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Microevolution Changing Allele Frequencies Evolution • Evolution is defined as a change in the inherited characteristics of biological populations over successive generations. • Microevolution involves the change in allele frequencies that occur over time within a population. • This change is due to four different processes: mutation, selection (natural and artificial), gene flow, and genetic drift. 2 Determining Allele Frequency • Examine the frog population presented here. • Their color is determined by a single gene, which has two alleles and phenotypically exhibits incomplete dominance. • CGCG is green, CG CR is blue, and CR CR is red • Calculate the allele frequency of the gene pool in the diagram. 3 Determining Allele Frequency • These frogs are diploid, thus have two copies of their genes for color. Allele: CG CR Green (11) 22 0 Blue (2) 2 2 Red (3) 0 6 Total: 24 8 Frequency: p = 24 ÷ 32 p = ¾ = 0.75 q = 8 ÷ 32 q = ¼ = 0.25 • If allelic frequencies change, then evolution is occurring. • Let’s suppose 4 green frogs enter the population (immigration). How do the frequencies change? 4 Immigration: Determining Allele Frequency Recall that currently: CG = 0.75 & CR = 0.25 Allele: Green (15) Blue (2) Red (3) Total: Frequency: CG 30 2 0 32 p = 32 ÷ 40 p = 8/10 = 0.80 CR 0 2 6 8 q = 8 ÷ 40 q = 2/10 = 0.20 5 Determining Allele Frequency How do the allelic frequencies change if 4 green frogs leave the population instead of enter the population? (emigration) 6 Emigration: Determining Allele Frequency Recall that originally: CG = 0.75 & CR = 0.25 Allele: Green (7) Blue (2) Red (3) Total: Frequency: CG 14 2 0 16 p = 16 ÷ 24 p = 2/3 = 0.67 CR 0 2 6 8 q = 8 ÷ 24 q = 1/3 = 0.33 7 Genetic Drift Small populations can experience changes in allele frequencies more dramatically than large populations. In very large populations the effect can be insignificant. Also in small populations genes can be lost more easily. When there is only one allele left for a particular gene in a gene pool, that gene is said to be fixed , thus there is no genetic diversity. 8 Genetic Drift • Genetic drift or allelic drift is the change in the frequency of a gene variant (allele) in a population due to random sampling in the absence of a selection pressure. • Genetic drift is important when populations are dramatically reduced. Genes are lost and deleterious genes can also increase. • When there are few copies of an allele, the effect of genetic drift is larger, and when there are many copies the effect is smaller. 9 Genetic Drift • Genetic drift can be most profound in populations that are dramatically reduced (bottle neck populations) usually due to some environmental catastrophe. • Also genetic drift occurs when a small population arrives at a new habitat such as an island. 10 Bottleneck Example In 1900, the population of prairie chickens in Illinois was 100 million but by 1995, the population was reduced to around 50 in Jasper County due to over hunting and habitat destruction which caused the bottleneck to occur. Summarize the data collected and draw a conclusion from that data 11 Founder Effect • The founder effect is the loss of genetic variation that occurs when a new population is established by a very small number of individuals from a larger population and is a special case of genetic drift. • Founder effects are very hard to study! 12 Founder Effect • Biologist got their chance after a hurricane wiped out all the lizard species on certain islands in the Bahamas, scientists repopulated the small islands with two lizard pairs, one having long limbs and one having short limbs. 13 Founder Effect • Before the hurricane, these islands supported populations of a Caribbean lizard, the brown anole, Anolis sagrei. • After the hurricane, seven of the islands were thoroughly searched. No lizards were found. 14 Founder Effect • In May 2005, the researchers randomly selected one male and one female brown anole from lizards collected on a nearby larger island to found new anole populations on seven small islands. • They then sat back and watched how those lizards evolved to get an up-close look at the Founder Effect. 15 Founder Effect • During the next four years, the researchers repeatedly sampled lizards from the source island, from the seven experimental founder islands, and from 12 nearby islands that served as a control. • The team found that all lizard populations adapted to their environment, yet retained characteristics from their founders. 16 A Human Founder Effect Example • The Amish community was founded by a small number of colonist. • The founding group possessed the gene for polydactyly (extra toes or fingers). • The Amish population has increased in size but has remained genetically isolated as few outsiders become a part of the population. • As a result polydactyly is much more frequent in the Amish community than it is in other communities. 17 Impact of Nonrandom Mating • Nonrandom mating also changes allele frequency. • Let’s revisit our adorable frogs and suppose that 4 frogs migrate to a pond some distance from the main pond. • It is likely that these 4 frogs will mate with one another, leaving the rest of the population in the main pond behind to also mate with one another. • Nonrandom mating implies a choice of mates which is more prevalent in animals. 18 Two Types of Sexual Selection • Darwin wrote: “The sexual struggle is of two kinds; in the one it is between individuals of the same sex, generally the males, in order to drive away or kill their rivals, the females remaining passive; whilst in the other, the struggle is likewise between the individuals of the same sex, in order to excite or charm those of the opposite sex, generally the females, which no longer remain passive, but select the more agreeable partners.” 