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GlobAl GlobAl edITIon edITIon eG dl ITo IobA nl Concepts of Genetics C For these Global editions, the editorial team at Pearson has o collaborated with educators across the world to address a wide range n c of subjects and requirements, equipping students with the best possible e p elevenTh edITIon learning tools. This Global edition preserves the cutting-edge approach t s and pedagogy of the original, but also features alterations, customization, o William S. Klug • Michael R. Cummings and adaptation from the north American version. f G Charlotte A. Spencer • Michael A. Palladino e n e t i c s e l e v e n T h e d I T I o n S K p l e u n g c • e r C • u m P a m l la in d g in s This is a special edition of an established title widely o used by colleges and universities throughout the world. Pearson published this exclusive edition for the benefit of students outside the United States and Canada. If you purchased this book within the United States or Canada, you should be aware that it has been imported without the approval of the Publisher or Author. Pearson Global Edition Klug_1292077263_mech.indd 1 27/08/15 6:42 PM CO N C EpT s Of E L E V E N T H E D I T I O N G L O B A L E D I T I O N William S. Klug THE COLLEGE Of NEw JErsEy Michael R. Cummings ILLINOIs INs TITuTE Of TECHNOLOGy Charlotte A. Spencer uNIVErsIT y Of ALBEr TA Michael A. Palladino MONMOuTH uNIVErsIT y with contributions by Darrell Killian COLOrADO COLLEGE Boston Columbus Indianapolis New York San Francisco Amsterdam Cape Town Dubai London Madrid Milan Munich Paris Montréal Toronto Delhi Mexico City São Paulo Sydney Hong Kong Seoul Singapore Taipei Tokyo A01_KLUG7260_11_GE_FM.indd 1 27/08/15 4:02 PM Credits and acknowledgments for materials borrowed from other Editor-in-Chief: Beth Wilbur sources and reproduced, with permission, in this textbook appear Senior Acquisitions Editor: Michael Gillespie on the appropriate page within the text. Executive Director of Development: Deborah Gale Executive Editorial Manager: Ginnie Simione-Jutson Pearson Education Limited Project Editor: Dusty Friedman Edinburgh Gate Editorial Assistant: Chloe Veylit Harlow Text Permissions Project Manager: Timothy Nicholls Essex CM20 2JE Text Permissions Specialist: PreMedia Global USA, Inc. 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Manager, Media Production, Global Edition: Vikram Kumar Senior Manufacturing Controller, Production, Global Edition: Authorized adaptation from the United States edition, entitled Trudy Kimber Concepts of Genetics, 11th edition, ISBN 978-0-321-94891-5, by Design Manager: Marilyn Perry William S. Klug, Michael R. Cummings, Charlotte A. Spencer, and Interior Designer: Cenveo Publisher Services Michael A. Palladino, published by Pearson Education © 2015. Cover Designer: Lumina Datamatics Ltd. All rights reserved. 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Where those designations appear in this book, and the publisher was aware of a trademark claim, the designations have been printed in initial caps or all caps. MasteringGenetics is a trademark, in the U.S. and/or other countries, of Pearson Education, Inc. or its affiliates. ISBN 10: 1-292-07726-3 ISBN 13: 978-1-292-07726-0 British Library Cataloguing-in-Publication Data A catalogue record for this book is available from the British Library 10 9 8 7 6 5 4 3 2 1 Typeset by Cenveo® Publisher Services Printed and bound by Vivar in Malaysia A01_KLUG7260_11_GE_FM.indd 2 27/08/15 4:02 PM Dedication To Kathy, Lee Ann, Bob, and Cindy, who mean the very most to us, and serve as our respective foundations when we are writing, and when we are not. WSK, MRC, CAS, and MAP 33 A01_KLUG7260_11_GE_FM.indd 3 27/08/15 4:02 PM About the Authors William S. Klug is an Emeritus Professor of Biology at Charlotte A. Spencer is a retired Associate Professor from The College of New Jersey (formerly Trenton