High fitness During selection Increase in variation After selection Only the extremes survived Evolution of Populations This is fueled by genetic variation For individuals, can be quantified using average heterozygosity (average % of genes for which an individual is heterozygous) For populations, you can directly compare individual karyotypes or gene sequences from each population Sometimes, the difference is dramatic, and sometimes the difference is a cline (gradual difference) This often exists b/c of geographic variation (isolation) Genetic Variation occurs for 2 reasons Sexual Reproduction Mutation is the ultimate source for most new genetic variations. Often these mutations are neutral, but occasionaly, you get an adaptive mutation. The rates at which mutation occurs varies between species. Hardy-Weinberg Useful for testing whether or not a population is evolving This is a mathematical model: p2+2pq+q2=1 p=frequency of the dominant allele q=frequency of the recessive allele When a population is in Hardy-Weinberg equilibrium, the equation works, but when populations are evolving, it is an inequation. Figure 24-10 Figure 25-1-1 A NUMERICAL EXAMPLE OF THE HARDY-WEINBERG PRINCIPLE Allele frequencies in parental generation: Allele A1 = p = 0.7 1. Suppose allele frequencies Allele A2 = q = 0.3 in the parental generation were 0.7 and 0.3. Gene pool (gametes from parent generation) 2. 70% of gametes in the gene pool carry allele A1, and 30% carry allele A2. A1 A1 0.7 0.7 = 0.49 p p = p2 A1 A2 0.7 0.3 = 0.21 q p = pq A2 A1 0.3 0.7 = 0.21 q p = pq 0.21 + 0.21 = 0.42 A2 A2 0.3 0.3 = 0.09 q q = q2 3. Pick two gametes at random from the gene pool to form offspring. You have a 70% chance of picking allele A1 and a 30% chance of picking allele A2. Figure 25-1-2 A NUMERICAL EXAMPLE OF THE HARDY-WEINBERG PRINCIPLE Frequency of A1A1 genotype is p2 = 0.49 Frequency of A1A2 genotype is 2pq = 0.42 Frequency of A2A2 genotype is q2 = 0.49 4. Three genotypes are possible. Calculate the frequencies of these three combinations of alleles. 5. When the offspring breed, 49% of offspring have the A1 A1 genotype. All will contribute A1 alleles to the new gene pool. 42% of offspring have the A1 A2 Genotype. Half of their gametes will carry the A1 allele and the other half will carry the A2 allele. 9% of offspring have the A2 A2 genotype. All will contribute A2 alleles to the new gene pool. imagine their gametes entering a gene pool. Calculate the frequencies of the two alleles in this gene pool. 6. The frequencies of A1 and A2 Allele frequencies in offspring gene pool 1 p = 0.49 + 2 (0.42) = 0.7 p = frequency of allele A1 q = 12 (0.42) + 0.09 = 0.3 q = frequency of allele A2 have not changed from parental to offspring generation. Evolution has not occurred. Genotype frequencies will be given by p2 : 2 p q : q2 as long as all Hardy-Weinberg assumptions are met. Conditions for Hardy-Weinberg No mutation Random Mating No natural selection Large population size No gene flow Rarely do all of these conditions exist at any given moment, but over time, populations tend to be in equilibrium Altering Gene Frequencies Genetic Drift-caused by small population size or random changes that make predicting gene frequency difficult. 2 examples: The founder effect-a small number of individuals are isolated from the larger group and have to reestablish a gene pool The bottleneck effect-catastrophic incidents drop population size quickly and dramatically In either case, genetic variation is lost, and harmful alleles can persist Gene Flow-occurs when genes transfer in and out of populations. Usually, this is negligible unless something causes any of the following factors to change dramatically: Immigration Emigration Figure 25-6 Figure 25-8 Lupines colonize sites and form populations. Gene flow reduces genetic differences among populations. Year 1: Seed establishes new population Year 2: Gene flow between source population and new population New population Source population Seed A1A1 A1A1 A1A1 A1A1 A1A2 New population Source population Gene flow A1A2 A1A1 A1A1 A1A2 A1A1 A1A2 A1A1 A1A1 A1A2 A1A2 Frequency of A1 = 0.90 Frequency of A1 = 0.50 Frequency of A1 = 0.83 Frequency of A1 = 0.67 Frequency of A2 = 0.10 Frequency of A2 = 0.50 Frequency of A2 = 0.17 Frequency of A2 = 0.33 Initially, allele frequencies are very different Gene flow