Biology of sensory systems

Since publication of the first edition, huge developments have taken place in sensory biology research and new insights have been provided in particular by molecular biology. These show the similarities in the molecular architecture and in the physiology of sensory cells across species and across sensory modality and often indicate a common ancestry dating back over half a billion years.

Biology of Sensory Systems has thus been completely revised and takes a molecular, evolutionary and comparative approach, providing an overview of sensory systems in vertebrates, invertebrates and prokaryotes, with a strong focus on human senses.

Written by a renowned author with extensive teaching experience, the book covers, in six parts, the general features of sensory systems, the mechanosenses, the chemosenses, the senses which detect electromagnetic radiation, other sensory systems including pain, thermosensitivity and some of the minority senses and, finally, provides an outline and discussion of philosophical implications.

New in this edition:

Greater emphasis on molecular biology and intracellular mechanisms New chapter on genomics and sensory systems Sections on TRP channels, synaptic transmission, evolution of nervous systems, arachnid mechanosensitive sensilla and photoreceptors, electroreception in the Monotremata, language and the FOXP2 gene, mirror neurons and the molecular biology of pain

Updated passages on human olfaction and gustation.

Over four hundred illustrations, boxes containing supplementary material and self-assessment questions and a full bibliography at the end of each part make Biology of Sensory Systems essential reading for undergraduate students of biology, zoology, animal physiology, neuroscience, anatomy and physiological psychology. The book is also suitable for postgraduate students in more specialised courses such as vision sciences, optometry, neurophysiology, neuropathology, developmental biology.

Praise from the reviews of the first edition:

"An excellent advanced undergraduate/postgraduate textbook." ASLIB BOOK GUIDE

"The emphasis on comparative biology and evolution is one of the distinguishing features of this self-contained book. .... this is an informative and thought-provoking text..." TIMES HIGHER EDUCATIONAL SUPPLEMENT

Author Notes

Dr. Christopher Upham Murray Smith . Honorary Visiting Fellow, Vision Sciences, Aston University, Birmingham, UK.

