Table Of ContentKB010lipshitz-FM Lipshitz (Typesetbyspipublisherservices,Delhi)iofxxii September12,2007 18:10
GENES,DEVELOPMENT,ANDCANCER
THELIFEANDWORKOFEDWARDB.LEWIS
KB010lipshitz-FM Lipshitz (Typesetbyspipublisherservices,Delhi)iiofxxii September12,2007 18:10
E.B.Lewis,1996
KB010lipshitz-FM Lipshitz (Typesetbyspipublisherservices,Delhi)iiiofxxii September12,2007 18:10
GENES, DEVELOPMENT, AND CANCER
THELIFEANDWORKOFEDWARDB.LEWIS
Editedwithcommentaryby
HOWARDD.LIPSHITZ
ProfessorandChair
DepartmentofMedicalGeneticsandMicrobiology
GraduateDepartmentofMolecularandMedicalGenetics
CanadaResearchChairinDevelopmentalBiology
UniversityofToronto
SeniorScientist
PrograminDevelopmentalandStemCellBiology
HospitalforSickChildrenResearchInstitute
Toronto,Canada
123
KB010lipshitz-FM Lipshitz (Typesetbyspipublisherservices,Delhi)ivofxxii September12,2007 18:10
AC.I.P.CataloguerecordforthisbookisavailablefromtheLibraryofCongress.
ISBN 978–1–4020–6343–5(PB)
ISBN 978–1–4020–6345–9(e-book)
PublishedbySpringer,
P.O.Box17,3300AADordrecht,TheNetherlands.
www.springer.com
Printedonacid-freepaper.
Allrightsreserved.
(cid:1)C 2007Springer
Nopartofthisworkmaybereproduced,storedinaretrievalsystems,ortransmitted
in any form or by any means, electronic, mechanical, photocopying, microfilming,
recording,orotherwise,withoutthewrittenpermissionfromthePublisher,withthe
exceptionofanymaterialsuppliedspecificallyforthepurposeofbeing enteredand
executedonacomputersystem,forexclusiveusebythepurchaserofthework.
KB010lipshitz-FM Lipshitz (Typesetbyspipublisherservices,Delhi)vofxxii September12,2007 18:10
ThiseditionisdedicatedtoPamelaHarrahLewis:artist,naturalist,opera
connoisseur,friend.
KB010lipshitz-FM Lipshitz (Typesetbyspipublisherservices,Delhi)viiofxxii September12,2007 18:10
The study of the fundamental problems of embryology by experimental methods had almost
come to a standstill until two new methods of procedure appeared above the horizon—one
thedirectapplicationofphysico-chemicalmethodstothedevelopingorganism;theother,the
application of genetics to problems of development. The combination of these two methods
holds for us, at present, I believe the most promising mode of attack on the problems of
developmentalphysiology.
ThomasHuntMorgan(1926)
It would therefore be strange if occasionally a mutation did not arise which rendered its cell
relatively irresponsive to some influence in the surrounding medium that normally exercised
an inhibiting action on its proliferation...We should then have a cancer cell. It is no mere
hypothesisgenemutationsoccasionallyoccurspontaneouslyinsomaticcells...Itisequallywell
known that irradiation enormously increases the frequency of these somatic mutations...it
is but a logical step to conclude that the carcinomas, sarcomas and leukemias arising after
irradiationrepresentmutationsinducedbythelatter...Moreoverthestudyofthemannerand
conditionsofmutationproductionbyirradiationshould...beofsomevalueinrelationtothe
problemofcancerproduction.
HermannJosephMuller(1937)
KB010lipshitz-FM Lipshitz (Typesetbyspipublisherservices,Delhi)ixofxxii September12,2007 18:10
CONTENTS
Permissions xiii
Preface xvii
PrefacetoSecondEdition xix
INTRODUCTION
E.B.LewisandHisScience
BiographicalMemoir 11
SECTIONI:GENES
LewisandtheNatureoftheGene 27
Background 27
Inventionofthecis–transtestforpositioneffects 28
Positionpseudoalleles 30
Thebithoraxpseudoallelesandtheconceptofdevelopmentalcontrol 32
Geneevolutionbytandemduplication 35
Fromcisvectiontotransvection 36
Extendingthebithoraxpseudoalleleicseries 37
Geneticversusfunctionalmodels 39
Fromgenestogenecomplexes 41
ix
KB010lipshitz-FM Lipshitz (Typesetbyspipublisherservices,Delhi)xofxxii September12,2007 18:10
x Contents
PAPERS
Lewis,E.B.(1939).Star-recessive,aspontaneousmutationinDrosophilamelanogaster.
