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Grzimek’s Animal Life Encyclopedia Evolution Michael Hutchins, Series Editor Valerius Geist and Erik Pianka, Advisory Editors • • • • In Association with The Wildlife Society ’ Grzimek s Animal Life Encyclopedia Evolution •••• (c) 2011 Cengage Learning. All Rights Reserved. ’ Grzimek s Animal Life Encyclopedia Evolution •••• Michael Hutchins, Series Editor Valerius Geist and Eric R. Pianka, Advisory Editors •••• In Association with The Wildlife Society (c) 2011 Cengage Learning. All Rights Reserved. Grzimek’sAnimalLifeEncyclopedia ©2011Gale,CengageLearning Evolution ALLRIGHTSRESERVED.Nopartofthisworkcoveredbythecopyrightherein ProjectEditor:DeirdreS.Blanchfield maybereproduced,transmitted,stored,orusedinanyformorbyany meansgraphic,electronic,ormechanical,includingbutnotlimitedto Editorial:JasonEverettandMelissaMcDade photocopying,recording,scanning,digitizing,taping,Webdistribution, RightsAcquisitionandManagement:Robyn informationnetworks,orinformationstorageandretrievalsystems,except Young aspermittedunderSection107or108ofthe1976UnitedStatesCopyright Composition:EviAbou-El-Seoud Act,withoutthepriorwrittenpermissionofthepublisher. Manufacturing:WendyBlurton Imaging:JohnWatkins Forproductinformationandtechnologyassistance,contactusat ProductDesign:KristineJulienandJennifer GaleCustomerSupport,1-800-877-4253. Wahi Forpermissiontousematerialfromthistextorproduct, submitallrequestsonlineatwww.cengage.com/permissions. Furtherpermissionsquestionscanbeemailedto [email protected] CoverphotographreproducedbypermissionofChristianZiegler/Minden Pictures/NationalGeographicStock(pictureofsixspeciesofbutterflyand twospeciesofmoth). Whileeveryefforthasbeenmadetoensurethereliabilityofthe informationpresentedinthispublication,Gale,apartofCengageLearning, doesnotguaranteetheaccuracyofthedatacontainedherein.Galeaccepts nopaymentforlisting;andinclusioninthepublicationofanyorganization, agency,institution,publication,service,orindividualdoesnotimply endorsementoftheeditorsorpublisher.Errorsbroughttotheattentionof thepublisherandverifiedtothesatisfactionofthepublisherwillbe correctedinfutureeditions. LIBRARYOFCONGRESSCATALOGING-IN-PUBLICATIONDATA Grzimek,Bernhard.[Animallifeencyclopediaevolution]Grzimek’s animallifeencyclopediaevolution/MichaelHutchins,serieseditor; ValeriusGeistandEricPianka,advisoryeditors;inassociationwithThe WildlifeSociety. p.cm. Includesbibliographicalreferencesandindex. ISBN978-1-4144-8669-7(hardcover)–ISBN978-1-4144-8670-3(e-book) 1.Evolution(Biology)–Encyclopedias.2.Evolution(Biology)–Studyand teaching.I.Hutchins,Michael.II.Geist,Valerius.III.Pianka,EricR.IV. WildlifeSociety.V.Title.VI.Title:Animallifeencyclopediaevolution. QH360.2.G652011 576.8—dc22 2010038259 Gale 27500DrakeRd. FarmingtonHills,MI,48331-3535 ISBN-13:978-1-4144-8669-7 ISBN-10:1-4144-8669-3 Thistitleisalsoavailableasane-book. ISBN-13:978-1-4144-8670-3 ISBN-10:1-4144-8670-7 ContactyourGale,apartofCengageLearningsalesrepresentativefor orderinginformation. Printed in the United States of America 1 2 3 4 5 6 7 15 14 13 12 11 (c) 2011 Cengage Learning. All Rights Reserved. ••••• Editorial board EditorinChief MichaelHutchins,Ph.D. ExecutiveDirector/CEO TheWildlifeSociety AssociateEditors ValeriusGeist,Ph.D. ProfessorEmeritusof EnvironmentalScience UniversityofCalgary EricR.Pianka,Ph.D.,D.Sc. DentonA.CooleyCentennial ProfessorofZoology SectionofIntegrativeBiology UniversityofTexasatAustin Grzimek’sAnimalLifeEncyclopedia v (c) 2011 Cengage Learning. All Rights Reserved. ••••• Contents Preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ix Sexual selection. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 179 The relevance of evolution to contemporary society . . . . xi Kin selection. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 187 Contributing writers . . . . . . . . . . . . . . . . . . . . . . . . . . xvii Coevolution. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 193 Optimal reproductive tactics and life history theory. . . . 201 Topic overviews Evolutionary physiology. . . . . . . . . . . . . . . . . . . . . . . . 