Chapter 23

The Evolution of Populations Lecture Outline

Overview: The Smallest Unit of Evolution One common misconception about evolution is that organisms evolve, in a Darwinian sense, during their lifetimes. Natural selection does act on individuals. Each individuals combination of inherited traits affects its survival and its reproductive success relative to other individuals in the population. However, the evolutionary impact of natural selection is only apparent in the changes in a population of organisms over time. It is the population, not the individual, that evolves. onsider the e!ample of bent grass "#grostis tenuis$ growing on the tailings of an abandoned mine. %hese tailings are rich in to!ic heavy metals. &hile many bent grass seeds land on the mine tailings each year, the only plants that germinate, grow, and reproduce are those that possess genes enabling them to tolerate metallic soils. %hese plants tend to produce metal'tolerant offspring. Individual plants do not evolve to become more metal' tolerant during their lifetimes. Concept 23.1 Population enetics provi!es a foun!ation for stu!"in evolution Darwin proposed a mechanism for change in species over time. &hat was missing from Darwins e!planation was an understanding of inheritance that could e!plain how chance variations arise in a population while also accounting for the precise transmission of these variations from parents to offspring. %he widely accepted hypothesis of the time(that the traits of parents are blended in their offspring(would eliminate the differences in individuals over time. )ust a few years after Darwin published On the Origin of *pecies, +regor ,endel proposed a model of inheritance that supported Darwins theory. ,endels particulate hypothesis of inheritance stated that parents pass on discrete heritable units "genes$ that retain their identities in offspring.
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 #lthough +regor ,endel and harles Darwin were contemporaries, Darwin never saw ,endels paper, and its implications were not understood by the few scientists who did read it at the time. ,endels contribution to evolutionary theory was not appreciated until half a century later. The mo!ern evolutionar" s"nthesis inte rate! #arwinian selection an! $en!elian inheritance. &hen ,endels research was rediscovered in the early -.th century, many geneticists believed that his laws of inheritance conflicted with Darwins theory of natural selection. Darwin emphasi/ed 0uantitative characters, those that vary along a continuum. %hese characters are influenced by multiple loci. ,endel and later geneticists investigated discrete 1either' or2 traits. It was not obvious that there was a genetic basis to 0uantitative characters. &ithin a few decades, geneticists determined that 0uantitative characters are influenced by multiple genetic loci and that the alleles at each locus follow ,endelian laws of inheritance. %hese discoveries helped reconcile Darwins and ,endels ideas and led to the birth of population enetics% the study of how populations change genetically over time. # comprehensive theory of evolution, the mo!ern s"nthesis% too3 form in the early 456.s. It integrated discoveries and ideas from paleontology, ta!onomy, biogeography, and population genetics. %he first architects of the modern synthesis included statistician 7. #. 8isher, who demonstrated the rules by which ,endelian characters are inherited, and biologist ). 9. *. Haldane, who e!plored the rules of natural selection. :ater contributors included geneticists %heodosius Dob/hans3y and *ewall &right, biogeographer and ta!onomist Ernst ,ayr, paleontologist +eorge +aylord *impson, and botanist +. :edyard *tebbins. %he modern synthesis emphasi/es; %he importance of populations as the units of evolution. %he central role of natural selection as the most important mechanism of adaptive evolution. %he idea of gradualism to e!plain how large changes can evolve as an accumulation of small changes over long periods of time. &hile many evolutionary biologists are now challenging some of the assumptions of the modern synthesis, it has shaped our ideas about how populations evolve.
