13.4 Darwi propo: d natural selection ARWIN’S THEORY AND THE MODERN SYNTHESIS mechanism of evolution ee a Darwin devoted much of The Origin of Species to the ways that organisins become adapted to th theory of how this happens arose from several ke tions. First ofall, Darwin recognized that all spec produce excessive numbers of offspring. He had Influential essay on human popalation written in British economist Thomas Malthus. Malthus conte much of human suffering—disease, famine, homelessness, and war—was the inescapable consequence of the humn population's potential to grow much faster than the rate at Which supplies of food and other resources could be pro- duced. 1t was apparent to Darwin that Malthus's concepts applied to all species. Darwin deduced that because natural resources are limited, the production of more individuals han the environment can support leads to a struggle for existence among the individuals of a population, with only a percentage of offspring surviving in each generation. Many, eggs are laid, young born, and seeds spread, but only a tiny fraction complete their development and leave offspring of, their own. The rest are starved, eaten, frazen, diseased, tunmated, or unable to reproduce for other reasons In addition to the overproduction of offspring, two other important observations by Darwin were that individuals of a population vary extensively in thelr charactenstics and that many of the varying truts are inhented—that is, passed, from one generation to the next. What do overproduction of offspring, limited natural resources, and heritable variations have to do with organ- isms becoming adapted to their environment? Darwin saw that every environment has only a limited supply of ir environment. His obsena- tendto 264 UNIT IL Concepts of Eeolution resources and that survival ina limited environmen depends in part on the features the organisms inherit from thuir parents. He concluded that within a varied populg fiom, Qadividuals whose characteristics adapt them best ty theie environment are most likely to survive and reproduce these individuals thus tend to leave more offspring than ey fit individuals do, Reproduction is central to what Darwin saw as the basic mechanism of evolution, the process he falled natural selection. Darwin perceived that the essence of natural selection is differential, or unexpal, suc cess in reproduction. Darwin’ insight was both simple and profound. Repm duction is unequal, with the individuals that best meet specific environmental demands having the grestet reproductive success. Put another way, differential ductive success (natural selection) is the means by which the environment filters variations, favoring some over oth- ers, As Darwin reasoned, natural selection results in favored traits being represented more and more and tnfs. vored ones less and less in ensuing generations. Thus, the unequal ability of individuals to survive and reproduce leads to a gradual change in the characteristics of a populs tion of organisms, with favored characteristics accumu ing over the generations. Darwin found convincing evidence for bis ideas inthe results of artificial selection, the selective breeding of domesticated plants and animals. He saw that by selecting individuals with the desired traits as breeding stock, humans were playing the role of the environment and bringing aboot differential reproduction. They were, in fact, modifying species. We see evidence of what Danwin was talking aboat in the vegetables we cat. For example, broccoli, cauliflower cabbages, Brussels sprouts, and kale (Figure 134A) are al varieties of a single species of wild mustard, and all were produced by artificial selection. These and many other domesticated plants and animals bear little resemblance to the wild species from which they were derived. Figure 13.4B shows some of the enormous diversity that breeders have produced in a few thousand years within a single species, the damestic dog Darwin reasoned that if so much change could be achieved in a relatively short period of time by artiaad selection, then over hundreds or thousands of generations, natural selection should be able to modify species consider ably. With natural selection operating over vast spans of time, heritable changes would gradually accumulate, Such changes would account for the evalution af new species— for exainple, the five species of canines in Figure 13 4C— from an ancestral species. Let's summarze the two main features of Darwin the ony: The diverse forms of life have arisen by descent wi: modification from ancestral species, and the mechanism of modification has been natural selection working over enor mous spans of time. In the nest module, we examine some documented cases of naturil selection. jAncettal dog Figure 1248 Fre breeds of dogs (al members ofthe samme species), the resuits of hundreds to th Ancestral canine Figure 13.4C Five different speces of canines, the results of thousands to millons of years of natural selection Wel/CD Thinking as a Scientist Hote Do Enrironmental Changes Affect a Population? Ditlerental reproductive success among a population's varying individuals in thei natural enwronment a called Chapter 13 How Populations E 265CONNECTION 13.5. Scientists can observe natural selection in action The bive-footed boobies described al the boginning of this, ‘chapter exhibit traits such as webbed feet and sall glands that, are evolutionary adaptations to their ocean-based lle, The ‘exquisite camouflage adaptations shown in Figure 13.5A by insects that evolved in difforent environments ara also exam ples of tho results of natural selection. But do we have exam- ples of natural selection in action? Indeed, over 100 cases of natural selection in nature have bbeon documented. For example, over a period of 20 years, Petor and Rosemary Grant found changes in beak size in a Population of ground finches in the Galdpagos Islands (see ‘Module 14.9) In dry years, when smal seeds aro in short sup- ply, bitds must eat more large seeds. Birds with larger, strongor ‘beaks have a feeding advantage and graater reproductive suc- ‘cess, and the average beak depth for the population increases. During wat years, smaller beaks are more efficient for eating the now abundant small seeds, and the average beak depth decreases. ‘A classic and unsettling example of natural selection in action is the evolution of insecticide resistance in hundreds of ingect species. Insecticides are poisons used to kill insect, pests in farmlands, swamps, backyards, and homes. Whon: ever a new type of insecticide is used to contro agi Pests, the story is similar (Figuie 13.88): A relatively small amount of poison dusted onto a crop may kill 99% of the insects, but subsequent sprayings are less and less etfective. Figure 13.5A Camouflage as an example of evolutionary ‘adaptation, 266 UNIT LIL Concepts of Eeulution ‘The few survivors of the first insecticide wave are insects {genes that make them resistant to the chemical attack Soihy poison kills most members of the population, leaving ihe tant individuals to roproduce and pass the genes jg, ‘cide resistance to their offspring. The proportion of insecticide-resistant individuals increases in each Like the finches, the ingect population has adapted to eng. ronmental change through natural selection. These two examples of evolutionary adaptation threo key points about natural selection. First, notice that na- tral selection is more an editing process than a creative mech: ‘nian. An insecticide does not create resistant individuals, by selects