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Chapter 15 Evolution on a Small Scale 15.1 Natural Selection • Process resulting in adaptation of a population to the biotic (living) and abiotic (nonliving) environments • Darwin’s mechanism for evolution • Most fit individuals become more prevalent in a population • Leads to change in population over time • Most fit individuals reproduce more than others because they are better adapted. • Types of selection • • • Most traits are polygenic and are controlled by more than one pair of alleles located at different gene loci. • Such traits have a range of phenotypes resembling a bell-shaped curve. • Directional selection • Stabilizing selection • Disruptive selection Directional selection • Occurs when an extreme phenotype is favored • Distribution curve shifts in that direction • Can occur when a population is adapting to a changing environment • Industrial melanism • Drug resistance in bacteria • Pesticide resistance in insects • Malaria—Plasmodium becoming resistant to chloroquine and mosquitoes resistant to DDT • Equus adapting from forest conditions to grassland conditions Stabilizing selection • Occurs when an intermediate phenotype is favored • Extreme phenotypes selected against • Individuals near the average are favored • • • Most common form of selection because the average individual is well adapted to its environment • Swiss starlings lay 4–5 eggs because this has the highest survival rate for young Disruptive selection • 2 or more extreme phenotypes are favored over any intermediate phenotype • British land snails are found in fields and forests • In fields, thrushes eat the snails with dark shells that lack light bands • In forests, thrushes feeds mainly on snails with light-banded shells Adaptations are not perfect • Natural selection does not produce perfectly adapted organisms. • Evolution is constrained by the available variations. • Imperfections are common because of necessary compromises. • • Success of humans attributed to freeing hands but walking upright puts stress on the spine Maintenance of variations • A population always shows some genotypic variations. • Population with limited variation may not be able to adapt to changing environmental conditions • Forces promoting variation constantly at work • Mutations, recombination, independent assortment, and fertilization create new combinations • Gene flow • Natural selection favors certain phenotypes but other remain • • Diploidy and the heterozygote The heterozygote advantage • Only alleles that are expressed are subject to natural selection. • Expressed = cause phenotypic differences • Heterozygotes can protect recessive alleles. • Recessive allele might have greater fitness in a changing environment • Balanced polymorphism—when natural selection favors the ratio of two or more phenotypes in generation after generation • • Sickle-cell disease Sickle-cell disease • Individuals with sickle cell disease HbSHBS • • Tend to die early Heterozygotes carry the trait HbAHBS • Red blood cells only sickle at low oxygen concentrations • Ordinarily the normal genotype is most fit HbAHBA • Recessive allele HbS has a higher frequency in regions in Africa where malaria is present • Malaria is caused by parasite that invades and destroys normal red blood cells • Parasite unable to live in heterozygote red blood cells • Each of the homozygotes is selected against but is maintained because the heterozygote is favored in those parts of Africa 15.2 Microevolution • Individuals do not evolve. • As evolution occurs, genetic and phenotypic changes occur within a population. • A population is all the members of a single species occupying a particular area at the same time and reproducing with one another. • Microevolution—small measurable evolutionary changes within a population from generation to generation • Darwin stressed that members of a population vary. • Each gene in sexually reproducing organisms has many alleles. • Reshuffling of alleles during sexual reproduction can result in a range of phenotypes. • Evolution in a genetic context • Gene pool—the various alleles at all the gene loci in all individuals of a population • • Described in terms of genotype and allele frequency Peppered moth color example • D = dark color d = light color • • • From the genotype frequencies, you can calculate the allele frequencies of a population. • Assuming random mating, we can use allele frequencies (gamete frequencies) to calculate the ratio of genotypes in the next generation using a Punnett square. • Allele frequencies remain the same—sexual reproduction alone does not bring about a change in allele frequencies • Dominance does not cause an allele to become a common allele. G. H. Hardy and W. Weinberg used the binomial equation to calculate the genotypic and allele frequencies of a population. • p = frequency of dominant allele • q = frequency of recessive allele • p2 + 2pq + q2 = 1 Hardy-Weinberg principle states that an equilibrium of allele frequencies in a gene pool will remain in equilibrium as long as 5 conditions are met • No mutations • No gene flow • Random mating • No genetic drift • No selection • These conditions are rarely if ever met. • Allele frequencies do change from one generation to the next. • Therefore microevolution occurs • Hardy-Weinberg equation is significant because it tells us what factors cause evolution • Evolution can be detected and measured by noting the amount of deviation from a HardyWeinberg equilibrium of allele frequencies in the gene pool of a population. • Industrial melanism example • Increase in the frequency of a dark phenotype due to pollution • Before soot darkened tree trunks, light moths escaped detection of birds and were more common. • After the advent of industry, dark-colored moths became more common as light moths were detected and eaten. • Natural selection can occur within a short time frame. • • • Change in gene pool frequencies occurs as microevolution occurs Causes of microevolution • Any condition that deviates from the list of conditions for allelelic equilibrium causes evolutionary change • Genetic mutation • Gene flow • Nonrandom mating • Genetic drift • Natural selection Genetic mutations • Ultimate source for allele differences • Without mutation there would be no new variations among members of a population for natural selection to act on • Adaptive value of mutation depends on current conditions 2. Gene flow Also called gene migration Movement of alleles among populations by migration of breeding individuals Can increase variation within a population by introducing novel alleles from another population Continued gene flow reduces differences among populations—can prevent speciation 3. Nonrandom mating Selection of mate according to genotype or phenotype (not chance) Assortative mating—tend to mate with individuals with the same phenotype • Homozygotes increase in frequency Sexual selection—favors characteristics that increase the likelihood of obtaining mates 4. Genetic drift Refers to changes in the allele frequencies of a gene pool due to chance Allele frequencies “drift” over time depending on which members die, survive, or reproduce More likely in small populations More likely to lose rare alleles Two types Bottleneck effect Founder effect Bottleneck effect Species suffers a near extinction and only a few survivors go on to produce the next generation Cheetahs—extreme similarity Infertility due to inbreeding Founder effect Rare alleles occur at a higher frequency in a population isolated from the general population. Alleles carried by founders are dictated by chance alone. Amish—1 in 14 carries recessive allele for unusual form of dwarfism compared to 1 in 1000 in most populations