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& population's ene pool is !efine! (" its allele fre)uencies. # population is a locali/ed group of individuals that belong to the same species. One definition of a species is a group of natural populations whose individuals have the potential to interbreed and produce fertile offspring. <opulations of a species may be isolated from each other and rarely e!change genetic material. ,embers of a population are far more li3ely to breed with members of the same population than with members of other populations. Individuals near the populations center are, on average, more closely related to one another than to members of other populations. %he total aggregate of genes in a population at any one time is called the populations ene pool. It consists of all alleles at all gene loci in all individuals of a population. If only one allele e!ists at a particular locus in a population, that allele is said to be fi!ed in the gene pool, and all individuals will be homo/ygous for that gene. If there are two or more alleles for a particular locus, then individuals can be either homo/ygous or hetero/ygous for that gene. Each allele has a fre0uency in the populations gene pool. 8or e!ample, imagine a population of =.. wildflower plants with two alleles " 7 and &$ at a locus that codes for flower pigment. *uppose that in the imaginary population of =.. plants, -. "6>$ are homo/ygous for the & allele " & &$ and have white flowers. Of the remaining plants, ?-. "@6>$ are homo/ygous for the 7 allele " 7 7$ and have red flowers. %hese alleles show incomplete dominance. 4@. "?->$ of the plants are hetero/ygous " 7 &$ and produce pin3 flowers. 9ecause these plants are diploid, the population of =.. plants has 4,... copies of the gene for flower color. %he dominant allele " 7$ accounts for A.. copies "?-. B 7 7 for C 4@. B 4 for 7 &$. %he fre0uency of the 7 allele in the gene pool of this population is A..D4,... E ..A, or A.>. %he & allele must have a fre0uency of 4.. F ..A E ..-, or -.>. &hen there are two alleles at a locus, the convention is to use p to represent the fre0uency of one allele and 0 to represent the fre0uency of the other.
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 %hus p, the fre0uency of the 7 allele in this population, is ..A. %he fre0uency of the & allele, represented by 0, is ..-. The *ar!"+,ein(er Theorem !escri(es a nonevolvin population. %he Hardy'&einberg theorem describes the gene pool of a nonevolving population. %his theorem states that the fre0uencies of alleles and genotypes in a populations gene pool will remain constant over generations unless acted upon by agents other than ,endelian segregation and recombination of alleles. %he shuffling of alleles by meiosis and random fertili/ation has no effect on the overall gene pool of a population. In our imaginary wildflower population of =.. plants, A.> "..A$ of the flower color alleles are 7, and -.> "..-$ are &. How will meiosis and se!ual reproduction affect the fre0uencies of the two alleles in the ne!t generationG &e assume that fertili/ation is completely random and all male'female mating combinations are e0ually li3ely. 9ecause each gamete has only one allele for flower color, we e!pect that a gamete drawn7from the gene pool at random has a ..A chance of bearing an allele and a ..- chance of bearing an & allele. *uppose that the individuals in a population not only donate gametes to the ne!t generation at random, but also mate at random. In other words, all male'female matings are e0ually li3ely. %he allele fre0uencies in this population will not change from one generation to the ne!t. Its genotype fre0uencies, which can be predicted from the allele fre0uencies, will also remain unchanged. 8or the flower'color locus, the populations genetic structure is in a state of *ar!"+,ein(er e)uili(rium. Hsing the rule of multiplication, we can determine the fre0uencies of the three possible genotypes in the ne!t generation. %he probability of pic3ing two 7 alleles "to obtain a 7 7 genotype$ is ..A B ..A E ..@6, or @6>. %he probability of pic3ing two & alleles "to obtain a & & genotype$ is ..- B ..- E ...6, or 6>. Hetero/ygous individuals are either 7 & or & 7, depending 7 on whether the allele arrived via sperm or egg. %he probability of being hetero/ygous "with a 7 & 7 & genotype$ is ..A B ..- E ..4@ for , ..- B ..A E ..4@ for & 7 , and ..4@ C ..4@ E ..?-, or ?->, for 7 & C & 7.
