The Evolution of Populations Chapter 23

What is the difference between a gene and an allele? Quick refresher- A gene is the basic instruction, a sequence of DNA, an allele is one variant of that instruction. Example: The cat fell from the roof. The cat fell off the roof. The snake fell from the roof. All 3 sentences are the equivalent of a gene, they represent different alleles. Sentences 1 &2 have nearly identical meanings, however sentence 3 has a very different meaning.

Genetic variation makes evolution possible Mutations are the only source of new genes and new alleles. Only mutations in gametes can be passed to offspring. Point mutations are changes in one base in a gene. They can have significant impact on phenotype, as in sickle-cell disease. Chromosomal mutations delete, disrupt, duplicate, or rearrange many loci at once. They are usually harmful. Most of the genetic variations within a population are due to the sexual recombination of alleles that already exist in a population. Sexual reproduction rearranges alleles into new combinations in every genera­tion. Recall there are three mechanisms for this shuffling of alleles: Crossing over during prophase I of meiosis. Independent assortment of chromosomes during meiosis Fertilization (223 X 223 different possible combinations for human sperm and egg) 3

Population Genetics Population genetics is the study of how populations change genetically over time. Population: A group of individuals of the same species that live in the same area and interbreed, producing fertile offspring. Gene pool: All of the alleles at all loci in all the members of a population. In diploid species, each individual has two alleles for a particular gene, and the individual may be either heterozygous or homozygous. If all members of a population are homozygous for the same allele, the al­lele is said to be fixed.

Gene Pools A gene pool is the combined genetic information of all the members of a particular population Recall that a population is a collection of individuals of the same species in a given area which share a common group of genes A gene pool typically contains 2 or more alleles (or forms of certain genes) The relative frequency of an allele is the number of times that allele occurs in a gene pool compared to the number of times other alleles occur.

Population Genetics Hardy-Weinberg Principle – states that allele frequencies tend to remain constant in populations unless something happens OTHER THAN Mendelian segregation and sexual recombination. This situation in which allele frequencies remain constant is called genetic equilibrium If allele frequencies do not change, the population will not evolve Hardy-Weinberg is a mathematical model that describes the changes in allele frequencies in a population Allows us to predict allele and genotype frequencies in subsequent generations (testable) 6

Hardy-Weinberg Theorem G.H. Hardy mathematician Hardy-Weinberg Theorem Model conditions required to maintain genetic equilibrium from generation to generation: Random mating population Large population size No emigration or immigration (no movement into or out of the population) No mutations No natural selection (all genotypes have an equal chance of survival and reproduction) If all 5 conditions are met, there should be NO EVOLUTION Describes a NON-EVOLVING POPULATION 7

Hardy-Weinberg Equation Let p= frequency of allele A Let q= frequency of allele a Let p2= frequency of genotype AA Let 2pq= frequency of genotype Aa Let q2= frequency of genotype aa p2+2pq+q2=1 (genotype frequencies) p+q=1 (allele frequencies) W. Weinberg physician The Hardy-Weinberg Equation can show that evolution IS OCCURING within a population 8

Causes of Microevolution Natural selection, genetic drift, and gene flow can alter allele frequencies in a population and cause MOST evolutionary changes. Microevolution –Evolution on its smallest scale. Generation to generation change in a population’s allele frequencies Three main causes: genetic drift natural selection gene flow

Genetic Drift Genetic drift is the unpredictable fluctuation in allele frequencies from one generation to the next. The smaller the population, the greater the chance is for genetic drift. This is a random, non- adaptive change in allele frequencies. Can and does lead to allele fixation Allele fixation means that a population changes (evolves) from many alleles represented, to only 1 allele represented 10

Consequences of Genetic Drift Fixation of alleles Effect of chance is different from population to population Small populations are effected by genetic drift more often than larger ones Given enough time, even in large populations genetic drift can have an effect Genetic drift reduces variability in populations by reducing heterozygosity REAL WORLD EXAMPLES OF GENETIC DRIFT: The Bottleneck Effect The Founder Effect 11