causes allele frequencies to become more similar Preserving Genetic Variation Diploidy-Since organisms get 2 copies of each gene, recessive alleles can be preserved Balancing Selection-occurs when natural selection maintains 2 forms of a trait in a population The heterozygote advantage-sickle cell disease and malaria symptom resistance Frequency-Dependent selection-as a phenotype becomes more common, it loses its advantage Neutral Variation-occurs when mutation has little to no effect on phenotype or on reproductive success Why isn’t there a “perfect” organism Selection can only act on existing variations (and each intermediate step between phenotypes must be adaptive) You can’t scrap ancestral anatomy to build something new (see above statement) Adaptations are often compromises (multifunctionality means you have to choose a primary function. Ex: seals don’t have legs b/c they also swim) Chance, natural selection, and the environment have to interact Types of evolution Microevolution-evolution of allele frequencies within gene pools Macroevolution-patterns of evolution over long time spans (like the emergence of new species) The Biological Species Concept This is the classic definition of the term “species” put forth by Ernst Mayr A species is a group of populations whose members interbreed in nature to produce fertile offspring Species are held together by proximity and interbreeding Making new species Requires Reproductive isolation-barriers that prevent the production of viable offspring (remember that hybrids can exist, but are sterile: ligers, mules, etc) Prezygotic barriers-block fertilization Blocking mating Blocking the successful completion of mating Preventing successful fertilization Postzygotic barriers-prevent a hybrid from mating successfully Types of Prezygotic Mechanisms Habitat Isolation-2 species occupy different habitats Temporal Isolation-species breed at different times Behavioral Isolation-courtship rituals differ Mechanical Isolation-differences in shape/form prevent mating Gametic Isolation-the gametes may not be able to fuse Types of postzygotic Mechanisms Reduced Hybrid viability-parental genes prevent the hybrid’s survival Reduced Hybrid Fertility-sterility due to inability to produce normal gametes Hybrid Breakdown-Some hybrids can mate with one another, but their offspring are not viable Limitations of Biological Species It’s hard to evaluate the reproductive isolation of fossils, nor does it address species that reproduce asexually Other species definitions Morphological species concept-characterizes species by body shape and structure (can be applied to sexual and asexual reproducers, however this relies on subjective criteria) Ecological species concept-characterizes a species based on its ecological niche (also can be applied to sexual and asexual reproducers, and emphasizes the role of disruptive selection in species definition) Phylogenetic species concept-a species is defined by the smallest group of individuals that share a common ancestor (difficult to deterime the degree of difference required to separate one species from another) Allopatric Speciation “other country” speciation-occurs when species are geographically isolated Populations become divided and evolve differently because of different environments, genetic drift, and different mutations Remember that they’re not different species until they’re reproductively isolated. If the populations are put back together and can still mate, they’re not different species Figure 26-5 DISPERSAL AND COLONIZATION CAN ISOLATE POPULATIONS. Island Continent 1. Start with one continuous 2. Island population begins 3. Finish with two populations population. Then, colonists float to an island on a raft. to diverge due to drift and selection. isolated from one another. VICARIANCE CAN ISOLATE POPULATIONS. River 1. Start with one continuous 2. Isolated populations begin 3. Finish with two populations population. Then a chance event occurs that changes the landscape (river changes course.) to diverge due to drift and selection. isolated from one another. Sympatric Speciation “same country” speciation-occurs when organisms are in the same area but speciate Can occur via several mechanisms: Polyploidy-having an extra set of chromosomes Autopolyploid-more than 2 sets of chromosomes from a single species (failure of