State Col- the Department of Oncology at the University of Alberta lege) in Ewing, New Jersey, where he served as Chair of in Edmonton, Alberta, Canada. She has also served as a the Biology Department for 17 years. He received his B.A. faculty member in the Department of Biochemistry at the degree in Biology from Wabash College in Crawfordsville, University of Alberta. She received her B.Sc. in Microbi- Indiana, and his Ph.D. from Northwestern University in ology from the University of British Columbia and her Evanston, Illinois. Prior to coming to The College of New Ph.D. in Genetics from the University of Alberta, followed Jersey, he was on the faculty of Wabash College as an As- by postdoctoral training at the Fred Hutchinson Cancer sistant Professor, where he first taught genetics, as well as Research Center in Seattle, Washington. Her research general biology and electron microscopy. His research in- interests involve the regulation of RNA polymerase II tran- terests have involved ultrastructural and molecular genetic scription in cancer cells, cells infected with DNA viruses, studies of development, utilizing oogenesis in Drosophila and cells traversing the mitotic phase of the cell cycle. She as a model system. He has taught the genetics course as has taught courses in biochemistry, genetics, molecular well as the senior capstone seminar course in Human and biology, and oncology, at both undergraduate and graduate Molecular Genetics to undergraduate biology majors for levels. In addition, she has written booklets in the Prentice over four decades. He was the recipient in 2001 of the first Hall Exploring Biology series, which are aimed at the un- annual teaching award given at The College of New Jersey, dergraduate nonmajor level. granted to the faculty member who “most challenges students to achieve high standards.” He also received the 2004 Michael A. Palladino is Dean of the School of Science and Outstanding Professor Award from Sigma Pi International, Professor of Biology at Monmouth University in West and in the same year, he was nominated as the Educator of Long Branch, New Jersey. He received his B.S. degree in the Year, an award given by the Research and Development Biology from Trenton State College (now known as The Council of New Jersey. College of New Jersey) and his Ph.D. in Anatomy and Cell Biology from the University of Virginia. He directs Michael R. Cummings is Research Professor in the Depart- an active laboratory of undergraduate student researchers ment of Biological, Chemical, and Physical Sciences at studying molecular mechanisms involved in innate im- Illinois Institute of Technology, Chicago, Illinois. For more munity of mammalian male reproductive organs and genes than 25 years, he was a faculty member in the Department involved in oxygen homeostasis and ischemic injury of the of Biological Sciences and in the Department of Molecular testis. He has taught a wide range of courses for both majors Genetics at the University of Illinois at Chicago. He has and nonmajors and currently teaches genetics, biotechnol- also served on the faculties of Northwestern University ogy, endocrinology, and laboratory in cell and molecular and Florida State University. He received his B.A. from St. biology. He has received several awards for research and Mary’s College in Winona, Minnesota, and his M.S. and teaching, including the 2009 Young Investigator Award of Ph.D. from Northwestern University in Evanston, Illinois. the American Society of Andrology, the 2005 Distinguished In addition to this text and its companion volumes, he Teacher Award from Monmouth University, and the 2005 has also written textbooks in human genetics and general Caring Heart Award from the New Jersey Association for biology for nonmajors. His research interests center on the Biomedical Research. He is co-author of the undergradu- molecular organization and physical mapping of the het- ate textbook Introduction to Biotechnology, Series Editor erochromatic regions of human acrocentric chromosomes. for the Benjamin Cummings Special Topics in Biology At the undergraduate level, he teaches courses in Mende- booklet series, and author of the first booklet in the series, lian and molecular genetics, human genetics, and general Understanding the Human Genome Project. biology, and has received numerous awards for teaching excellence given by university faculty, student organiza- tions, and graduating seniors. 