lor resistant insects that were already presert in the population. Second, natural selection is contingent on tire ‘and place: It favors those characteristics in a varying pope tion that fit the current, local environment. For instance, maz. tions that endow houselfies with resistance to the pesticce DDT also reduce their growth rate. Belore DDT was intr: duced, such mutations were a handicap to the fies that had them. But once ODT was part of the environment, the mutart alleles were advantageous, and natural selection increased their frequency in fly populations. Finally, these examples show that significant evolutionary change can occur in a short time, In what sense is natuta selection more of an “edting” process than a “creating” process? Chrcmesome win gene — conferring resistance to insecucida CS Aiton ‘appications of tne ‘same insecticide wil bbe lass effective, and ge ¢ Survivor oe plies pares pores NS ae par Gs ¢ ¢ € € Figure 13.58 Evolution of insecticide resistance in insect populationi re the units of evolution ih era tion Is «group of individuals of living in the same place ut the same time, Itis the sreallea unit that can evolve. We enn in fat, measure evalu ge change in the prevalence of certain heritable tris inn pop ulation over it span of generations, The increasing ing propor. one exumple. Natural selection favored inseets with genes nsects left more offspring, and the population changed, or evolved. Note that the indi Vidual insects did not evolve. It is true that natural selection acts on individuals; their characteristics affect their chances of survival and their reproductive success within a local environment. But the evolutionary impact of natural selee- tion is only for insecticide resistance: the apparent when tracking how a population changes over tine sen aaa Danvin understood that itis populations that evolve. He saw that natural selection, in favoring some heritable traits cover others, changes populations over successive generations, What eluded him was the genetic basis of population change Without this basis, he could not explain the cause of variation among the individuals account for the perpetuation of parents’ trits in their off spring, Today, we know that heritable traits are carried by genes on chromasomes and that mutations may produce new traits. Also, we understand how Mendel principles of segre gation and independent assortiment of alleles operate during meiosis to prodlce genetic variation in gametes and resulting offspring (see Module 9.17). In Darwin’ time, however, litle was known about chroinasomes—and nothing about’ gene mutations or meiosis. Although Mendel and Darwin were contemporaries, the significance of Mendel’ work did not come to light until 20 years after Danwin’s death, some 40 ‘years after the publication of The Origin of Species. An important turning point for evolutionary theory was the birth in the 1920s of population genetics—the science of genetic change in populations. A theory of evolution that xenctics was developed in the early 1940s, Known as the modern synthesis, this theory focuses om popula tions as the units of evolution and includes mast of Darwin's ideas. Most importantly, it melds population genetics with the theory of natural selection, Central to the inodem synthesis is the relationship berween populations and species. We will have more to say about this in Chapter 14, but for now let’ limit our discus sion to organisms that reproduce sexually. In this context, pecies as a group of populations whose individuals have the potential to interbreed and produce fertile offspring, Each species is distributed over a geo- graphic range, where individuals usually concentrate in localized populations. One population may be isolated from ‘others of the same species. If individuals in the isolated pop- ulation interbreed only rarely with those in other popula- tions, there will be little exchange of genes. Such isolation is common for populations confined to widely separated. islands, unconnected lakes, or mountain ranges separated by includes Figure 13.6 Hunan population centers in North America| lowlands. However, populations are not always isolated, nor do they necessarily have sharp boundaries. One papulat center may blur into another in an intermediate region where members of both populations occur but are less Figure 136 illustrates the tendency for humans to con centrate locally. This is a nighttime satellite photograph showing the lights of population centers in North America, We know that these populations are not realy isolated: peo- ple move around, and there are suburban und nities between cities. Nevertheless, people ure As a result, for humans and other ost likely to choose mates locally species, individuals ii on population center are, on wer age, more closely related to one another than ta members of other populations Understanding what populations are and how they behave sets the stage for understanding how populations and species evolve To analyze populations in an evolution ary context, we must consider the genes in the population, Is to focus on population genetics. Before we begin, you might wish to review Mendelian genetics in Chapter 9 (especially Modules 9.2 and 9.4 Pp] 12 dem evtstonay syinensol he 19403 brovght fogether Darwin's theory of natural selection and theory of inhentance. 267 Chapter 13 How Populations Evolve13.7 Microevolution i change in a population’s gene pool over time Meena eee eee eee eee Oe eee In studying evolution at the po ‘on what is called the gene pool, ina population at tion level, geneticists focus total collection of genes y One time. The gene pool [s the reservoir from which members of the next generation of that pops tion derive their genes; it consists of all alleles (alternative forms of genes) in all the individuals making up a population. For most gene loci, there are two or more alleles in the gene pool, and individuals may be either homazygous ( Ing two identical alleles) or heterozygous (having two diffe ent alleles) for each locus. For example, in an insect population there may be two alleles for a particular enzyme. One allele codes for an enzyme that breaks down a certain Insecticide, and the other codes for an enzyine that does not The relative frequencies of these and other alleles in a gene pool may change over time. In an untreated field, the allele pane may have a higher frequency iy than the allele for the insectdde: But in Belds sprayed with insect destroying enzym' x the allele for the enzyme conferring resistance will increas fn frequency and the other allele will decrease in frequen When the relative frequencies of alleles in a population hunge like this ever a number of generations, evan is ‘ccurring on its smallest scale. Such a change in a gene penis failed microevolution. To understand how microevolution sworks, it helps to begin by examining the genetics ofa Iypothetical population whose gene pool is not changing BY sroouion ns charge inva frequency t___ na, population's gene pool. a _ 13.8 The gene pool of a nonevolving population remains constant over the generations Lets consider an imaginary, nonevelving population of blue- footed boobies with two varieties that differ in foot webbing, (Figure 138A), Lets also imagine that foot webbing is con: trolled by a single gene. We'll assume that the allele for non- webbed fect (W) Is completely dominant to the allele for webbed feet (w) The term dominant (see Module 9.3) inay scem to suggest that over many generations of sexval repro- duction, the W allele will somehow