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 #s you can see, the processes of meiosis and random fertili/ation have maintained the same allele and genotype fre0uencies that e!isted in the previous generation. %he Hardy'&einberg theorem states that the repeated shuffling of a populations gene pool over generations does not increase the fre0uency of one allele over another. %heoretically, the allele fre0uencies in our flower population should remain at ..A for 7 and ..- for & forever. %o generali/e the e!ample, in a population with two alleles with fre0uencies of p and 0, the combined fre0uencies must add to 4..>. %herefore p C 0 E 4. If p C 0 E 4, then p E 4 F 0 and 0 E 4 F p. 7In the wildflower e!ample, p is the fre0uency of red alleles " $ and 0 is the fre0uency of white alleles " &$. %he probability of generating an 7 7 offspring is p- "an application of the rule of multiplication$. In our e!ample, p E ..A and p- E ..@6. %he probability of generating a & & offspring is 0-. In our e!ample, 0 E ..- and 0- E ...6. %he probability of generating a 7 & offspring is -p0. In our e!ample, - B ..A B ..- E ..?-. %he genotype fre0uencies must add up to 4..; p- C -p0 C 0- E 4.. 8or the wildflowers, ..@6 C ..?- C ...6 E 4... %his general formula is the *ar!"+,ein(er e)uation. Hsing this formula, we can calculate fre0uencies of alleles in a gene pool if we 3now the fre0uency of genotypes, or the fre0uency of genotypes if we 3now the fre0uencies of alleles. -ive con!itions must (e met for a population to remain in *ar!"+,ein(er e)uili(rium. %he Hardy'&einberg theorem describes a hypothetic population that is not evolving. However, real populations do evolve, and their allele and genotype fre0uencies do change over time. %hat is because the five conditions for nonevolving populations are rarely met for long in nature. # population must satisfy five conditions if it is to remain in Hardy'&einberg e0uilibrium; 1. E!tremely large population si/e. In small populations, chance fluctuations in the gene pool can cause genotype fre0uencies to change over time. %hese random changes are called enetic !rift.
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2. No gene flow. +ene flow, the transfer of alleles due to the migration of individuals or gametes between populations, can change the proportions of alleles. 3. No mutations. Introduction, loss, or modification of genes will alter the gene pool. 4. 7andom mating. If individuals pic3 mates with certain genotypes, or if inbreeding is common, the mi!ing of gametes will not be random. 5. No natural selection. Differential survival or reproductive success among genotypes will alter their fre0uencies. Evolution usually results when any of these five conditions are not met. #lthough natural populations are rarely, if ever, in true Hardy'&einberg e0uilibrium, the rate of evolutionary change in many populations is so slow that they appear to be close to e0uilibrium. In such cases, we can use the Hardy'&einberg e0uation to estimate genotype and allele fre0uencies. &e can use the theorem to estimate the percentage of the human population that carries the allele for the inherited disease phenyl3etonuria "<IH$. #bout 4 in 4.,... babies born in the Hnited *tates is born with <IH, a metabolic condition that results in mental retardation and other problems if left untreated. %he disease is caused by a recessive allele. Is the H.*. population in Hardy'&einberg e0uilibrium with respect to the <IH geneG 1. %he H.*. population is very large. 2. <opulations outside the Hnited *tates have <IH allele fre0uencies similar to those seen in the Hnited *tates, so gene flow will not alter allele fre0uencies significantly. 3. %he mutation rate for the <IH gene is very low. 4. <eople do not choose their partners based on whether or not they carry the <IH allele, and inbreeding "marriage to close relatives$ is rare in the Hnited *tates. 5. *election against <IH only acts against the rare hetero/ygous recessive individuals. 8rom the epidemiological data, we 3now that fre0uency of homo/ygous recessive individuals "0- in the Hardy'&einberg theorem$ E 4 in 4.,..., or .....4. %he fre0uency of the recessive allele "0$ is the s0uare root of .....4 E ...4. %he fre0uency of the dominant allele "p$ is p E 4 F 0, or 4 F ...4 E ..55. %he fre0uency of carriers "hetero/ygous individuals$ is -p0 E - B ..55 B ...4 E ...45A, or about ->. %hus, about -> of the H.*. population carries the <IH allele.
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Concept 23.2 $utation an! se.ual recom(ination pro!uce the variation that ma/es evolution possi(le 0ew enes an! new alleles ori inate onl" (" mutation. # mutation is a change in the nucleotide se0uence of an organisms DN#. ,ost mutations occur in somatic cells and are lost when the individual dies. Only mutations in cell lines that form gametes can be passed on to offspring, and only a small fraction of these spread through populations and become fi!ed. # new mutation that is transmitted in a gamete to an offspring can immediately change the gene pool of a population by introducing a new allele. # point mutation is a change of a single base in a gene. <oint mutations can have a significant impact on phenotype, as in the case of sic3le'cell disease. However, most point mutations are harmless. ,uch of the DN# in eu3aryotic genomes does not code for protein products. However, some noncoding regions of DN# do regulate gene e!pression. hanges in these regulatory regions of DN# can have profound effects. 9ecause the genetic code is redundant, some point mutations in genes that code for proteins may not alter the proteins amino acid composition. On rare occasions, a mutant allele may actually ma3e its bearer better suited to the environment, increasing reproductive success. %his is more li3ely when the environment is changing. *ome mutations alter gene number or se0uence. hromosomal mutations that delete or rearrange many gene loci at once are almost always harmful. In rare cases, chromosomal rearrangements may be beneficial. 8or e!ample, the translocation of part of one chromosome to a different chromosome could lin3 genes that act together to positive effect. +ene !uplication is an important source of new genetic variation.