4 A01_KLUG7260_11_GE_FM.indd 4 27/08/15 4:02 PM Brief Contents pArT ONE pArT fOur GENEs, CHr OMOsOMEs, GENOMICs AND HErEDIT y 20 Recombinant DNA Technology 523 1 Introduction to Genetics 35 21 Genomics, Bioinformatics, and Proteomics 556 2 Mitosis and Meiosis 50 22 Applications and Ethics of Genetic Engineering 3 Mendelian Genetics 74 and Biotechnology 603 4 Extensions of Mendelian Genetics 104 5 Chromosome Mapping in Eukaryotes 138 pArT fIVE 6 Genetic Analysis and Mapping in Bacteria and GENETICs Of Or GANIsMs Bacteriophages 168 AND pOpuLATIONs 7 Sex Determination and Sex Chromosomes 198 23 Quantitative Genetics and Multifactorial 8 Chromosome Mutations: Variation Traits 638 in Number and Arrangement 222 24 Neurogenetics 659 9 Extranuclear Inheritance 248 25 Population and Evolutionary Genetics 681 pArT TwO spECIAL T OpICs DNA: sTruCTurE, rEpLICATION, IN MODErN GENETICs AND VArIATION 1 Epigenetics 708 10 DNA Structure and Analysis 265 2 Emerging Roles of RNA 718 11 DNA Replication and Recombination 295 3 DNA Forensics 735 12 DNA Organization in Chromosomes 322 4 Genomics and Personalized Medicine 746 5 Genetically Modified Foods 758 pArT THrEE 6 Gene Therapy 772 GENE ExprEssION, rEGuLATION, AND DEVELOpMENT Appendix A Selected Readings 787 13 The Genetic Code and Transcription 342 Appendix B Answers to Selected Problems 799 14 Translation and Proteins 371 Glossary 841 15 Gene Mutation, DNA Repair, and Credits 863 Transposition 401 Index 867 16 Regulation of Gene Expression in Prokaryotes 430 17 Regulation of Gene Expression in Eukaryotes 451 18 Developmental Genetics 479 19 Cancer and Regulation of the Cell Cycle 503 5 A01_KLUG7260_11_GE_FM.indd 5 27/08/15 4:02 PM Explore Cutting Edge Topics EXPANDED! Six Special Topics Special TopicS in Modern GeneTicS 5 in Modern Genetics Genetically Modified Foods mini-chapters concisely explore cutting-edge, engaging, relevant topics, and three are new to the Throughout the ages, humans have used selective violence. On August 8, 2013, 400 protesters broke through breeding techniques to create plants and animals security fences surrounding a field trial of Golden Rice in Eleventh Edition: with desirable genetic traits. By selecting organisms the Bicol region of the Philippines (ST Figure 5–1). Within with naturally occurring or mutagen-induced variations 15 minutes, they had uprooted and trampled most of the and breeding them to establish the phenotype, we have GM rice plants. The attackers argued that Golden Rice was evolved varieties that now feed our growing populations a threat to human health and biodiversity and would lead to and support our complex civilizations. Western corporate control of local food crops. Although we have had tremendous success shuffling Opposition to GM foods is not unique to Golden Rice. ■ Epigenetics genes through selective breeding, the In 2013, approximately two million people process is a slow one. When recombinant marched against GM foods in rallies held ■ New! Emerging Roles DNA technologies emerged in the 1970s “genetic engineering in 52 countries. Some countries have out- and 1980s, scientists realized that they of animals and right bans on all GM foods, whereas others of RNA could modify agriculturally significant plants promised embrace the technologies. Opponents cite ■ DNA Forensics C 5 owragya—nibsmy si dienn tai fyminogr e apnrde ccilsoen ainngd greanpieds pahna seex