207 Scientific methods and human knowledge. . . . . . . . . . . . .1 Industrial melanism in moths . . . . . . . . . . . . . . . . . . . . 215 Creation stories: The human journey to understand the Galápagos finches . . . . . . . . . . . . . . . . . . . . . . . . . . . . 221 origins of life on Earth . . . . . . . . . . . . . . . . . . . . . . . .9 Artificial selection . . . . . . . . . . . . . . . . . . . . . . . . . . . . 229 History of evolutionary thought. . . . . . . . . . . . . . . . . . .23 Convergent evolution and ecological equivalence. . . . . . 235 Charles Darwin: A life of discovery. . . . . . . . . . . . . . . . .31 Evolution of the animal eye . . . . . . . . . . . . . . . . . . . . . 243 Mechanisms of evolutionary change . . . . . . . . . . . . . . . .45 Evolution of flight. . . . . . . . . . . . . . . . . . . . . . . . . . . . 253 Origins of the universe. . . . . . . . . . . . . . . . . . . . . . . . . .55 Evolution of limblessness. . . . . . . . . . . . . . . . . . . . . . . 263 Origins of life on Earth . . . . . . . . . . . . . . . . . . . . . . . . .63 Canid evolution: From wolf to dog. . . . . . . . . . . . . . . . 271 Speciation: The origins of diversity. . . . . . . . . . . . . . . . .73 Dinosaurs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 279 Sexual reproduction. . . . . . . . . . . . . . . . . . . . . . . . . . . .89 Megafauna. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 291 Species nomenclature, classification and problems of Non-human primate and human evolution . . . . . . . . . . 299 application . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .95 Evolution of the brain and nervous system . . . . . . . . . . 311 The fossil record: A window to the past . . . . . . . . . . . . 103 Evolution of language . . . . . . . . . . . . . . . . . . . . . . . . . 319 What does the fossil record tell us about evolution?. . . . 111 Evolutionary medicine and the biology of health. . . . . . 325 Biogeography: The distribution of life . . . . . . . . . . . . . 123 Evolution and biodiversity conservation . . . . . . . . . . . . 335 Genetics: The blueprint of life. . . . . . . . . . . . . . . . . . . 133 Further reading. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 347 Genes and development. . . . . . . . . . . . . . . . . . . . . . . . 141 Organizations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 351 Evolutionary ecology. . . . . . . . . . . . . . . . . . . . . . . . . . 151 Glossary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 353 Adaptation and evolutionary change . . . . . . . . . . . . . . . 169 Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 359 Grzimek’sAnimalLifeEncyclopedia vii (c) 2011 Cengage Learning. All Rights Reserved. ••••• Preface The idea that species were divinely created and immutable westerncoastofSouthAmerica.Herecordedseveraldifferent persisted for centuries. Then, a studious young man decided species of giant tortoises, each inhabiting different islands in to leave his comfortable home in England and take a long, thearchipelago, whichsuggested thatthey allhadacommon difficult voyage around the world. Charles Darwin’s now- ancestor but had somehow changed over time. Darwin famous trip on H.M.S. Beagle from 1831–1836 would observed a new and unique species of iguana, which, unlike ultimately alter our view of the origin of species. itsmainland ancestor,swam underwaterandgrazedonalgae. He documented a new species of cormorant that possessed Of course others—such as Darwin’s grandfather, Erasmus under-developed wings and had completely lost theability to Darwin, and the well-known French naturalist, Jean-Baptiste fly.Furthermore,Darwinstudiedthemanyspeciesoffinches Lamarck—hadspeculatedthatspeciesmightchangeovertime, that,althoughtheylookedverysimilar,haddifferentlyshaped but they could not identify the mechanism. Darwin’s keen beaksthatallowedthemtoexploitdifferentfoodsourcesmore observations of nature and his landmark 1859 book—The efficiently.Oneevenusedcactusspinesasatooltopryinsects