come to “dominate” population, becoming more and more common at te expense of the recessive w allele, In fact, this is not what happens. The shuffling of genes that accompanies sexual reproduction Joes hot alter the genetic makcup of the population. In other words, sexual reproduction alone does not lea to microevolu- tion, No matter how many times alleles are segregated into different gametes by melosis and united in differen binations by fertilization, the frequency of each alle gene pool will remain constant unless acted on by other agents This. principle is known as Hardy-Weinberg equ Librium, named for the two scientists who derived jt independently in 19€8. To test Hardy-Weinberg equilibrium, let's look at two generations of our booby population. Figure I38B shows the geactic situation in the original population, We have atotal of 500 boobies; of these, 820 birds have the geno- have the heterozygous genotype, Wie (nonwebbed feet, sage IW is dominant). The proportions ofthe three poxble am: types (the genotype frequencies) are 0.64 for WWW (= 0.64), 0.04 for ww (2%; = 0.04), and 0.32 for Ww (ge = 022), From the genotype frequencies, we can calculate the fre quency of cach allele in this population. Each booby cares two genes far foot type, so the population has 1,000 genes for this characteristic. To find the number of W alleles, we add the number carried by the WW boobies, 2 x 320 = 640, tothe number carried by the Wi boobies, 160. The total nusaber of W alleles is thus $00. The frequency of the W allele, which we will callp, is 788, or 08, We can calculate the frequency fhe ww allele in a similar way; this frequency, called q, is 02. (The letters p and q are often uscd to represent allele frequencies) type WW (nonwebbed feet), 20 have the genotype tww (webbed feet), and 160 No webbing webbing Figure 13.88 Imaginary blue-footed boobies, with and without foot webbing UNIT III Concepts of Ecolution : 268Figure 13.8 Gone poo! of nest goneration of boobies What happens when the boobies of this parent popula tion form gametes? At the end of meiosis, each gamete has one allele for foot type, either W or w. The frequency of the two alleles in the gametes is the same as its in the parental population, 0.8 for Wand 0.2 forw. Figure 13.8C uses these gamete allele frequencies and the rule of multiplication (see Module 9.7) to ealeulate the fre- quencies of the three possible genotypes in the next genera- tion. The probability of producing u WW individual (by combining two W alleles from the pool of gametes) isp X p = p?, or 08 X 0.8 = 0.64. Thus, the frequency of WW boobies in the next generation would be 0.64, or 64%. Likewise, the frequency of wow individuals would be q? = 0.04, or 4%, For eterazygous individuals, Wie, the genotype ean form in two ways, depending on whether the sperm or egg supplies the dominant allele. In other words, the frequency of Wio would CONNECTION 13.9 The Hardy-Weinberg equation be 2pq = 2 x 08 x 0.2 = 032, oF 32%. Thus, the three pas- sible genotypes have the same frequency in the nest genera- tion as they did in the parent generation. Finally, what about the frequencies of the alleles in this next generation? Since the genotype frequencies are the ‘sume as inthe parent population, the allele frequencies, p and 4, are the sume, too, In fact, we could follow the frequencies through many generations, and the results would continue to, be the same. Thus, the gene pool of this population is in a state of equilibrium—Hardy-Weinbery equilibrium. Now let's write a general formula for calculating the fre- ‘quencies of alleles in a gene pool from the frequencies of genotypes, and vice versa In our iinaginary blue-footed booby population, the frequency ofthe W allele (p) is 0.8, and the frequency of the w allele (q) is 0.2. Note that p +9 = 1; this is always true, since the combined frequencies of all al leles must be 100% of the alleles for that gene in the popula- tion. If there are only two alleles and we know the frequency of one, we can calculate the frequency of the other one: Pi 1=p Notice in Figures 13 8B and 13.8C that the frequencies of all possible genotypes in the populations also add up to 1 (Whats, 0.64 + 0.32 + 0.04 = 1). We can represent this symn- bolically with the Hardy-Weinberg equation: =q and P+ lt 1 Frequency Frequeticy Frequency of WW of Ww of ww WebvCD Thinking asa Scientist How Can Frequency of Alleles ‘Be Calculated? IBY 2 ersaray booty popaton oe sare space shown Figure 138Ais in Hardy Wernberg equilbrium. The frequency ofthe recessive alele for wobbed feat 0.4, What the frequoncy of ndviduals that have nomwebbed foot? (rat am ero pura a ac r0 useful in public health science The Hardy-Weinberg equation has broad application. For instance, public health scientists use it to estimate how many people carry alleles for certain inherited diseasos. Consider the case of phenylketonuria (PKU), which is an inherted inability to break down a certain amino acid, PKU occurs in about one out of 10,000 babies born in the United States ‘and, i untreated, results in severe mental retardation. New- bom babies are now routinely tested for PKU, and symptoms ‘can be prevented by following a strict diet. PKU is due to a recessive allele, $0 the frequency of indi \iduals in the U.S. population bom with PKU corresponds to the q? term in the Hardy-Weinberg equation. Given one PKU ‘occurrence per 10,000 births, g2 = 0.0001. Therefore, the frequency of the recessive allele for PKU in the population, 9, cequala the square root of 0,0001, or 0.01. And the frequency, of the dominant allele, p, equals 1 ~ ¢, or 0.99, The frequency of carriers, heterozygous people who are normal but may pass the PKU allele on to offspring, is 2pq, which equals 2 0.99 x 0.01, or 0.0198. Thus, the equation tells us that about 296 (actually 1.98%) of the U.S. population carries the PKU alee, Estimating the frequency of a harmful allele is essential for any public health program dealing with genetic diseases. EY ich term in iho Hardy Weirborg equation-p?, 2p, oF @?-corresponds tothe lrequency of individuals who have ‘no alloles for the disease PKU? é Chapter 13 How Populations Evolve 26913.10. Five conditions are required for Hardy-Weinberg equilibrium ee Hardy-Weinberg equilibrium (ells us that something oth than the reshuflling processes of sexual reproduction is, required to alter a gene pool—that is, to change allele frequencies in a population from one generation to the next. One way to find out what factors can change a gene pool is to identify the conditions that must be met if genetic equilibrium is to be maintained. For a population to be in Hardy-Weinberg equilibrium, it must satisly five main conditions: 1. The population is very large. 2 The population is slated that Is, there is no migration of individuals or gametes into or out ofthe population 3. Mutations (changes in genes) do not alter the gene pool. 13.11 There are several poten! Deviations from the conditions for Hardy-Weinberg equl- librium can cause changes in gene pools, oF microevolution. The two main causes of microevolution are genetic drift and natural selection, although gene flow and mutation may also, change allele frequencies, Nonrandom mating can affect genotype frequences, but it doesn’t change allele frequen- ies and thus is not a cause of microevolution, Genetic drift is a change in the gene pool of a small population due to chance. Flip a coin ten times, and an out come of seven heads and three tails would seem within rea- son. But fip a coin a thousand times, and a result of 700 heads and 300 tails would make you very suspicious about that coin. The smaller the sample, the greater the chance of deviation from an idealized result—an equal number of heads and tails, in the case of a sample of coin tosses. Let's apply this logic to a population's gene pool. Ifa new genera- tion draws its alleles at random from better the new generation wi the previous generation.