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Chapter 23: The Evolution of Populations Things to Know: 1. All those bold-faced words again 2. The first part of the chapter, pgs 445-452, will be covered in a lab situation. But basically these pages are asking and answering the question, “How does the allelic frequency in a population change over time?” 3. The second part of the chapter emphasizes how the genetic variation or the number of different alleles in a population produces a large format upon which the environment can act in order to select the most fit organisms. 4. Hopefully the concept map will help. 1 Figure 23.0 Shells 2 Concept Map EVOLUTION may occur through Mutation Migration may result in Genetic Drift Natural Selection Speciation occurs as Genetic Variation Over-reproduction Competition for Limited Resources from Mutations acts on populations 3 Gene Pool vs. Genotype Contrast the meanings of the terms “gene pool” and “genotype.” • Gene pool is the total genetic information within the population (Campbell). It can also be defined as “is a collection of all the alleles possessed by all the reproductive members of a population of sexually producing organisms. • Genotype is the genetic makeup of an individual within that population. So it includes only those alleles that the individual carries and thus carries a maximum of two alleles at each gene locus. • So in a gene pool you have many more alleles at a given locus. Take blood type alleles for example. If you are type A, then you have a maximum of two possible alleles because you can be IAIA or Iai. But within the population you could also have IB allele. • So imagine if you will a population where a few individuals have the allele for a receptor that will prevent HIV from infecting cells. . . 4 • A research team in 1996 looked at thousands of blood samples and found that a few of the samples had cells with DNA that had a mutation to the gene that makes the CCR5 receptor. This receptor is needed for viral binding to the WBC of humans. • So people who carried two copies of this allele for the gene made no CCR5 receptors and those with one copy made fewer receptors than average. • So people carrying two copies of the mutant CCR5 gene almost never became infected. • Who carries this mutation? • Relatively common in Europe, 20% of population carry one or two copies • Most common in Sweden • As you move towards southeast Europe, the frequency of carriers decreases. 5 • Only a small fraction of Greeks • Even smaller number of Central Asians do. • From the rest of the world, this particular mutation is missing altogether. • So why do we find the distribution pattern for CCR5 in this fashion? Could “it” have been valuable to European ancestors in some way and therefore favored by natural selection? • Could the CCR4 receptor been selected for during an earlier infectious condition where millions of others were killed and those possessing the receptor were afforded some advantage? • 700 years ago The Black Death, one of several bubonic plague occurrences, killed over 25% of all Europeans between 1347 and 1350. • Bubonic plague is caused by a bacterium Yersinia pestis, which binds to white blood cells, injects toxin, depresses the immune system, multiplies. 6 • The hypothesis is: does this bacterium bind to the same receptor? If so, Europeans who were born without CCR5, were protected from the plague and their descendants are protected from HIV. • “The fact that the AIDS epidemic has been far more destructive in Africa and Southeast Asia than in Europe or the United States might be due in part to the different evolutionary histories of the continents.” • Extra Credit questions??? 7 Equilibrium in a Gene Pool? Can complete equilibrium in a gene pool exist in real situations? • What does “equilibrium” or “genetic equilibrium” mean? • H-W states that a population at equilibrium will have genotypic frequencies remain the same from generation to population. • Of the necessary H-W conditions that need to be met: • large populations can be met in real situations • the absence of mutations can never be met because spontaneous mutations always occur. • Most gene probably undergo spontaneous mutations once in every 50,000 to 1,000,000 duplications and the rate of mutation is probably different for different genes. But this rate is very low and is usually insignificant in altering gene frequencies in a large population. 8 • Immigration and emigration of individuals would change the gene pool in a population and a high percentage of natural populations do experience some amount of migration. • Random mating? An organism’s genotype does influence its mate selection, the physical efficiency and frequency of mating, its fertility so random mating just doesn’t exist! • No natural selection. All alleles have equal chance of existing. 