OriginofSpeciesbyMeansofNaturalSelection,orthePreservation from cracks and crevices. of Favoured Races in the Struggle for Life—would provide that answerandrockthescientificworld.Hiswritingsandthoseof IhadtheprivilegeofvisitingtheGalápagosIslandsin1979 the many scientists that followed would forever alter our and of observing these natural wonders firsthand. For perceptions of ourselves and the other living creatures that biologists like me, a visit to the Galápagos is analogous to a sharethisplanet. pilgrimage,butunlikeDarwin,Iknewwhattoexpectbasedon alonglineofscientistswhohadgonebeforeme.Whilesailing Darwin’s most significant contribution to the biological through these unique desert islands, I often wondered how scienceswashistheoryofevolutionthroughnaturalselection. Darwin felt during his first encounters with these unusual The basic concept is simple and elegant: Individual animals naturalphenomena.Iassumehewasthrilledbythediscoveries, possessing heritable traits that allow them to survive and butIalsosuspectthatitchallengedhispreconceivedviews,and reproduce will pass those traits on to the next generation, began his remarkable intellectual journey. It also shaped his while those that do not survive and reproduce will not pass philosophy of science and life. Once a deeply religious man along their traits. The result is a gradual adaptation of a who accepted what he was told purely on the basis of faith, species to its environment, and the evolution of traits that Darwingraduallybecamemoreskeptical,listeningtowhatthe promote survival and reproduction. natural world “told” him through direct observation and experimentation.Heoncesaid:“Greatisthepowerofsteady Ofcourse,environmentsdonotremainstatic,afactoralso misrepresentation, but the history of science shows how, seen asfueling evolutionary change and ultimately leading to fortunately,thispowerdoesnotlongendure.” the creation of new species. Amazingly, this concept was “discovered”independentlybyanotheravidnaturalist,Alfred Itisalsoremarkablethat neitherDarwinnorWallacehad RussellWallace,acontemporary ofDarwin’s,whoalsospent anyideaofhowphysical(phenotypic)traitswerepassedfrom manyyears inthefieldobservingandrecordingdetails about one generation to the next. That had to wait until the the natural world. disciplineofgeneticsdeveloped,whichbeganwithworkofan obscureAustrianmonk,GregorMendel,whostudiedpatterns Darwin and Wallace based their theories on direct of inheritance in garden peas. The development of this observationsofnature.However,itwasDarwinwhoamassed discipline eventually led, in the late1930s and early1940s, the largest amount of evidence. This, and his prolific writing on the topic, explains why his name, not Wallace’s, has to another great synthesis between modern genetics and evolutionary theory. Led by such notables as Theodosius become most closely associated with the discovery of Dobzhansky and Ernst Mayer, this period produced the evolution through natural selection. Hardy-Weinberg Equation, which provided a greater under- ManyofDarwin’smostsignificantandperceptiveobserva- standing of how species change as the result of population- tionsoccurredduringhisvisittotheGalápagosIslandsoffthe level changes in gene frequencies that code for particular Grzimek’sAnimalLifeEncyclopedia ix (c) 2011 Cengage Learning. All Rights Reserved. Preface Evolution traits.Theseconceptswerefurtherconfirmedbythediscovery Evolutionarytheoryisoftenmisinterpreted.Inthepast,some of DNA—the building block of all life—by James Watson, peoplehaveusedittopromoteandjustifysocialdominanceof FrancisCrick and RosalindFranklin in 1953. onegroupoveranother,soitisimportantthatweremainvigilant. However, it is also important to note the positive message of Since then, the field of evolutionary biology and all of its evolution:thatalllivingorganismsarerelatedanddependenton sister disciplines—including geology, paleontology, genetics, manyofthesamethingsfortheirsurvival.Thisshouldbeaneven ecology, animal behavior, zoology, botany and others—have more powerful argument that humans should show greater