*If a population of organisms is sinall, its gene pool may not be accurately represented in the next generation, It is analogous to the erratic outeome from 1 small sunple of coin tosses. Genetic daft is u case of microevolution caused solely by chance, Natural selection is not involved. A population must be infinitely large for genetic drift to be ruled out com- pletely as an aent of microevolution; however, the popula tionis in which genetic drift is most likely to play a major role typically have 100 or fewer individuals, TWo situations that, can shrink populations down to a small size are known as the bottleneck effect and the founder effect. ‘The bottleneck effect is genetic drift resulting from an event that drastically reduces population size, Eveats such. * as earthquakes, floods, or fires may kill large numbers of individuals unselectively, producing a small surviving popu: lation thut is unlikely to have the sume genetic makeup as 270 UNIT 111 Concepts of Evolution 4, Mating is random. 5. All individuals are equal in reproductive success, that ig sural selection does not occur. “These five conditions are rarely met, and thus, we dong realy expecta natural population to be in Hardy-Weinke equilibrium, But Hardy-Weinberg equilibrium gives is 3 esis for comparing idealized, nonevolving populations yi ‘actual ones in which gene pools are changing, Lets look soy at how real populations evolve. rot Hardy-Weinberg equilbrium describes a population that g Been 1 causes of microevolution the original population. ‘The analogy of shaking Justa few marbles through a bottleneck illustrates how a bottleneck. ing event works (Figure 13.114). Certain alleles (blue rar- bles) may be present at higher frequency in the sur population than in the orginal population, others (white marbles) may be present at lower frequency, and some (gd marbles) inay not be present at all. In areal example, hurnan hunters in the 1890s reduced the population of northern elephant seals in California to about 20 individuals, Since then, this mammal (pictured in Figure 19.L1B) has become a protected species, and the populition has grown back to aver 30,000 members. However, in eam ining 24 gene loci in a representative sample of the seal researchers found no variation. For each of the 24 genes they found only one allele, probably because of bottlenecking In contrast, genetic variation abounds in populations ofa clasly related species, the southern elephant seal, which wat tot bottlenecked. Orginal population Figure 13.11A The bottleneck effectA second sit ean pro- hoc popallon Sal Cg SP genetic drift Is the colonization of a new location by a individuals. The smaller the sample size, the less the genetic makeup of the colonists will represent the gene pod! ofthe langer population they left In the mast extreme case, a single pant animal or a single plant seed found a new population. Ifthe colony is successful, random changes in allele frequencies will contin until the population is lange enough for genetic drift to be minimal. Such genetic dnft in a small colony is called the founder effect. The founder effect undoubtedly contnbuted to the evolutionary divergence of the finches and other South American organisms that arnved as strays on the Salipagos Islands, The founder effect explains the relatively high frequency of certain inherited disorders among some human popula- tions established by small numbers of colonists. In 1514, 15, people founded a Bntish colony on Tristan da Cunha, a group of smal islands in the middle of the Atlantic Ocean, (Figure 13.11C). Apparently, one of the colonists carried a recessive allele for retinitis pigmentosa, a progressive form of blindness. OF the 240 descendants who stil lived on the {slands inthe 1960s, 4 had retinitis pigmentosa, and at least, 9 others were know to be heterozygous carers of the allele The frequency of this allele is much higher in this popula. tion than in the population from which the founders came xy be an agent of microevolution Gene fertile individuals mene into or out of a population or when gametes (such as the sperm of plant remote os Sa Figure 13.11¢ Residents of Tristan da Cunha in the early 19008, Figure 13.118 Elephant seals descended {rem bolleneck survivors pollen) are transferred between populations. Gene flow tends to reduce genetic differences between populations. Over the history of our own species, for example, the isola- tion of local groups has reduced gene flow, resulta genetic distinctions among groups of people living in differ- ent parts of the world. Reflecting these genetic differences may be phenotypic variations in skin color and facial charac- teristics. Working against reproductive isolation has been the influence of migrations and wars, which tend to increas interbreeding among groups. Today, itis possibl all over the globe to interact, and there is mor among geographically isolated populations than ever before Mutation may also be an agent of microevolution. A mutation is a random change in an organisms DNA that may ereate a new allele (see Module 10.16). Mutations of a given gene are rare events, typically occurring only about once per gene locus per 10% or 10° gametes. Asa result, in a large population, mutation alone does not have much effect in ration. Over the long term, however, muta~ tion is vital to evolution because it is the only force that actu- ally generates new alleles. ‘Thus, mutation is the ultimate source of the genetic variation that serves as ra evolution Natura selection, or differential success in reproduction, is the only cause of microevolution that is likely to result in adaptive changes in a gene pool, Let's consider this next, material for Web/CD Activity 19D Causes of Microevolution EY = fr causes of mcroovliton. Which two wv he ‘most important factors that alter allele frequencies in a population? senescent ate sus jul 2m ope pin eg ng wore fms poe nts my we Chapter 13 How Populations Evolve 27113.12 Adaptive change results when natural selection upsets genetic equilibrium One condition for Hardy-Weinberg equilibrium—that all individuals in a population be equal in their ability to repro- duce—is probably never met in nature, Populations of sexi ally reproducing organisms consist of varied individuals, and some variants leave more offspring than others. In our imag- inary blue-footed booby population, birds with webbed feet (genotype ww) might produce more offspring because they are more efficient at finding food than birds without webbed feet (genotype Ww or IVW). Genetic equilibrium would be disturbed as the frequency of the w allele increased in the gene pool, In this way, natural selection results in the accu- mulation and maintenance of traits that adapt a population to its environment. If the environment should change, nat selection would favor traits adapted to the new eng tions. The degree of adaptation that can occur is limited by the amount and kind of genetic variation in the population, Fa & polation of ecto apa art act arc Sayemeenltioay Gemeente Sirin te poplin gore ello ae 13.13 Variation is extensive in most populations We have no trouble recognizing our friends in a crowd. We are very