Building brains : an introduction to neural development PDF
Preview Building brains : an introduction to neural development
Building Brains Building Brains: An Introduction to Neural Development, First Edition. David Price, Andrew Jarman, John Mason and Peter Kind. © 2011 John Wiley & Sons, Ltd. Published 2011 by John Wiley & Sons, Ltd. ISBN: 978-0-470-71230-6 Building Brains: An Introduction to Neural Development David Price, Andrew Jarman, John Mason and Peter Kind Thiseditionfirstpublished2011#2011byJohnWiley&Sons,Ltd. Wiley-BlackwellisanimprintofJohnWiley&Sons,formedbythemergerofWiley’sglobalScientific,Technicaland MedicalbusinesswithBlackwellPublishing. Registeredoffice: JohnWiley&SonsLtd,TheAtrium,SouthernGate,Chichester,WestSussex,PO198SQ,UK Editorialoffices: 9600GarsingtonRoad,Oxford,OX42DQ,UK TheAtrium,SouthernGate,Chichester,WestSussex,PO198SQ,UK 111RiverStreet,Hoboken,NJ07030-5774,USA Fordetailsofourglobaleditorialoffices,forcustomerservicesandforinformationabouthowtoapplyforpermission toreusethecopyrightmaterialinthisbookpleaseseeourwebsiteatwww.wiley.com/wiley-blackwell. TherightoftheauthortobeidentifiedastheauthorofthisworkhasbeenassertedinaccordancewiththeUK Copyright,DesignsandPatentsAct1988. Allrightsreserved.Nopartofthispublicationmaybereproduced,storedinaretrievalsystem,ortransmitted,inany formorbyanymeans,electronic, mechanical,photocopying,recordingorotherwise,exceptaspermittedbytheUKCopyright,DesignsandPatentsAct 1988,withoutthepriorpermissionofthepublisher. Designationsusedbycompaniestodistinguishtheirproductsareoftenclaimedastrademarks.Allbrandnamesand productnamesusedinthisbookaretradenames,servicemarks,trademarksorregisteredtrademarksoftheir respectiveowners.Thepublisherisnotassociatedwithanyproductorvendormentionedinthisbook.This publicationisdesignedtoprovideaccurateandauthoritativeinformationinregardtothesubjectmattercovered.Itis soldontheunderstandingthatthepublisherisnotengagedinrenderingprofessionalservices.Ifprofessionaladvice orotherexpertassistanceisrequired,theservicesofacompetentprofessionalshouldbesought. LibraryofCongressCataloging-in-PublicationData Buildingbrains:anintroductiontoneuraldevelopment/DavidPrice...[etal.]. p.;cm. Includesindex. ISBN978-0-470-71230-6(cloth)–ISBN978-0-470-71229-0(pbk.) 1. Developmentalneurobiology.2. Nervoussystem–Growth. I.Price,DavidJ.,1957Nov.20- [DNLM:1. Neurons–physiology.2. NervousSystem–anatomy&histology. 3. Neurogenesis. WL102.5] QP363.5.B852011 612.8–dc22 2010053724 AcataloguerecordforthisbookisavailablefromtheBritishLibrary. Thisbookispublishedinthefollowingelectronicformats:ePDF:978-0-470-97963-1; WileyOnlineLibrary:978-0-470-97962-4;ePub:978-0-470-97988-4 Setin10/12ptOUPSwift-RegularbyThomsonDigital,Noida,India. FirstImpression 2011 Contents Preface xi Chick 27 ConventionsandCommonlyusedAbbreviations xiii Mouse 29 2.5 Secondaryneurulationin 1 Modelsand MethodsforStudyingNeural vertebrates 36 Development 1 2.6 Formationofinvertebrateandvertebrate 1.1 Whatisneuraldevelopment? 1 peripheralnervoussystems 37 1.2 Whyresearchneuraldevelopment? 2 Invertebrates 37 Vertebrates:theneuralcrestandtheplacodes 38 Theuncertaintyofcurrentunderstanding 2 Vertebrates:developmentofsenseorgans 40 Implicationsforhumanhealth 2 2.7 Summary 41 Implicationsforfuturetechnologies 3 1.3 Majorbreakthroughsthathave contributedtounderstanding 3 NeuralInduction:AnExampleofHow developmentalmechanisms 4 IntercellularSignallingDetermines 1.4 Invertebratemodelorganisms 4 CellFates 43 Fly 4 Worm 5 3.1 Whatisneuralinduction? 