accumulatedvaststoresofknowledgeaboutthenaturalworld. respectfornatureandnotdriveotherspeciestowardirreversible The power of evolutionary theory has been demonstrated extinction.Giventheinterdependenciesthatexistinnature,such time and time again through both observation and predictive attitudesmaybecriticalforthefutureofHomosapiens. experimental studies. Grzimek’s Animal Life Encyclopedia is a 17-volume set, co- Does this mean that Darwinian natural selection can veringtheentireanimalkingdom,fromprotozoatomammals. explain everything about the natural world? No. There are Originally edited by the famous Frankfurt Zoo director and manynuancestoscience,andweadmittedlystillhavemuchto conservationist, Bernhard Grzimek, and translated into learn. As an example: Niles Eldredge and Stephen J. Gould Englishintheearly1970s,thefirstEnglishedition,consisting introduced the concept of “punctuated equilibrium” in 1972, of 11 volumes, was published by Van Nostrand Reinhold in a comparatively rapid form of evolution that can occur in both England and the United States. That version was very response to environmental change and lead to speciation. popular, but after 30 years the series was in urgent need of Though it did not change the basic concept behind natural updating. During that time, thousands of biologists, working selection,itdidchallengetheideathatallevolutionisuniform inboth thefield andlaboratory,had generated atremendous and gradual. amount ofnew information covering awiderangeofspecies. Recognizing an urgent need to include the new information, This volume is an attempt to summarize our current theentirecollectionwasupdated,reformatted,andcomplete- knowledge about the evolutionary process and the evidence ly rewritten by English-speaking experts from 2001 to 2003 that has accumulated over the past century and a half since under the direction of new owner and publisher, Gale. I had Darwin’s writings. With the discovery of genetics and DNA, the privilege of serving as the primary Series Editor for that which is common to all living organisms, evolution through project. However, the project was not complete, as the natural selection has become the unifying theory in all of original series also included three additional topical volumes: biology. As such, it is a powerful tool with which to Ethology (animal behavior), Ecology, and Evolution. understand the origin of many of nature’s wonders, whether it be fish that change sex, flowers that resemble bees, or the WiththepublicationofGrzimek’sEvolution,SecondEdition,we colorful plumage and bizarre courtship behavior of New havenowupdated,completelyrewritten,andreformattedoneof Guinea’s birds of paradise. thesetopicalvolumes.Yetdespitethemanyscientificadvances detailedinthisbook,thebasicgoalsandperspectiveremainthe Darwin’s revelation challenged prevailing non-scientific sameasthoseexpressedbytheeditorofthe1976firstedition, explanationsfortheoriginoflifeonEarth,includinghumans. Herbert Wendt, who said: “In the individual articles here the His theory generates public controversy even to this day. readerwillfindmaterialhehasnotencounteredinotherworks. Some U.S. states have attempted to ban the teaching of However,sciencenevercomestoastandstill;itknowsnofinal evolutioninelementaryschools,harkeningbacktotheheady conclusion, and each new work brings forth new information days of the Scopes Monkey Trial and the spirited debates of compiledfromthelatestresearchinthatfield....Thereadercan WilliamJenningsBryantandClarenceDarrow.However,the become acquainted with current theoretical thinking about courts have consistently upheld the differences between evolution in reading this volume.” This volume is the next science-based knowledge and religion, which is not testable iterationinthecontinuingaccumulationofknowledgeaboutthe via the scientific method. That being said, controversy originsandinnerworkingsofournaturalworld. persists, and education is key, making the dissemination of information about evolution vitallyimportant.Indeed, evolu- I’dliketoexpressmydeepappreciationtopublisherGale; tion is not just about dinosaurs