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Genes Within Populations Chapter 20

Genetic Variation and Evolution Differences in alleles of genes found within individuals in a population Raw material for natural selection Evolution How an entity changes through time Development of modern concept traced to Darwin “Descent with modification”

“Through time, species accumulate differences; as a result, descendants differ from their ancestors. In this way, new species arise from existing ones.” Charles Darwin

Darwin was not the first to propose a theory of evolution Unlike his predecessors, however, Darwin proposed natural selection as the mechanism of evolution Rival theory of Jean-Baptiste Lamarck was evolution by inheritance of acquired characteristics

giraffes has characteristics of modern-day okapi. Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Stretching Stretching Proposed ancestor of giraffes has characteristics of modern-day okapi. The giraffe ancestor lengthened its neck by stretching to reach tree leaves, then passed the change to offspring. Reproduction a. Lamarck’s theory: acquired variation is passed on to descendants.

Some individuals born happen to have longer necks due to Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Some individuals born happen to have longer necks due to genetic differences. Reproduction Individuals pass on their traits to next generation. Natural selection Reproduction Over many generations, longer-necked individuals are more successful, perhaps because they can feed on taller trees, and pass the long-neck trait on to their offspring. b. Darwin’s theory: natural selection or genetically-based variation leads to evolutionary change.

Genetic variation is the raw material for selection Population genetics Study of properties of genes in a population Evolution results in a change in the genetic composition of a population Genetic variation is the raw material for selection In nature, genetic variation is the rule

Polymorphic variation More than one allele at frequencies greater than mutation alone SNPs Used to assess patterns in human and natural populations.

Hardy–Weinberg principle Changes in allele frequency Hardy–Weinberg equilibrium Proportions of genotypes do not change in a population if No mutation takes place No genes are transferred to or from other sources (no immigration or emigration) Random mating is occurring The population size is very large No selection occurs

Principle can be written as an equation Used to calculate allele frequencies For 2 alleles, p and q p = B for black coat color Black cat is BB or Bb q = b for white coat color White cats are bb p2 + 2pq + q2 = 1 BB + Bb + bb = 1

genotype in population 0.36 0.48 0.16 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Generation One Phenotypes 84% 16% Genotypes BB Bb bb Frequency of genotype in population 0.36 0.48 0.16 Frequency of gametes 0.36 + 0.24 = 0.60B 0.24 + 0.16 = 0.40b

Generation Two B p = 0.60 b q = 0.40 Eggs B p = 0.60 BB p2 = 0.36 Bb Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Generation Two B p = 0.60 b q = 0.40 Eggs B p = 0.60 BB p2 = 0.36 Bb pq = 0.24 Sperm b q = 0.40 Bb pq = 0.24 bb q2 = 0.16 p2 + 2 pq + q2 = 1

If all 5 assumptions for Hardy-Weinberg equilibrium are true, allele and genotype frequencies do not change from one generation to the next In reality, most populations will not meet all 5 assumptions To determine this, look for changes in frequency Suggest hypotheses about what process or processes are at work to cause changes to the frequencies

5 agents of evolutionary change Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Mutation Rates generally low Other evolutionary processes usually more important in changing allele frequency Ultimate source of genetic variation Makes evolution possible Mutation Mutagen DNA C G T G C G A G a. The ultimate source of variation. Individual mutations occur so rarely that mutation alone usually does not change allele frequency much.

Gene flow Movement of alleles from one population to another Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Gene flow Movement of alleles from one population to another Animal physically moves into new population Drifting of gametes or immature stages into an area Mating of individuals from adjacent populations Gene Flow b. A very potent agent of change. Individuals or gametes move from one population to another.

Nonrandom mating Assortative mating Disassortative mating Phenotypically similar individuals mate Increases proportion of homozygous individuals Disassortative mating Phenotypically different individuals mate Produces excess of heterozygotes Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Nonrandom Mating Self-fertilization c. Inbreeding is the most common form. It does not alter allele frequency but reduces the proportion of heterozygotes.

Copyright © The McGraw-Hill Companies, Inc Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Genetic Drift Genetic drift In small populations, allele frequency may change by chance alone Magnitude of genetic drift is negatively related to population size Founder effect Bottleneck effect d. Statistical accidents. The random fluctuation in allele frequencies increases as population size decreases.

