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Microevolution Chapter 16 Selective Breeding & Evolution • Evolution is genetic change in a line of descent through successive generations • Selective breeding practices yield evidence that heritable changes do occur Domestication of Dogs • Began about 50,000 years ago • 14,000 years ago - artificial selection – Dogs with desired forms of traits were bred • Modern breeds are the result Results of Artificial Selection • Extremes in size – Great Dane and Chihuahua • Extremes in form – Short-legged dachshunds – English bulldog • Short snout and compressed face • Extreme traits lead to health problems Evolutionary Theories • Widely used to interpret the past and present, and even to predict the future • Reveal connections between the geological record, fossil record, and organismal diversity Early Scientific Theories • Hippocrates - All aspects of nature can be traced to their underlying causes • Aristotle - Each organism is distinct from all the rest and nature is a continuum or organization Confounding Evidence • Biogeography • Comparative anatomy • Geologic discoveries Biogeography • Size of the known world expanded enormously in the 15th century • Discovery of new organisms in previously unknown places could not be explained by accepted beliefs – How did species get from center of creation to all these places? Comparative Morphology • Study of similarities and differences in body plans of major groups • Puzzling patterns: – Animals as different as whales and bats have similar bones in forelimbs – Some parts seem to have no function Comparative Anatomy Python Human Geological Discoveries • Similar rock layers throughout world • Certain layers contain fossils • Deeper layers contain simpler fossils than shallow layers • Some fossils seem to be related to known species 19th Century - New Theories • Scientists attempt to reconcile evidence of change with traditional belief in a single creation event • Two examples – Georges Cuvier - multiple catastrophes – Jean Lamark - inheritance of acquired characteristics The Theory of Uniformity • Lyell’s Principles of Geology • Subtle, repetitive processes of change, had shaped Earth • Challenged the view that Earth was only 6,000 years old Darwin’s Voyage • At age 22, Charles Darwin began a fiveyear, round-the-world voyage aboard the Beagle • In his role as ship’s naturalist he collected and examined the species that inhabited the regions the ship visited Voyage of the Beagle EQUATOR Galapagos Islands Galapagos Islands Darwin Wolf Volcanic islands far off coast of Ecuador Pinta Genovesa Marchena All inhabitants are descended from species that arrived on islands from elsewhere Santiago Bartolomé Fernandia Seymour Baltra Rabida Pinzon Santa Cruz Santa Fe Tortuga San Cristobal Isabela Española Floreana Glyptodonts & Armadillos • In Argentina, Darwin observed fossils of extinct glyptodonts • Animals resembled living armadillos Malthus - Struggle to Survive • Thomas Malthus, a clergyman and economist, wrote essay that Darwin read on his return to England • Argued that as population size increases, resources dwindle, the struggle to live intensifies and conflict increases Galapagos Finches • Darwin observed finches with a variety of lifestyles and body forms • On his return he learned that there were 13 species • He attempted to correlate variations in their traits with environmental challenges Darwin’s Theory A population can change over time when individuals differ in one or more heritable traits that are responsible for differences in the ability to survive and reproduce Alfred Wallace • Naturalist who arrived at the same conclusions Darwin did • Wrote to Darwin describing his views • Prompted Darwin to finally present his ideas in a formal paper On the Origin of Species • Darwin’s book • Published in 1859 • Laid out in great detail his evidence in support of the theory of evolution by natural selection Missing Links • If one species can evolve into another, there should be transitional forms • When Darwin published his work, no such forms were known • First fossil Archaeopteryx found in 1860 Populations Evolve • Biological evolution does not change individuals • It changes a population • Traits in a population vary among individuals • Evolution is change in frequency of traits The Gene Pool • All of the genes in the population • Genetic resource that is shared (in theory) by all members of population Variation in Phenotype • Each kind of gene in gene pool may have two or more alleles • Individuals inherit different allele combinations • This leads to variation in phenotype • Offspring inherit genes, NOT phenotypes What Determines Alleles in New Individual? • Mutation • Crossing over at meiosis I • Independent assortment • Fertilization • Change in chromosome number or structure Genetic Equilibrium • Allele frequencies at a locus are not changing • Population is not evolving Five Conditions • No mutation • Random mating • Gene doesn’t affect survival or reproduction • Large population • No immigration/emigration Microevolutionary Processes • Drive a population away from genetic equilibrium • Small-scale changes in allele frequencies brought about by: – Natural selection – Gene flow – Genetic drift Gene Mutations • • • • • Infrequent but inevitable Each gene has own mutation rate Lethal mutations Neutral mutations Advantageous mutations Hardy-Weinberg Rule At genetic equilibrium, proportions of genotypes at a locus with two alleles are