cinit isncgie nnetiwfi c ssaofmetey sacniedn etinstvsi roanndm ecnotmalm ceorncciaelr nisn, tewrhesiltes ■ Genomics and OPI tinhgat tchoensfee rg deneseisr aibnlteo t roarigtsa,n tihsemns i.n Gtreondeutcic- agriculture, efoxotodls . tThhe e atlompiocs to fl GimMit lfeosos dv aitrttruaecst so hfy pGeMr- Personalized Medicine AL T eisnegdi naene reinxgci toifn agn nimewal sp ahnadse p ilnan stsc ipenrotimfic- prowduitcht iivnictyr,e raesdeudc ed btioolne, aanndd m eixsiangfgoerrmataetdio nrh—eotonr ibco, thin sfiodrems oa-f ■ New! Genetically SPECI arflegadrviuoccrue ladtnu drep n,e uswttirictiihtdi oein n.curseea,s edan pdr oednuhcatinvciteyd, enpheastnicciedde fluasveo, ra nadnd tfohoe ddSse?ob ,a Itnwe .hthati s aSrpe ectihael Ttrouptihcs cahbaoputet r,G wMe Modified Foods Beginning in the 1990s, scientists cre- nutrition.” will introduce the science behind GM foods ated a large number of genetically modi- and examine the promises and problems ■ New! Gene Therapy fied (GM) food varieties. The first one, approved for sale in 1994, was the Flavr Savr tomato—a tomato that stayed firm and ripe longer than non-GM toma- toes. Soon afterward, other GM foods were developed: papaya and zucchini with resistance to virus infection, canola containing the tropical oil laurate, corn and cotton plants with resistance to insects, and soybeans and sugar beets with tol- erance to agricultural herbicides. By 2012, more than 200 different GM crop varieties had been created. Worldwide, GM crops are planted on 170 million hectares of arable land, with a global value of $15 billion for GM seeds. Although many people see great potential for GM foods—to help address malnutrition in a worl d wdIiSthCU aS SgIrOonw Q-UESTIOnS 771 ing human population and climate change—others question NEW! review Questions the technology, oppose GM food development, and some- times resort to violence to stop the introduction of GM vari- ST Figure 5–1 Anti-GM protesters attacking Golden Rice End-of-Chapter 1. Hismows cdroea gteedn etthicroalulyg hm soedleicfiteivde o brrgeaendievisntmigitea?s smc.o iEmnvp eaArne pwGrietohcl duorergsnaon Rr- iacned— wa6. a vWcsia edhrniaecetiv etmeys l ieonoa psfdu eerrvdeiecs l oeohp nativhn eaga btch eouceunomn nttraatikenaseii?nnt T astoro ti dahaalneltee v, ihaotew fitv hseiutrlacodmcu.e isgOnsh fnAu a l A hdsuaeefivgce-uu srtit 8y ,f e2n0c1e3 a, npdro dteessttreorsy eind tahne e Pxhpielirpimpiennetsa bl firoeklde of 2. Can current GM crops be consideredn aosn trpanrosgfient ibc aosr icsi stgoen hice?l p alleviathtees ve isttaramteginie sA be dene?ficiencies in the Golden Rice plants. Questions are provided 3. WOfh tyh?e approximately 200 GM cropd evvareielotiepsi nthga t whaovrel db—eenh as be78e.. nWD ehtshactre iib se tGa orhglodewetn Rpolifac neo t2sp, apcnaodns hitobiweo nwtr aaasn nistf dcor remateedd (?Puhsiontgo cobuiorltiesstyic P hilippine Department of Agriculture Regional Field Unit 5) developed, only a few are widely used. What are these varieties, methods. How does this method compare with Agrobacterium for each Special Topic and how prevalent are they? 724 tumefaciens-mediated transformation? 4. How does glyphosate work, and how has it been used with GM 9. How do positive and negative selection techniques contribute crops to increase agricultural yields? to the development of GM crops? chapter to help students 5. Describe the mechanisms by which the Cry proteins from 10. Describe how the Roundup-Ready soybean variety was devel- Bacillus thuringiensis act as insecticides. oped, and what genes were used to transform the soybean plants. review key ideas and to M30_KLUG8915_11_ST05_pp724-737.indd 724 20/05/14 5:58 PM facilitate class discussions. discussion Questions Questions are assignable 1. What are the laws regulating the development, approval, and 3. One of the major objections to GM foods is that they may be use of GM foods in your region and nationally? harmful