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 *mall pieces of DN# can be introduced into the genome through the activity of transposons. *uch duplicated segments can persist over generations and provide new loci that may eventually ta3e on new functions by mutation and subse0uent selection. New genes may also arise when the coding subsections of genes 3nown as e!ons are shuffled within the genome, within a single locus or between loci. *uch beneficial increases in gene number appear to have played a maJor role in evolution. 8or e!ample, mammalian ancestors carried a single gene for detecting odors that has been duplicated though various mutational mechanisms. ,odern humans have close to 4,... olfactory receptor genes. @.> of these genes have been inactivated in humans, due to mutations. ,ice, who rely more on their sense of smell, have lost only -.> of their olfactory receptor genes. ,utation rates vary from organism to organism. ,utation rates are low in animals and plants, averaging about 4 mutation in every 4..,... genes per generation. In microorganisms and viruses with short generation spans, mutation rates are much higher and can rapidly generate genetic variation. Se.ual recom(ination also pro!uces enetic variation. On a generation'to'generation timescale, se!ual recombination is far more important than mutation in producing the genetic differences that ma3e adaptation possible. *e!ual reproduction rearranges alleles into novel combinations every generation. 9acteria and viruses can also undergo recombination, but they do so less regularly than animals and plants. 9acterial and viral recombination may cross species barriers. Concept 23.3 0atural selection% enetic !rift% an! ene flow can alter a population's enetic composition #lthough new mutations can modify allele fre0uencies, the change from generation to generation is very small. 7ecombination reshuffles alleles but does not change their fre0uency.
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 %hree maJor factors alter allele fre0uencies to bring about evolutionary change; natural selection, genetic drift, and gene flow. 0atural selection is (ase! on !ifferential repro!uctive success. Individuals in a population vary in their heritable traits. %hose with variations better suited to the environment tend to produce more offspring than those with variations that are less well suited. #s a result of selection, alleles are passed on to the ne!t generation in fre0uencies different from their relative fre0uencies in the present population. Imagine that in our imaginary wildflower population, white flowers are more visible to herbivorous insects and thus have lower survival. Imagine that red flowers are more visible to pollinators. *uch differences in survival and reproductive success would disturb the Hardy'&einberg e0uilibrium. %he fre0uency of the & allele would decline and the fre0uency of the 7 allele would increase. 1enetic !rift results from chance fluctuations in allele fre)uencies in small populations. 1enetic !rift occurs when changes in gene fre0uencies from one generation to another occur because of chance events "sampling errors$ that occur in small populations. 8or e!ample, you would not be too surprised if a thrown coin produced seven heads and three tails in ten tosses, but you would be surprised if you saw K.. heads and ?.. tails in 4,... tosses(you would e!pect close to =.. of each. %he smaller the sample, the greater the chance of deviation from the e!pected result. In a large population, allele fre0uencies will not change from generation to generation by chance alone. However, in a small wildflower population with a stable si/e of only ten plants, genetic drift can completely eliminate some alleles. +enetic drift at small population si/es may occur as a result of two situations; the bottlenec3 effect or the founder effect. %he (ottlenec/ effect occurs when the numbers of individuals in a large population are drastically reduced by a disaster. 9y chance, some alleles may be overrepresented and others underrepresented among the survivors. *ome alleles may be eliminated altogether.
