Building brains an introduction to neural development

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adam_text Titel: Building brains Autor: Price, David J. Jahr: 2011 Preface xi Chick 27 Conventions and Commonly iised Abbreviations xiii Mouse 29 2.5 Secondary neurulation in 1 Models and Methods for Studying Neural vertebrates 36 Development 1 2.6 Formation ofinvertebrate and vertebrate 1.1 What is neural development? 1 peripheral nervous Systems 37 1.2 Why research neural development? 2 Invertebrates 37 The uncertaintyof current understanding 2 Vertebrates: the neural crest and the placodes 38 Implicationsforhumanhealth 2 Vertebrates: developmentofsenseorgans 40 Implicationsforfuture technologies 3 2.7 Summary 41 1.3 Maj or breakthroughs that have contributed to understanding developmentalmechanisms 4 3 Neural Induction: An Example of How 1.4 Invertebratemodelorganisms 4 Intercellular Signalling Determines Ry 4 CellFates 43 vYorm 5 3.1 What is neural induction? 43 Other invertebrates 9 3.2 Specification and commitment 44 1.5 Vertebrate model organisms 9 3.3 The discoveryof neural induction 44 Frog 9 3.4 A more recent breakthrough: Chick 10 identifying molecules that mediate Zebrafish 10 neural induction 46 Mouse 12 35 Conservation of neural induction Humans 16 mechanisms in Drosophüa 49 Other vertebrates 16 36 ßeyond the default model - other 1.6 Observation and expenment: methods for signalling pathways involved in neural studying neural development 17 induction 49 1.7 Summary 18 37 Signal transduction: how cells respond to intercellular Signals 54 2 The Anatomy of Developing Nervous 3g Intercellular signalling regulates gene Systems 19 expression 55 2.1 The nervous system develops from the Generd mechanisms oftranscriptionai embryomc neuroectoderm 19 regulation 55 2.2 Anatomical terms used to describe Transcription factors involved in neural locations in embryos 20 induction 59 2.3 Development of the neuroectoderm of What genes do transcription factors invertebrates 21 control? 60 C. elegans 21 Genejunction can also be controlled by other Drosophüa 21 mechanisms 60 2.4 Development of the neuroectoderm of 3.9 The essence of development: a complex vertebrates and the process of interplay of intercellular and intracellular neurulation 24 signalling 62 Frog 25 3.10 Summary 63 VI • CONTENTS Patterning the Neuroectoderm 65 5.4 The regulationof neuronal subtype 4.1 Regional patterning of the nervous identity 99 system 65 Neural precursors already have intrinsic Patterns ofgene expression are set up by identity 99 morphogens 65 Different proneural genes - different programmes Patterning occurs within a monolayer epithelium 66 ofneurogenesis 100 Patterning happens progressively 66 Combinatorial control by transcription factors 4.2 Patterning the anteroposterior (AP) axis of creates neuronal diversity WO the Drosophüa CNS 68 5.5 The regulationof cell proliferation during Creating domains oftranscription factor neurogenesis 102 expression 68 Signals that promote proliferation 102 Dividing the ectoderm into segmental units 70 Cell division patterns during neurogenesis 103 Assigning segmental identity - the Hox code 70 Asymmetrie cell division in Drosophüa 4.3 Patterning the AP axis of the vertebrate requiresNumb 103 CNS 71 Control ofasymmetric cell division in vertebrate Hox genes are highly conserved 71 neurogenesis 106 Initial AP information is imparted by the ln vertebrates, division patterns are regulated to mesoderm 73 generatevastnumbersofneurons 107 Mesoderm Signals set up domains of transcription 5.6 Temporal regulation of neural factor expression 75 identity 109 The hindbrain is organized into segments called A neural cell s time ofbirth is importantfor neural rhombomeres 76 identity 109 How rhombomeres are speeified 77 Time ofbirth can generate spatial patterns 4.4 Refining AP axis patterning within ofneurons 110 regions and segments 79 ^ow aoes bifth date influence a neuron sfate? 112 Rhombomere cell populations are kept separate by Intrinsic mechanism of temporal control Eph-ephrin signalling 79 in Drosophüa neuroblasts 112 Boundaries organize local patterning in Drosophüa ßirth date- lamination and competence segments 80 in the mammalian cortex 114 In the vertebrate brain, boundaries organize local 5.7 Why do we need to know about patterning 82 neurogenesis? 117 4.5 Patterning the dorsoventral (DV) axis of 5.8 Summary 117 the nervous system 83 Patterns ofneurons in the DV oxis 6 Neuronal Migration 119 ofthe spinal cord 83 6.1 Many neurons migrate long distances Embryonic origin ofthe DV axis 84 during formation ofthe nervous DV neural patterning in Drosophüa 84 system 119 DVpatterntng in vertebrates 86 62 How can neuronal migration be Morphogens set up DVprogenitor domains 89 observed? 