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UNIT sj Evolution by Natural Selection The rise of Whats the evidence for evolion? )_(Cyolton n acon (Comments about natural salactan and evalu ‘is chapter is about one of the groat ideas in selonco: the theory of evolution by natural selectio populations—individuals of the same species that live in the same area at the same time—have come to the five key attributes of life: Populations of organisms evolve (Ch. 1, Section 1.1). In other words, the her table characteristics of populations change overtime. Evolution by natural selection is one ofthe best supported and most important theories inthe history of scientific research. But like most scientific breakthroug! seTharein and Ared Rineesl Wallace. Tha thorny empl ree dd to environments ranging from arcti tundra to tropical wet forest. It revealed one of ts, this one did not come easily. When Darwin published his theory in 1859 in a book called On the Origin of Specico by Meara af Natural Selection, it unleashed fireotorm of protest throughout Europe. In everyday English, the word “theory” suggests a thought- fl guess, hut a scientific theary is an explanation for» broad lass af ahsorvations that i widoly supparted hy averwhelming evidence. (For help with problematic words, see BioSkills 17.) Scientific theories usually have two components: a pattern and a process. 1. The pattorn eompanent is a statement that summarizes a se- ries of observations about the natural world. The pattern component. about facts~about how things arein nature. 2. The nrocess component isa mechanism that produces that nat tern orsetof observations "This chaptor bopine by givingyou an overview of the dovelop- Hanary thenight ‘Then you'll examine the pattern and pmcess components of the theory of evolution hy natural selection. This chapter also clarifies several common misconcep- ‘tions about evolution. For example, did you know there are other processes of evolution besides natural selection? mont of eve 22.1 The Rise of Evolutionary Thought People often describe the theory of evolution by natural selec- ‘ton as revolutionary. A scientific revolution overturns an exist- ng dea about tow nature works and replaces I wit radically different idea. Revolutionary sclontifle theories include Copernicus’s theory of the Sun a8 the center of our solar system, Newton's laws of motion and theory of "afer vou complete this section, you should be able to. 1 Contrast typological thinking, Lamarekn evolution, ane Darwinian oreo (0) Plato: (0) Aristotie: Typologice! ‘Typologicel thinking ‘thinking “+ scale of nature h Change through time + scale of nature Higher srovitation, the germ theory of discase, the theory of plete tee tonies, and Rinstain’ ganoral thoory of ralativity, Thoce thaoriog are the foundation of modern seience. ‘The advance ofthe theory of evolution by natural section rep- resented a profound scientific revolution. The idea that Darwin and Wallace overturned-—that species were supernaturally, not naturally, created—had dominated thinking about the nature of| organisms in Western civilization for over 2000 years. ‘Tohelp you understand the conceptual change that occurred, Jet's walk through a time sequence of visual models ofthe diver- sity af life, summarized in Figura 294 Plato and Typological Thinking ‘The Greek philosopher Plato, who lived over 2400 years ago in ancient Greece, claimed that every organisin was an example ofa perfect cosence, or type, arcated by Cod, and thatthooe typea were tinchanging (igure 22 1a) Plato acknowledged that individiale of a species sometimes varied slightly from one another. but that these were just trivial deviations around a “perfect essence.” ‘Today, philosophers and biologists refer to ideas lke thisasty- pologial thinking, Typological thinking also occursin the Bible's ook of Genesis and In the ereation stories of many other reli lon, where a dvine being creates eae type of organist. Aristotle and the Scale of Nature One of Plato's students, Aristotle, organized typological thinking {nto a linear scheme called the great chain of being, or scale of, nature, where "seale” means a ladder or stairway (Figure 22. 