9 • So, since it is unlikely that all of H-W conditions will be met, there will always be a continuous change to the genetic makeup of a population and this is evolution. • So H-W is a mathematical model to determine if a population is at equilibrium. It also resolves the issue of why recessive genes do not disappear in a population over time because without the stated parameters, there is nothing to change the genetic frequencies. • What will change those frequencies? •Natural selection •Mutation •Migration •Genetic drift •Nonrandom mating or sexual selection 10 How can you determine whether or not a given population is in genetic equilibrium? • Simply compare the genotypic frequencies between two generations • You are studying a population’s blood groups from one generation to the next. You find: Phenotype Generation (G) M MN N Total 1 241 604 195 1040 2 183 460 154 797 11 • So recall that MN blood groups can be determined by two alleles, LM and LN with some codominance. So we can have LMLM, LMLN, and LNLN. Frequency G1 So: G2 LMLM 241/1040 = 0.231 183 / 797 = 0.230 LMLN 604 / 1040 = 0.581 460 / 797 = 0.577 LNLN 195 / 1040 = 0.188 154 / 797 = 0.193 1.0 1.0 So the genotypic frequencies are found to be essentially the same. You could also have proven this using allelic frequencies. Can you do t this? 12 Figure 23.3a The Hardy-Weinberg theorem 13 Figure 23.3b The Hardy-Weinberg theorem 14 Figure 23.4 Genetic drift 15 Figure 23.5 The bottleneck effect: an analogy 16 Genetic Variation Genetic variations are the key to natural selection and they can occur: • Within a population • What is meant by polymorphism? • All the genes in the gene pool allow for some variability but this variability doesn’t prevent the organisms from interbreeding. That is, they look different (different phenotype) but still interbreed. • Examples: blood groups of A, B, AB and O and sickle-cell trait 17 Figure 23x2 Polymorphism 18 • Balanced Polymorphism is when different forms of the polymorphic genotype are in equilibrium in the population. This is usually achieved through complex geneticenvironmental interactions. •Heterozygote Superiority: So the Aa state has a survival advantage over both the dominant and recessive homozygous condition. So both alleles are maintained in the population. Sickle cell anemia in a malarial environment. AA: sickle cell anemia aa: susceptible to malaria Aa: sickle-cell trait but resistant to malaria. 19 Figure 23.12x Normal and sickled cells 20 Figure 23.10 Mapping malaria and the sickle-cell allele 21 •How is balanced polymorphism important in evolution? It maintains variability to permit continued adaptive changes. But it seems like only a certain number of different forms can increase the fitness of a population. Too many different forms, lowers the fitness and this “genetic load” reduces the overall fitness of the population. 22 • Between Populations • Geographic Polymorphism or clines •Here you simply have a variety of forms but each is specifically adapted to its environmental conditions. This way the species maintains itself in a variety of environmental conditions and allowing it to become more widespread. 23 Figure 23.8 Clinal variation in a plant 24 Figure 23.9 Geographic variation between isolated populations of house mice 25 What is “fitness?” • It seems to give us a mindset of “males battling it out” but may include producing and releasing more sperm, better camouflage, more efficient attractors of pollinators, more resistant to attack by a predator (a plant may contain a toxic substance that tastes repulsive). • So Campbell defines “fitness” as the contribution an individual, and therefore its genotype, makes to the gene pool of the next generation as compared to other individuals.” So if one individual, plant or animal is able to reproduce more efficiently than other, its fitness is greater. •So it certainly is a combination of surviving and reproducing. The individual exposes itself through its phenotype to its environment and survives based on that phenotype. Its reproductive ability allows it pass on those “survivability traits” and this increases its genes in the gene pool. 26 Figure 23.12 Modes of selection Natural selection can favor a phenotype in three fashions. All modes pass on traits that represent differential reproductive success To one extreme Phenotype favored is at both extremes of the range Favored phenotype is more intermediate 27 Figure 23.13 Directional selection for beak size in a Galápagos population of the medium ground finch 28 Figure 23.14 Diversifying selection in a finch population The beaks on these birds represent two extremes where the smaller beaks on the left feed efficiently on soft seeds while the larger- beaked birds feed efficiently on the hard seeds. It’s easier to see why an intermediate sized beak would not be able to crack the hard seeds but the soft seeds? Maybe the intermediate beak has trouble getting the seed out of the shell once it is cracked. I really don’t know. 29 Figure 23.15 The two-fold disadvantage of sex 30 Figure 23.16x1 Sexual selection and the evolution of male appearance 31 Figure 23.16x2 Male peacock 32 Figure 23.x1 Edaphic Races of Gaillardia pulchella 33 Figure 23.2 Population distribution 34 Figure 23.1 Individuals are selected, but populations evolve 35 Figure 23.5x Cheetahs, the bottleneck effect 36 Figure 23.6 Gene flow and human evolution 37 Figure 23.7 A nonheritable difference within a population •These butterflies differ in coloration despite having the same gene at the same locus for coloration. •This coloration difference is simply due to when the butterflies emerged from their cocoons. Hormone levels are thought to be the factor. 38 Figure 23.11 Frequency-dependent selection in a host-parasite relationship 39