43 Otherinvertebrates 9 3.2 Specificationandcommitment 44 1.5 Vertebratemodelorganisms 9 3.3 Thediscoveryofneuralinduction 44 Frog 9 3.4 Amorerecentbreakthrough: Chick 10 identifyingmoleculesthatmediate Zebrafish 10 neuralinduction 46 Mouse 12 3.5 Conservationofneuralinduction Humans 16 mechanismsinDrosophila 49 Othervertebrates 16 3.6 Beyondthedefaultmodel–other 1.6 Observationandexperiment:methodsfor signallingpathwaysinvolvedinneural studyingneuraldevelopment 17 induction 49 1.7 Summary 18 3.7 Signaltransduction:howcellsrespondto intercellularsignals 54 2 TheAnatomyofDevelopingNervous 3.8 Intercellularsignallingregulatesgene Systems 19 expression 55 2.1 Thenervoussystemdevelopsfromthe Generalmechanismsoftranscriptional embryonicneuroectoderm 19 regulation 55 2.2 Anatomicaltermsusedtodescribe Transcriptionfactorsinvolvedinneural locationsinembryos 20 induction 59 2.3 Developmentoftheneuroectodermof Whatgenesdotranscriptionfactors invertebrates 21 control? 60 C.elegans 21 Genefunctioncanalsobecontrolledbyother Drosophila 21 mechanisms 60 2.4 Developmentoftheneuroectodermof 3.9 Theessenceofdevelopment:acomplex vertebratesandtheprocessof interplayofintercellularandintracellular neurulation 24 signalling 62 Frog 25 3.10 Summary 63 vi (cid:1) CONTENTS 4 PatterningtheNeuroectoderm 65 5.4 Theregulationofneuronalsubtype 4.1 Regionalpatterningofthenervous identity 99 system 65 Neuralprecursorsalreadyhaveintrinsic Patternsofgeneexpressionaresetupby identity 99 morphogens 65 Differentproneuralgenes–differentprogrammes Patterningoccurswithinamonolayerepithelium 66 ofneurogenesis 100 Patterninghappensprogressively 66 Combinatorialcontrolbytranscriptionfactors 4.2 Patterningtheanteroposterior(AP)axisof createsneuronaldiversity 100 theDrosophilaCNS 68 5.5 Theregulationofcellproliferationduring Creatingdomainsoftranscriptionfactor neurogenesis 102 expression 68 Signalsthatpromoteproliferation 102 Dividingtheectodermintosegmentalunits 70 Celldivisionpatternsduringneurogenesis 103 Assigningsegmentalidentity–theHoxcode 70 AsymmetriccelldivisioninDrosophila 4.3 PatterningtheAPaxisofthevertebrate requiresNumb 103 CNS 71 Controlofasymmetriccelldivisioninvertebrate neurogenesis 106 Hoxgenesarehighlyconserved 71 InitialAPinformationisimpartedbythe Invertebrates,divisionpatternsareregulatedto generatevastnumbersofneurons 107 mesoderm 73 5.6 Temporalregulationofneural Mesodermsignalssetupdomainsoftranscription factorexpression 75 identity 109 Thehindbrainisorganizedintosegmentscalled Aneuralcell’stimeofbirthisimportantforneural rhombomeres 76 identity 109 Howrhombomeresarespecified 77 Timeofbirthcangeneratespatialpatterns 4.4 RefiningAPaxispatterningwithin ofneurons 110 regionsandsegments 79 Howdoesbirthdateinfluenceaneuron’sfate? 112 Rhombomerecellpopulationsarekeptseparateby Intrinsicmechanismoftemporalcontrol inDrosophilaneuroblasts 112 Eph–ephrinsignalling 79 Birthdate,laminationandcompetence BoundariesorganizelocalpatterninginDrosophila inthemammaliancortex 114 segments 80 Inthevertebratebrain,boundariesorganizelocal 5.7 Whydoweneedtoknowabout patterning 82 neurogenesis? 117 4.5 Patterningthedorsoventral(DV)axisof 5.8 Summary 117 thenervoussystem 83 PatternsofneuronsintheDVaxis 6 NeuronalMigration 119 ofthespinalcord 83 6.1 Manyneuronsmigratelongdistances EmbryonicoriginoftheDVaxis 84 duringformationofthenervous DVneuralpatterninginDrosophila 84 system 119 DVpatterninginvertebrates 86 6.2 Howcanneuronalmigrationbe MorphogenssetupDVprogenitordomains 89 observed? 119 4.6 Bringingitalltogether 89 Watchingneuronsmoveinlivingembryos 119 4.7 Summary 90 Observingmigratingneuronsincultured tissues 121 5 Neurogenesis:GeneratingNeural Trackingcellmigrationbyindirectmethods 122 Cells 91 6.3 Majormodesofmigration 125 5.1 Generatingneuralcells 91 Somemigratingneuronsareguidedby 5.2 NeurogenesisinDrosophila 92 ascaffold 125 Proneuralgenespromoteneuralcommitment 92 Someneuronsmigrateingroups 126 Lateralinhibition:Notchsignallinginhibits Someneuronsmigrateindividually 128 commitment 94 6.4 Initiationofmigration 130 5.3 Neurogenesisinvertebrates 96 Initiationofneuralcrestcellmigration 130 Proneuralgenesareconserved 96 Initiationofneuronalmigration 131 InthevertebrateCNS,neurogenesisinvolves 6.5 Howaremigratingcellsguidedtotheir radialglialcells 96 destinations? 