Biology of Sensory Systems, Second Edition

514 Pages·2009·32.8 MB·English

by  C. U. M. Smith(auth.)

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Biology
of
Sensory
Systems
BiologyofSensorySystems,SecondEdition C. U. M.Smith
© 2008JohnWiley&Sons,Ltd. ISBN: 978-0-470-51862-5
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For Jenny
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Biology
of
Sensory
Systems
Second Edition
C. U. M. SMITH
VisionSciences,AstonUniversity,
Birmingham,UK
John Wiley&Sons,Ltd
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Thiseditionfirstpublished2008(cid:1)C 2008byJohnWiley&SonsLtd
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LibraryofCongressCataloguing-in-PublicationData
Smith,C.U.M.(ChristopherUphamMurray)
Biologyofsensorysystems/C.U.M.Smith.–2nded.
p.; cm.
Includesbibliographicalreferencesandindex.
ISBN978-0-470-51862-5(alk.paper)–ISBN978-0-470-51863-2(alk.paper)
1.Sensesandsensation. I.Title.
[DNLM:1.SenseOrgans. 2.Evolution. 3.MolecularBiology.
4.Physiology,Comparative.WL700S644b2008]
QP431.S5362008
612.8–dc22 2008021435
ISBN:978-0-470-51862-5
AcataloguerecordforthisbookisavailablefromtheBritishLibrary.
Setin10/11.5ptTimesbyAptara
PrintedinSingaporebyMarkonoPrintMediaPteLtd
FirstImpression 2008
Cover:Imageofahorizontalsectionthroughtheommatidiaofaninsect’seye(Chrysopa;thegreenlacewing).Courtesy
ofProfessorAdrianHorridge,AustralianNationalUniversity.
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CONTENTS
PrefacetoSecondEdition ................. xi Chapter3 GeneralFeaturesofSensory
Systems................................... 31
PrefacetoFirstEdition.................... xiii
3.1 ClassificationoftheSenses........... 33
3.2 Modality............................ 34
3.3 Intensity............................ 34
PARTI: PRELIMINARIES............... 1
3.4 Adaptation.......................... 36
3.5 ReceptiveFields..................... 36
Chapter1 Elements........................ 3
3.6 MapsofSensorySurfaces............ 37
1.1 AllostericEffectors.................. 4
3.7 HierarchicalandParallelDesign ..... 38
1.2 Membranes......................... 5
3.8 FeatureExtractionandTriggerStimuli 38
1.2.1 Lipids........................... 5
3.9 ConcludingRemarks................ 39
1.2.2 Proteins......................... 6
Box3.1 HermannvonHelmholtz .. 32
1.2.3 MobilityofProteins............. 8
1.3 MembraneSignallingSystems........ 8
1.3.1 ReceptorMolecules.............. 9 Chapter4 ClassificationandPhylogeny..... 41
1.3.2 G-proteins...................... 10 4.1 Systematics.......................... 41
1.3.3 EffectorsandSecondMessengers. 12 4.2 ClassificationintoSixKingdoms..... 42
1.4 ChannelsandGates................. 14 4.3 Unicellularity ....................... 42
1.4.1 TRPChannels................... 14 4.4 Multicellularity...................... 42
1.4.2 Ligand-GatedIonChannels 4.5 ProtostomesandDeuterostomes..... 43
(LGICs)......................... 16 4.6 ClassificationoftheMetazoa......... 44
1.4.3 Voltage-GatedIonChannels 4.7 EvolutionofNervousSystems........ 49
(VGICs)......................... 16 4.7.1 Cnidaria........................ 50
1.5 ConcludingRemarks................ 18 4.7.2 Platyhelminthes................. 50
4.7.3 Nematoda....................... 50
Chapter2 Membranes,ActionPotentials, 4.7.4 Annelida........................ 51
Synapses.................................. 19 4.7.5 Arthropoda..................... 52
2.1 TheMeasurementofRestingPotentials 20 4.7.6 Mollusca........................ 53
2.2 TheIonicBasesofRestingPotentials. 21 4.7.7 Echinodermata.................. 55
2.3 ElectrotonicPotentialsandCable 4.7.8 Chordata........................ 55
Conduction......................... 23 4.8 ConcludingRemarks................ 55