1b). Aristolle proposed Unat species were fxed types ongantized Into f sequence based on Increased size and complexity. The scale started with minerals and lower plants at the bottom, then rose through higher plants, lower and higher invertebrates (animals without backbones), and lower and higher vertebrates (animals with backbones) to humans atthe top, surpassed only byangels and God. (€) Lamarck: {(@) Derwin and Wallace: ‘Change through time ++.common ancestry Figure 22.1 Modelo ofthe Diversity of Life Have Changed through Time. Vicual modele arc helpful for ‘comparing ideas. Tha mesial shown hrm inci enly fu ving species, for simplicity Fach movil ‘explained in the text (se also BioSkils 12), CHAPTER 22 Evolution by Natural Selection 449) Aristotle's dens remained popular in sefentifie and religious iro wo inta the 1700, Singa many cultures had ambraced Aristotle's model for so long—over 20 centuries—the notion of “lower” and “higher” species, including the separation of hu- ‘mans from other animals lingers as a cultural habit even today (discussed in Section 22.5). Lamarck and the Idea of Evolution as Change through Time ‘Typologieal thinking eventually began to break down. In 1809) the biologist Jean-Baptiste de Lamarck proposed the first fur- ‘mal theory of evolution—that species are not static but change through time. However, the pattern component of Lamarck’s {theory was initially based on the scale of nature. ‘When he started nis Work on evolution, Lamarck claimed that siuple vganisiis vigiale atthe base uf Use stale by spuutanee ‘ous generation (Ch. Section 1.2) and then evolve by moving up the seale over time (Figura 29.16), Thus, Lamarekian evslistion is progressive in the sense of always producing larger and more complex, or “better” species. Lamarck also contended that species change through time vila the inheritance of acquired characters. The idea here is that, as an individual develops, its phenotype changes in response ‘w challenges posed by the environment, and It passes on these phenotypic changes to offspring. classic Lamorckian scenario is that giraffes develop long necks as they stretch to reach leaves hhigh in treetops, and they then produce offspring with elon- gated necks. Darwin and Wallace and Evolution by Natural Selection AAs his thinking matured, Lamarck eventually ubandoned his, linear and progressive view of life, Darwin and Wallace agreed. [But more important, they emphasized that the process respan- sible for change through time—natural selection—occurs be- cause traits vary among the individuals in a population, and because individuals with certain traits leave more offspring than others do. Darwin claimed that instead of being unimportant, variation among individuals in a population was the key to understanding, the nature of species. Biologists roferto thie view as population think’ ‘The theory of evolution by natural selection was revolution- ary for several reasons: ‘+ Ttoverturned the idea that species are static and unchanging. Instead, it suggested that species change through time and are related by common ancestry (Figure 22.14) ‘ Itreplaced tvpological thinkine with population thinking. ‘* Itwas seiontifie. It proposed a mochanism that could account for change through time and madle predictions that could he tested through observation and experimentation, Plata and his followers emphasized the existence of fixed ‘vpes. whereas evolution by natural selection is all about change 450 _ UNIT4 Evolutionary Patters and Processes and diversity, With the advent of evolutionary thought, new ‘question aroee: What evidence backs tho claim that specioe are not fixed types? What data support the theary of evalutian by natural selection? [iehecn vous unsensmNeine ‘Ifyou understood this section, you snould be able to Imagine you are standing in a meadow of sunflowers, Compare your view of sunflowers from the perspective of typalngica thinking and frm that of papain thinking 22.2 The Pattern of Evolution: Have Species Changed, and Are They Related? In On the Origin of Species, Darwin repeatedly described evolu- tion as deseent with modifieation, In essence, the pattern component of the theory of evolution by natural selec tion makes two predictions shout the nature af species: ‘ter you complete this section, you should beable to 11 Summarize evidence that supports the theory of 1, Specica change through youton, time. 2. Speciesare related by common ancestry. Let's consider the evidence for each prediction in turn. he ‘diferent types of evidence ate numbered to ielp you keep track ts you read along: 1. § for epecieo change through time, 6 8 for speriee are related hy common ancestry Evidence for Change through Time When Darwin began hio work, blologiots and geologits had Just begun to assemble and interpret the fossil record. A fossil is anv trae ofan organism that lived in the past. These traces range from bones, branches, shells and dung to tracks or im= pressions left by organisms in soft sediments such as sand and clay (Ch, 25, Section 25.2). The fossil record consists of all the {ossis that have been found on Barth and described inthe sel- cenlifieterature. Why did data in the fsll record support the hypothests that species have