132 ProneuralfactorsandNotchsignallinginthe DirectionalmigrationofneuronsinC.elegans 132 vertebrateCNS 98 Guidanceofneuralcrestcellmigration 133 CONTENTS (cid:1) vii Guidanceofneuralprecursorsinthedeveloping 8 AxonGuidance 165 laterallineofzebrafish 135 8.1 Manyaxonsnavigatelongandcomplex Guidancebyradialglialfibres 136 routes 165 6.6 Locomotion 137 8.2 Thegrowthcone 165 6.7 Journey’send–terminationof 8.3 Howmightaxonsbeguidedtotheir migration 138 targets? 166 6.8 Themechanismsthatgovernmigration 8.4 Breakingthejourney–intermediate ofimportantpopulationsofcortical targets 168 neuronsremainunknown 141 8.5 Contactguidance 169 6.9 Summary 143 Contactguidanceinaction:pioneersandfollowers, fasciculationanddefasciculation 170 7 HowNeuronsDevelopTheirShapes 145 Extracellularmatrixprovidesasubstratefor 7.1 Neuronsformtwospecializedtypesof navigatingaxons 170 Ephsandephrins:versatilecellsurfacemolecules outgrowth 145 withrolesincontactguidance 171 Axonsanddendrites 145 8.6 Guidanceofaxonsbydiffusible Thecytoskeletoninmatureaxonsand cues–chemotropism 173 dendrites 147 7.2 Thegrowingneurite 148 Netrin–achemotropiccueexpressedatthe ventralmidline 174 Aneuriteextendsbygrowthatitstip 148 Slits 174 Mechanismsofgrowthconedynamics 149 Semaphorins 174 7.3 Stagesofneuriteoutgrowth 150 Otheraxonguidancemolecules 177 Neuriteoutgrowthinculturedhippocampal 8.7 Howdoaxonschangetheirbehaviourat neurons 150 choicepoints? 177 Neuriteoutgrowthinvivo 151 7.4 Neuriteoutgrowthisinfluencedbya Commissuralaxonslosetheirattraction tonetrinoncetheyhavecrossedthefloor neuron’ssurroundings 151 plate 177 Theimportanceofextracellularcues 151 Puttingitalltogether–guidancecuesand Extracellularsignalsthatpromoteorinhibit theirreceptorschoreographcommissuralaxon neuriteoutgrowth 152 pathfindingattheventralmidline 180 7.5 Molecularresponsesinthegrowth Aftercrossingthemidline,commissuralaxons cone 153 projecttowardsthebrain 183 Keyintracellularsignaltransduction 8.8 Howcansuchasmallnumberof events 153 cuesguidesuchalargenumberof SmallGproteinsarecriticalregulators axons? 183 ofneuritegrowth 154 Thesameguidancecuesaredeployedinmultiple Effectormoleculesdirectlyinfluenceactin axonpathways 184 filamentdynamics 155 Interactionsbetweenguidancecues Regulationofotherprocessesintheextending andtheirreceptorscanbealteredby neurite 156 co-factors 185 7.6 Activetransportalongtheaxonis 8.9 Some axons form specific importantforoutgrowth 157 connections over very short 7.7 Thedevelopmentofneuronal distances, likely using different polarity 158 mechanisms 185 Signallingduringaxonspecification 158 8.10 Thegrowthconehasautonomy Ensuringthereisjustoneaxon 160 Whichneuritebecomestheaxon? 160 initsabilitytorespondtoguidance 7.8 Dendrites 161 cues 186 Regulationofdendritebranching 161 Growthconescanstillnavigatewhensevered Dendritebranchesundergo fromtheircellbodies 186 self-avoidance 162 Localtranslationingrowthcones 186 Dendritesandothersensorystructuresbased 8.11 Transcriptionfactorsregulateaxon onmodifiedcilia 163 guidancedecisions 187 7.9 Summary 164 8.12 Summary 189 viii (cid:1) CONTENTS 9 MapFormation 191 10.4 Synaptogenesis 227 9.1 Whataremaps? 