Preface to Second Edition	p. xi
Preface to First Edition	p. xiii
Part I Preliminaries	p. 1
Chapter 1 Elements	p. 3
1.1 Allosteric Effectors	p. 4
1.2 Membranes	p. 5
1.2.1 Lipids	p. 5
1.2.2 Proteins	p. 6
1.2.3 Mobility of Proteins	p. 8
1.3 Membrane Signalling Systems	p. 8
1.3.1 Receptor Molecules	p. 9
1.3.2 G-proteins	p. 10
1.3.3 Effectors and Second Messengers	p. 12
1.4 Channels and Gates	p. 14
1.4.1 TRP Channels	p. 14
1.4.2 Ligand-Gated Ion Channels (LGICs)	p. 16
1.4.3 Voltage-Gated Ion Channels (VGICs)	p. 16
1.5 Concluding Remarks	p. 18
Chapter 2 Membranes, Action Potentials, Synapses	p. 19
2.1 The Measurement of Resting Potentials	p. 20
2.2 The Ionic Bases of Resting Potentials	p. 21
2.3 Electrotonic Potentials and Cable Conduction	p. 23
2.4 Receptor and Generator Potentials	p. 24
2.5 Sensory Adaptation	p. 25
2.6 Action Potentials	p. 26
2.7 Synapses and Synaptic Transmission	p. 28
2.8 Concluding Remarks	p. 30
Chapter 3 General Features of Sensory Systems	p. 31
3.1 Classification of the Senses	p. 33
3.2 Modality	p. 34
3.3 Intensity	p. 34
3.4 Adaptation	p. 36
3.5 Receptive Fields	p. 36
3.6 Maps of Sensory Surfaces	p. 37
3.7 Hierarchical and Parallel Design	p. 38
3.8 Feature Extraction and Trigger Stimuli	p. 38
3.9 Concluding Remarks	p. 39
Box 3.1 Hermann von Helmholtz	p. 32
Chapter 4 Classification and Phylogeny	p. 41
4.1 Systematics	p. 41
4.2 Classification into Six Kingdoms	p. 42
4.3 Unicellularity	p. 42
4.4 Multicellularity	p. 42
4.5 Protostomes and Deuterostomes	p. 43
4.6 Classification of the Metazoa	p. 44
4.7 Evolution of Nervous Systems	p. 49
4.7.1 Cnidaria	p. 50
4.7.2 Platyhelminthes	p. 50
4.7.3 Nematoda	p. 50
4.7.4 Annelida	p. 51
4.7.5 Arthropoda	p. 52
4.7.6 Mollusca	p. 53
4.7.7 Echinodermata	p. 55
4.7.8 Chordata	p. 55
4.8 Concluding Remarks	p. 55
Chapter 5 Genes, Genomics and Neurosensory Systems	p. 57
5.1 Introduction	p. 57
5.2 Comparative Genomics	p. 60
5.3 Genomes and Neurosensory Systems	p. 62
5.4 Concluding Remarks	p. 64
Box 5.1 Nomenclature of Genes and Proteins	p. 59
Part I Self Assessment	p. 65
Part I Notes, References and Bibliography	p. 69
Part II Mechanosensitivity	p. 73
Chapter 6 Mechanosensitivity of Cell Membranes	p. 75
6.1 Mechanosensitive Channels in E. coli	p. 76
6.2 Detection of Osmotic Swelling by Hypothalamic Cells in Mammals	p. 80
6.3 Concluding Remarks	p. 83
Chapter 7 Kinaesthesia	p. 85
7.1 Kinaesthetic Mechanisms in Arthropods	p. 85
7.1.1 Stretch Receptors in Crustacean Muscle	p. 86
7.1.2 Insect Sensilla	p. 87
7.1.3 Maintenance of Equilibrium in Flight	p. 90
7.2 Kinaesthetic Mechanisms in Mammals	p. 91
7.2.1 Intrafusal Spindle Endings	p. 91
7.2.2 Golgi Tendon Organs	p. 94
7.2.3 Joint Receptors	p. 96
7.3 Concluding Remarks	p. 97
Chapter 8 Touch	p. 99
8.1 Mechanoreception in Caenorhabditis Elegans	p. 99
8.2 Spiders	p. 103
8.2.1 Tactile Hairs	p. 103
8.2.2 Trichobothria	p. 103
8.2.3 Slit Sensilla and Lyriform Organs	p. 104
8.3 Insects	p. 106
8.3.1 Acoustic Sensilla and Tympanic Organs	p. 109
8.4 Tactile Receptors in Mammalian Skin	p. 112
8.4.1 Fast Adapting Receptors	p. 112
8.4.2 Slow Adapting Receptors	p. 114
8.5 Cerebral Analysis of Touch	p. 114
8.6 Plasticity of the Somaesthetic Cortex	p. 118
8.7 Concluding Remarks	p. 120
Chapter 9 Equilibrium and Hearing: The Uses of Hair Cells	p. 123
9.1 Anatomy and Physiology of Hair Cells	p. 123
9.2 Lateral Line Canals	p. 128
9.3 Evolution of the Vertebrate Ear	p. 129
9.3.1 Equilibrium	p. 129
9.3.2 Phonoreception	p. 132