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 +enetic drift will continue to change the gene pool until the population is large enough to eliminate the effect of chance fluctuations. %he bottlenec3 effect is an important concept in conservation biology of endangered species. <opulations that have suffered bottlenec3 incidents have lost genetic variation from the gene pool. %his reduces individual variation and may reduce adaptation. 8or e!ample, in the 4A5.s, hunters reduced the population of northern elephant seals in alifornia to -. individuals. Now that it is a protected species, the population has increased to more than ?.,.... However, a study of -6 gene loci in a representative sample of seals showed no variation. One allele had been fi!ed for each gene. <opulations of the closely related southern elephant seal, which did not go through a bottlenec3, show abundant genetic variation. %he foun!er effect occurs when a new population is started by only a few individuals who do not represent the gene pool of the larger source population. #t an e!treme, a population could be started by a single pregnant female or single seed with only a tiny fraction of the genetic variation of the source population. +enetic drift would continue from generation to generation until the population grew large enough for sampling errors to be minimal. 8ounder effects have been demonstrated in human populations that started from a small group of colonists. & population ma" lose or ain alleles (" ene flow. 1ene flow is genetic e!change due to migration of fertile individuals or gametes between populations. 8or e!ample, if a nearby wildflower population consisted entirely of white flowers, its pollen " & alleles only$ could be carried into our target population. %his would increase the fre0uency of & alleles in the target population in the ne!t generation. +ene flow tends to reduce differences between populations. If e!tensive enough, gene flow can amalgamate neighboring populations into a single population with a common gene pool. Humans today migrate much more freely than in the past, and gene flow has become an important agent of evolutionary change in human populations that were previously isolated.
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Concept 23.2 0atural selection is the primar" mechanism of a!aptive evolution Of all the factors that can change a gene pool, only natural selection leads to adaptation of an organism to its environment. Natural selection accumulates and maintains favorable genotypes in a population. ,ost populations have e!tensive genetic variation. Not all variation is heritable. 8or e!ample, body builders alter their phenotypes but do not pass on their huge muscles to their children. Only the genetic component of variation can have evolutionary conse0uences as a result of natural selection. %his is because only heritable traits pass from generation to generation. 1enetic variation occurs within an! (etween populations. 9oth 0uantitative and discrete characters contribute to variation within a population. Luantitative characters are those that vary along a continuum within a population. 8or e!ample, plant height in a wildflower population ranges from short to tall. Luantitative variation is usually due to polygenic inheritance in which the additive effects of two or more genes influence a single phenotypic character. Discrete characters, such as flower color, are usually determined by a single locus with different alleles that produce distinct phenotypes. Phenot"pic pol"morphism occurs when two or more discrete phenotypes are represented in high enough fre0uencies to be noticeable in a population. %he contrasting forms are called morphs, as in the red' flowered and white'flowered morphs in our wildflower population. Human populations are polymorphic for a variety of physical "e.g., frec3les$ and biochemical "e.g., blood types$ characters. <olymorphism applies only to discrete characters, not 0uantitative characters. Human height, which varies in a continuum, is not a phenotypic polymorphism. <opulation geneticists measure genetic variation by determining the amount of hetero/ygosity at the level of whole genes "gene variability$ and at the molecular level of DN# "nucleotide variability$.
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 &vera e hetero3" osit" measures gene variability, the average percent of gene loci that are hetero/ygous. In the fruit fly "Drosophila$, about A@> of their 4?,... gene loci are homo/ygous "fi!ed$. #bout 46> "4,A.. genes$ are hetero/ygous. 0ucleoti!e varia(ilit" measures the mean level of difference in nucleotide se0uences "base pair differences$ among individuals in a population. In fruit flies, about 4> of the bases differ between two individuals. %wo individuals differ, on average, at 4.A million of the 4A. million nucleotides in the fruit fly genome. &hy does average hetero/ygosity tend to be greater than nucleotide diversityG %his is because a gene can consist of thousands of bases of DN#. # difference at only one of these bases is sufficient to ma3e two alleles of that gene different and count toward average hetero/ygosity. Humans have relatively little genetic variation. Nucleotide diversity is only ..4>. Mou and your neighbor probably have the same nucleotide at 555 out of every 4,... nucleotide sites in your DN#. 1eo raphic variation results from differences in phenotypes or genotypes between populations or between subgroups of a single population that inhabit different areas. Natural selection contributes to geographic variation by modifying gene fre0uencies in response to differences in local environmental factors. +enetic drift can also lead to variation among populations through the cumulative effect of random fluctuations in allele fre0uencies. +eographic variation can occur on a local scale, within a population, if the environment is patchy or if dispersal of individuals is limited, producing subpopulations. %his is termed spatial variation. +eographic variation in the form of graded change in a trait along a geographic a!is is called a cline. lines may represent intergrade /ones where individuals from neighboring, genetically different, populations interbreed. #lternatively, clines may reflect the influence of natural selection based on gradation in some environmental variable. 8or e!ample, the average body si/e of many North #merican species of birds and mammals increases gradually with increasing latitude, allowing Northern populations to conserve heat in cold environments by decreasing the ratio of surface area to volume.