Genetic drift Alters allele frequencies in small populations Population must be large Small number of individuals drift from population Can lead to the loss of alleles in isolated populations

Genetic drift can lead to the loss of alleles in isolated populations Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Parent population Bottleneck (drastic reduction in population) Surviving individuals Next generation Genetic drift can lead to the loss of alleles in isolated populations Alleles that initially are uncommon are particularly vulnerable

Bottleneck effect If organisms do not move from place to place their populations may be drastically reduced Survivors may constitute a random genetic sample of the original population Results in loss of genetic variability

Northern Elephant Seal Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. population in 1890, reduced to inhabiting Guadalupe only UNITED STATES current population Guadalupe MEXICO Northern Elephant Seal Bottleneck case study Nearly hunted to extinction in 19th century As a result, species has lost almost all of its genetic variation Population now numbers in tens of thousands

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Selection Some individuals leave behind more progeny than others, and the rate at which they do so is affected by phenotype and behavior Artificial selection Natural selection

3 conditions for natural selection to occur and to result in evolutionary change Variation must exist among individuals in a population Variation among individuals must result in differences in the number of offspring surviving in the next generation Variation must be genetically inherited

Natural selection and evolution are not the same Natural selection is a process Only one of several processes that can result in evolution Evolution is the historical record, or outcome, of change through time Result of evolution driven by natural selection is that populations become better adapted to their environment

Pocket mice come in different colors Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Light coat color pocket mouse is vulnerable on lava rock Light coat color favored by natural selection because it matches sand color Dark coat color favored by natural selection because it matches black lava color Pocket mice come in different colors Population living on rocks favor dark color Populations living on sand favor light color

Housefly has pesticide resistance alleles at Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Housefly has pesticide resistance alleles at pen gene decreases insecticide uptake kdr and dld-r genes decrease target sites for insecticide Pesticide molecule Target site Resistant target site Insect cell membrane a. Insect cells with resistance allele at pen gene: decreased uptake of the pesticide. Target site b. Insect cells with resistance allele at kdr gene: decreased number of target sites for the pesticide.

Fitness and its measurement Individuals with one phenotype leave more surviving offspring in the next generation than individuals with an alternative phenotype Relative concept; the most fit phenotype is simply the one that produces, on average, the greatest number of offspring

Fitness has many components Survival Sexual selection – some individuals more successful at attracting mates Number of offspring per mating Traits favored for one component may be a disadvantage for others Selection favors phenotypes with the greatest fitness Phenotype with greater fitness usually increases in frequency

Larger female water striders lay more eggs per day Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. 8 50 200 40 6 150 Number of Eggs Laid per Day 30 4 Life Span of Adult Female (days) Number of Eggs Laid During Lifetime 100 20 2 50 10 12 13 14 15 16 12 13 14 15 16 12 13 14 15 16 Length of Adult Female Water Strider (mm) Length of Adult Female Water Strider (mm) Length of Adult Female Water Strider (mm) Larger female water striders lay more eggs per day Large females survive for a shorter period of time As a result, intermediate-sized females produce the most offspring over the course of their entire lives and thus have the highest fitness

Interactions Mutations and genetic drift may counter selection In nature, mutation rates are rarely high enough to counter selection Selection is nonrandom but genetic drift is random Drift may decrease an allele favored by selection Selection usually overwhelms drift except in small populations

Gene flow can be Constructive Constraining Spread beneficial mutation to other populations Constraining Can impede adaptation by continual flow of inferior alleles from other populations

Maintenance of variation Frequency-dependent selection Fitness of a phenotype depends on its frequency within the population Negative frequency-dependent selection Rare phenotypes favored by selection Rare forms may not be in “search image” Positive frequency-dependent selection Favors common form Tends to eliminate variation “Oddballs” stand out

Taken by Fish Predators Color Type Frequency in Population Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Negative frequency-dependent selection In water boatman, fish eat the most common color type more than they would by chance alone 100 80 60 Percent of Color Type Taken by Fish Predators 40 Color type of water boatman 20 dark brown medium brown light brown 20 40 60 80 100 Color Type Frequency in Population

Positive frequency-dependent selection

Oscillating selection Selection favors one phenotype at one time and another phenotype at another time Effect will be to maintain genetic variation in the population Medium ground finch of Galápagos Islands Birds with big bills favored during drought Birds with smaller bills favored in wet conditions

Heterozygote advantage Heterozygotes are favored over homozygotes Works to maintain both alleles in the population Sickle cell anemia Hereditary disease affecting hemoglobin Causes severe anemia Homozygotes for sickle cell allele usually die before reproducing (without medical treatment)

Why is the sickle cell allele not eliminated? Leading cause of death in central Africa is malaria Heterozygotes for sickle cell allele do not suffer anemia and are much less susceptible to malaria Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Normal red blood cells Sickled red blood cells Sickle cell allele in Africa 1–5% 5–10% 10–20% Geographic distribution of P. falciparum

Selection Many traits affected by more than one gene Selection operates on all the genes for the trait Changes the population depending on which genotypes are favored Types of selection Disruptive Directional Stabilizing

Acts to eliminate intermediate types Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Disruptive selection Acts to eliminate intermediate types Different beak sizes of African black-bellied seedcracker finch Available seeds fall into 2 categories Favors bill sizes for one or the other Number of Individuals 25 50 75 100 125 Body Size (g) Selection for small and large individuals Two peaks form Number of Individuals 25 50 75 100 125 Body Size (g) a. Disruptive selection

Birds with intermediate-sized beaks are at a disadvantage with both seed types – they are unable to open large seeds and too clumsy to efficiently process small seeds Fix runover style in em dash entry?