119 4.6 Bringingitalltogether 89 Watching neurons move in Uvingembryos 119 4./ Summary 90 Observingmigrating neurons in eultured tissues 121 Neurogenesis: Generating Neural Traädng cell migration by indirect methods 122 Cells 91 6.3 Major modesof migration 125 5.1 Generating neural cells 91 Some migrating neurons are guidedby 5.2 Neurogenesis in Drosophüa 92 a scaffold 125 Proneural genes promote neural commitment 92 Some neurons migrate in groups 126 Lateral Inhibition: Notch signalling inhibits Some neurons migrate individually 128 commitment 94 6.4 Initiation of migration 130 5.3 Neurogenesis in vertebrates 96 Initiation of neural crest cell migration 130 Proneural genes are conserved 96 Initiation of neuronal migration 131 In the vertebrate CNS, neurogenesis involves 6.5 How are migrating cells guided to their radial glial cells 96 destinations? 132 Proneural factors and Notch signalling in the Diredional migration ofneurons in C. elegans 132 vertebrate CNS 98 Guidance of neural crest cell migration 133 CONTENTS • VII Guidance of neural precursors in the developing 8 AxonGuidance 165 lateral lineofzebrafish 135 8.1 Many axons navigate long and complex Guidanceby radial glialfibres 136 routes 165 6.6 Locomotion 137 82 The growth cone 165 6.7 Journey s end - termination of 83 How might axons be guided to their migration 138 targets? 166 6.8 The mechanisms that govern migration 84 ßreaking the journey - intermediäre ofimportantpopulationsofcortical targets 168 neurons remain unknown 141 85 Contact guidance 169 6.9 Summary 143 Contact guidance in action: pioneers and followers, fasciculation and defasciculation 170 How Neurons Develop Their Shapes 145 Extracellular matrix provides a Substrate for 7.1 Neurons form two specialized types of navigathig axons 170 nllfrrr nwrh 145 ^^s and eP^nns: versatile cell surface molecüles Axnm and dendrites 145 with roles in contact guidance 171 The cytoskeleton in mature axons and 8-6 Guidance of axons by drffusible dendrites 147 cues - chemotropism 173 7 2 The erowing neurite 148 Netrin - a chemotropic cue expressed at the A neunte extendsby growth atitstip 148 ventral midline 174 Mechanisms of growth cone dynamics 149 7.3 Stagesof neurite outgrowth 150 Semaphorins 174 ir ° ^ ., . ,4. ji- , Other axon guidance molecüles 177 Neurite outgrowth in culturedhippocampal TT ö , , . , , neurons 150 w axons change their behaviour at Neurite outgrowth in vivo 151 choice points? 177 7 4 Neurite outgrowth is influenced by a Commissural axons lose their attraction neuron s surroundings 151 tonetrinoncetheyhavecrossedthefioor The importance of extracellular cues 151 Pae . ,_ . ^ * « i ¦ . i *!. * *. ™ • i.-uj* Putting it all together - guidance cues and Extracellular Signals that promote or inhibit , . * * ö , . , ., ir_ their receptors Choreograph commissural axon neunte outgrowth 152 f +¦,¦*,¦ __ ., , , -+.1. ^-v. pathnnding at the ventral midline 180 7.5 Molecular responses in the growth *, J ? ., . . , *• *J (Xttov /-» / »cciMOr tV 0 minima mmmiccumt n cone 153 After crossing the midline, commissural axons project towards the brain 183 Key intracellular Signal transduction gg How can such a small number of 6V£TltS T.S3 _ ... , , + cues guide such a large number of Small G proteins are cntical regulators ö ö of neurite growth 154 ff° .J3 , , , Effector molecüles directly influence actin ^ same Suldance mes are dePIo^d in multiPle filament dynamics 155 axonpathways 184 Regulation of other processes in the extending Interactions between guidance cues neurite 156 and t^r recePtors can ^e altered by 7.6 Active transport along the axon is nn co-factors 185 important for outgrowth 157 89 Some axons form specific 7.7 The development of neuronal connections oyer very short polarity 158 distances, hkely using different Signalling during axon specification 158 mechanisms 185 Ensuring there isjust one axon 160 810 The growth cone has autonomy Which neurite becomes the axon? 160 m its abihty to respond to guidance 7.8 Dendrites 161 cues 186 Kegularion ofdendrite branching 161 Growth cones can still navigate when severed Dendrite branches undergo from lheir cell bodies 186 self-avoidance 162 Local translation in growth cones 186 Dendrites and other sensory structures based 8.11 Transcription factors regulate axon onmodifiedcilia 163 guidance decisions 187 7.9 Summary 164 8.12 Summary 189 VIII • CONTENTS Map Formation 191 10.4 Synaptogenesis 227 9.1 What are maps? 191 The Synapse 227 9.2 Typesofmaps 191 Stagesof synaptogenesis 227 Coarsemaps 192 Synaptic speäfication and induction 229 Finemaps 195 Synapse formation 233 9.3 Principles of map formation 196 Synapse selection: stabilization and Axon order during development 196 withdrawal 234 Theories of map formation 197 10.5 Spinogenesis 235 9.4 Development ofcoarse maps: cortical Spine shape and dynamics 237 areas 198 Theories of spinogenesis 238 Protomapvs.protocortex 198 Mouse modeis