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Let's ta/e a closer loo/ at natural selection. %he terms 1struggle for e!istence2 and 1survival of the fittest2 are misleading because they suggest that individuals compete directly in contests. In some animal species, males do compete directly for mates. 7eproductive success is generally subtler and depends on factors other than battle for mates. 8or e!ample, a barnacle may produce more eggs than its neighbors because it is more efficient at filtering food from the water. &ildflowers may be successful because they attract more pollinators. %hese e!amples of adaptive advantage are all components of evolutionary fitness. 8itness is defined as the contribution an individual ma3es to the gene pool of the ne!t generation, relative to the contributions of other individuals. <opulation geneticists define relative fitness as the contribution of a genotype to the ne!t generation compared to the contribution of alternative genotypes for the same locus. onsider our wildflower population. :ets assume that individuals with red flowers produce fewer offspring than those with white or pin3 flowers, which produce e0ual numbers of offspring. %he relative fitness of the most successful variants is set at 4.. as& a basis for comparison, so the relative fitness of & white " $ and pin3 " 7 &$ plants is 4... If plants with red flowers " 7 7$ produce only A.> as many offspring, their relative fitness is ..A. #lthough population geneticists measure the relative fitness of a genotype, it is important to remember that natural selection acts on phenotypes, not genotypes. %he whole organism is subJected to natural selection. %he relative fitness of an allele depends on the entire genetic and environmental conte!t in which it is e!pressed. *urvival alone does not guarantee reproductive success. 7elative fitness is /ero for a sterile organism, even if it is robust and long'lived. On the other hand, longevity may increase fitness if long' lived individuals leave more offspring than short'lived individuals. In many species, individuals that mature 0uic3ly, become fertile at an early age, and live for a short time have greater relative fitness than individuals that live longer but mature later.
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There are three mo!es of selection: !irectional% !isruptive% an! sta(ili3in . Natural selection can alter the fre0uency distribution of heritable traits in three ways, depending on which phenotypes in a population are favored. %he three modes of selection are called directional, disruptive, and stabili/ing selection. #irectional selection is most common during periods of environmental change or when members of a population migrate to a new habitat with different environmental conditions. Directional selection shifts the fre0uency curve for a phenotypic character in one direction by favoring individuals who deviate from the average. 8or e!ample, fossil evidence indicates that the average si/e of blac3 bears in Europe increased during each glacial period, only to decrease again during the warmer interglacial periods. :arge bears have a smaller surface'to'volume ratio and are better at conserving body heat during periods of e!treme cold. #isruptive selection occurs when environmental conditions favor individuals at both e!tremes of the phenotypic range over those with intermediate phenotypes. 8or e!ample, two distinct bill types are present in ameroons blac3'bellied seedcrac3ers. :arger'billed birds are more efficient in feeding on hard seeds and smaller' billed birds are more efficient in feeding on soft seeds. 9irds with intermediate bills are relatively inefficient at crac3ing both types of seeds and thus have lower relative fitness. Disruptive selection can be important in the early stages of speciation. Sta(ili3in selection favors intermediate variants and acts against e!treme phenotypes. *tabili/ing selection reduces variation and maintains the status 0uo for a trait. Human birth weight is subJect to stabili/ing selection. 9abies much larger or smaller than ?N6 3g have higher infant mortality than average'si/ed babies. #iploi!" an! (alancin selection preserve enetic variation. %he tendency for natural selection to reduce variation is countered by mechanisms that preserve or restore variation, including diploidy and balanced polymorphisms. Diploidy in eu3aryotes prevents the elimination of recessive alleles via selection because recessive alleles do not affect the phenotype in hetero/ygotes.