2.4 ReceptorandGeneratorPotentials... 24
2.5 SensoryAdaptation.................. 25 Chapter5 Genes,Genomicsand
2.6 ActionPotentials.................... 26 NeurosensorySystems..................... 57
2.7 SynapsesandSynapticTransmission.. 28 5.1 Introduction......................... 57
2.8 ConcludingRemarks................ 30 5.2 ComparativeGenomics.............. 60
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5.3 GenomesandNeurosensorySystems. 62 Chapter9 EquilibriumandHearing:
5.4 ConcludingRemarks................ 64 TheUsesofHairCells..................... 123
Box5.1 NomenclatureofGenesand 9.1 AnatomyandPhysiologyofHair
Proteins........................... 59 Cells................................ 123
PartI:SelfAssessment..................... 65 9.2 LateralLineCanals.................. 128
PartI:Notes,ReferencesandBibliography.. 69 9.3 EvolutionoftheVertebrateEar...... 129
9.3.1 Equilibrium..................... 129
9.3.2 Phonoreception................. 132
9.3.3 BatSonar....................... 145
PARTII: MECHANOSENSITIVITY...... 73
9.4 ConcludingRemarks................ 145
Box9.1 BiophysicsofOuterHair
Chapter6 MechanosensitivityofCell
Cells.............................. 141
Membranes................................ 75
Box9.2 GeneticsandDeafness..... 143
6.1 MechanosensitiveChannelsin
E.coli............................... 76
Chapter10 CerebralAnalysis.............. 147
6.2 DetectionofOsmoticSwellingby
10.1 The Mammalian Vestibular Pathway
HypothalamicCellsinMammals..... 80
andReflexes........................ 148
6.3 ConcludingRemarks................ 83
10.2 TheMammalianAuditoryPathway. 150
10.2.1 CochlearFibres................ 151
Chapter7 Kinaesthesia.................... 85
10.2.2 CochlearNucleus .............. 152
7.1 KinaestheticMechanismsin
10.2.3 SuperiorOlivaryNuclei......... 153
Arthropods.......................... 85
10.2.4 InferiorColliculus.............. 153
7.1.1 StretchReceptorsinCrustacean
10.2.5 MedialGeniculateNucleus..... 153
Muscle.......................... 86
10.3 TheAvianAuditoryPathwayandthe
7.1.2 InsectSensilla................... 87
MappingofAuditorySpacebythe
7.1.3 MaintenanceofEquilibrium
BarnOwl.......................... 153
inFlight ........................ 90
10.4 TheMammalianAuditoryCortex... 157
7.2 KinaestheticMechanismsinMammals 91
10.5 TheBatAuditorySystemand
7.2.1 IntrafusalSpindleEndings....... 91
Echolocation....................... 158
7.2.2 GolgiTendonOrgans............ 94
10.6 TheHumanAuditoryCortex
7.2.3 JointReceptors.................. 96
andLanguage...................... 162
7.3 ConcludingRemarks................ 97
10.7 LateralizationandtheNeuroanatomy
ofLanguage........................ 167
Chapter8 Touch.......................... 99 10.8 LanguageandtheFOXP2Gene..... 169
8.1 MechanoreceptioninCaenorhabditis 10.9 CallosectomyandAfter............. 170
Elegans.............................. 99 10.10 ConcludingRemarks.............. 172
8.2 Spiders.............................. 103 Box10.1 BrocaandWernicke...... 166
8.2.1 TactileHairs.................... 103 PartII:SelfAssessment.................... 173
8.2.2 Trichobothria................... 103 PartII:Notes,ReferencesandBibliography. 179
8.2.3 SlitSensillaandLyriformOrgans. 104
8.3 Insects.............................. 106
8.3.1 AcousticSensillaandTympanic
Organs.......................... 109 PARTIII: CHEMOSENSITIVITY........ 187
8.4 TactileReceptorsinMammalianSkin 112
8.4.1 FastAdaptingReceptors......... 112 Chapter11 Chemosensitivityin
8.4.2 SlowAdaptingReceptors........ 114 Prokaryocytes............................. 189
8.5 CerebralAnalysisofTouch........... 114 11.1 ChemosentivityinE.coli............ 189
8.6 PlasticityoftheSomaestheticCortex. 118 11.1.1 MolecularGenetics............. 192