to human health. Do you agree or disagree, and why? through MasteringGenetics 2. Do you think that foods containing GM( ain)gredients should be (b) labeled as such? What would be the advantages and disadvan- tages to such a strategy? S P E C R oAdAsV7 IAL ■ New! Photos and AAV8 TO iblleuesntr aadtidoends have R PAAAArhEAAAA8VVVVR2789 C AoAnrh64VeR1rhs98R AAAAVV98 AAAAVV27 PIC 5 throughout the text. rh64R1 A01_KLUG7260_11_GE_FM.indd 6 27/08/15 4:02 PM M30_KLUG7260_11_GE_ST05.indd 771 14/08/15 1:52 PM Scion TATA G F P Rootstock TATA GF FG Graft junction Dicer siRNA RISC Explore Classic and Modern Approaches NEW! Chapter 10 Chapter 21 Evolving Concept of the Gene sections, integrated in key chapters, highlight how scientists’ understanding of the gene has changed over time. Chapters 3, 4, 5, 6, 10, 12, 14, 16, 21 NEW! Modern Approaches to understanding Gene function feature introduces the impact of modern gene targeting approaches on our understanding of gene function. Each entry explores experimental approaches, analyzes data, and relates to a concept discussed in the chapter. Includes discussion questions. Topics include: ■ Identifying Mendel’s Gene for Regulating White Flower ■ MicroRNAs Regulate Ovulation in Female Mice Color in Peas (Ch. 3) (Ch. 17) ■ Drosophila Sxl Gene Induces Female Development ■ Single-Gene Signaling Mechanism Reveals Secrets to (Ch. 7) Head Regeneration in Planaria ■ Mouse Models of Down Syndrome (Ch. 8) (Ch. 18) ■ Lethal Knockouts of DNA Ligase Genes (Ch. 11) ■ RbAp48 and a Potential Molecular Mechanism for ■ Transposon-Mediated Mutations Reveal Genes Age-Related Memory Loss (Ch.24) Involved in Colorectal Cancer (Ch. 15) A01_KLUG7260_11_GE_FM.indd 7 27/08/15 4:02 PM 13.10 rna pOlyMEraSE dirEctS rna SynthESiS 321 In an elegant experiment using the E. coli–phage system, phosphodiester bonds (see Figure 10–12). The energy cre- the results of which were reported in 1961, Sydney Brenner, ated by cleaving the triphosphate precursor into the mono- François Jacob, and Matthew Meselson clarified this question. phosphate form drives the reaction, and inorganic phos- They labeled uninfected E. coli ribosomes with heavy isotopes phates (PPi) are produced. and then allowed phage infection to occur in the presence A second equation summarizes the sequential addition of of radioactive RNA precursors. By following these compo- each ribonucleotide as the process of transcription progresses: nents during translation, the researchers demonstrated that DNA the synthesis of phage proteins (under the direction of newly (NMP)n + NTP e¡nzyme (NMP)n+1 + PPi synthesized RNA) occurred on bacterial ribosomes that were As this equation shows, each step of transcription involves present prior to infection. The ribosomes appeared to be the addition of one ribonucleotide (NMP) to the growing nonspecific, strengthening the case that another type of RNA polyribonucleotide chain (NMP)n+1, using a nucleoside serves as an intermediary in the process of protein synthesis. triphosphate (NTP) as the precursor. That same year, Sol Spiegelman and his colleagues RNA polymerase from E. coli has been extensively reached the same conclusion when they isolated 32P-labeled characterized and shown to consist of subunits designated RNA following the infection of bacteria and used it in a, b, b′, v, and s. The active form of the enzyme, the holo- molecular hybridization studies. They tried hybridizing enzyme, contains the subunits a2bb′s and has a molecular this RNA to the DNA of both phages and bacteria in sepa- weight of almost 500,000 Da. While there is some variation rate experiments. The RNA hybridized only with the phage in the subunit composition of other bacteria, it is the b and DNA, showing that it was complementary in base sequence b′ polypeptides that provide the catalytic mechanism and to the viral genetic information. active site for transcription. As we will see, the s (sigma) The results of these experiments agree with the concept factor [Figure 13–9(a)] plays a regulatory function in the of a messenger RNA (mRNA) being made on a DNA tem- initiation of RNA transcription. plate and then directing the synthesis of specific proteins While there is but a single form of the enzyme in E. in association with ribosomes. This concept was formally coli, there are several different s factors, creating variations proposed by François Jacob and Jacques Monod in 1961 as of the polymerase holoenzyme. On the other hand, eukary- part of a model for gLene eregualatiorn inn bac teraia. Sinnce tdhen, potesr disaplayc threte diisctincet for msp of RrNAo polybmeralsee, eacmh solving mRNA has been isolated and thoroughly studied. There is consisting of a greater number of polypeptide subunits no longer any question about its role in genetic processes. than in bacteria. In this section, we will discuss the process of transcription in prokaryotes. We will return to a discussion of eukaryotic transcription later in this chapter. 64 3 Mendelian Genetics 13.10 rna polymerase directs rna insiGhts and solutions Synthesis 13–3 The following represent deoxyribonucleotide sequences As a student, you will be asked to demonstrate your knowledge Apply the approach we just studied to the following problems. 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Using these proportions to complete a forked-line diagram M13_KLUG8915_11_C13_pp308-336.indd 321 hyoinutrs ,w aonrdk achgaecinksint gth e 24/03/14 3:49 PM btrBBhabbbtebi ostFn11 1mem//a22aal leHHpesseo altamennerd-odo z zffyebyesgmgmooauaul lesyse s mbsg,, l bra-Babcbyksotedpecrtphaortaon itlobfi w asrrebmeeip lasoir rttediehes eaes:np 9lpt: s3lu yp:i3irn:o1ngp pttohhreeti inpoooIrntosydn pouifncc 9 trs /la1at6wisio:3 .ag /(s1 R 6we:ah3em /c1eo6mtnm:1bp/se1ur6 tdt.e h) taNht eot htfiein s al Answers Appendix. and Solutions. M03_KLUG8915_11_C03_pp040-069.indd 64 15/01/14 4:36 PM MasteringGenetics™ MasteringGenetics helps students master key genetics concepts while reinforcing problem solving skills with hints and feedback specific to their misconceptions. Tutorial topics include: ■ Pedigree analysis ■ DNA replication ■ Sex linkage ■ RNA processing ■ Gene interactions ■ Genomics A01_KLUG7260_11_GE_FM.indd 8 27/08/15 4:02 PM 162 6 Genetic analySiS and MappinG in Bacteria and BacteriophaGeS 19. In an analysis of rII mutants, complementation testing yielded 21. Using mutants 2 and 3 from the previous problem, following the following results: mixed infection on E. coli B, progeny viruses were plated in a series of dilutions on both E. coli B and K12 with the following results. Mutants Results (1/2 lysis) (a) What is the recombination frequency between the two 1, 2 1 mutants? 1, 3 1 1, 4 2 Strain Plated Dilution Plaques 1, 5 2 E. coli B 10-5 2 Predict the results of testing 2 and 3, 2 and 4, and 3 and 4 together. E. coli K12 10-1 5 20. If further testing of the mutations in Problem 19 yielded the following results, what would you conclude about mutant 5? (b) Another mutation, 6, was tested in relation to mutations Mutants Results 1 through 5 from the previous problems. In initial testing, 2, 5 2 mutant 6 complemented mutants 2 and 3. In recombination 3, 5 2 testing with 1, 4, and 5, mutant 6 yielded recombinants 4, 5 2 wmiuthta 1ti oannd 6 ?5, but not with 4. What can you conclude about extra-Spicy problems Visit for instructor-assigned tutorials and problems. “How Do We 22. During the analysis of seven rII mutations in phage T4, 25. An Hfr strain is used to map three genes in an interrupted mat- Know?” questions 5m wutearne tisn 1 c, i2st, raonnd B 6. Owef rteh einse c, imsturotann At 4, wwhaisl ea mdeulteatniotns 3o,v 4e,r laanpd- irnifgr .e (xNpeor immaepn to. rThdeer icsr oimssp ilsie dH firn tah+eb +licst+irnifg *of Fth-e a a-lble-lecs-; ping mutant 5. The remainder were point mutations. Nothing rifr is resistance to the