conscious of human diversity, but individuality in populations of other animals and plants may escape our notice. Nonetheless, individual variation occurs in popula- tions of all species that reproduce sexually In addition to ‘anatomical differences, most populations have a great deal of variation that can only be observed at the molecular level. Not all variation in a population is heritable. The pheno- type results from a combination of the genotype, which is inherited, and many environmental influences, For instance, fa strength-training program can build up your muscle mass, but you would not pass this environmentally procuced physique on to your offspring. Only the genetic component of variation is relevant to natural selection. Many of the variable traits of the individuals in « popula: tion result from the combined effect of several genes, As we saw in Module 9.16, polygenic inheritance produces traits that vary more or less continuously—in human height, for instance, from very short individuals to very tall ones. By contrast, other fea tures, “such — as human ABO blood groups (see Module 9.13), are deter. mined by a single gene locus, with dif: ferent alleles pro- ducing only distinct phenotypes. In such cases, when a popu- lation includes: two ‘or more forms of 4 phenotypic charac. teristic, the differ- cent forms are called morphs, A popula- Figure 13.13 Polymorphism in a ‘population of king snakes 272 UNIT LIL Concepts of Evolution tion is said to be polymorphic for a characteristic if two or more morphs are present in noticeable numbers. Polymor phism is extensive in human populations, both in physical charicteristics, such as the presence or absence of freckles and in biochemical features, such as the ABO blood groups Figure 13.13 illustrates a striking example of polymorphism within a population of California king snakes, The two morphs differ markedly in their color patterns. In addition to variation within populations, most species cethibit geographic variation between populations, Sometimes this variation occurs in what i called a eline, a graded change in an inherited characteristic along a geographic continuum, For example, the body size of many birds and mamunal tends, to increase with increasing latitude in North America. Large size is adaptive in colder latitudes because it reduces the ratio of body surface area to volume and helps conserve body heat How is genetic variation measured? Population genetics look at both gene diversity and nucleotide diversity: Cene diversity is the average percent of gene loci that are hetero Zygous In a population. Nucleotide diversity is determined by comparing the nucleotide sequences of DNA samples Humans have less genetic variation than mast other species Gene diversity is 14%; that is, we are heterazygous at aboot 14% of our gene loci, Our nucleotide diversity is only about 0.1%, So while you and yourneighbor have the same nucle — {ide at 999 out of every 1,000 nucleotide sites in your DNA, there is still enough variation to account for the genetic com — Ponent of the enormous individuality we observe in people How do inbented vuriations arise? We address this que- tion in the next module, 4CONNECTION 13.14 Mutation and sexual recombination generate variation CUAL LeCOmUINA LON gene rate,variat On ‘As we saw in Module 10.16, mutations, or changes in the flucleotide sequence of DNA, can create new allolos. A muta- tion that substitutes one nucleotide for another will be harm- less it it does not affect the function of the protein the DNA encodes. However, if it does affect the protein's function, the mutation will probably be harmful. An organism is a refined product of thousands of generations of past selection, and a random change in ts DNA is not likely to improve its genome ‘any more than shooting a bullet through the hood of a car is likely to improve engine performance. (On rare occasions, however, a mutant allela may actually improve the adaptation of its bearer to the environment and enhance reproductive success. This kind of elfect is more likely when the environment is changing in such a way that mutations that were once disadvantageous are favorable Under the new conditions. The evolution of DDTesistant houseflies (see Module 13.6) illustrates this point. In another ‘example, mutations that make the HIV virus resistant to antiv- ral drugs also slow the reproductive rate of the virus (see Module 10.21). Howaver, once antiviral drugs are used to treat HIV-infected individuals, these mutant alleles are advan- tageous because they allow the virus to survive, and natural selection increases their frequency in the HIV population, ‘On a larger scale are so-called chromosomal mutations, which are changes involving stretches of DNA long enough to be detected microscopically (see Figures 8.23A and 8.238), A single chromosomal mutation affects many genes and is, almost certain to be harmful. Very rarely a duplication of a chromosome segment might bring benefits. If the repeated segment can persist over the generations, it may provide a & i Parents MEIOSIS i) 19 ye FERTILIZATION Ottspring. with now combinations of alleles Figure 13.14 Shuffing alleles by sosual recombination, bigger genome with extra genes that may eventually take on new functions by mutation In microorganisms with very short generation spans, muts- tion genorates genetic variation very rapidly, For example, HIV has a generation span of about 2 days. In an AIDS patient, the HIV infection produces 101° or more new viruses per day. Each replication provides a chance for mutations to occur In addition, HIV has an RNA genome, which has a much higher mutation rate than DNA genomes. Because of these high replication and mutation rates, single-drug treatments will probably never be effective for long against HIV. Even double- drug treatments do not remain effective, because individual, Viruses with double mutations conferring resistance to both rugs arise daily. This explains why the most effective treat- ments for HIV infection are drug “cocktails? combinations of more than two drugs. Bacteria also multiply s0 rapidly thal a beneficial mutation can increase its frequency in descendant populations in a matter of hours or days. Because bacteria, like viruses, are generally haploid, with only a single gene for each inherited tat, a newly created allele can have an eect immediately. ts ‘expression cannot be obscured by another alele for the same trait. For most animals and plants, however, their long genera- tion times and generally diploid condition prevent most muta- tions from significantly affecting genetic variation from one {generation to the next. Consequently, animals and plants depend mainly on sexual recombination for the genetic varia- tion that makes adaptation possible, As we saw in Modules 8.14 and 8.15, fresh assortments of existing alleles arise ‘every generation from three random components of sexual recombination: crossing aver, independent assortment of homologous chromosomes, and random fertilization. During ‘meiosis, homologous chromosomes, one inherited from each parent, trade some of their genes by crossing over, and then the homologous chromosomes separate independently into ‘gametes, Gametes from one individual vary extensively in their {Genetic makeup, and each zygote made by a mating pair has {8 unique assortment of alleles resulting from the random union of a sperm and an ovum. In Figure 13.14, we see the results of a mating of parents with the genotypes A’A’ and A2A3, whore A', A2, and A? are three different alleles for a ‘gene. Even without considering independent assortment or ‘crossing over, the offspring have different combinations of alleles than wore present in their parents. Web/CD Activity 13E Genetic Variation from Sesual Recombination 9 la the ulimate source of genetic variation in a £4 population, but in a serual population witha relatively long, ‘generation span, most ofthe variation we observe is due to omens runt Lome Chapter 13 How Populations Evclve 27313,15 Overview: How natural selection affects variation Natural selection acting on the variations within a pop tion adapts onganisis to their environment. But what pre vents natural selection from eliminating this variation as it selects against unfavorable genotypes? Why aren’ the less adaptive alloles eliminated as the best alleles are passed on to the nest generation? The tendency for natural selection to reduce variation in a population is countered by mecha- nisms that maintain variation, Most eukaryotes are diploid, and having two sets of chro- mosomes helps to prevent populations from becom genetically uniform, The effects of recessive alleles are not often displayed in diploid organisms. A recessive allele is subject to natural selection only wien it influences the phe- notype, and this occurs only when two copies of it appear in a homozygous individual, In a heterozygote, a recessive allele fs, in effect, hidden, or protected, from natural selec- tion. This hiding of recessive alleles in the presence of dom: Inant ones can allow a large number of recessive alleles to remain in a gene pool, and these alleles may prove advanta- ‘geous in later generations, Individuals with a trut resulting from two copies ofa recessive allele might be eliminated by natural selection in one environment, but if the environ- ment changes, such individuals might have greater repro- ductive success Genetic variability in diploid organisms can also be pre- served by natural selection, the very force that generally reduces it. The ability of natural selection to maintain stable frequencies of two or more phenotypic forms in a popula- tion is called balanced polymorphism, Sometimes there is a heterozygote advantage—thi is, heterozygous indhidy. als have greater reproductive success than homozygotes iq which case two or more alleles for a trait will be maintained ral selection. One such example is the resistance ip conferred by the recessive sickle-cell allele (see Module 9.14), In areas where malaria is a major cause of death, natural selection favors heterozygotes (carriers ofthe si¢kle-cell allele) because they are resistant to malaria ‘second mechanism promoting balanced polymorphism is frequency-dependent selection, in which the survival and reproduction of any one morph declines if that phenotype form becomes too common in the population. Perhaps a species of butterfly has several different coloration patterns. Birds may more easily locate and feed on any morph that becomes too common. The frequency of the other colar pat- tens would then increase. In other cases, a patchy environ. ment may favor different phenotypes in different areas. For example, protective coloration suited to different back. grounds may help explain the morphs of the king snakes shown in Figure 13.13. BBY 7 wou natural soection tnd to recice genetic ‘aration more in populations ot haploid organisms than in populatona of diploid organisms? ere prem fe pam rx om put pears genic tt aan pace em 13.16 Not all genetic variation may be subject to natural selection Some genetic variations in populations seem to have a trivial impact on reproductive success and therefore may not be subject to natural selection, The diversity of human finger- prints, for example, seems to be an eample of neutral varlation—variation in a heritable characteristic that pro- vides no apparent selective advantage for some individuals over others. Some of these supposedly neutral alleles will Increase their frequency in the gene pool and others will decrease by the chance effects of genetic drift, but natural selection will not affect them. Figure 13.16 Human firgerprns, probably an example of neutral variation 274 UNIT LIT Concepts of Evolution There is no consensus among evolutionary biologists about how much genetic vanation is neutral, Some researchers do not believe that any variation is truly neutral, They point out that variations appearing to be neutral may | inlluence reproductive success in ways that are difficult to | measure. We may be able to show that a particular allele is harmful or beneficial, but we cannot demonstrate thal a allele has no effect at all on an organism. Also, a ‘ay be neutral in one environment but not in another. We) can never know the degree to which genetic variation is tral, But we'can be certain that even if only a fraction off ‘extensive variation in a gene pool significantly affects ‘Oongunisi, that is still an enormous resource of raw mater for natural selection and the adaptive evolution it PA 7 iso t possible to detormire with certainty Sa 10019 ae a abesmromen 0 sDeimge maceCONNECTION 13.17 Endangered species often have reduced variation The topic of genetic vanation has direct bearing on one of our most pressing problems, the unprecedented rate of the worldwide loss of species. Largoly because human activities have reduced their ling space, many species are in danger ‘of becoming extinct. Without significant efforts to curb habitat destruction, the rate will climb even more, Endangered species typically havo low genetic variability, ‘As their populations are severely reduced, the diversity of their gene pools also deciines. One such species is the chee- tah (Acinonyx jubatus), pictured in Figure 13.17. The fast of all running animals, the cheetah can run down the swaftest ‘antelope. These magnificent cals were formerly widespread "In Arica and Asia. Their numbers fell drastically during the last "ce age some 10,000 years ago. At that time, the species may _ have suffered a severe bottleneck, possibly as a result of dis- "aso, human hunting, and periodic droughts. Today, only three ‘srall cheetah populations exist in the wild, one in East AMrica, ‘ene in South Africa, and a third in northern tran (which may ‘number less than 50). ‘Studios of tho African cheetah populations show very low = genetic variation. Today, only about 0.04% of the South "Arican population's gene loci are heterozygous. This repre- ‘ents extreme genetic uniformity—far greater than average for "mammals and even greater than for some highly inbred vari ‘ties of laboratory mice. Some researchers think that the ‘South African cheetah population suffered a second bottle- ‘Neck during the nineteenth century, when South AVrican farm: ‘ers hunted the animals to near extinction. Figure 13.17 The cheetah, a species with low genete vanabiity This lack of variability, coupled with an increasing loss of habitat, makes the cheetah’s future precarious. The cheetahs remaining in Africa are being crowded into nature preserves and parks as human demands on the land increase. Along wih crowding comes greater opportunity for predation by other large cats, such as lions, and increased potential for the spread of disease. With s0 little variabilly, the cheetah may have a reduced capacity to adapt to such environmental chal- lenges. Captive breeding programs are already under way ard may be required for the cheatah’s long-lorm survival Ep) wiry might new strains of pathogens pose a greater threat to cheetah populations than to mammalian populations having more genetic variation? “pemen au gen ou pape