8.7 ConcludingRemarks................ 120 11.2 ConcludingRemarks............... 194
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Chapter12 MammalianChemo- 15.2.2 ThePinholeEyeofNautilus
Enteroreceptors............................ 195 Pompilius...................... 266
12.1 LocationofMammalian 15.2.3 TheMirrorEyeofPecten,
ChemoreceptorsforPaO andPaCO 196 theScallop..................... 267
2 2
12.2 Structure........................... 197 15.2.4 TheVesicularEyeofOctopus... 267
12.3 Physiology......................... 197 15.2.5 LateralEyesofLimulus,theKing
12.4 Biochemistry....................... 200 orHorseshoe‘Crab’............ 271
12.5 ConcludingRemarks............... 201 15.2.6 TheAdvancedOcellarEyesof
JumpingSpiders(Salticidae).... 275
Chapter13 Gustation...................... 203 15.2.7 The Advanced Compound Eyes
13.1 GustationinInsects................ 204 ofFlies(Diptera)............... 277
13.1.1 TachinidFlies.................. 204 15.3 ConcludingRemarks............... 279
13.1.2 Drosophila..................... 206 Box15.1 TheEvolutionofOpsins.. 254
13.2 GustationinMammals............. 208 Box15.2 EarlyGeneticsofEyes.... 263
13.2.1 TasteBuds..................... 208
13.2.2 CentralProjections............. 216 Chapter16 TheHumanEye............... 281
13.2.3 LabelledLinesorPopulation 16.1 Anatomy........................... 282
Profile?......................... 217 16.1.1 Eyeball........................ 282
13.3 ConcludingRemarks............... 217 16.1.2 Adnexa........................ 283
16.2 Embryology........................ 287
Chapter14 Olfaction...................... 219 16.3 DetailedAnatomyandPhysiology... 293
14.1 InsectOlfactorySystems............ 220 16.3.1 Cornea......................... 293
14.1.1 Hygroreceptors................. 220 16.3.2 Sclera.......................... 297
14.1.2 OlfactorySensilla.............. 221 16.3.3 Lens........................... 297
14.1.3 Central Processing in Drosophila 16.3.4 UvealTract.................... 303
AntennalLobes................ 224 16.3.5 Iris............................ 309
14.2 MammalianOlfactorySystems...... 225 16.3.6 VitreousHumour............... 312
14.2.1 OlfactoryEpitheliumoftheNasal 16.4 MovementsoftheEyeball .......... 313
Cavity......................... 225 16.5 ConcludingRemarks............... 314
14.2.2 OlfactoryNeurosensoryCells... 226 Box16.1 GeneticsofCataract...... 302
14.2.3 MolecularBiology.............. 227
Chapter17 TheRetina..................... 315
14.2.4 CentralProcessing.............. 230
17.1 RetinalPigmentEpithelium(RPE).. 316
14.3 The Vertebrate Vomeronasal Organ
17.2 Retina............................. 317
(VNO)andPheromones............ 232
17.2.1 PhotoreceptorCells............. 319
14.4 ConcludingRemarks............... 235
17.2.2 HorizontalCells................ 319
PartIII:SelfAssessment................... 237
17.2.3 BipolarCells................... 337
PartIII:Notes,ReferencesandBibliography 240
17.2.4 Mu¨llerCells.................... 340
17.2.5 InterplexiformCells............ 341
17.2.6 AmacrineCells................. 341
PARTIV: PHOTOSENSITIVITY......... 245 17.2.7 GanglionCells................. 341
BoxI4.1 Bacteriorhodopsin........ 249 17.2.8 WiringDiagrams............... 345
17.2.9 Colour......................... 348
Chapter15 InvertebrateVision............. 253
17.3 ConcludingRemarks............... 350
15.1 DesignsofInvertebrateEyes........ 253
Box17.1 RetinitisPigmentosa...... 330
15.1.1 EvolutionoftheVesicularEye.. 257
Box17.2 MacularDegeneration.... 342
15.1.2 EvolutionoftheCompoundEye 258
15.1.3 ScanningEyes.................. 261 Chapter18 VisualPathwaysandCortices... 358
15.2 ExamplesofInvertebrateEyes...... 262 18.1 VisualPathwaysintotheBrain...... 354