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 Even recessive alleles that are unfavorable can persist in a population through their propagation by hetero/ygous individuals. 7ecessive alleles are only e!posed to selection when both parents carry the same recessive allele and combine two recessive alleles in one /ygote. %his happens only rarely when the fre0uency of the recessive allele is very low. %he rarer the recessive allele, the greater the degree of protection it has from natural selection. Hetero/ygote protection maintains a huge pool of alleles that may not be suitable under the present conditions but may become beneficial when the environment changes. Natural selection itself preserves variation at some gene loci. 9alancing selection occurs when natural selection maintains stable fre0uencies of two or more phenotypes in a population, a state called (alance! pol"morphism. One mechanism producing balanced polymorphism is hetero3" ote a!vanta e. In some situations, individuals who are hetero/ygous at a particular locus have greater fitness than homo/ygotes. In these cases, natural selection will maintain multiple alleles at that locus. Hetero/ygous advantage maintains genetic diversity at the human gene for one chain of hemoglobin. Homo/ygous recessive individuals suffer from sic3le'cell disease. Homo/ygous dominant individuals are vulnerable to malaria. Hetero/ygous individuals are resistant to malaria. %he fre0uency of the sic3le'cell allele is highest in areas where the malarial parasite is common. In some #frican tribes, it accounts for -.> of the gene pool, a very high fre0uency for such a harmful allele. Even at this high fre0uency, only 6> of the population suffers from sic3le'cell disease "0- E ..- B ..- E ...6$, while ?-> of the population is resistant to malaria "-p0 E - B ..A B ..- E ..?-$. %he aggregate benefit of the sic3le'cell allele in the population balances its aggregate harm. # second mechanism promoting balanced polymorphism is fre)uenc"+!epen!ent selection. 8re0uency'dependent selection occurs when the fitness of any one morph declines if it becomes too common in the population.
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 <redators may develop 1search images2 of the most common forms of prey. # prey morph that becomes too common may become disproportionately vulnerable to predation. 8re0uency'dependent selection has been observed in a number of predator'prey interactions in the wild. *ome genetic variations, neutral variations% have negligible impact on fitness, and thus natural selection does not affect these alleles. 8or e!ample, the diversity of human fingerprints seems to confer no selective advantage to some individuals over others. ,ost of the base differences between humans that are found in untranslated parts of the genome appear to confer no selective advantage. Pseu!o enes% genes that have become inactivated by mutations, accumulate genetic variations. Over time, some neutral alleles will increase and others will decrease by the chance effects of genetic drift. %here is no consensus among biologists on how much genetic variation can be classified as neutral or even if any variation can be considered truly neutral. It is almost impossible to demonstrate that an allele brings no benefit at all to an organism. #lso, variant alleles may be neutral in one environment but not in another. Even if only a fraction of the e!tensive variation in a gene pool significantly affects an organism, there is still an enormous reservoir of raw material for natural selection and adaptive evolution. Se.ual selection ma" lea! to pronounce! secon!ar" !ifferences (etween the se.es. harles Darwin was the first scientist to investigate se!ual selection, which is natural selection for mating success. *e!ual selection results in se.ual !imorphism% mar3ed differences between the se!es in secondary se!ual characteristics not directly associated with reproduction. ,ales and females may differ in si/e, coloration, and ornamentation. In vertebrates, males are usually the larger and showier se!. It is important to distinguish between intrase!ual and interse!ual selection. 4ntrase.ual selection is direct competition among individuals of one se! "usually males$ for mates of the opposite se!.
