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Chapter 23 The Evolution of Populations Original population Evolved population Directional selection Disruptive selection Stabilizing selection Overview: The Smallest Unit of Evolution • Student misconception = organisms evolve during their lifetimes – Reminder- natural selection acts on individuals, but only populations evolve • Genetic variations in populations contribute to evolution • Microevolution is a change in allele frequencies in a population over generations Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Concept 23.1: Mutation and sexual reproduction produce the genetic variation that makes evolution possible • 2 processes produce variation in gene pools that contributes to differences among individuals = mutation and sexual reproduction – Variation in individual genotype leads to variation in individual phenotype – Not all phenotypic variation is heritable • Diet/environmentally-induced – Natural selection can only act on variation with a genetic component Mutation • Mutations are changes in the nucleotide sequence of DNA • Mutations cause new genes and alleles to arise • Only mutations in cells that produce gametes can be passed to offspring Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Point Mutations • A point mutation is a change in one base in a gene • Effects can vary: – Mutations in noncoding regions often harmless – Mutations in gene might not affect protein production b/c of redundancy in genetic code – Mutations that result in change in protein production are often harmful, but can sometimes increase the fit between organism and environment Mutations That Alter Gene Number or Sequence • Chromosomal mutations that delete, disrupt, or rearrange many loci are typically harmful • Duplication of large chromosome segments is usually harmful • Duplication of small pieces of DNA is sometimes less harmful and increases the genome size • Duplicated genes can take on new functions by further mutation Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Mutation Rates • Mutation rates are low in animals and plants • The average is about one mutation in every 100,000 genes per generation – lower in prokaryotes (but short generations so mutations accumulate quickly) – higher in viruses (and short generations so mutations really accumulate quickly) • RNA Sexual Reproduction • Sexual reproduction can shuffle existing alleles into new combinations – Crossing-over – Independent assortment – Random fertilization • In organisms that reproduce sexually, recombination of alleles is more important than mutation in producing the genetic differences that make adaptation possible Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Population genetics: study of how populations change genetically over time Population: group of individuals that live in the same area and interbreed, producing fertile offspring Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings • Gene pool: all of the alleles for all genes in all the members of the population – Diploid species: 2 alleles for a gene (homozygous/heterozygous) • Fixed allele: all members of a population only have 1 allele for a particular trait – The more fixed alleles a population has, the LOWER the species’ diversity Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Hardy-Weinberg Theorem Hardy-Weinberg Theorem: The allele and genotype frequencies of a population will remain constant from generation to generation …UNLESS they are acted upon by forces other than Mendelian segregation and recombination of alleles Equilibrium = allele and genotype frequencies remain constant from generation or generation Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Conditions for Hardy-Weinberg equilibrium 1. No mutations. 2. Random mating. 3. No natural selection. 4. Extremely large population size. 5. No gene flow. If at least one of these conditions is NOT met, then the population is EVOLVING! Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings • The frequency of an allele in a population can be calculated: – For diploid organisms, • total # of alleles at a locus = total # of individuals x 2 • total # of dominant alleles at a locus = 2 alleles for each homozygous dominant individual plus 1 allele for each heterozygous individual • same logic applies for recessive alleles Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings • By convention, if there are 2 alleles at a locus, p and q are used to represent their frequencies – p = dominant – q = recessive • The frequency of all alleles in a population will add up to 1 – For example, p + q = 1 Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Hardy-Weinberg Principle Allele Frequencies: • Gene with 2 alleles : p, q p = frequency of dominant allele (A) q = frequency of recessive allele (a) p+q=1 Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Note: 1–p=q 1–q=p Hardy-Weinberg Equation Genotypic Frequencies: • 3 genotypes (AA, Aa, aa) 2 p + 2pq + 2 q =1 p2 = AA (homozygous dominant) 2pq = Aa (heterozygous) q2 = aa (homozygous recessive) Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Allele frequencies Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Genotypic frequencies Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Strategies for solving H-W Problems: 1. If you are given the genotypes (AA, Aa, aa), calculate p and q by adding up the total # of A and a alleles. 2. If you know phenotypes, then use “aa” to find q2, and then q. (p = 1-q) 3. To find out if population is evolving, calculate p2 + 2pq + q2. – If in equilibrium, it should = 1. – If it DOES NOT = 1, then the population is evolving! Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Hardy-Weinberg practice problem #1 The scarlet tiger moth has the following genotypes. Calculate the allele and genotype frequencies (%) for a population of 1612 moths. AA = 1469 Aa = 138 Allele Frequencies: A= a= Genotypic Frequencies: AA = Aa = aa = Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings aa = 5 Hardy-weinberg practice problem #2: PTC Tasters • Taster = AA or Aa Nontaster = aa • Tasters = ____ Nontasters = ___ q2 = q= p+q=1 p=1–q= p2 + 2pq + q2 = 1 Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Applying the Hardy-Weinberg Principle • We can assume the locus that causes phenylketonuria (PKU) is in Hardy-Weinberg equilibrium given that: – The PKU gene mutation rate is low – Mate selection is random with respect to whether or not an individual is a carrier for the PKU allele – Natural selection can only act on rare homozygous individuals who do not follow dietary restrictions – The population is large – Migration has no effect as many other populations have similar allele frequencies Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings • The occurrence of PKU is 1 per 10,000 births – q2 = 0.0001 – q = 0.01 • The frequency of normal alleles is – p = 1 – q = 1 – 0.01 = 0.99 • The frequency of carriers is – 2pq = 2 x 0.99 x 0.01 = 0.0198 – or approximately 2% of the U.S. population • Homework- p.475 #2 & 3, p.486 #1; check Appendix CAUSES OF EVOLUTION Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Conditions for Hardy-Weinberg equilibrium 1. No mutations. 