15.2.1 EyespotsoftheProtista......... 264 18.1.1 TheRetino-TectalPathway..... 354
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18.1.2 TheRetino-Geniculo-Striate Chapter21 MinoritySenses................ 423
(RGS)Pathway................. 355 21.1 InfraredRadiation.................. 424
18.2 PrimaryVisualCortex.............. 358 21.2 PolarizedLight..................... 425
18.2.1 Structure....................... 358 21.3 ElectricFields...................... 428
18.2.2 Functioning.................... 360 21.3.1 Fish........................... 428
18.2.3 Plasticity....................... 369 21.3.2 Monotremata.................. 432
18.3 ExtrastriateCortices................ 372 21.4 MagneticFields.................... 434
18.4 FaceRecognition................... 372 21.5 ConcludingRemarks............... 436
18.5 Prosopagnosia...................... 376
18.6 ConcludingRemarks............... 378 Chapter22 Pain........................... 437
Box18.1 TheRealityofCortical 22.1 TheBiologicalSignificanceofPain.. 438
Columns.......................... 367 22.2 NeurophysiologyofPain............ 440
Box18.2 Blindsight................ 374 22.2.1 NociceptorFibres.............. 440
22.2.2 NociceptorIonChannels....... 443
Chapter19 OtherVertebrateVisualSystems 379 22.2.3 MolecularBiology.............. 444
19.1 VisualPigments.................... 380 22.2.4 CentralPathways............... 446
19.2 Photoreceptors..................... 380 22.3 NeuropharmacologyofPain
19.3 Tapeta............................. 383 Pathways........................... 449
19.4 Retinae............................ 385 22.4 ReferredPain...................... 450
19.4.1 DeepSeaFish.................. 385 22.5 GateTheory....................... 451
19.4.2 Frog........................... 385 22.6 ConcludingRemarks............... 452
19.4.3 AreaeCentralesofMammalsand PartV:SelfAssessment .................... 455
Birds........................... 386 PartV:Notes,ReferencesandBibliography. 458
19.5 DioptricApparatus................. 388
PARTVI: CODA.......................... 463
19.6 MedianEyes....................... 391
19.7 VisualPathways.................... 393
Chapter23 SummingUp.................. 465
19.8 VisualCentresintheBrain.......... 394
23.1 MolecularThemes.................. 466
19.8.1 Amphibia...................... 394
23.2 CellularThemes.................... 468
19.8.2 Reptiles........................ 394
23.3 SenseOrgans....................... 469
19.8.3 Birds........................... 395
23.4 CentralAnalysers.................. 470
19.9 ConcludingRemarks............... 396
23.5 Homeostasis....................... 472
PartIV:SelfAssessment.................... 397
23.6 DifferentSensoryWorlds........... 473
PartIV:Notes,ReferencesandBibliography. 403
23.7 FromAbiotictoBiotic:
Communication.................... 474
PARTV: OTHERSENSES................ 413 23.8 FromBiotictoSocial
Communication:MirrorNeurons... 475
Chapter20 Thermosensitivity.............. 415 23.9 ConcludingRemarks............... 477
20.1 MolecularBiology.................. 416
20.1.1 CaenorhabditisElegans......... 416 Chapter24 PhilosophicalPostscript........ 479
20.1.2 DorsalRootGanglia(DRG).... 416 24.1 Descartes.......................... 479
20.1.3 MouseThermosensoryNeurons. 417 24.2 Qualia............................. 480
20.2 Poikilotherms...................... 417 24.3 TabulaRasa?....................... 481
20.3 Homeotherms...................... 418 24.4 EpigeneticEpistemology............ 482
20.3.1 ThermoreceptorsinTheSkin... 419 24.5 EvolutionaryEpistemology......... 485
20.3.2 DeepThermoreceptors......... 419 24.6 BeyondDescartes.................. 487
20.3.3 HypothalamicThermoreceptors. 420 24.7 ConcludingRemarks............... 490