Real World Examples of Genetic Drift The Bottleneck Effect Occurs when only a few individuals survive a random event, resulting in a shift in allele frequencies within the population Small population sizes facilitate inbreeding and genetic drift, both of which decrease genetic variation Reduces genetically variability because at least some alleles are likely to be lost from gene pool Northern elephant seals have reduced genetic variation probably because of a population bottleneck humans inflicted on them in the 1890s. Hunting reduced their population size to as few as 20 individuals at the end of the 19th century. Their population has since rebounded to over 30,000—but their genes still carry the marks of this bottleneck: they have much less genetic variation than a population of southern elephant seals that was not so intensely hunted. 12

Figure 23.5 The bottleneck effect: an analogy 13

Real World Examples of Genetic Drift Founder effect: A few individuals become isolated from a larger popu­lation and establish a new population whose gene pool is not reflective of the source 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. For example, the Afrikaner population of Dutch settlers in South Africa is descended mainly from a few colonists. Today, the Afrikaner population has an unusually high frequency of the gene that causes Huntington’s disease, (which causes nerve cells in certain parts of the brain waste away, or degenerate.) because those original Dutch colonists just happened to carry that gene with unusually high frequency. This effect is easy to recognize in genetic diseases. 14

The Founder Effect Descendants Founding Population A Founding Population B 15

Gene Flow Gene flow can also change allele frequencies Gene flow is the physical flow of alleles into or out of a population. Immigration – alleles coming in (added) Emigration – alleles moving out (lost) Gene flow counteracts differences that arise through mutation, natural selection, and genetic drift. Gene flow helps keep separated populations genetically similar – reduces differences between populations 16

Natural selection is the only mechanism that consistently causes adaptive evolution Relative fitness refers to the contribution an organism makes to the gene pool of the next generation relative to the contributions of other members. Fitness does not indicate strength or size. It is measured only by reproductive success. Natural selection acts more directly on the phenotype and indirectly on the genotype and can alter the frequency distribution of heritable traits in three ways.

Examples 1. Directional selection Example: Large black bears survived periods of extreme cold better than smaller ones, and so became more common during glacial periods. 2. Disruptive selection Example: A population has individuals with ei­ther large beaks or small beaks, but few with the intermediate beak size. Apparently the intermediate beak size is not efficient in cracking either the large or small seeds that are common. 3. Stabilizing selection Example: Birth weights of most humans lie in a narrow range, as those babies who are very large or very small have higher mortality

Directional selection Shifts the overall makeup of the population by favoring variants that are at one extreme of the distribution. In this case, darker mice are favored because they live among dark rocks, and a darker fur color conceals them from predators.

Disruptive selection Favors variants at both ends of the distribution. These mice have colonized a patchy habitat made up of light and dark rocks, with the result that mice of an intermediate color are at a disadvantage.

Stabilizing selection Removes extreme variants from the population and preserves intermediate types. If the environment consists of rocks of an intermediate color, both light and dark mice will be selected against

Sexual Selection Sexual selection is a form of natural selection in which individuals with certain in­herited characteristics are more likely than other individuals to obtain mates. It can result in sexual dimorphism, a difference between the two sexes in secondary sex­ual characteristics such as differences in size, color, ornamentation, and behavior.

Figure 23.16x1 Sexual selection and the evolution of male appearance

Types of Sexual Selection Intrasexual Selection means selction within the same sex – typically males Individuals of one sex compete directly for mates of the opposite sex Often it is based on rituals and displays that don’t risk injury Intersexual Selection is also called “mate choice” – typically females Female choice is typically based on showiness of the male’s appearance and/or behavior Males will often weight the attraction of predators versus the attraction of mates

How is genetic variation preserved in a population? Diploidy: (refers to organisms carrying genes in pairs) Because most eukaryotes are diploid, they are capable of hiding genetic variation (recessive alleles) from selection. Heterozygote advantage: Individuals who are heterozygous at a certain locus have an advantage for survival. Example: sickle-cell disease, individuals homozygous for normal hemoglobin are more susceptible to malaria, whereas homozygous recessive individuals suf­fer from the complications of sickle-cell disease. Heterozygotes benefit from protection from malaria and do not have sickle-cell disease, so the mutant allele remains relatively common.