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ompetition may ta3e the form of direct physical battles between individuals. %he stronger individuals gain status. ,ore commonly, rituali/ed displays discourage lesser competitors and determine dominance. Evidence is growing that intrase!ual selection can ta3e place between females as well. 4nterse.ual selection or mate choice occurs when members of one se! "usually females$ are choosy in selecting their mates from individuals of the other se!. 9ecause females invest more in eggs and parental care, they are choosier about their mates than males. # female tries to select a mate that will confer a fitness advantage on their mutual offspring. In many cases, the female chooses a male based on his showy appearance or behavior. *ome male showiness does not seem to be adaptive e!cept in attracting mates and may put the male at considerable ris3. 8or e!ample, bright plumage may ma3e male birds more visible to predators. Even if these e!travagant features have some costs, individuals that possess them will have enhanced fitness if they help an individual gain a mate. Every time a female chooses a mate based on appearance or behavior, she perpetuates the alleles that caused her to ma3e that choice. *he also allows a male with that particular phenotype to perpetuate his alleles. How do female preferences for certain male characteristics evolveG #re there fitness benefits to showy traitsG *everal researchers are testing the hypothesis that females use male se!ual advertisements to measure the males overall health. ,ales with serious parasitic infections may have dull, disheveled plumage. %hese individuals are unli3ely to win many females. If a female chooses a showy mate, she may be choosing a healthy one, and her benefit is a greater probability of having healthy offspring. Se. is an evolutionar" eni ma. #s a mechanism of rapid population growth, se! is far inferior to ase!ual reproduction. onsider a population in which half the females reproduce only ase!ually and half the females reproduce only se!ually. #ssume that both types of females produce e0ual numbers of offspring each generation.
Lecture Outline for Campbell/Reece Biology, 7th Edition, Pearson Education, Inc. 2 !"7

 %he ase!ual condition will increase in fre0uency, because; #ll offspring of ase!ual females will be reproductive daughters. Only half of the offspring of se!ual females will be daughtersO the other half will necessarily be males. *e! is maintained in the vast maJority of eu3aryotic species, even those that also reproduce ase!ually. *e! must confer some selective advantage to compensate for the costs of diminished reproductive output. Otherwise, migration of ase!ual individuals or mutation permitting ase!ual reproduction would outcompete se!ual individuals and the alleles favoring se!. %he traditional e!planation for the maintenance of se! was that the process of meiosis and fertili/ation generate genetic variation on which natural selection can act. However, the assumption that se! is maintained in spite of its disadvantages because it produces future adaptation in a variable world is difficult to defend. Natural selection acts in the present, favoring individuals here and now that best fit the current, local environment. :et us instead consider how the genetic variation promoted by se! might be advantageous in the short term, on a generation'to'generation timescale. +enetic variability may be important in resistance to disease. <arasites and pathogens recogni/e and infect their hosts by attaching to receptor molecules on the hosts cells. %here should be an advantage to producing offspring that vary in their resistance to different diseases. One offspring may have cellular mar3ers that ma3e it resistant to virus #, while another is resistant to virus 9. %his hypothesis predicts that gene loci that code for receptors to which pathogens attac3 should have many alleles. In humans, there are hundreds of alleles for each of two gene loci that give cell surfaces their molecular fingerprints. #t the same time, parasites evolve very rapidly in their ability to use specific host receptors. However, se! provides a mechanism for changing the distribution of alleles and varying them among offspring. %his coevolution in which host and parasite must evolve 0uic3ly to 3eep up with each other has been called a 17ed Lueen race.2 0atural selection cannot fashion perfect or anisms.
Lecture Outline for Campbell/Reece Biology, 7th Edition, Pearson Education, Inc. 2 !"&

 %here are at least four reasons natural selection cannot produce perfection. 1. Evolution is limited by historical constraints. Evolution does not scrap ancestral features and build new comple! structures or behavior from scratch. Evolution co'opts e!isting features and adapts them to new situations. 8or e!ample, birds might benefit from having wings plus four legs. However, birds descended from reptiles that had only two pairs of limbs. o'opting the forelimbs for flight left only two hind limbs for movement on the ground. 2. #daptations are often compromises. Each organism must do many different things. 9ecause the flippers of a seal must allow it to wal3 on land and also swim efficiently, their design is a compromise between these environments. *imilarly, human limbs are fle!ible and allow versatile movements, but are prone to inJuries, such as sprains, torn ligaments, and dislocations. 9etter structural reinforcement would compromise agility. 3. hance and natural selection interact. hance events affect the subse0uent evolutionary history of populations. 8or e!ample, founders of new populations may not necessarily be the individuals best suited to the new environment, but rather those individuals that were carried there by chance. 4. *election can only edit e!isting variations. Natural selection favors only the fittest variations from those phenotypes that are available. New alleles do not arise on demand. Natural selection wor3s by favoring the best variants available. %he many imperfections of living organisms are evidence for evolution.

Lecture Outline for Campbell/Reece Biology, 7th Edition, Pearson Education, Inc.

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