20.4 ConcludingRemarks............... 421 Box24.1 SensorySubstitution..... 484
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PartVI:SelfAssessment................... 491 AcronymsandAbbreviations................ 501
PartVI:Notes,ReferencesandBibliography 493
Glossary................................... 505
Appendix:SomeTechniques................. 497 Index...................................... 511
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PREFACE TO SECOND EDITION
Eightyearshavepassedsincethefirsteditionofthis themolecularbiologyofpain,mirrorneuronsandso
text was published. Eight years in which the sub- on. One of the most interesting developments since
ject matter of sensory biology has seen huge devel- the first edition was published has been the Nobel-
opments. In particular, the molecular revolution of Prize-winning work on olfaction in mammals and
ourtimeshascontinuedapaceandculminatedinthe insects and the chapters on olfaction have been ex-
publicationofnotonlythehumangenomebutalso panded to take this work into account. In addition
those of many other living forms. Although to the anumberofnewBoxeshavebeenincorporatedinto
innocenteyethelivingworldseemstobepopulated the text: on the genetics of hearing loss; on macu-
byanalmostinfinitevarietyofformspossessingan lardegeneration;onthenomenclatureofgenesand
equallyunimaginablevarietyofsensorysystems,the proteins;ontherealityofcorticalcolumns.
explosive development of molecular biology shows As I wrote in the preface to the first edition, the
thatthereisadeepunderlyingunity.Inrevisingthis studyofsensorysystemscanformabridgebetween
book I have attempted to bring out this unity to theworldsofbiophysics,molecularbiologyandneu-
an even greater extent than in the first edition. Not rophysiologyandtheworldsofcognitivescienceand
onlyhaseachchapterbeenthoroughlyrevisedwith psychology.Myhoperemainsthattheoverviewpre-
thisinmindbutanadditionalchapter,Chapter5,on sented in the pages of this second edition, as in the
genesandgenomicshasbeeninserted.Otherwisethe first, can form a basis on which special studies can
overallorganizationofthebookremainslargelyun- be built. Although the book provides a wide sweep
changed.Themajorthemesremainasbefore:foun- through the world of animal sensory systems the
dationsinmolecularbiologyandbiophysics;astrong major focus remains on mammalian and especially
emphasisonevolutionarydevelopmentandcompar- on human systems. The ubiquity of the underlying
ativeanatomy;afocusonhumansystems;reference molecular biology often means that the molecular
tothe‘hardproblem’oftherelationofoursensory disordersresponsibleforhumandiseaseanddisabil-
experiencetotheanatomyandphysiologyofsensory ity can be studied in infrahuman species. Wherever
systems. possibletheopportunityistakentobringoutthese
Although the overall structure of the volume re- connections and to outline the molecular bases of
mains much as it was in thefirst edition,each page human sensory disorders and pathology. In spite
has been subjected to detailed revision and updat- of this focus on human sensory systems and their
ingandoverfortynewillustrationshavebeenadded. pathologies I hope that the wide coverage of other
Several new sections have been inserted. Amongst animalsbringswithitanothermessage:thatinstudy-
these are sections on transient receptor potential ingsensorysystemsandtheirfunctioningweshould
channels (TRP channels), synaptic transmission, bearinmindthathumansystemsarebutoneamong
evolutionofnervoussystems,arachnidmechanosen- many.
sitive sensilla and photoreceptors, electroreception Inconcluding,Icanonlyreiteratethethanksof-
intheMonotremata,languageandtheFOXP2gene; fered in the preface to the first edition to all those
xi