and other extinct species. On toTheWildlifeSocietyCouncilforallowingmyparticipation the contrary, it is highly relevant to today’s society and we in this project; to my topic editors Valerius Geist and Eric ignoreitatourperil.Ourunderstandingofmodernmedicine, Pianka; and to the many expert authors who contributed agriculture, and conservation—just to name a few—are essays that make up this new volume. dependent on our knowledge of biological evolution. How else can we combat disease organisms that quickly adapt to Bethesda,Maryland,2010 antibiotics, ensure the future of domestic animals and plants, MichaelHutchins andconservesomesemblanceofnatureforfuturegenerations SeriesEditor in a world dominated by human influences? x Grzimek’sAnimalLifeEncyclopedia (c) 2011 Cengage Learning. All Rights Reserved. ••••• The relevance of evolution to contemporary society A lot of time and energy has been consumed debating the transmitted,andtodesigneffectiveprogramstohaltepidemics. question of whether evolution has happened or not, but to Several fields of study outside of biology have benefited from peoplewhoareneitherevolutionarybiologistsnorcreationists,a theinsightthatself-replicatingsystemsevolveandhaveactually better question might be “why does it matter?” In many harnessedtheevolutionaryprocesstoimprovethedesignof,for branchesofscience,includingevolutionarybiology,theimpacts example, computer programs. Finally, evolutionary biology ofaparticulartheoreticaladvanceornewbitofdatacanseem may become a key tool to confront the challenge of a rapidly obscure, orevenirrelevant, to manypeople.Toillustratethis, changingclimate. considerthefollowingtitlesfromtheprestigiousjournalNature on the topic of evolution from the first half of 2010: “Allelic This entry offers examples of some of the benefits that VariationinaFatty-AcylReductaseGeneCausesDivergencein evolutionary biology has bestowed on humanity. Many such MothSexPheromones”;“Post-copulatorySexualSelectionand examplesexist,butonlyalimitednumberhavebeenhighlighted Sexual Conflict in the Evolution of Male Pregnancy”; “Co- here, including some of the major contributions evolutionary optionoftheHormone-SignallingModuleDafachronicAcid– biology has made to contemporary life. Some of the ways in DAF-12 in Nematode Evolution”; “The Ectocarpus Genome whichevolutionarybiologyisexpectedtoassisthumankindinthe and the Independent Evolution of Multicellularity in Brown futurearealsodiscussed. Algae”;and“EvolutionofSelf-CompatibilityinArabidopsisbya MutationintheMaleSpecificityGene.”Asimplesearchofany Topic overviewsFirst, however, it is critical to define the recentscientificjournalwilllikelyrevealequallydensetitles. term evolution. Evolution simply refers toapatternofchange in the phenotype of members of a population (e.g., any These topics are certainly interesting to scientists who characteristic of its behavior or anatomy) through time. work on similar questions, and they may even be inherently Evolution refers only to changes in traits that are inherited interesting to people with a desire to understand the way the fromonegenerationtothenext,usuallybecausesuchchanges world works, but it can be hard to see how these discoveries are the result of genetic differences, though cultures can also will have a material impact on people’s day-to-day lives. If evolvebyprocessesanalogoustobiologicalevolution(seethe there is no impact—if the study of evolution is merely entry “Evolution oflanguage”).Several processes canlead to interesting but not important—then why is the topic subject evolution. Genetic drift, the change in the genetic composi- tosuchimpassioneddebate?Whydomanyindividualsdevote tion of a population due to random variation in birth and theirentirecareerstostudyingtheevolutionaryprocess?Why death, is one such process. Mutation, genetic changes due to do taxpayers invest hundreds of thousands of dollars funding errors during DNA replication, is another. However, the evolutionary research? process that most evolutionary biologists credit with produc- ing broad evolutionary patterns is