Notes – Ch. 23 Evolution of Populations Individuals are selected Populations evolve • One misconception is that organisms evolve, in the Darwinian sense, during their lifetimes • Genetic variations in populations contribute to evolution 2007-2008 Population genetics provides a foundation for studying evolution • Microevolution is change in the genetic makeup of a population from generation to generation • Population genetics is the study of how populations change genetically over time • Population genetics integrates Mendelian genetics with the Darwinian theory of evolution by natural selection Gene Pools and Allele Frequencies • WHAT IS A SPECIES? – A species is a group of organisms that are able to interbreed and produce fertile offspring – (there are always exceptions – some organisms of different species, like a dog and wolf – can interbreed and produce fertile offspring even through they are different species) • A population is a localized group of individuals capable of interbreeding and producing fertile offspring • The gene pool is the total aggregate of genes in a population at any one time The Hardy-Weinberg Theorem • The Hardy-Weinberg theorem describes a population that is not evolving (this is rarely the case) • It states that frequencies of alleles and genotypes in a population’s gene pool remain constant from generation to generation Hardy-Weinberg Equilibrium • Hardy-Weinberg equilibrium describes a population where allele frequencies do not change • The five conditions are required in order for a population to remain at Hardy-Weinberg equilibrium are: – 1.A large breeding population – 2.Random mating – 3.No change in allelic frequency due to mutation – 4.No immigration or emigration – 5.No natural selection *The five conditions for non-evolving populations are rarely met in nature • If p and q represent the relative frequencies of the only two possible alleles in a population at a particular locus, then – p2 + 2pq + q2 = 1 – And p2 and q2 represent the frequencies of the homozygous genotypes and 2pq represents the frequency of the heterozygous genotype LE 23-5 Gametes for each generation are drawn at random from the gene pool of the previous generation: 80% CR (p = 0.8) 20% CW (q = 0.2) Sperm CR CW (20%) p2 pq 64% CRCR 16% CRCW (20%) CR (80%) CW Eggs (80%) qp 4% CWCW 16% CRCW q2 Conditions for Hardy-Weinberg Equilibrium • The Hardy-Weinberg theorem describes a hypothetical population • In real populations, allele and genotype frequencies do change over time Population Genetics and Human Health • We can use the Hardy-Weinberg equation to estimate the percentage of the human population carrying the allele for an inherited disease Mutation and sexual recombination produce the variation that makes evolution possible • Two processes, mutation and sexual recombination, produce the variation in gene pools that contributes to differences among individuals Mutation • Mutations are changes in the nucleotide sequence of DNA • Mutations cause new genes and alleles to arise Point Mutations • A point mutation is a change in one base in a gene • It is usually harmless but may have significant impact on phenotype Mutations That Alter Gene Number or Sequence • Chromosomal mutations that delete, disrupt, or rearrange many loci are typically harmful (frameshift) • Gene duplication is nearly always harmful Mutation Rates • Mutation rates are low in animals and plants • The average is about one mutation in every 100,000 genes per generation • Mutations are more rapid in microorganisms Sexual Recombination • Sexual recombination is far more important than mutation in producing the genetic differences that make adaptation possible Natural selection, genetic drift, and gene flow can alter a population’s genetic composition • Three major factors alter allele frequencies and bring about most evolutionary change: – Natural selection – Genetic drift – Gene flow Natural Selection • Differential success in reproduction results in certain alleles being passed to the next generation in greater proportions • “survival of the fittest” – those with the traits best suited to the environment will survive and pass on their DNA to their offspring • Natural selection accumulates and maintains favorable genotypes in a population Genetic Drift • The smaller a sample, the greater the chance of deviation from a predicted result – Genetic drift describes how allele frequencies fluctuate unpredictably from one generation to the next – Genetic drift tends to reduce genetic variation through losses of alleles • • • • • The process of genetic drift can be illustrated using 20 marbles in a jar to represent 20 organisms in a population. Consider this jar of marbles as the starting population. Half of the marbles in the jar are red and half blue, and both colors correspond to two different alleles of one gene in the population. In each new generation the organisms reproduce at random. To represent this reproduction, randomly select a marble from the original jar and deposit a new marble with the same color as its "parent" into a new jar. (The selected marble remains in the original jar.) Repeat this process until there are 20 new marbles in the second jar. The second jar then contains a second generation of "offspring", consisting of 20 marbles of various colors. Unless the second jar contains exactly 10 red marbles and 10 blue marbles, a random shift occurred in the allele frequencies. Repeat this process a number of times, randomly reproducing each generation of marbles to form the next. The numbers of red and blue marbles picked each generation fluctuates: sometimes more red, sometimes more blue. This fluctuation is analogous to genetic drift – a change in the population's allele frequency resulting from a random variation in the