changed through time? And what data fom extant species—those living today—support the claim that they are ‘modified forms of ancestral species? Evidence 1: The Vastness of Geologic Time Evidence for the Earth's vast age began to mount in the late 1700s with James Hutton’s proposal of the principle of untformitarianism—me ea hat yeulugical processes uicutring tulay are sina to ‘what occurred in the past, Hutton reached hie incight by evalu iting ovidenre—he traveled anand Ranrape and masanved gene logical patterns and rates of rock formation and erosion, Figure 222 Seamentary Rocks Reveal the vastness of Geologlc Time. The relative ages of seaimentary rocks ‘ar© used to éctormine the relative agea of fea rganiama because younger layers aro deposited on top of older ‘ones. The dearest rock layerin the Grand Canyon is ove blion years old. and the top layer is 270 milion years ‘ig. (mya lion years ago) Sedimentary rocks form from sand or mud or other mate- Hlals deposited In layers at locations such as beaches or river ‘mouths. Hutton calculated that sedimentary rocks form at quite a slow rate. When he extrapolated this rate to determine how Jong it would take for maccive rock formation to form, it wa ‘lear that Barth was very old. It was much, much older than the {6000 vears claimed by many proponents of special creation. Hutton’s ideas were popularized by Darwin's close friend, the ‘geologist Charles Lyell. Sedimentary rocks, along with rocks de- rived from episodic lava flows, form with younger layers depos- ted on top of older layers Lyell and otters used this information lw place fossils ia youuper-to-ukder sequence, based oi Use fs ails relative position in layers of sedimentary rock (Figure 22.2). [Ac the scientists ohesrved similarities in rocks and fossile at «aifferent sites, they began to create a geologic time scale: &se- ‘quence of named intervals ealled eons, eras, and periods thatrep- resented the major events in Earth history (see Ch. 25, Section 25.2), This geologic time scale was a relative one, however. The absolute age of Harth wasstill unknown, After Marte Curie’ discovery of radioactivity in the tate 1000s, researchers realized that radioactive decay—the steady rate at which unstable *parent” atoms are converted into more ‘tule “Aaughter” atone fiimlshed a way ta acsign aha ge, in ears tothe relative azesin the weolovic time scale. Radioactive decay functions as @ “natural clock" in which the decay of different elements can be used to measure different time scales. For example, the half-life of uranum-238 18 about 4.5 bi- ion years, whieh means tat 50 percent of wraniun-288 atom wil decay to Ica 206 atomo during thi time. Knowing the half life geologists can use the ratio of uranium to Tend ina rock sample ta Inferthe aze ofthe sample. ‘According to data from radiometric dating, arth is about 4.6 billion years old, and the earliest signs of life appear in rocks that formed 3.43.8 billion years ago. Data from relative and absolute ‘ating techniques agree: Life on Barth is ancient, A great deal of, Une has gue by for eng bo veut Evidence 2: Extinction Changes the Species Present over Time In the early nineteenth century, researchers began dis- covering fossil bones, leaves, and shells that were unlike struc- tures from any known animal or plant. At frst, many scentists ‘insisted that living examples of these species would be found in unexplored regions of the globe. But as research continued and ‘the number and aiversity of ssl collections grew, the argument Luecan essai ese plausible, "The jestuo wae finally settled in 1812 when Baron Georges Cuvier published a dotailed analyte of several extinet apwclon— that is. snecies that no longer exist. Cuvier intentionally focused CHAPTER 22 Evolution by Natural Selection 451 Figure 22:3 Evidence of Extintion. Ths 19th-century drawing pics Cuvier fuss evince, Suiits ayrowd Una Ue sot, ko othor giant focel vertbratee, wae toe largo and unique to be ‘overlooked iit were ave: it must have cone extinct. oon the fossils of large terrestrial animals such as mammoths, ‘mastodons, glant armadillo, giant deer, and glant slots. He rea- soned that unlike many plants or marine animals, those large, tstinetive terrestrial unas probubly would have been discov- cred if they were all live (Figure 82.0). Cuvier's overwhelming, evidence convinced setentists af the fact af oxtinetion Danwin interpreted extinet forms as evidence that species are not stati, immutable entities, unchanged sinee the moment of special creation. He reasoned that if species have gone extinct, then the array of species ivingon Barth has changed through time. ‘Kecent analyses of the ossi record suggest that over 9 per- ‘ent of all the species that have ever lived are now extinet. The data also Indicate that species have gone extinct continuously ‘throughout Rarth’s history—not just in one or even a fow cata- strophie events (Ch. 25, Section 25.4). Evidence 3: Transitional Features Link Older and Younger Species Long before Darwin published his theory, researchers reported striking resemblances between the fossils found in the rocks underlying certain regions and the living species succeed- ng tea In Ue Sanne geugrapile areas, sult us ue extlet lant othe of South America and the slothe that occur there today. "The pattorn wae cn wirasprend that ithecama known ae the “law of succession.” 