19 Sexual Selection • Sexual selection of mates also affects allele frequency. • The peacock provides a particularly well known example of intersexual selection, where ornate males compete to be chosen by females. • The result is a stunning feathered display, which is large and unwieldy enough to pose a significant survival disadvantage. 20 Sexual Selection • Female birds of many species choose the male. • Males that are “showier” will better attract females. • These males have a selective advantage even though they are more susceptible to predators. 21 Sexual Selection • Females that are drab, blend in to their surroundings and as a result, avoid predators which giving females a survival advantage. • This illustrates that the importance of mating with the correct male outweighs the importance of being preyed upon. • Sexual selection can lead to sexual dimorphism where there is a distinct difference between males and females. 22 Hardy-Weinberg Equilibrium So, when is there no change in the allele frequency? When the population is said to be in Hardy-Weinberg Equilibrium, thus no evolution is occurring. FIVE Conditions of Hardy-Weinberg Equilibrium: 1. Population must be large so chance is not a factor. (No genetic drift). 2. Population must be isolated to prevent gene flow. (No immigration or emigration) 3. No mutations occur. 4. Mating is completely random with respect to time and space. 5. Every offspring has an equal chance of survival without regard to phenotypes. (No natural selection) 23 Hardy-Weinberg Equilibrium • Condition #1 can be met. It is important to have large populations in order that the loss or addition of genes is not a factor. By contrast, small populations experience genetic drift. Additionally, if a small population moves to another area or becomes isolated, the gene pool will be different from the original gene pool. And the founder effect comes into play. • Condition #2 can only be met if the population is isolated. If individuals immigrate or emigrate from the population, the allele frequencies change and evolution occurs. • Condition #3 cannot ever be met since mutations always occur. Thus mutational equilibrium can never be met. 24 Hardy-Weinberg Equilibrium Condition #4 can never be met. Mating is never random. Pollen from an apple tree in Ohio is more likely to pollinate a tree in Ohio than one in Washington state. Condition #5 can never be met. There will always be variation. Variation can help organisms survive longer and/or reproduce more effectively. Since 3 out of the 5 H-W conditions can never be met, evolution DOES occur and allele frequencies do indeed change. 25 Applying the H-W Model Here we go with our frogs again! Let’s suppose that in a population of 100 frogs, 36 were green (CGCG), 48 were blue (CGCR) and 16 were red (CRCR) and there was total random mating. Allele: CG CR Green (36) 72 0 Blue (48) 48 48 Red (16) 0 32 120 80 Total: Frequency: p = 120 ÷ 200 p = 3/5 = 0.60 q = 80 ÷ 200 q = 2/5 = 0.40 Thus, it can be assumed that 60% of all the gametes (eggs and sperm) should carry the CG allele and 40% of the gametes should carry the CR allele. 26 Applying the H-W Model CR 0.40 CG 0.60 CG 0.60 CGCG 0.36 CGCR 0.24 CR 0.40 CGCR 0.24 CRCR 0.16 A population Punnett square is shown above. It indicates that the next generation should have the following offspring distribution: 36% green (CGCG), 48% blue(CGCR), 16% red (CRCR). When the second generation gets ready to reproduce, the results will be the same as before. Allele: CG CR Green (36) 72 0 Blue (48) 48 48 Red (16) 0 32 120 80 Total: Frequency: p = 120 ÷ 200 p = 3/5 = 0.60 q = 80 ÷ 200 q = 2/5 = 0.40 27 Applying the H-W Model So, the allele frequency remains at 0.40 CG and 0.60 CR thus no evolution is taking place. Let’s suppose that there is an environmental change that makes red frogs more obvious to predators. How is the population affected and now the population consists of 36 green, 48 blue, and 6 red frogs? Allele: CG CR Green (36) 72 0 Blue (48) 48 48 Red (6) 0 12 Total: 120 60 Frequency: p = 120 ÷ 180 p = 2/3 = 0.66 q = 60 ÷ 180 q = 1/3 = 0.33 Now, allele frequencies are changing and there is an advantage to being green or blue but NOT red. Evolution is indeed occurring. 28 Deriving the H-W Model CR 0.40 CG 0.60 CG 0.60 CGCG 0.36 CGCR 0.24 CR 0.40 CGCR 0.24 CRCR 0.16 Examine this Punnett square again. If p represents the allele frequency of CG (dominant) and q represents the allele frequency of CR (recessive) then two equations for a population in Hardy-Weinberg equilibrium can be derived where the following genotypes are represented by: CGCG = p2 CRCR = q2 CGCR = 2pq Mathematically then p + q = 0.60 + 0.40 = 1 (1st H-W equation) So, the Punnett square effectively crossed (p + q ) (p + q ) which gives p2 + 2pq + q2 = 1 (2nd H-W equation) 29 Natural Selection Natural Selection is the only mechanism that consistently causes adaptive evolution. •Evolution by natural selection is a blend of chance and “sorting”. – Chance in the context of mutations causing new genetic variations – Sorting in the context of natural selection favoring some alleles over others •This favoring process causes the outcome of natural selection to be anything but random! •Natural Selection consistently increases the frequencies of alleles that provide reproductive advantage and thus leads to adaptive evolution. •Write a specific example of natural selection occurring Relative Fitness • There are animal species in which individuals, usually males, lock horns or otherwise compete through combat for mating privileges. • Reproductive success is usually far more subtle! • Relative fitness is defined as the contribution an individual makes to the gene pool of the next generation relative to the contributions of other individuals.