ProceedingsoftheMinnesotaAcademyofScience7:23–26. 45
Lewis,E.B.(1941).AnothercaseofunequalcrossingoverinDrosophilamelanogaster.
ProceedingsoftheNationalAcademyofSciencesUSA27:31–34. 49
Lewis,E.B.(1945).TherelationofrepeatstopositioneffectinDrosophilamelanogaster.
Genetics,30:137–166. 55
Lewis,E.B.(1952).ThepseudoallelismofwhiteandapricotinDrosophilamelanogaster.
ProceedingsoftheNationalAcademyofSciencesUSA38:953–961. 85
Lewis,E.B.(1951).Pseudoallelismandgeneevolution.ColdSpringHarborSymposiaon
QuantitativeBiology,16:159–174. 93
Lewis,E.B.(1954).Thetheoryandapplicationofanewmethodofdetectingchromosomal
rearrangementsinDrosophilamelanogaster.AmericanNaturalist,88:225–239. 117
Lewis,E.B.(1955).Someaspectsofpositionpseudoallelism.AmericanNaturalist,
89:73–89. 133
Lewis,E.B.(1967).Genesandgenecomplexes.HeritagefromMendel,pp.17–47.
EditedbyR.A.Brink.UniversityofWisconsinPress,Madison,Wisconsin. 151
SECTIONII:GENESANDDEVELOPMENT
LewisandtheGeneticControlofDevelopment 179
Developmentalgenetics 179
Gain-of-functionandloss-of-functionbithoraxalleles,andtheoperonmodel 182
Geneticmosaics:spatialandtemporalaspectsofbithoraxfunction 184
The1978paper:Aparadigmforthegeneticcontrolofdevelopment 186
Trans-actingregulators:positiveandnegativecontrolofthehomeoticcomplexes 191
Cis-regulationwithintheBX-C:therulesofengagement 193
Summary 197
PAPERS
Lewis,E.B.(1963).Genesanddevelopmentalpathways.AmericanZoologist,
3:33–56. 199
Lewis,E.B.(1978).AgenecomplexcontrollingsegmentationinDrosophila.
Nature,276:565–570. 229
Duncan,Ian,andLewis,E.B.(1982).Geneticcontrolofbodysegmentdifferentiationin
Drosophila.In:DevelopmentalOrder:ItsOriginandRegulation.40thAnnualSymposium
oftheSocietyforDevelopmentalBiology,Boulder,Colorado,June1981.Editedby
S.SubtelnyandP. B.Green.AlanR.Liss,Inc.,NewYork,1982,pp.533–554. 243
Lewis,E.B.(1982).ControlofbodysegmentdifferentiationinDrosophilabythebithorax
genecomplex.In:EmbryonicDevelopment,PartA:GeneticAspects.EditedbyM.M.Burger
andR.Weber.AlanR.Liss,NewYork,pp.269–288. 263
Lewis,E.B.(1985).RegulationofthegenesofthebithoraxcomplexinDrosophila.Cold
SpringHarborSymposiaonQuantitativeBiology,50:155–164. 279
SECTIONIII:MOLECULESANDDEVELOPMENT
LewisandtheMolecularControlofDevelopment 299
PositionalcloningoftheBX-C 299
Completingthebithoraxmutantseries 301
KB010lipshitz-FM Lipshitz (Typesetbyspipublisherservices,Delhi)xiofxxii September12,2007 18:10
Contents xi
Themolecularbasisofdevelopmentalcontrol 302
Bithoraxandtheevolutionofsexuallydimorphictraits 305
DNAsequencemotifsinthecis-regulatoryregionsoftheBX-C 307
EvolutionoftheHOXcomplexes 309
Summary 310
PAPERS
Bender,W.,Akam,M.,Karch,F.,Beachy,P.A.,Peifer,M.,Spierer,P.,Lewis,E.B.,and
Hogness,D.S.(1983).MoleculargeneticsofthebithoraxcomplexinDrosophila
melanogaster.Science,221:23–29. 311
Karch,F.,Weiffenbach,B.,Peifer,M.,Bender,W.,Duncan,I.,Celniker,S.,Crosby,M.,
andLewis,E.B.(1985).Theabdominalregionofthebithoraxcomplex.