452 UNIT4 Evolutionary Patterns and Processes, Oder Eusthenooteron Figure 22.4 Transitional Features during the Evolution of the Totrapod Limb. Fossil species similar to today's lungfish ‘and tetrapods have fn and limb bones that are transitional features. Evsttenopteron was aquatic, Tulerpeton was probably ‘comiaquatic. ¥ Contract haw the tranaitians chown here would ft into Lamarei’ ‘arly model of evolution (see Figure 22.1c) versus Darwin and ‘Wallace's model of evolution (see Figure 22.14), Darwin pointed out that this pattern provided strong evidence in favor of the hypothesis that epeciae had changed through ‘time. He proposed that the extinet forms and living forms were related—that they represented ancestors and descendants, ‘As the fossil record expanded, researchers discovered spe- cles with characteristics that broadened the scope of the law of succession, A transitional feature isa trait ina fossil species, ‘that is intermediate between those of ancestral (older) and de- ved (yuunyer) species. Fur exp, Intensive work over Ue past several decades hae yielded forilsthat document a gradual [change aver time from aqiatic animals that had fins ta terwas- ‘rial animals that had limbs (Floure 22.4). Overa period of about 26 million years, the fing of species similar to today’s lungfish transitioned int. limbs cimitar ta those find in taday"e ame phibians. reptiles, and mammals—a group called the tetrapods. (literally, “four-footed”).. Note that such evolutionary transitions are not goal oriented ‘or purposeful. Rather, some individuals with favorable traits managed to survive and reproduce in the new environment, re- sulting in change In the population over time (explained further InSection 22.5), Similar sequence of transitional features document changes ‘that led to the evolution af feathers and flight in hid; stomata and vascular tissue in plants; upright posture, flattened faces, and large brains in humans; jaws in vertebrates; the loss of limbs in snakes; and other traits. Data like these are consistent with predictions from the theory of evolution: If the traits observed in ‘more recent species evolved from traits in more ancient species, ‘ent austen forms ate expected tu uxcur I the appropriate ‘time sequence. Note, however, that individual foscla of tranct ‘tonal forms are not necessarily diret ancestors of lator spoclos— ‘they may be relatives ofthe direct ancestor. Evidence 4: Vestigial Traits Are Evidence of Change through Time Darwin was the first to provide a widely accepted inter- ‘pretaton of vestigial Urals. A vestigial tralt Isa reduced or It ‘completely developed structure that has no function, or reduced function, but is clearly similar to functioning organs or struc- ‘tures in ancestral or closely related species. Biologists have documented thousands of examples of vesti- sia traits + Some whales and snakes have tiny hip and leg bones that do ‘ot help them swim or slither. * Ostriches and kiwis have reduced wings and cannot fy *+ Blind cave-dwelling fish have eve sockets but no eyes. ‘+ Monkeys aud many other primates have loug tals bul vur tiny tailbone, orcoceys, illustrated in Figure 22.5, too small tohelp us maintain balance or grab tree limbs for support ‘The change in species over time is logical because the envi- ronments of epeciee change dramatically over time, Change can ‘occur du to alate factors, ch we 9 ree fn temperatire or drop in sea level, ond biotic factor, auch a3 arsival of a new par asita or tho axtinetinn af a predatae (Ch. 49, Section 49.2), Rar example, a tall can be a beneficial. heritable trait for life in the forest canopy, but may become less beneficial if changes n raln- fall patterns cause the forest to transition tograssland, Evidence 5: Species Gan Be Ubserved Changing Today Biologists have documented hundreds of contemporary popull- tions that are evolving in response to changes in their environ ‘ment, In Section 22-4 you will find a detailed analysis of research fn twa examples of eunlitian in action: the evolution of antie biotic