2. Random mating. 3. No natural selection. 4. Extremely large population size. 5. No gene flow. If at least one of these conditions is NOT met, then the population is EVOLVING! Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Minor Causes of Evolution: #1 - Mutations • Rare, very small changes in allele frequencies #2 - Nonrandom mating • Affect genotypes, but not allele frequencies Major Causes of Evolution: • Natural selection, genetic drift, gene flow (#3-5) Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Major Causes of Evolution #3 – Natural Selection • Individuals with variations better suited to environment pass more alleles to next generation • See Chapter 22 Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Major Causes of Evolution #4 – Genetic Drift • Small populations have greater chance of fluctuations in allele frequencies from one generation to another • Tends to reduce genetic variation through losses of alleles • Examples: • Founder Effect • Bottleneck Effect Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Genetic Drift Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Founder Effect • A few individuals become isolated from larger population • Certain alleles under/over represented Polydactyly in Amish population Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Bottleneck Effect • Sudden change in environment drastically reduces population size • Resulting gene pool no longer reflective of the original population’s gene pool Northern elephant seals hunted nearly to extinction in California Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Effects of Genetic Drift: A Summary 1. Genetic drift is significant in small populations 2. Genetic drift causes allele frequencies to change at random 3. Genetic drift can lead to a loss of genetic variation within populations 4. Genetic drift can cause harmful alleles to become fixed Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Major Causes of Evolution #5 – Gene Flow • Movement of fertile individuals or gametes between populations • Gain/lose alleles • Reduce genetic differences between populations • Can increase or decrease the fitness of a population Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings • Gene flow can decrease the fitness of a population • In bent grass, alleles for copper tolerance are beneficial in populations near copper mines, but harmful to populations in other soils • Windblown pollen moves these alleles between populations • The movement of unfavorable alleles into a population results in a decrease in fit between organism and environment Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings • Gene flow can increase the fitness of a population • Insecticides have been used to target mosquitoes that carry West Nile virus and malaria • Alleles have evolved in some populations that confer insecticide resistance to these mosquitoes • The flow of insecticide resistance alleles into a population can cause an increase in fitness Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Relative Fitness • The phrases “struggle for existence” and “survival of the fittest” are misleading as they imply direct competition among individuals – Reproductive success is generally more subtle and depends on many factors • Relative fitness is the contribution an individual makes to the gene pool of the next generation, relative to the contributions of other individuals • Selection favors certain genotypes by acting on the phenotypes of certain organisms Directional, Disruptive, and Stabilizing Selection • Three modes of selection: – Directional selection favors individuals at one end of the phenotypic range – Disruptive selection favors individuals at both extremes of the phenotypic range – Stabilizing selection favors intermediate variants and acts against extreme phenotypes Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Directional Selection: eg. larger black bears survive extreme cold better than small ones Disruptive Selection: eg. small beaks for small seeds; large beaks for large seeds Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Stabilizing Selection: eg. narrow range of human birth weight The Key Role of Natural Selection in Adaptive Evolution • Natural selection increases the frequencies of alleles that enhance survival and reproduction • Adaptive evolution occurs as the match between an organism and its environment increases – Because the environment can change, adaptive evolution is a continuous process • Genetic drift and gene flow do not consistently lead to adaptive evolution as they can increase or decrease the match between an organism and its environment Sexual Selection • Sexual selection is natural selection for mating success • It can result in sexual dimorphism, marked differences between the sexes in secondary sexual characteristics Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings • Intrasexual selection is competition among individuals of one sex (often males) for mates of the opposite sex http://abcnews.go.com/GMA/video/violentgiraffe-fight-video-viral-giraffes-slam-necks-18099718 • Intersexual selection, often called mate choice, occurs when individuals of one sex (usually females) are choosy in selecting their mates http://www.youtube.com/watch?v=L54bxmZy_NE • Male showiness due to mate choice can increase a male’s chances of attracting a female, while decreasing his chances of survival (remember the guppies from Ch 22) Preserving genetic variation • Diploidy: hide recessive alleles that are less favorable • Heterozygote advantage: greater fitness than homozygotes • eg. Sickle cell disease Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Why Natural Selection Cannot Fashion Perfect Organisms 1. Selection can act only on existing variations 2. Evolution is limited by historical constraints -descent with modification 3. Adaptations are often compromises 4. Chance, natural selection, and the environment interact Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings You should now be able to: 1. Explain why the majority of point mutations are harmless 2. How does sexual recombination generate genetic variability? 3. Define: population, species, gene pool, relative fitness, neutral variation, gene flow 4. List the five conditions of Hardy-Weinberg equilibrium 5. Apply the H-W equation to population genetics problems 6. Explain why natural selection is the only mechanism that consistently produces adaptive change 7. Explain the role of population size in genetic drift; 2 types? 8. Distinguish among the following sets of selection terms: directional/disruptive/stabilizing; intrasexual/intersexual 9. Why can’t natural selection produce perfect organisms?