antibiotic ri>fampicin.) The a>+ gene is ask students to identify was known about mutant 7. Predict the results of complemen- required for the biosynthesis of nutrient A, the b+ gene for tation (1 or –) between 1 and 2; 1 and 3; 2 and 4; and 4 and 5. nutrient B, and c+ for nutrient C. The minus alleles are auxo- 23. In studies of recombination between mutants 1 and from trophs for these nutrients. The cross is initiated at time=0, and examine the the previous problem, the results shown in the following table and at various times, the mating mixture is plated on three were obtained. types of medium. Each plate contains minimal medium experimental basis (MM) plus rifampicin plus specific supplements that are Strain Dilution Plaques Phenotypes indicated in the following table. (The results for each time underlying important E. coli B 10-7 4 r interval are shown as the number of colonies growing on E. coli K12 10-2 8 1 each plate.) concepts. (a) Calculate the recombination frequency. Time of Interruption (b) When mutant 6 was tested for recombination with mutant 5 min 10 min 15 min 20 min 1, the data were the same as those shown above for strain B, Nutrients A and B 0 0 4 21 but not for K12. The researcher lost the K12 data, but remem- Nutrients B and C 0 5 23 40 bered that recombination was ten times more frequent than when mutants 1 and 2 were tested. What were the lost values Nutrients A and C 4 25 60 82 (dilution and colony numbers)? (c) Mutant 7 (Problem 22) failed to complement any of the (a) What is the purpose of rifampicin in the experiment? other mutants (1–6). Define the nature of mutant 7. (b) Based on these data, determine the approximate location 24. In Bacillus subtilis, linkage analysis of two mutant genes affect- on the chromosome of the a, b, and c genes relative to one ing the synthesis of two amino acids, tryptophan (trp2-) and another and to the F factor. NE W! Concept Etthyreo xfsoinlloet w(tirynrg1- ad),a wta-a sa npSde rdfoprarmw eiadllc upsoiynssgib t lreaP ncosfnorcrlmuosaitoinonbs .r Eegxlaarmediinngme s (ecx)p Cerainm tehnet ?l oIcf antoiot,n d oefs itghne arinf geexnpee rbime deentte trom dineteedr mini nthei tsh e linkage. What is the purpose of Part B of the experiment? [Ref- location of rif relative to the F factor and to gene b. Questions ask students to cerhencae: El.l Neesnter,g Me. Sc hsafteru, andd J.e Lendertbesrg (t19o63) .]solve 26. A plaque assay is performed beginning with 1 mL of a solution containing bacteriophages. This solution is serially check their understanding of cD oonmor DNpAlex R epcipirenot CeblllTterrpa+nm s ftoyrrm-sa,nt smN19oa6. ny basdteiiallu ddteidlu ttihorne ew tiitmh e9s.9 b my Lc oomf lbiiqnuiindg m0e.1d imumL .o Tf heeanc h0 .s1e mquLe no-f Key Concepts. oA n. trp 2+d tyar1+ta tdrp2- e tyrr1-ivettdrrpp-+ fttyyrrr++om3326 87primarytphl aeq fuineas.l Wdilhuatti oins tihs ep ilnatietida li nd etnhsei tpyl aoqf ubea catsesraiyo panhdag yeise lidns t 1h7e g B e. ttarrnnpp22+-d e ttyytrr11-+ics trlpi2- t teyr1-rattttrrruppp+-+ r ttteyyyrrr-++. 12 95206 27. oItDinroe intgaesi rn omcaofl itg nr1eae nm nwesLshf o?tiwcrmho gaatetin oaen ts i maerxeep, “etlrhiinemk feeondllt”o, wtoui snwignh gdic ahvta ao rwtihoeeurress .pcroomdubicnead-. M06_KLUG8915_11_C06_pp134-163.indd 162 18/03/14 1:47 PM MasteringGenetics™ N E W! 140 Prepare students Additional for the challenging problems they will Practice see on tests and Problems offer exams: question more opportunities types include to develop problem sorting, labeling, solving skills. These entering numerical questions appear only information, in MasteringGenetics, multiple choice, and they include and fill-in-the- targeted wrong answer blank. feedback to help students learn from their mistakes. Complete the Problems and Discussion Questions at the end of each chapter. 90% of questions are now available in MasteringGenetics . A01_KLUG7260_11_GE_FM.indd 9 27/08/15 4:02 PM