resistance in bacteria and the evolution of beak shape in finches. One key insight from these examples is that evolution canbe observed onthe scale of days, weeks, and months, not just thousands or millions of years. ‘To summarize, evidence from the fossil record and living spe- es Iniales Ue fe io cle aid Ul spectes have eueiged through the course of Barth's history. Purther, change through ‘ime continues today and ean bo measured diretly Evidence of Descent froma Gommon Ancestor Data from the fossil record and contemporary species refute the ‘dea that species are statie, unchanging types. What about the claim that species were ereated independently—meaning that they are unrelated to each other? Evidence 6: Similar Species Are Found in the Same Geographic ‘Area Charles Darwin began to realize that apeciea are related by common ancestry during a five-year voyage he took aboard the English naval ship HMS Beagle. While fulfilling ts mission to explore and map the coast of South America, the Beagle spent a few weeks in the Galépagos Islands off the coast of present-day Ecuador (see Figure 22.6a on page 454). Darwin had taken over the role of ship's naturalist and, as the ‘rst sclentist to study the area, gathered extensive collections of the plants and animals found in these islands. Most famous ‘among the birds he collected were the Galépagos mockingbird snd finchon, Figure 22:5 Vestigial Traits Are Reduced Versions of Traits in Other Species. Te talbune isa hua Ua that has veuuced function. fig a larger sal for halanea and lacomation ‘How could vestigial tits auch as the tty human talon be explained via inheritance of acquired characters? CHAPTER 22 Evolution by Natural Selection 453 (a) Mainland birdo colonized the Galspages. (0) Qaleyayye flies share a won areestor. ‘Tanagors of South ancestor of Gasspagos
Matick at te er an alte tt In rocks in or noar the layors whore the fossils wore found, also agrees with the order of species indicated in the phylogeny. © A piplogeny) off living wholes and dolphins, extiranted from similarities and differences in DNA sequences. indi- cates that hippos—which spend much of their time in shallow Ielonye (50-87 nya) ceracens ‘Anbulocets (88 ya) common tnecator of Fenthoratie (87 mys) ‘onion (-40 mya) Pnphingntonss tart cmc ete ona Gio spate 106m Hoote, semiaquatic soem ‘Semiacuatic, ea Bh sepamatc, ae Fly equane, tai-powered 2 swimmer * vest ars Modem ‘whales [Figure 22.7 Data on Evolution from Independent Sources Aro Consistant. This phylogeny of fossil cotzcoans is ‘consistent with data from relative dating, absolute datina, and phviogenies estimated from molecular tats in ving ‘species ~all agree that whales evolved from terrestrial ancestors that also gave rise to hippos. ‘water—are the closes living relative of cetaceans (shown in (Ch. 25, Figure 25.4). This observation supports the hypothesis that cetaceans and hippos shared a common ancestor that was semlaquatic. * Vestigial hip and hindlimb bones occur in some whales as adults, and vestigial hindlimb buds—outgrowths where legs form in other mammals—occur in some dolphin ‘embryos. “rhe general message here is that many independent ties ot evidence converge on the same conclusion: Whales gradually ‘evolved from a terrestrial ancestor about 50 million years ago. [As you evaluate the evidence eupporting the pattern com- ponent of the theory of evolution, though, i's Important to recognize that no single observation or experiment instantly, “proved! the fact of evolution. Rather, data from many differ- ‘ent sources are much more consistent with evolution than with special creation, Descent with modification is a successful and, [Powerful scientific theory because it explains observations— Such a vest Luts wud Une eluse sitar tes aanwiyg species ‘on neighboring ielands- that special creation does not. What ahont the pracese componant af tha thenry af avnition bby natural selection? If hippos and whales were not created inde- ‘pendently, how did they come tobe? [check vour uNDensTaNDING ‘¥ Ifyou understood this section, you should be able to 1. Determine what kind of evidence would suppor the hrypothesis that beds evoved trom dinosaurs. 