ae aoe ms Bs promed« ayy none Sonn on on ne Gens em 13.18 The perpetuation of genes defines evolutionary fitness phrases “struggle for existence” and “survival of the are misleading if we take them to mean direct compet- ‘contests between individuals, There are animal species Individuals lock horns or otherwise do combat to deter- mating privilege. But reprdluctive success is generally subile and passive. In a varying population of moths, Individuals may average more oflspring than others their wing colors hide them from predators better. na wildflower population may differ in reproductive because some are better able to attract pollinators, to slight variations in flower colar, shape, or fragrance. A may produce more egys than her neighbors because she is er ut catching insects for food, These examples point to a definition of fitness. Darwinian fitness is the con- an individual makes to the gene poo! of the next gen- relative to the contributions of other individuals. Thus, individuals in the context of evolution are those that the greatest number of genes to the next generation. alone does not guarantee reproductive success be biggest, fastest, toughest frog in the pond has a fitness ifit is sterile, Production of fertile offspring is the Evolutionary fitness has to do with genes, but its the phe- notype of an organism—its physical traits, metabolism, and behavior—that is directly exposed to the environment. Act- ing on phenotypes, sclection indirectly adapts a population to its environment by increasing or maintaining favorable geno- types in the gene pool. The fitness of any one allele, however, depends on the entire genetic context in which it works, For example, alleles that enhance the growth af the trunk of a tree may be useless or even detrimental in the absence of alleles at other loci that enhance the growth of roots required ta support the tree. On the other hand, alleles that con- trbute nothing to an organism's success or may even be inal- adaptive may be perpetuated because they are present in individuals whose overall finess is high. The whole basehall team wins the World Series, even the player with the worst batting average and the most errors. ‘What determines an organism's Darwinian fitness? Chapter 13 How Populations Evohe13.19 There are three general outcomes of natural selection Let's sve how the culling effects of natural selection can affect the frequencies of phenotypic variants using an imaginary deer mouse population. The bell-shaped curve in the top graph of Figure 13.19 depicts the frequencies of Individuals that could result from a polygenic inheritance pattern for variation in fur color. In this starting popula- ton, fur color varies along a continuum from very light only @ few individuals) through various intermediate shades (many individuals) to very dark (few individuals). ‘The other three graphs show three different ways in which natural selection could alter the phenotypic vuriation in the idealized population, The large downward arrows sym- bolize the pressure of natural selection working against certain phenotypes. Stabilizing selection favors intermediate variants. It typically occurs in relatively stable environments, where ‘conditions tend to reduce phenotypic variation. In the ‘mouse population depicted in the graph on the bottom left, stabilizing selection has eliminated the extremely light and ark individuals, and the population has a greater number of intermediate phenotypes, which are best suited to a stable ‘environment. Stabilizing selection probably prevails most of the time in most populations. For example, this type of selection keeps the majority of human birth weights in the range of 3-4 kg (6.5-9 Ib). For babies much smaller or larger than this size, infant mortality is greater. Frequency of individuals = Phenotypes (tur color) ‘Stabilizing selection Figure 13.19 Tivee possibe eects of natural selection on a phenotypic charactor 276 UNIT 111 Concepts of Esolution Directional selection shifts the overall makeup of the population by acting against individuals at one of the phe- nolypie extremes. For the mouse population in the bot. tom center graph, the trend is toward darker fur color, as night occur if the landscape has become shaded by the growth of trees. Directional selection is most common uring periods of environmental change or when.mem- bers of a species migrate to some new habitat with difer- ‘ent environmental conditions. The changes we described in populations of insects exposed to insecticides and HIV exposed to antiviral drugs are examples of directional selection. Diversifying selection typically occurs when environ- ‘mental conditions are varied in 1 way that favors individuals at both extremes of a phenotypic range. For the mice in the graph on the bottom right, individuals with light and dark fur have increased their numbers relative to intermediate variants. Perhaps the mice had recently colonized a patchy habitat where a background of light soil was studded with dark rocks. Diversifying selection can lead to balanced poly- ‘morphisin with two or more contrasting morphs, such as in the population of king snakes in Figure 13.13. Nest we consider a special case of selection, one that leads to phenatypic dif ferences between males and females. BBY] 0 the three modes of natura selection, “which ia most common? noon a13.20 s. The nimal species obviously have different reproductive organs. But they may also have other marked differences, not directly associated with reprodue- tion, called secondary sexual characteristics, This distinction in appearance is called sexual dimorphism. It is often manifested in a size difference, but can also be evident in the form of male adornment, such as colorful plumage in birds, manes on lions, or antlers on deer, Males are the showier sex, at least among vertebrates. Danwin considered sexual selection, the determining of ‘who mates with whom, to be a separate selection process that produces sexual dimorphism, In some species, sec- ondary sex structures may be used to compete with mem- bers of the same sex for mates (Figure 13.20A). Contests may involve physical combat, but are more often ritualized displays (see Chapter 37). This so-called intrasexval sele tion (within the same sex) is common in species where the ‘winning male gamers a harem of females. In.a more common type of sexual selection, called inter- ‘sexual selection or mate choice, individuals of one sex (usually females) are choosy in selecting their mates. Apparently males with the most impressive features are the most attrac- tive to females, A peacock strutting in front of hens with his tail feathers spread is an example of this “choose me” state- ment (Figure 13.20B), What intrigued Darwin about such behavior is that some of these mate-attracting features do not seem to be otherwise adaptive and may in fact pose some risks. For example, showy plumage may make male birds more visible to predators, But if such secondary sexual char- acteristics help a male gain a mate, then they will be rein- forced over the generations for the most Darwinian of ual selecti es and females of an may produce sexual dimorphism Figure 13.208 A contest for access to mates reasons—because they enhance reproductive Every time a female chooses a mate based on a certain appearance orbehavior, she perpetuates the alleles that caused her to make that choice and allows a male with a particular phenotype to perpetuate his alleles. E59 Mes wih bo mast elaborate omamentation may gamer the most mates. How might such a mate choice