9.3.3 Bat Sonar	p. 145
9.4 Concluding Remarks	p. 145
Box 9.1 Biophysics of Outer Hair Cells	p. 141
Box 9.2 Genetics and Deafness	p. 143
Chapter 10 Cerebral Analysis	p. 147
10.1 The Mammalian Vestibular Pathway and Reflexes	p. 148
10.2 The Mammalian Auditory Pathway	p. 150
10.2.1 Cochlear Fibres	p. 151
10.2.2 Cochlear Nucleus	p. 152
10.2.3 Superior Olivary Nuclei	p. 153
10.2.4 Inferior Colliculus	p. 153
10.2.5 Medial Geniculate Nucleus	p. 153
10.3 The Avian Auditory Pathway and the Mapping of Auditory Space by the Barn Owl	p. 153
10.4 The Mammalian Auditory Cortex	p. 157
10.5 The Bat Auditory System and Echolocation	p. 158
10.6 The Human Auditory Cortex and Language	p. 162
10.7 Lateralization and the Neuroanatomy of Language	p. 167
10.8 Language and the FOXP2 Gene	p. 169
10.9 Callosectomy and After	p. 170
10.10 Concluding Remarks	p. 172
Box 10.1 Broca and Wernicke	p. 166
Part II Self Assessment	p. 173
Part II Notes, References and Bibliography	p. 179
Part III Chemosensitivity	p. 187
Chapter 11 Chemosensitivity in Prokaryocytes	p. 189
11.1 Chemosentivity in E. coli	p. 189
11.1.1 Molecular Genetics	p. 192
11.2 Concluding Remarks	p. 194
Chapter 12 Mammalian Chemo-Enteroreceptors	p. 195
12.1 Location of Mammalian Chemoreceptors for PaO 2 and PaCO 2	p. 196
12.2 Structure	p. 197
12.3 Physiology	p. 197
12.4 Biochemistry	p. 200
12.5 Concluding Remarks	p. 201
Chapter 13 Gustation	p. 203
13.1 Gustation in Insects	p. 204
13.1.1 Tachinid Flies	p. 204
13.1.2 Drosophila	p. 206
13.2 Gustation in Mammals	p. 208
13.2.1 Taste Buds	p. 208
13.2.2 Central Projections	p. 216
13.2.3 Labelled Lines or Population Profile?	p. 217
13.3 Concluding Remarks	p. 217
Chapter 14 Olfaction	p. 219
14.1 Insect Olfactory Systems	p. 220
14.1.1 Hygroreceptors	p. 220
14.1.2 Olfactory Sensilla	p. 221
14.1.3 Central Processing in Drosophila Antennal Lobes	p. 224
14.2 Mammalian Olfactory Systems	p. 225
14.2.1 Olfactory Epithelium of the Nasal Cavity	p. 225
14.2.2 Olfactory Neurosensory Cells	p. 226
14.2.3 Molecular Biology	p. 227
14.2.4 Central Processing	p. 230
14.3 The Vertebrate Vomeronasal Organ (VNO) and Pheromones	p. 232
14.4 Concluding Remarks	p. 235
Part III Self Assessment	p. 237
Part III Notes, References and Bibliography	p. 240
Part IV Photosensitivity	p. 245
Box 14.1 Bacteriorhodopsin	p. 249
Chapter 15 Invertebrate Vision	p. 253
15.1 Designs of Invertebrate Eyes	p. 253
15.1.1 Evolution of the Vesicular Eye	p. 257
15.1.2 Evolution of the Compound Eye	p. 258
15.1.3 Scanning Eyes	p. 261
15.2 Examples of Invertebrate Eyes	p. 262
15.2.1 Eyespots of the Protista	p. 264
15.2.2 The Pinhole Eye of Nautilus Pompilius	p. 266
15.2.3 The Mirror Eye of Pecten, the Scallop	p. 267
15.2.4 The Vesicular Eye of Octopus	p. 267
15.2.5 Lateral Eyes of Limulus, the King or Horseshoe 'Crab'	p. 271
15.2.6 The Advanced Ocellar Eyes of Jumping Spiders (Salticidae)	p. 275
15.2.7 The Advanced Compound Eyes of Flies (Diptera)	p. 277
15.3 Concluding Remarks	p. 279
Box 15.1 The Evolution of Opsins	p. 254
Box 15.2 Early Genetics of Eyes	p. 263
Chapter 16 The Human Eye	p. 281
16.1 Anatomy	p. 282
16.1.1 Eyeball	p. 282
16.1.2 Adnexa	p. 283
16.2 Embryology	p. 287
16.3 Detailed Anatomy and Physiology	p. 293
16.3.1 Cornea	p. 293
16.3.2 Sclera	p. 297
16.3.3 Lens	p. 297
16.3.4 Uveal Tract	p. 303
16.3.5 Iris	p. 309
16.3.6 Vitreous Humour	p. 312
16.4 Movements of the Eyeball	p. 313
16.5 Concluding Remarks	p. 314
Box 16.1 Genetics of Cataract	p. 302