given by the equation: p2 AA + 2pq Aa + q2 aa = 1 Frequency of allele A = p Frequency of allele a = q Punnett Square p A q a p A AA(p2) Aa(pq) q a Aa(pq) aa(q2) Frequencies in Gametes F1 genotypes: Gametes: 0.49 AA A 0.42 Aa A A 0.09 aa a a a 0.49 + 0.21 0.21 + 0.09 0.7A 0.3a STARTING POPULATION 490 AA butterflies Dark-blue wings No Change Through Generations 420 Aa butterflies Medium-blue wings 90 aa butterflies White wings THE NEXT GENERATION 490 AA butterflies 420 Aa butterflies 90 aa butterflies NO CHANGE THE NEXT GENERATION 490 AA butterflies 420 Aa butterflies 90 aa butterflies NO CHANGE Natural Selection • A difference in the survival and reproductive success of different phenotypes • Acts directly on phenotypes and indirectly on genotypes Reproductive Capacity & Competition • All populations have the capacity to increase in numbers • No population can increase indefinitely • Eventually, the individuals of a population will end up competing for resources Variation in Populations • All individuals have the same genes that specify the same assortment of traits • Most genes occur in different forms (alleles) that produce different phenotypes • Some phenotypes compete better than others Change Over Time • Over time, the alleles that produce the most successful phenotypes will increase in the population • Less successful alleles will become less common • Change leads to increased fitness – Increased adaptation to environment Results of Natural Selection Three possible outcomes: • A shift in the range of values for a given trait in some direction • Stabilization of an existing range of values • Disruption of an existing range of values Number of individuals in the population Number of individuals in the population • Allele frequencies shift in one direction Range of values for the trait at time 1 Range of values for the trait at time 2 Number of individuals in the population Directional Selection Range of values for the trait at time 3 Peppered Moths • Prior to industrial revolution, most common phenotype was light colored • After industrial revolution, dark phenotype became more common Pesticide Resistance • Pesticides kill susceptible insects • Resistant insects survive and reproduce • If resistance has heritable basis, it becomes more common with each generation Antibiotic Resistance • First came into use in the 1940s • Overuse has led to increase in resistant forms • Most susceptible cells died out and were replaced by resistant forms • Intermediate forms are favored and extremes are eliminated Number of individuals in the population Stabilizing Selection Range of values for the trait at time 1 Range of values for the trait at time 2 Range of values for the trait at time 3 Selection for Gall Size • Gall-making fly has two major predators • Wasps prey on larvae in small galls • Birds eat larvae in large galls • Flies that cause intermediate-sized galls have the highest fitness Number of individuals Number of individuals in the population in the population • Forms at both ends of the range of variation are favored • Intermediate forms are selected against Range of values for the trait at time 1 Range of values for the trait at time 2 Number of individuals in the population Disruptive Selection Range of values for the trait at time 3 African Finches 60 Number of individuals • Selection favors birds with very large or very small bills • Birds with intermediate-sized bill are less effective feeders 50 40 30 20 10 10 12.8 15.7 18.5 Widest part of lower bill (millimeters) Sexual Selection • Selection favors certain secondary sexual characteristics • Through nonrandom mating, alleles for preferred traits increase • Leads to increased sexual dimorphism Balanced Polymorphism • Polymorphism - “having many forms” • Occurs when two or more alleles are maintained at frequencies greater than 1 percent Sickle-Cell Trait: Heterozygote Advantage • Allele causes sickle-cell anemia when heterozygous • Heterozygotes are more resistant to malaria than homozygotes HbS Malaria case Sickle cell trait less than 1 in 1,600 1 in 400-1,600 1 in 180-400 1 in 100-180 1 in 64-100 more than 1 in 64 Gene Flow • Physical flow of alleles into a population • Tends to keep the gene pools of populations similar • Counters the differences that result from mutation, natural selection, and genetic drift Genetic Drift • Random change in allele frequencies brought about by chance • Effect is most pronounced in small populations • Sampling error - Fewer times an event occurs, greater the variance in outcome Computer Simulation 1.0 AA in five populations 0.5 allele A lost from four populations 0 1 5 10 15 20 25 30 35 40 45 50 Generation (25 stoneflies at the start of each) Computer Simulation 1.0 0.5 allele A neither lost nor fixed 0 1 5 10 15 20 25 30 35 40 45 Generation (500 stoneflies at the start of each) 50 Bottleneck • A severe reduction in population size • Causes pronounced drift • Example – Elephant seal population hunted down to just 20 individuals – Population rebounded to 30,000 – Electrophoresis revealed there is now no allele variation at 24 genes Founder Effect • Effect of drift when a small number of individuals start a new population • By chance, allele frequencies of founders may not be same as those in original population • Effect is pronounced on isolated islands Inbreeding • Nonrandom mating between related individuals • Leads to increased homozygosity • Can lower fitness when deleterious recessive alleles are expressed • Amish, cheetahs