be advantageous to a female? ariye vn ot pepe a prose img min po sympa me pod ae Ss ep we 13.21 Natural selection cannot fashion perfect organisms There are atleast four reasons why natural selection cannot produce perfection: L. Organisms are locked into historical constraints. Each species has a legacy of descent with modification from a Jong line of ancestral forms, Evolution does not scrap ‘ancestral unatomy and build each new complex structure from scratch, but co-opts existing structures and adapts them to new situations, The wings of birds are fushioned from bones that supported the walking legs of their ancestors. Adaptations are often compromises. Each organism must do many different things. A blue-footed booby uses its webbed feet to swim as It dives into the ocean for prey, but these same feet make for clumsy travel on lan. 3. Not all evolution is adaptice, Chance probably affects the genetic structure of populations to a greater extent than was once believed. For instance, when a storm blows insects hundreds of miles over an ocean to un island, the wind does not necessarily transport the speci- Pr mens that are best suited to the new environment And not all alleles fixed by genetic drift in the gene pool of a small population are better suited to the environment than alleles that are lost 4, Selection can only edit existing variations. Natural selec~ tion favors only the fittest yariations from the pheno- types that are available, which may not be the ideal traits. New alleles do not arise on demand, With ull these constraints, we cannot expect evolution to ceraft perfect organisms. Natural selection operates on a “better than” basis We can see evidence for evolution in the subtle imperfections of the organisms it produces. Humans owe much oftheir versatility and athleticism to their lexble imbs and joints, But we are prone to spraine, tor ligaments, and dislocations. Why aren't our bodies erect? * saogenj suey reas mmaasaane oe ee Chapter 13 How Populations Evalve 27 Figure 13.208 A male peacock’s adverisement lor matesCONNECTION 13.22, The evolution of antibiotic resistance in bacteria health concern a serious publig Antibiotics aro drugs that disable or kil infectious microorgar ‘ems. Most antibiotics are naturally occurring chemicals orived from other microorganisms, Panicilin, for oxamplo, ‘was originally isolated from a mold and has been widoly pro- scribed sinca the 1940s. A revolution in human health rapidly followed its introduction, rendering many proviously fatal di ‘eases easily curable (such as sirop throat and surgical infoc: tions). During the 1950s, some doctors evon predicted the end cf human infectious disease. Why hasn't this optimistic prediction come tuo? Bocauso it {did not take into account the force of evolution. In the same way that pesticides select for resistant insects (soa Modula 13.5), antibiotics select for resistant bacteria. The genes thal confor ‘Such antibiote resistance ara often carried on R plasmids (see Module 12.2), which are passod on to bacterial offspring and may even be transferred to othor bacteria. For nealy every — ‘antibiotic that has boon developed, a <2 resistant strain of bactoria has appeared within afew decades. For example, somo strains of the tuberculosis-causing bac- terium (Figure 13.22) aro now resistant to all threo of the antibiotics commonly Used 10 treat the disease. Tn what ways do we contribute to the problem of antibiotic rosistance? Lve- stock producers add antibiotics to ani- mal foed as a growth promoter, As a gure 1322 result, much of the packaged meat for yoobacterium tuberculosis sale in supermarkets contains bactena Chapter Review that aro resistant to standard antibiotics. Doctors con to tho problom by overprescnbing antibiotics—for example patients with wral infections, which do not respond to ani otic troaiment. And patients contribute to this through the misuse of prescribed aniboics~for ramp promaturely stopping the medication because they fel bese? Tis alow mutant actora that may be kiled mor howl tho drug to survive and multiply. Subsequent such bacteria may lead to full-blown antibiotic During the anthrax crisie of 2001, pubs health offal peg parched czoa to avoid unoacassa taking the drug used to treat the deadliest form of anthrax ain do ao erste eetan nec a Diffcullyin treating Common human infections i a seraug public health concern. Peneiin was effective agama may ‘all bacterial infections in the 1940s but is virtually useey today in its eriginal form. Increasingly powertul drag hag sinca boon developed, but they continue to be rendered gg fective a3 resistant bacteria evolve. The medical ‘and pharmaceutical companies are engaged in an ong race against the powerful force of bacterial evolution, WelvCD Thinking aya Scientist Connection: What Are the Pte of Antibiotic Resistance? PY e102 wy ho fotowng stterartsnconect havo croated resistant bactena” . ‘mmc paeg peed pee pee 1 comme 21 tee Lrg moma om oman pe ————— CHAPTER SUMMARY Evidence of Evolution (Introduction-13.9) __Allongaisms hs evoltionary adaptations inherited characteristics that -guhanee their ability to survive and reproduce in a partculur etn troniment {ntroduction), Aristotle ad the Judeo-Christian culture held that species are fined Fossils spgrstel that Me forms change, a concept ‘ermbrace! by J, B.Lavnarek i the early 1500s. While on the vwyage of “MS gle in ths 153, Charles Darwin ober sinatieg between vdng and fossil organisms and the diversity of life on the Galipagea Islands, Darwin breamne gousiace that Earth was ol aa entinally changing, He concluded that living things abo change, or evolve, ver ‘generations und thal lining species arv descended (rom caricr life-forms: this concept he called descent with modification (1: Fensils (the rev. ‘ants of extinct organisms) und the fs recor (the order in which fa “sly appear Io the strata of sedimentary rocks) strongly supp the theory ‘of evolution. The fesl record shows that rganiams have appeared in a starial sequence, and many fsa link carly extinct species with species living today (13.2). Other evidence fur evolution comes froin bio- geography, companitive anatomy, comparative embryology, and molecular biology: Conclusions baie on comparisons of DNA and proteins gener- ally agree with those hased on fosils ere] anatainy (13.3), 278 UNIT IIT Concepts of Evolution Darwin Theory and the Modern Synthesis (134-Ek13) Darain obuerwel tha organisms proaluce move offpeing than the eae hat gains vary, and that thet arian ea be uals best sit fra peter environment are more likely to surive an repeveluce than tae areal faapeed. Darwin aw natural selection, the essere of which i ila (unesqual) repretactive succes, 48 the Bask ronment ean supp Inherited, Me copctuded that the result of natural ylection, the propution uf characteitis increases, and populations gr the envtmament, Darwin abo sw that whew with denne characteristics an breeling sch tary are taking the pled the emvuinent,bnong about leet prec tom al me species (134), Evolutionary adaptations prelaced by natural een lave been olserved in poulatons of bir, leet, end may okey Oonganisns (13.5) ‘The madera snes coments Darwin theory with poplin r0etic Aspcies i 2 grup of populations whee Ilion fa ie ‘bee aad prose fet fspring (13.6). Agee peal the ttl ede the of ges ina population at any ane tine, Mcroeodution ace | tthe relative frequencies of alleles in a gene pool (13.7) As prea: by Hanh Weinberg cquiibrium, the shulfing of genes ding seal repmaluction doesnot alter the proportions af diferent alee 4