natural selection. First The answer is that the study of evolution has important elucidated by Charles Darwin in 1859 in his book On the implicationsforhumanwelfare.Thehumanspecies(andevery Origin of Species, natural selection is the process by which otherspeciesontheplanet)istheoutcomeofmillionsofyearsof individualswithadvantageousvariations(forexample,animals evolution;everyaspectofourbodiesandmindshasbeenhoned that are faster or stronger) succeed in producing more bynaturalselection.Understandinghumanevolutionarybiolo- offspring, while individuals with harmful variations fail to gyallowsustocareforourbodiesandmindsmoreeffectively. survive or reproduce. For natural selection to lead to Learningabouttheevolutionaryprocesshasalsoenabledusto evolution, individual variation must be passed from parent harnessthisforceforourownends.Evolutionaryprincipleshave to offspring; in other words, it must be heritable. Over time, unwittinglybeeninuseforthousandsofyearsinthepracticeof many small advantageous variations can accumulate, leading agriculture; more recently, an explicit understanding of evolu- tolargerpatternsofchangewithinthepopulation.Evolution- tion has allowed us to advance agricultural science. Pathogens ary biologists propose that these processes acting together— areconstantlyevolvingtothwartoureffortstocontrolthem,but naturalselection,geneticdrift,andmutation—canexplainthe evolutionary studies of pathogens have enabled scientists to diversity and complexity of all life on Earth. identifynewemergingdiseases,todeterminehowdiseasesare Grzimek’sAnimalLifeEncyclopedia xi (c) 2011 Cengage Learning. All Rights Reserved. Therelevanceofevolutiontocontemporarysociety Evolution A variety of methods are used to study the evolutionary increaseinthesizeandnumberofkernelsperear,adecrease process. Paleontologists study evolution using fossils to in the height and number of branches of the plant, and a observe some of the different life-forms that have existed reduction in the strength of the protective coat surrounding throughhistory,includingtheancestorsofplantsandanimals each kernel. alive today (see the entries “The fossil record: A window to the past” and “What does the fossil record tell us about Research conducted since the early twentieth century has evolution?). Phylogeneticists use similarities and differences helped to identify the genes that are responsible for these between organisms toarrangespeciesinto phylogenetic trees changes. For example, it appears that a single amino acid that reflect the historical relationships among groups. change in the gene teosinte glume architechture1 (tga1) is Evolutionary ecologists and experimental evolutionary biolo- responsibleforthereductionofthethicknessoftheseedcoat gists measure selection within or among natural or experi- in corn. Changes in the gene Teosinte branched1 are responsi- mental populations in real time (see the entry ”Evolutionary ble for a reduction in the height and number of branches in ecology“).Evolutionary geneticistsusemolecularmethods to domestic corn versus wild corn. Genes have also been tracethegeneticchangesthatallowanorganismtotakeona identified that control kernel color, starch content, inflores- new appearance (see the entry ”Genetics: The blueprint of cence structure, and the circadian rhythms of domestic corn. life“). Finally, some evolutionary biologists use computer simulationstopredictwhattheoutcomeofselectionmightbe These studies have increased our understanding of how under given conditions. Together, these methods have been domestic corn differs from teosinte; they may also help usedtoadvanceourunderstandingoftheprocessofevolution. scientists accelerate the development of new varieties. For This body of knowledge has been put to use for human example,plantbreedingcompaniesareworkingtoidentifythe benefit in a variety of ways. genetic changes that are responsible for their best varieties— several of these changes could then be combined in a single variety. They are also screening wild populations of teosinte (and other wild progenitors of domestic crops) for