distribution of alleles from one generation to the next. It is even possible that in any one generation no marbles of a particular color are chosen, meaning they have no offspring. In this example, if no red marbles are selected the jar representing the new generation contains only blue offspring. If this happens, the red allele has been lost permanently in the population, while the remaining blue allele has become fixed: all future generations are entirely blue. In small populations, fixation can occur in just a few generations. LE 23-7 CWCW CR CR CR CR CR CW Only 5 of 10 plants leave offspring CR CR CWCW CR CR C R CW CWCW CR CR C R CW CR CW CR CR CWCW CR CW CR C R CR CR CR CW Generation 1 p (frequency of CR) = 0.7 q (frequency of CW) = 0.3 Only 2 of 10 plants leave offspring CR CR CR CR CR CR CR CR CR CR CR CR CR CR CR CR CR CW CR C W Generation 2 p = 0.5 q = 0.5 CR CR CR CR Generation 3 p = 1.0 q = 0.0 The Bottleneck Effect • The bottleneck effect is a sudden change in the environment (fire, drought – kills many in a single generation) that may drastically reduce the size of a population • The resulting gene pool may no longer be reflective of the original population’s gene pool – Ex – elephant seal (hunting reduced population to just 20 individuals; has since rebounded to 3000) – cheetahs are sufficiently closely related to one another that transplanted skin grafts do not provoke immune responses The Founder Effect • A founder effect occurs when a new colony is started by a few members of the original population. This small population size means that the colony may have: – reduced genetic variation from the original population. – a non-random sample of the genes in the original population. • The founder effect occurs when a few individuals become isolated from a larger population • It can affect allele frequencies in a population Gene Flow • Gene flow consists of genetic additions or subtractions from a population, resulting from movement of fertile individuals or gametes • Gene flow causes a population to gain or lose alleles • It tends to reduce differences between populations over time Genetic Variation • Genetic variation occurs in individuals in populations of all species • It is not always heritable (influenced by environment - fetal alcohol syndrome) LE 23-9 Map butterflies that emerge in spring: orange and brown Map butterflies that emerge in late summer: black and white Variation Within a Population • Both discrete and quantitative characters contribute to variation within a population • Discrete characters can be classified on an either-or basis – discreet trait is something that is either one way or the other, for example, trait for sickle cell anemia. if you have the gene, you have the trait. • Quantitative characters vary along a continuum within a population – quantitative trait is something that has a range, for example, height. there are many genes that play a role and there are not a discreet number of heights, there is a range. Polymorphism • Polymorphism is when two or more clearly different phenotypes exist in the same population of a species – in other words, the occurrence of more than one form or morph • Phenotypic polymorphism describes a population in which two or more distinct morphs for a character are represented in high enough frequencies to be readily noticeable (you can see their traits) • Genetic polymorphisms are the heritable components of characters that occur along a continuum in a population (genotypes) Directional, Disruptive, and Stabilizing Selection • Selection favors certain genotypes by acting on the phenotypes of certain organisms • Three modes of selection: – Directional- favors individuals at one end of the phenotypic range – Disruptive- favors individuals at both extremes of the phenotypic range – Stabilizing- favors intermediate variants and acts against extreme phenotypes The Preservation of Genetic Variation • Various mechanisms help to preserve genetic variation in a population Diploidy • Diploidy maintains genetic variation in the form of hidden recessive alleles – Diploid – pairs of genes Balancing Selection • Balancing selection occurs when natural selection maintains stable frequencies of two or more phenotypic forms in a population • Balancing selection leads to a state called balanced polymorphism Heterozygote Advantage • Some individuals who are heterozygous at a particular locus have greater fitness than homozygotes • Natural selection will tend to maintain two or more alleles at that locus • The sickle-cell allele causes mutations in hemoglobin but also confers malaria resistance LE 23-13 Frequencies of the sickle-cell allele 0–2.5% 2.5–5.0% 5.0–7.5% Distribution of malaria caused by Plasmodium falciparum (a protozoan) 7.5–10.0% 10.0–12.5% >12.5% 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 • Intrasexual selection is competition among individuals of one sex for mates of the opposite sex – occurs when individuals of one sex (usually females) are choosy in selecting their mates from individuals of the other sex • Selection may depend on the showiness of the male’s appearance The Evolutionary Enigma of Sexual Reproduction • Sexual reproduction produces fewer reproductive offspring than asexual reproduction, a socalled “reproductive handicap” • So why do it? LE 23-16 Asexual reproduction Female Sexual reproduction Generation 1 Female Generation 2 Male Generation 3 Generation 4 • Sexual reproduction produces genetic variation that may aid in disease resistance Why Natural Selection Cannot Fashion Perfect Organisms • Evolution is limited by historical constraints (what’s happened in the past to produce current products) • Adaptations are often compromises • Chance and natural selection interact • Selection can only edit existing variations