Directional selection Acts to eliminate one extreme Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Directional selection Acts to eliminate one extreme Often occurs in nature when the environment changes In Drosophila, artificially selected flies that moved toward the light Now fewer have that behavior Number of Individuals 25 50 75 100 125 Body Size (g) Selection for larger individuals Peak shifts Number of Individuals 25 50 75 100 125 Body Size (g) b. Directional selection

Directional selection for negative phototropism in Drosophila Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Light 11 10 9 8 7 Average Tendency to Fly Toward Light 6 5 4 3 2 Dark 1 2 4 6 8 10 12 14 16 18 20 Number of Generations Directional selection for negative phototropism in Drosophila

Stabilizing selection Acts to eliminate both extremes Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Stabilizing selection Acts to eliminate both extremes Makes intermediate more common by eliminating extremes In humans, infants with intermediate weight at birth have the highest survival rate Number of Individuals 25 50 75 100 125 Body Size (g) Selection for mid-size individuals Distribution gets narrower Number of Individuals 25 50 75 100 125 Body Size (g) c. Stabilizing selection

Stabilizing selection for birth weight in humans Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. births in population infant mortality 20 100 70 15 50 30 20 Percent of Births in Population Percent Infant Mortality 10 10 7 5 5 3 2 2 3 4 5 6 7 8 9 10 Birth Weight in Pounds Stabilizing selection for birth weight in humans

Experimental studies To study evolution, biologists have traditionally investigated what has happened in the past Fossils or DNA evidence Laboratory studies on fruit flies common for more than 50 years Only recently started with lab and field experiments

Guppy coloration Found in small streams in northeastern South America and Trinidad Some are capable of colonizing portions of streams above waterfalls Different dispersal methods Other species not able to make it upstream Dispersal barriers create 2 different environments Predators rare above waterfall

(Poecilia reticulata) (Poecilia reticulata) Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Pike cichlid (predator) rare above waterfall Killifish rarely eats guppies Guppy males larger and gaudier Predator common below waterfall Individuals more drab and reproduce earlier Killifish (Rivulus hartii) Guppy (Poecilia reticulata) Pikecichlid (Crenicichla alta) Guppy (Poecilia reticulata)

Guppy lab study Other explanations are possible for field results 10 large pools Added pike cichlids to 4, killifish to 4, and 2 left as controls 14 months and 10 guppy generations later Guppies in killifish and control pool – large and colorful Guppies in pike cichlid pools – smaller and drab

Duration of Experiment (months) Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. no predation low predation 14 high predation 13 12 11 Spots per Fish 10 9 8 4 8 12 Duration of Experiment (months)

Constraints on evolutionary change Mutation and genetic drift May counter or promote selection Natural in abandoned mine sites in Great Britain Copper tolerance in plants

Slender bent grass at copper mines Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Pollen Prevailing wind Non- mine Mine Site Nonmine 60 Index of Copper Tolerance 40 Bent grass (Agrostis tenuis) 20 20 20 40 60 80 100 120 140 160 Distance Upwind from Mine (m) Distance Away from Mine (m) Slender bent grass at copper mines Resistance allele occurs at intermediate levels in many areas Individuals with resistance gene grow slower on uncontaminated sites Gene flow between sites high enough to counteract selection

Limits of selection Multiple phenotypic effects of alleles Larger clutch size leads to thinner shelled eggs Lack of genetic variation Gene pool of thoroughbreds limited and performance times have not improved for more than 50 years Phenotypic variation may not have genetic basis Interactions between genes – epistasis Selective advantage of an allele at one gene may vary from one genotype to another

Kentucky Derby Winning Time (seconds) Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. 130 125 120 Kentucky Derby Winning Time (seconds) 115 110 1900 1920 1940 1960 1980 2000 Year Selection for increased speed in racehorses is no longer effective

Copyright © The McGraw-Hill Companies, Inc Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Right eye of insect Left eye of insect Ommatidia Differences in the number of ommatidia in fly eyes does not have a genetic basis