cell division) Allopolyploid-caused by an extra set of chromosomes via hybridization of 2 species (fertile when mating with one another only) Habitat Differentiation-when a subpopulation exploits a resource that the rest of the population doesn’t Sexual Selection-when different groups of females prefer different groups of males Figure 26-7 Soapberry bugs use their beaks to reach seeds inside fruits. Feeding on the fruit of a nonnative species Feeding on the fruit of a native species Nonnative fruits are much smaller than native fruits. Nonnative plant (small fruit) Native plant (large fruit) Evidence for disruptive selection on beak length Short-beaked population growing on nonnative plants Long-beaked population growing on native plants Figure 26-8 Diploid parent (Two copies of each chromosome) Tetraploid parent (Four copies of each chromosome) Meiosis Haploid gametes Diploid gametes (Two copies of (One copy of each chromosome) each chromosome) Mating Triploid zygote (Three copies of each chromosome) Meiosis Gametes The gametes of a triploid individual rarely contain the same number of each type of chromosome. When gametes combine, offspring almost always have an uneven (dysfunctional) number of chromosomes. Hybrid Zones When allopatric species come back into contact with one another, you get a hybrid zone There are several possibilities for what can happen in a hybrid zone Reinforcement-occurs when hybrids are less fit than the parent species Fusion-occurs when reproductive barriers are weak and the species become increasingly alike Stability-occurs when the hybrids persist Figure 26-11 Hybrids have intermediate characteristics. Hybrids inherit species-specific mtDNA sequences from their mothers. Townsend’s warbler All individuals have Townsend’s mtDNA Present range of Townsend’s warblers (in red) Townsend’s-hermit hybrid Pacific Ocean Individuals that look like Townsend’s warblers but have hermit mtDNA Some individuals have Townsend’s mtDNA, others have hermit mtDNA Present hybrid zones (where two ranges meet) Hermit warbler Present range of hermit warblers (in orange) All individuals have hermit mtDNA Speciation Rates Darwin originally believed that gradualism existed (species change at a slow, steady rate over time) From the fossil record, we now know that punctuated equilibrium exists (periods of equilibrium followed by periods of natural selection) This can happen very rapidly, and as little as 1 gene can make a species reproductively isolated Geologic Time This is a time scale that uses the fossil record to trace the major events in the planet’s history Dates are determined by dating fossils Relative Dating-accomplished via the law of superposition Absolute Dating-accomplished via radiometric dating Figure 27-5 HOW FOSSILIZATION OCCURS Earth’s History Earth is believed to have existed for 4.6 billion years Life on earth is believed to have originated 3.5 billion years ago. The first life forms were probably prokaryotes There have been 5 mass extinctions over the history of the planet, and in each, the dominant group of organisms was replaced by another group Figure 27-8a Figure 27-8b Figure 27-8c Figure 27-8d Figure 27-5 HOW FOSSILIZATION OCCURS Major events in Earth’s History: The earth cools 1st life forms Accumulation of atmospheric oxygen 1st eukaryotes Multicellular organisms Animals evolved Plants and fungi colonized land Land became colonized by other organisms Where would life come from? Recall that Miller and Urey tested Oparin’s primordial soup hypothesis and were able to create biochemicals Lab experiments since then have been able to form polymers in conditions similar to early earth RNA was probably the 1st genetic material These have been shown to be produced abiotically in the lab Recall that Ribozymes exist as well. Protobionts can self-assemble These are aggregates of abiotically produced molecules that form “membranes” and often can sustain chemical reactions (like a metabolism) What about Eukaryotic cells? One current hypothesis, the endosymbiant hypothesis tries to explain this Mitochondria have their own DNA, resemble bacteria, and replicate themselves for cell division This suggests that once the 1st primitive cells evolved, they “swallowed” other cells. It is believed that they could have become dependent on one another to carry out parts of their metabolism