Cell,43:81–96. 327
Celniker,S.E.,andLewis,E.B.(1987).Transabdominal,adominantmutantofthebithorax
complex,producessexuallydimorphicsegmentaltransformationofDrosophila.Genes
andDevelopment,1:111–123. 359
Celniker,S.E.,andLewis,E.B.(1993).MolecularbasisofTransabdominal—anovel
sexuallydimorphicmutantofthebithoraxcomplexofDrosophila.Proceedingsofthe
NationalAcademyofSciencesUSA,90:1566–1570. 383
Lewis,E.B.,Knafels,J.D.,Mathog,D.R.,andCelniker,S.E.(1995).Sequenceanalysisof
thecis-regulatoryregionsofthebithoraxcomplexofDrosophila.Proceedingsofthe
NationalAcademyofSciencesUSA,92:8403–8407. 393
VonAllmen,G.,Hogga,I.,Spierer,A.,Karch,F.,Bender,W.,Gyurkovics,H.,andLewis,
E. B.(1966).SplitsinfruitfulyHoxgenecomplexes.Nature,380:116. 403
Lewis,E.B.,Pfeiffer,B.,Mathog,D.R.,andCelniker,S.E.(2003).Evolutionofthe
homeoboxcomplexintheDiptera.CurrentBiology,13:R587–R588 407
SECTIONIV:RADIATIONANDCANCER
LewisandtheSomaticEffectsofIonizingRadiation 415
Geneticandsomaticeffectsofionizingradiation 415
Thepoliticalandsocialcontext 416
Leukemiaandionizingradiation 418
Politicalfallout 421
Thyroidradiationdosesfromweaponsfallout 427
Bestestimatesofrisk:leukemiainpopulationsexposedtolowdosesofradiation 428
Estimatesoflungcancerrisk:theBatelleBeagles 430
Summary 430
PAPERS
Lewis,E.B.(1957).Leukemiaandionizingradiation.Science,125:965–972. 433
Lewis,E.B.(1959).Thyroidradiationdosesfromfallout.ProceedingsoftheNational
AcademyofSciences,45:894–897. 453
Lewis,E.B.(1963).Leukemia,multiplemyeloma,andaplasticanemiainAmerican
radiologists.Science,142:1492–1494. 459
Lewis,E.B.(1970).Ionizingradiationandtumorproduction.In:GeneticConceptsand
Neoplasia,Williams&WilkinsCo,pp.57–73. 467
KB010lipshitz-FM Lipshitz (Typesetbyspipublisherservices,Delhi)xiiofxxii September12,2007 18:10
xii Contents
Lewis,E.B.(1971).Letters:Leukemia,radiation,andhyperthyroidism.Science,
174:454. 481
Lewis,E.B.(1976).AnalysisoflungtumormortalityintheBatelleBeagleLifespan
experiment.In:HealthEffectsofAlpha-EmittingParticlesintheRespiratoryTract,Reportof
AdHocCommitteeon“HotParticles”oftheAdvisoryCommitteeontheBiological
EffectsofIonizingRadiations,NationalAcademyofSciences–NationalResearch
Council.Publ.EPA520/4-76-013,U.S.EnvironmentalProtectionAgency,Washington,
DC. 483
Lewis,E.B.(1998).Ionizingradiation,cancerinductionandradioactivefallout.In:
ProceedingsoftheInternationalConference.TheDiscoveryofPoloniumandRadium—itsscientific
andphilosophicalconsequences,benefitsandthreatstomankind.PolishAcademyofSciences,
Warsaw,Poland,pp.227–237. 495
SECTIONV:HISTORICALPERSPECTIVES
Lewis,E.B.(1994).Homeosis:thefirst100years.TrendsinGenetics,10:341–343. 505
Lewis,E.B.(1995).RememberingSturtevant.Genetics,141:1227–1230. 511
Lewis,E.B.(2003).C.B.Bridges’repeathypothesisandthenatureofthegene.
Genetics,164:427–431. 517
Lewis,E.B.(2004).DidDemerecdiscoverintragenicrecombinationin1928?Genetics
168:1785–1786. 527
Lewis,E.B.(unpub).Autobiographicalsketch. 531
Lewis,E.B.(1996).Thebithoraxcomplex:thefirstfiftyyears.LesPrixNobel,1995.
Almquist&WiksellInternational:Stockholm,Sweden,pp.233–260. 539
APPENDIX
CurriculumvitaeofE.B.Lewis 565
References 575
Index 583