new gene Evolutionary agriculture variants that may prove useful. In some cases evolutionary ThousandsofyearsbeforeDarwinarticulatedthetheoryof geneticistshavebeenabletoincreasethenutritionalvalueofa evolution by natural selection, humans were intuitively using cropbymanipulatinggeneexpression.Forexample,scientists evolutionary principles to shape domestic plants and animals. havebeenabletoincreasetheprovitaminAcontentofseveral In fact, Darwin’s knowledge of animal husbandry (including varietiesofriceusingavariantknownfromwildpopulations. his own extensive work breeding pigeons) was integral to the development of his theory. Breeding experiments at home Thetransformationsthathumanshavewroughtinanimals helpedhimtoappreciatethevariationamongindividualsofa areinsomecasesfarmoredramaticthanthechangesobserved speciesandthepotentialtoharnessthatvariationforavariety in plants. For example, all breeds of the domestic dog are ofpurposes.Bykeepingandbreedinglivestockwithespecially believedtobetheresultofasingledomesticationofthegray usefulcharacteristics,earlyhumans(andDarwin)intentionally wolf (see the entry “Canid evolution: From wolf to dog”). or unintentionally preserved desirable variations and elimi- nated undesirable ones. Likewise, by saving only the largest In some cases, the domestication of wild animals has and most flavorful seeds to plant in the coming year, early resulted in genetic changes to both animals and humans. For humans were sure to gradually improve the quality of their example, very few human cultures have begun using dairy crops. The steady accumulation of variation in a particular products from cattle to supplement their nutritional needs. direction ultimately produced plants and animals so different Dairy products such as milk and cheese contain a unique from their wild counterparts that they could scarcely be sugar, lactose, than can be digested only with the enzyme recognizedasrelatives.Differentvarietiesofthesamespecies lactase. All people have the gene for lactase and produce became as different as distinct species, or even genera. copious quantities of the enzyme as infants when milk comprises the majority of their diet. The adult human diet, Although agriculture is no longer the primary occupation however,hasnothistoricallycontainedmilkormilkproducts, ofcitizensinmanydevelopedcountries,agriculturalproducts andinmostpeoplethelactasegeneisshutofflaterinlife.For are still one of the foundations on which civilization is built. adults who are not producing lactase (who are lactose Consider corn, one of the most successful and widespread intolerant), the consumption of dairy products can result in agricultural products in history. About 800 million tons of severe gastrointestinal distress. corn are produced annually; corn is consumed directly, modified to make a variety of food additives, used as feed Intheculturesinwhichdairycattleareused,adultswould fordomesticanimals,andfermentedtomakeethanolforfuel. clearlybenefitfrombeingabletoconsumelactose-containing Yet the wild progenitor of corn, teosinte grass, is a humble milkasadults.Thisability,alsocalledlactasepersistence,has plant with little nutritional or economic value. The transfor- infactevolvedindependentlyinseveralregionsinwhichdairy mation of corn began thousands of years ago when Central products are a part of the diet. Genetic mutations in the American people began to harvest and plant the seeds of regulatory region of the gene have resulted in lactase teosinte. In its wild form, teosinte has only five to ten small expression in adulthood, allowing comfortable digestion of kernels,andeachkernelisencasedinahardshellthatmakesit lactose.Thegeneticchangesthatcauselactasepersistenceare difficult and time consuming to extract. A relatively small currentlyunderpositiveselection,astheconsumptionofdairy number of changes resulted in modern corn, including an has spread worldwide. Consequently, these genetic changes xii Grzimek’sAnimalLifeEncyclopedia (c) 2011 Cengage Learning. All Rights Reserved.

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