191 Thesynapse 227 9.2 Typesofmaps 191 Stagesofsynaptogenesis 227 Coarsemaps 192 Synapticspecificationandinduction 229 Finemaps 195 Synapseformation 233 9.3 Principlesofmapformation 196 Synapseselection:stabilizationand withdrawal 234 Axonorderduringdevelopment 196 10.5 Spinogenesis 235 Theoriesofmapformation 197 9.4 Developmentofcoarsemaps:cortical Spineshapeanddynamics 237 areas 198 Theoriesofspinogenesis 238 Mousemodelsofspinogenesis:theweaver Protomapvs.protocortex 198 mutant 239 Spatialpositionofcorticalareas 200 Molecularregulatorsofspinedevelopment 239 9.5 Developmentoffinemaps: 10.6 Summary 241 topographic 200 Retinotectalpathways 200 Sperryandthechemoaffinityhypothesis 201 11 LifeandDeathintheDevelopingNervous Ephrinsactasmolecularpostcodes System 243 inthechicktectum 202 11.1 Thefrequencyandfunctionofcelldeath 9.6 Inputsfrommultiplestructures:when duringnormaldevelopment 243 mapscollide 205 11.2 Cellsdieinoneoftwomainways: Fromretinatocortexinmammals 206 apoptosisornecrosis 245 Activity-dependenteyespecificsegregation:arole 11.3 Studiesininvertebrateshavetaughtus forretinalwaves 207 muchabouthowcellskill Formationofoculardominancebands 209 themselves 247 Ocular dominance bands form by directed ingrowth Thespecificationphase 249 ofthalamocorticalaxons 210 Thekillingphase 249 Activityandtheformationofoculardominance Theengulfmentphase 250 bands 210 11.4 Mostofthegenesthatregulate 9.7 Developmentoffeaturemaps 211 programmedcelldeathinC.elegansare Featuremapsinthevisualsystem 211 conservedinvertebrates 250 Roleofexperienceinorientationanddirection 11.5 Examplesofneurodevelopmental mapformation 213 9.8 Summary 214 processesinwhichprogrammedcell deathplaysaprominentrole 252 10 MaturationofFunctional Programmedcelldeathinearlyprogenitor Properties 217 cellpopulations 252 Programmedcelldeathcontributestosexual 10.1 Neuronsareexcitablecells 218 differencesinthenervoussystem 253 Whatmakesacellexcitable? 218 Programmedcelldeathremovescellswithtransient Electricalpropertiesofneurons 218 functionsoncetheirtaskisdone 255 Typesofionchannels 219 Programmedcelldeathmatchesthenumbers 10.2 Neuronalexcitabilityduring ofcellsininteractingneuraltissues 259 development 220 11.6 Neurotrophicfactorsareimportant Neuronalexcitabilitychangesdramaticallyduring regulatorsofcellsurvivalanddeath 261 development 221 Growthfactors 261 EarlyactionpotentialsaredrivenbyCa2+, Cytokines 263 notNa+ 221 11.7 Aroleforelectricalactivityinregulating Neurotransmitterreceptorsregulateexcitability programmedcelldeath 265 priortosynapseformation 223 GABAergicreceptoractivationswitchesfrombeing 11.8 Summary 265 excitatorytoinhibitory 223 10.3 Developmentalprocessesregulatedby 12 Experience-DependentDevelopment 267 neuronalexcitability 225 12.1 Effectsofexperienceonvisualsystem Electricalexcitabilityregulatesneuronalproliferation development 268 andmigration 225 Seeingoneworldwithtwoeyes:oculardominanceof Neuronalactivityandaxonguidance 226 corticalcells 268 CONTENTS (cid:1) ix Visualexperienceregulatesoculardominance 269 Synapticchangesthatmediatetheexpression Competitionregulatesexperience-dependent ofLTP/LTDandexperience-dependent plasticity:theeffectsofdark-rearingand plasticity 286 strabismus 270 Metaplasticity 288 Physiologicalchangesinoculardominanceprior Spike-timingdependentplasticity 289 toanatomicalchanges 272 12.3 Cellularbasisofplasticity:development Cooperativebinocularinteractionsand ofinhibitorynetworks 292 visualcortexplasticity 275 Inhibitionmediatesexpressionoftheeffects Thetimingofdevelopmentalplasticity:sensitive ofmonoculardeprivation 292 orcriticalperiods 275 Developmentofinhibitorycircuitsregulatesthe Multiplesensitiveperiodsinthedeveloping time-courseofthesensitiveperiodformonocular visualsystem 277 deprivation 292 12.2 Howdoesexperiencechangefunctional 12.4 Homeostaticplasticity 294 connectivity? 279 12.5 Structuralplasticityandtheroleofthe Electricalpropertiesofdendrites 279 extracellularmatrix 295 Cellularbasisofplasticity:synapticstrengthening 12.6 Summary 297 andweakening 280 Thetime-courseofchangesinsynaptic SuggestionsforFurtherReading 299 weightinresponsetomonoculardeprivation 282 CellularandmolecularmechanismsofLTP/LTD Glossary 303 induction 284 Index 321
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