of spinogenesis: the weaver Spatial position of cortical areas 200 mutant 239 9.5 Development of finemaps: Molecvlar regulato^ ofspine development 239 topographic 200 10.6 Summary 241 Retinotectal pathways 200 Sperry and the chemoaffinity hypothesis 201 11 Life and Death in the Developing Nervous Ephrins act as molecülar postcodes System 243 in the chick tectum 202 11.1 The frequency and function of cell death 9.6 Inputs from multiple structures: when during normal development 243 maps collide 205 11.2 Cells die in one of two main ways: From retina to cortex in mammals 206 apoptosis or necrosis 245 Activity-dependent eye spetific segregation: arole 1 j .3 studies in invertebrates have taught us for retinal waves 207 much about how cells kill Formation ofoculardominancebands 209 themselves 247 Ocular dominance bands form by directed ingrowth ^ dficatim hase 249 ofthalamocortical axons 210 Thekülingphase 249 Acttvtty and the formation of ocular dominance ^ mgulftnmt phase 250 bands 2W 11.4 Most ofthe genes that regulate 9.7 Development offeature maps 211 ° iuwu-%»u» ,m D . ^ . ., . , . ^ _.. programmed cell death m C. elegans are conserved in vertebrates 250 Feature maps in the Visual system 211 Role ofexperience in orientation and direction . , . map formation 213 11.5 Examples of neurodevelopmental 9.8 Summary 214 processes in which programmed cell death plays a prominent role 252 10 Maturation of Functional Programmed cell death in early progenitor Properties 217 cell populations 252 10.1 Neurons are excitable cells 218 Programmed cell death contributes to sexual What makesa cell excitable? 218 differences in the nervous system 253 ui ^- 1 *• c -,-, , Programmed cell death removes cells -with transient Electrica! properties ofneurons 218 . * i7 . _ , . , ___ Typesof ton Channels 219 functions once their task is done 255 . ., , -^ v-,._ , • Programmed cell death matches the numbers 10.2 Neuronal excitabihty during o/cJk . Memcting nmml fou£S 259 development 220 11.6 Neurotrophic factors are important Neuronal exataMty changes dramatically during regulators of cell survival and death 261 development 221 ö Early action Potentials are drivenbyCa2+, (?m1thpCt0r* 261 notNa+ 221 Cytohnes 263 Neurotransmitter reeeptors regulate exatability n-7 Arole for electrical activity in regulatmg prior to Synapse formation 223 programmed cell death 265 GABAergicreeeptoractivationswitchesfrombeing WA Summary 265 excitatory to inhibitory 223 10.3 Developmental processes regulatedby 12 Experience-Dependent Development 267 neuronal excitabihty 225 12.1 Effects ofexperience on Visual system Electrica! excitability regulates neuronal proliferation development 268 and migration 225 Seeing one world with two eyes: ocular dominance of Neuronal activity and axon guidance 226 cortical cells 268 CONTENTS • IX Visual experience regulates ocular dominance 269 Synaptic changes that mediate the expression Competition regulates experience-dependent oflTP/LTD and experience-dependent plastiaty: the effects of dark-rearing and plastiaty 286 Strabismus 270 Metaplastidty 288 Physiological changes in ocular dominance prior Spike-timing dependent plastiaty 289 to anatomical changes 272 12.3 Cellular basis of plasticity: development Cooperative binocular interactions and of inhibitory networks 292 Visual cortex plasticity 275 Inhibition mediates expression ofthe effects The timing ofdevelopmental plasticity: sensitive ofmonocular deprivation 292 or critical periods 275 Development of inhibitory circuits regulates the Multiple sensitive periods in the developing time-course ofthe sensitive periodfor monocular Visual system 277 deprivation 292 12.2 How does experience change functional 12.4 Homeostatic plasticity 294 Connectivity? 279 12.5 Structural plasticity and the role ofthe Electrica! properties of dendrites 279 extracellular matrix 295 Cellular basis of plasticity: synaptic strengthening 12 6 Summarv 297 and wedkening 280 The time-course of changes in synaptic weight in response to monocular deprivation 282 Suggesüons for Further Reading 299 Cellular and molecular mechanisms of LTP/LTD Glossary 303 induction 284 Index 321
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title Building brains an introduction to neural development
title_auth Building brains an introduction to neural development
title_exact_search Building brains an introduction to neural development
title_full Building brains an introduction to neural development David Price ...
title_fullStr Building brains an introduction to neural development David Price ...
title_full_unstemmed Building brains an introduction to neural development David Price ...
title_short Building brains
title_sort building brains an introduction to neural development
title_sub an introduction to neural development
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