Chapter 17 The Retina	p. 315
17.1 Retinal Pigment Epithelium (RPE)	p. 316
17.2 Retina	p. 317
17.2.1 Photoreceptor Cells	p. 319
17.2.2 Horizontal Cells	p. 319
17.2.3 Bipolar Cells	p. 337
17.2.4 Müller Cells	p. 340
17.2.5 Interplexiform Cells	p. 341
17.2.6 Amacrine Cells	p. 341
17.2.7 Ganglion Cells	p. 341
17.2.8 Wiring Diagrams	p. 345
17.2.9 Colour	p. 348
17.3 Concluding Remarks	p. 350
Box 17.1 Retinitis Pigmentosa	p. 330
Box 17.2 Macular Degeneration	p. 342
Chapter 18 Visual Pathways and Cortices	p. 358
18.1 Visual Pathways into the Brain	p. 354
18.1.1 The Retino-Tectal Pathway	p. 354
18.1.2 The Retino-Geniculo-Striate (RGS) Pathway	p. 355
18.2 Primary Visual Cortex	p. 358
18.2.1 Structure	p. 358
18.2.2 Functioning	p. 360
18.2.3 Plasticity	p. 369
18.3 Extrastriate Cortices	p. 372
18.4 Face Recognition	p. 372
18.5 Prosopagnosia	p. 376
18.6 Concluding Remarks	p. 378
Box 18.1 The Reality of Cortical Columns	p. 367
Box 18.2 Blindsight	p. 374
Chapter 19 Other Vertebrate Visual Systems	p. 379
19.1 Visual Pigments	p. 380
19.2 Photoreceptors	p. 380
19.3 Tapeta	p. 383
19.4 Retinae	p. 385
19.4.1 Deep Sea Fish	p. 385
19.4.2 Frog	p. 385
19.4.3 Areae Centrales of Mammals and Birds	p. 386
19.5 Dioptric Apparatus	p. 388
19.6 Median Eyes	p. 391
19.7 Visual Pathways	p. 393
19.8 Visual Centres in the Brain	p. 394
19.8.1 Amphibia	p. 394
19.8.2 Reptiles	p. 394
19.8.3 Birds	p. 395
19.9 Concluding Remarks	p. 396
Part IV Self Assessment	p. 397
Part IV Notes, References and Bibliography	p. 403
Part V Other Senses	p. 413
Chapter 20 Thermosensitivity	p. 415
20.1 Molecular Biology	p. 416
20.1.1 Caenorhabditis Elegans	p. 416
20.1.2 Dorsal Root Ganglia (DRG)	p. 416
20.1.3 Mouse Thermosensory Neurons	p. 417
20.2 Poikilotherms	p. 417
20.3 Homeotherms	p. 418
20.3.1 Thermoreceptors in The Skin	p. 419
20.3.2 Deep Thermoreceptors	p. 419
20.3.3 Hypothalamic Thermoreceptors	p. 420
20.4 Concluding Remarks	p. 421
Chapter 21 Minority Senses	p. 423
21.1 Infrared Radiation	p. 424
21.2 Polarized Light	p. 425
21.3 Electric Fields	p. 428
21.3.1 Fish	p. 428
21.3.2 Monotremata	p. 432
21.4 Magnetic Fields	p. 434
21.5 Concluding Remarks	p. 436
Chapter 22 Pain	p. 437
22.1 The Biological Significance of Pain	p. 438
22.2 Neurophysiology of Pain	p. 440
22.2.1 Nociceptor Fibres	p. 440
22.2.2 Nociceptor Ion Channels	p. 443
22.2.3 Molecular Biology	p. 444
22.2.4 Central Pathways	p. 446
22.3 Neuropharmacology of Pain Pathways	p. 449
22.4 Referred Pain	p. 450
22.5 Gate Theory	p. 451
22.6 Concluding Remarks	p. 452
Part V Self Assessment	p. 455
Part V Notes, References and Bibliography	p. 458
Part VI Coda	p. 463
Chapter 23 Summing Up	p. 465
23.1 Molecular Themes	p. 466
23.2 Cellular Themes	p. 468
23.3 Sense Organs	p. 469
23.4 Central Analysers	p. 470
23.5 Homeostasis	p. 472
23.6 Different Sensory Worlds	p. 473
23.7 From Abiotic to Biotic: Communication	p. 474
23.8 From Biotic to Social Communication: Mirror Neurons	p. 475
23.9 Concluding Remarks	p. 477
Chapter 24 Philosophical Postscript	p. 479
24.1 Descartes	p. 479
24.2 Qualia	p. 480
24.3 Tabula Rasa?	p. 481
24.4 Epigenetic Epistemology	p. 482
24.5 Evolutionary Epistemology	p. 485
24.6 Beyond Descartes	p. 487
24.7 Concluding Remarks	p. 490
Box 24.1 Sensory Substitution	p. 484
Part VI Self Assessment	p. 491
Part VI Notes, References and Bibliography	p. 493
Appendix: Some Techniques	p. 497
Acronyms and Abbreviations	p. 501
Glossary	p. 505
Index	p. 511

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