2 Explain why the ONA eequances of chimpanzoee and humans ae about 96 percent similar 22.3 The Process of Evolution: How Does Natural Selection Work? Darwin's greatest contribution was not just recognizing the fact of evolution. Lamarck and other researchers, including Darwin's own grandfather Erasmus Darwin, had pro- After yoU complete ths section, posed evolution long betore you should be abt. Charles Darwin beyant lus # Expt ow rar selection work, Inctead, Darwin cru eatisas evnlition cla insight was to recognize 4 process, called natural selection, that could explain the pat= tern of descent with modification. CHAPTER 22 Evolution by Natural Selection 457. Figure 22.8 Diversity of Pigeon Breeds in Captivity. Darvin sed the Breeding of pigeons as a made! system to stuay now the charactortics of populations can change overtime. Darwin's Inspiration How did Darwin arrive at his insight? First, he had spent decades exploring and documenting the diversity of plants and animals, ols arouiad Use globe ad ia his sive Enghaud Su he lad ‘wealth of data on variation within and among apeciea, and he viowod this variation in the context of the anciont and changing, Earth as popularized by his geologist friend. Charles Lyell Allof this careful work gave Darwin an especially strong foun- dation in the pattern of evolution. To make sense of the process of evolution, Darwin turned in part to pigeon breeding—a model system that would be easier to study and manipulate than popu- lations tn the wild, Pigeon breeding was popular in England at the time, and in Darwin's words, “The diversity of the breeds is something actoniehing” (Figure 22.8) Darwin crossbred pigeons and observed how characteristics were passed on to offspring. He chose certain individuals with desirable traits to reproduce, thus manipulating the compost- tion of the population by a process called artificial selection. It ‘was cleat to Darwin and other breeders thatthe diverse varieties ‘were all descended from the wild rock pigeons. ‘Another influence on Darwin was the publication of a ‘book by Thomas Robert Malthus, An Essay on the Principle of Paniation, which inapived heated discnssion in Rngland at the time. Malthus's studies of human populations in England and elsewhere led him toa startling conclusion: Since many more in dividualsare born than can survive, a “struggle for existence” oc- ceursas people compete for food and places to live. Darwin combined his observations of artificial selection with this notion of “struggle for existence” in natural populations, whieh he knew=-fivn his countless studies—contained vaviation, From this eynthosie arose hie concept of natural selection. ‘Although hath Darwin and Wallace arrived atthe same iden, Darwin’s name fs more closelv associated with natural selection because of is extensive evidence fori in On the Origin of Species. 458 UNIT4 Evolutionary Patterns and Processes, ‘stn 79.9 8 gM Pon Drawing modelo can halp you understand andl romember abstact ideas such as Darwin's postulates. Simo crawings can be very ‘etfectv, so cus on the Kas rather than tying to make tee inustraons, Consiga his example rs & =® % ee MODEL Using the tower exami Darwin's ist postulate, in us Mastering Biology EE Darwin's Four Pustulales Darwin distilled the process evolution by natural selection into ‘tour simple postulates (criteria) that form a logical sequence (see Making Models 222)- 1. Variation exists among individual organisms that make up « population, suet ws variation in size and shape. 2, Some of the trait differences are heritable, meaning that ‘they are passed on to offspring, For example, parent flowers ‘with long petals tend to have offspring with long petals, 3. Survival and reproductive success are highly variable. Many more offspring are produce an can possibly survive ‘Thus, only some individuals in each generation survive long ‘enough to produce offspring, and among the individuals that produce offspring, some will produce more than athers, 4. The oubset of individuals that survive best and produce the most offepring is not a random sample of the popu- lation. Instead. individuals with certain heritable traits are more likely tosurvive and reproduce. Altogether, natural selection occurs when individuals with certain heritable traits produce more surviving offspring than do, Individuals without those traits. Thus, the frequency of the se- lected traits increases from one generation to the next. Biologists now know that traits are determined by alleles, particular ver- sons of genes (Chapter 14). So the outcome of evolution by natu- ral selection isa change in allele frequencies in a population over ‘time (Chapter 23). In studying these criteria, you should realize that variation among individuals in a population is essential for evolution to ‘occur. Darwin had to introduce population thinking into biology because itis populations that change over time. To come up with ‘these postulates and understand their consequences, Darwin had. ‘to think ina revolutionary way. Toly, Vologlsts usually condense Darwiars tur postulates Into two part ctatement that more forcefully communteates the ‘essence of evolution by natural selection: Evolution by natural selection aceurs when (1) heritable variation leads to (2) dlifer- ential reproductive success Biological Definitions of Fitness, Adaptation, and Selection ‘To explain the proces of natural selection, Darwin referred to successful individuals as *more ft” than other individuals, In ‘ing so, he gave the wurd “Mines” x definition diferent um its eveiye Engine usage Divlugical fitteas the ability weit Individual to produce surviving, fertile offspring relative to that ability in other individuals in the population. Note that ftness canbe measured, When researchers study a ‘population inthe labor inthe fel, they can measure the rela- tive fitness of individuals by comparing the number of surviving offspring each individual produces, ‘The oncepr or mess, n tur, provides a compact way orfor- tually defining adaptation. The biological meanlagof adaptation, like the biologiea! meaning of itnec, fo diferent frm ite every day Rnglishneage (ene Rate #7) th avaltianary hlalogy, en adaptation isa heritable trait that increases the fitness ofan in- dividual ina particular environment relative to individuals lack- Ing the trait, Adaptations increase ftness—the allt to produce viable, fertile offspring, (You can see the Big Picture of how adap- {ation and fitness relate to natural selection on pages 5#2-53,) Lasty, me term “selection” nas & commonsense meaning in the contextof artificial selection, Dreedera choose which charac- teristics they want to keep or get rd ofin their plant and animal breeds. However, selection has avery different meaning in the biological context of natural selection. Here, it refers to a pas- sive process—iferential reproduction as a result of heritable variation—nota purposeful choice. [I cHteck Your UNDERSTANDING ‘2 Ifyou understood this section, you should he ale to 1 THINK CAREFULLY Ing petals incrmacatha fines af certain flowers by increasing their ability to attract polinators. What does fitness mean inthis context? 2, MODEL Using a popuiton of lowers with citrent petal lengths 2s the example, aw a model of Dwi’ fourth postulate. 22.4 Evolution in Action: Measuring Natural Selection in Populations Today ‘The theory of evolution by natural selectionstestable, meaning bi- ‘ologlats can tet the validity ofeach of Darwin's postulates. Indood, researchers have documented heritable variation and differential, reproductive suceessin a wide array of natural populations. ‘This section summarizes two examples of evolution by natural selection being ob: ‘serve i tue, utes other cave studies are ava slo, imenlving a wide varaty of traits and anganeme Tats start ‘by exploring the evolution of drug resistance, one ofthe great chal- lenges of our time. ‘ter you complete this section, you should beable to 11 Apply Darwin's postulates of natural selection to a ret-wort seca. Case Study 1; How Did Mycobacterium luberculusis Become Resistant lo Antibiulics? Mycobacterium tuberculosis, the bacterium that causes the infec- {ious disease tuberculosis, or TB, has long been a scourge of hu- mankind Tusaly ects the Tunys und causes fever, coughing, seals, wig Io, end uften dea. In Buse and Ure Unite States, TB was once at groat a public health suo a8 cancer Is ‘now. Itreceded in importance during the early 1900s, though, for two reasons: Ay ere Yor los le pee Wile she bt Gt rost M uberculsisinfectionsauickv 2. Dovelopmentof antibiotic allowed phyicans testopinfestons lus Uhe late 1980s, however, tales uf Mf labereuluss infix tion aurged in many countrico, and in 1903 the World Health ‘irgvnnetton doclenedi a global hesttr emergent, Phyuletens were particularly alarmed because the strains of M. tuberculosis responsible for the increase were largely or completely resistant toantibiotics that were once extremely effective. How and why did the evolution of drug resistance occur? The case of a single patient—a young man who lived in Baltimore— IMlustrares whats happening. over the world. ‘APatient History The story begins when the man was admitted to the hospital with fever and coughing. Chest X-rays, followed by bacterial cultures of fluid coughed up from the lungs, showed that he had an active TB infection. He was given several antibiot- Jes for © Weeks, followed by twice-weekly doses of the antibiotic tani for an aditional $$ weeks. Ten nivulls after Usevapy started, bacterial cultures from the patients cheat fluid indicated 10 M tuberrasis call, His chest Xorays wore algo normal, The antibiotics seemed to have cleared the infection. Just two months after the TB tests proved normal, however, the young man was readmitted to the hospital with a fever, se- vere cough, and labored breathing. Despite being treated with a variely of antibiotics, including rifampin, he died of respiratory ‘lure 10 days later, Samples of ud from his lungs showed st ‘M. tuboreulsio was agein growing actively there. But this time the bbactoral colle wore completely resistant to rifampin. Tug-resistant M. therrulasic cells had killed this patient ‘Where did they come from? Could a strain that was resistant to antibiotic treatment have evolved within him? To answer this ques- tion, a research team analyzed DNA from the drug-resistant strain nd compared itwith stored DNA from M.tuberculosiscells that had been isolated a year earlier from the same patient. ARer examining ‘extensive stretches flom each genome, the biologists were able to find only one difference: a point mutation ina gene called rpoB, ‘A Mutation in a Bacterial Gene Confers Resistance The rpob gene codes for a component of RNA polymerase, This enzyme transcribes DNA to mRNA, so itis essential to the survival and reproduction of bacterta cells (Ch. 17, Section 17.1). In this case, the point mutation in the rpoD gene changed a cytosine to thy smne, forming a new alle for the paB gone (see Ch. 16, Section 164) This missense mutation cansed a change in the amina acid ‘sequence of the RNA polymerase (Irom a serine toa leucineat the 153rd amino acid)—and a change in its shape. CHAPTER 22 Evolution by Natural Selection 459) "Thi shape change proved ertical. Rifampin, the antibiotic ‘that was hoing need ta traat the pationt, works hy hinding tothe RNA polymerase of M. tuberculosis and interfering with transceip- tion, Bacterial cells with the C+ mutation continue to produce offspring efficiently even in the presence of the drug, because the ‘drug cannot bind efficiently to the mutant RNA polymerase. In ‘summary, the rpoB mutation, the shape of RNA polymerase, and, ‘drugresistance are related inthis way: DNA ————> Protein ————> Phenotype Mutation Shape change in Org innibts banding of antbitle ‘These results sugeest the steps that led to this patient's death (Figure 22.9): 1. Complotaly hy chance, ana ar a fow af the lange popsiiation of M. tuberculosis bacterial cells present in the patient before the onset of drug therapy had the rpoB allele with the C-sT ‘mutation, Under normal conditions, mutant forms of RNA polymerase do not work as well as the more common form, Lage popltiono ‘atients lungs makes fim sick, ‘ow ofthe bacteria cals happen fo have e point ‘tation inte po gene. Socells withthe C-+1 mutation would not produce many o orm pod gene spin std woul tay aw feguecy—even wile he over ———s {ll population grew t the polnt of inducing aymptomn that a sent the young man tothe hospital > IVIAHJOIVIR +2. Rifampin drug therapy began, killing most M. tuberculosis —— cells with normal RNA polymerac. Ava reelt the overall tell pegultion il in ice vray Bath vietssae wire nl deters ene y 2 Dug thay beg log beet cole rereont fn rorelar alien Borne wale ‘tome cred an ra heey Eventually the M. tuberculosis population regained its former Keune 4. A second ronnd of rine therany hegan hist had na effect an i the drug-resistant M, tuberculosis cells that then dominated ‘ofthe remaining call are ‘the population, so the patient died. ee Testing Darwin's Postulates Does the sequence of events illustrated in Figure 22.9 indicate that evolution by natural selection occurred? One way of answering this question is to The mutant cals prt, review Darwin's four postulates an test whether each one was evn verifed: pater becomes oak aga, 1 Dia veraon exist n the population? es. Dae et beh posencoedyetonaeet orday te es oc bre onsen ote dey Mow Me Eas tastes We naopelu ports oee il every 10" to 10° cells. d Z hy ‘4. .secone roune ot Sein gene Simtecsonte ae a iiretctit Figure 229 Alles That Confer Drug tsar i Fesetaeea cette Ffomoney Wren Nc oa AT tito aes 1 Ein (oe hes i sag 3 ai oe tir \ ; Spratt haaiec cae water aie gettaneniing paleot CL al step 3 or step 4 would respond to drug therapy, 460 UNIT 4 Evolutionary Patterns and Processes, 2 Was this variation heritable? Yes, The researchers showed that the variation in the phenotypes of the we strains—from drug susceptibility todrug resistance—was due ‘ovariation in their genotypes. Because the mutant rpoB gene is passed on to daughter cells when an M. tuberculosis cell replicates, the allele and the phenotype it produces—drug resistance—are passed on to offspring. 3. Was there variation in survival and reproductive success? Yes. Only a tiny fraction of M. tuberculosis cells {im the patient survived the first round of antibiotics long envy to reproduce. Must cells died anid Jet ny oF alinust uo offopring, 4. Were survivat and reproductton nonrandom? Yes. ‘When rifampin was present, certain cells—those with the

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