ReCombinatorics Dan Gusfield withcontributionsfrom CharlesH.Langley YunS.Songand YufengWu ReCombinatorics TheAlgorithmicsofAncestralRecombinationGraphs andExplicitPhylogeneticNetworks TheMITPress Cambridge,Massachusetts London,England (cid:2)c 2014MassachusettsInstituteofTechnology Allrightsreserved.Nopartofthisbookmaybereproducedinanyformbyanyelectronicormechanicalmeans (includingphotocopying,recording,orinformationstorageandretrieval)withoutpermissioninwritingfromthe publisher. MIT Press books may be purchased at special quantity discounts for business or sales promotional use. For information,[email protected]. ThisbookwassetinTimesRomanusingLATEXbyTEXnologyInc.andwasprintedandboundinthe UnitedStatesofAmerica. LibraryofCongressCataloging-in-PublicationData Gusfield,Dan. ReCombinatorics:thealgorithmicsofancestralrecombinationgraphsandexplicit phylogeneticnetworks/DanGusfield. p.cm Includesbibliographicalreferencesandindex. ISBN978-0-262-02752-6(hardcover:alk.paper) 1.Geneticrecombination.2.Evolution(Biology)I.Title.II.Title:Recombinatorics. QH443.G872014 572.8’77dc23 2013043434 10 9 8 7 6 5 4 3 2 1 Contents ix 8.3 AnEfficientAlgorithmfortheRoot-KnownGalled-TreeProblem 244 8.4 TheEssentialUniquenessofReducedGalledTrees 250 8.5 AnEfficientAlgorithmfortheRoot-UnknownGalled-TreeProblem 252 8.6 ExtensionstoOtherBiologicalPhenomenaandStructured Recombination 260 8.7 ACommentonGalledTreesandLowerBounds 262 8.8 AdvancedTopic:FurtherSpeedingUptheGalled-TreeAlgorithms 263 8.9 AdvancedTopic:FurtherLimitationsontheNumberofSite ArrangementsonaGall 268 8.10 AdvancedTopic:AConciseNecessaryandSufficientCondition foraGalledTree 270 8.11 AdvancedTopic:TheCharacter-Removal,Site-Compatibility ProblemonGalledTrees 279 8.12 AdvancedTopic:RelationofGalledTreestotheBack-Mutation Model 281 9 GeneralARGConstructionMethods 285 9.1 ARGConstructionMethodsthatDestroyM 285 9.2 ARGBuildingbyTree-Scanning 328 9.3 AdvancedMaterial:TheFastest(inWorst-Case)MinARGAlgorithm 354 10 TheHistoryandForestLowerBounds 361 10.1 TheHistoryLowerBound 361 10.2 TheForestBound 375 11 ConditionstoGuaranteeaFullyDecomposedMinARG 381 11.1 Introduction 381 11.2 SufficientConditionsforaFullyDecomposedMinARG 382 11.3 TheMostGeneralResult 387 11.4 AdditionalApplications 387 12 TreeandARG-BasedHaplotyping 389 12.1 Introduction 389 12.2 TheHaplotypeInference(HI)Problem 391 Idedicatethisbook tothememoryofmymother IrmaGusfield, andtothefutureofmydaughters TaliaandShira Contents Preface xi Acknowledgments xix 1 Introduction 1 1.1 CombinatorialGenomesandtheGrandChallenge 1 1.2 GenealogicalandPhylogeneticNetworks 2 1.3 TheCentralThesisoftheBook 14 1.4 FundamentalDefinitions 14 1.5 TheObservedData 22 1.6 AFewGraphDefinitions 23 1.7 TheBook 33 2 TreesFirst 35 2.1 TheRootedPerfect-PhylogenyProblem 35 2.2 TheCaseofaKnown,Non-ZeroAncestralSequence 47 2.3 TheRoot-UnknownPerfect-PhylogenyProblem 49 2.4 TheSplits-EquivalenceTheorem:TheFundamentalTheoremof Trees 54 2.5 GeneralReferencesonPhylogeneticTrees 59 3 ADeeperIntroductiontoRecombinationandNetworks 61 3.1 TheBiologicalandPhysicalContextofRecombination 61 3.2 TheAlgorithmicandMathematicalContextofRecombination 67 viii Contents 3.3 TheCoreAlgorithmicProblem:RecombinationMinimization 82 3.4 WhyDoWeCareaboutRmin(M)andMinARGs? 84 3.5 Non-MeioticRecombination,andanExtensionoftheModel beyondAnimalsandPlants 87 3.6 MindtheGap 88 4 ExploitingRecombination 97 4.1 HaplotypesandGenotypes 98 4.2 Problem/Solution1:GeneticMappingbyLinkage 99 4.3 Problem/Solution2:LocatingSignaturesofRecentPositiveSelection 110 4.4 Problem/Solution3:AnIdealizedIntroductiontoAssociation Mapping 118 5 FirstBounds 127 5.1 IntroductiontoBounds 127 5.2 LowerBoundsonRmin 128 5.3 AdvancedMaterial:ASharpAnalyticalUpperBoundonRmin(M) 173 6 FundamentalCombinatorialStructureandTools 177 6.1 IncompatibilityandConflictGraphs 177 6.2 ConnectedComponentsandtheStructureofM 179 6.3 SurprisinglyFastIdentificationoftheComponentsoftheConflict andIncompatibilityGraphs 186 7 FirstUsesofFundamentalStructure 197 7.1 TheConnected-ComponentLowerBoundonRmin 197 7.2 TheARGFull-DecompositionTheory 201 7.3 ProofoftheFull-DecompositionTheoremandItsReverse 208 7.4 TheUtilityofFullDecomposition 228 7.5 BroaderImplicationsandApplications 231 8 GalledTrees 235 8.1 IntroductiontoGalledTrees 235 8.2 FirstMajorResults 241
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