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COMPARATIVE VERTEBRATE NEUROANATOMY: EVOLUTION AND ANATOMY, 2nd Edition
By: Ann B. Butler and William Hoods Jon Wiley & Sons, Inc. FROM THE PUBLISHER The Second Edition of this landmark text presents a broad survey of comparative vertebrate neuroanatomy at the introductory level, representing a unique contribution to the field of evolutionary neurobiology. It has been extensively revised and updated, with substantially improved figures and diagrams that are used generously throughout the text. Through analysis of the variation in brain structure and function between major groups of vertebrates, readers can gain insight into the evolutionary history of the nervous system. The text is divided into three sections: 1) Introduction to evolution and variation, including a survey of cell structure, embryological development, and anatomical organization of the central nervous system; phylogeny and diversity of brain structures; and an overview of various theories of brain evolution, 2) Systematic, comprehensive survey of comparative neuroanatomy across all major groups of vertebrates, 3) Overview of vertebrate brain evolution, which integrates the complete text, highlights diversity and common themes, broadens perspective by a comparison with brain structure and evolution of invertebrate brains, and considers recent data and theories of the evolutionary origin of the brain in the earliest vertebrates, including a recently proposed model of the origin of the brain in the earliest vertebrates that has received strong support from newly discovered fossil evidence Ample material drawn from the latest research has been integrated into the text and highlighted in special feature boxes, including recent views on homology, cranial nerve organization and evolution, the relatively large and elaborate brains of birds in correlation with their complex cognitive abilities, and the current debate on forebrain evolution across reptiles, birds, and mammals. Comparative Vertebrate Neuroanatomy is geared to upper-level undergraduate and graduate students in neuroanatomy, but anyone interested in the anatomy of the nervous system and how it corresponds to the way that animals function in the world will find this text fascinating. ABOUT THE AUTHORS Ann B. Butler, Professor Kranow Institute for Advanced Study and Department of Psychology George Mason University Fairfax, Virginia William Hodos, Distinguished University Professor Department of Psychology University of Maryland College Park, Maryland
CONTENTS
Preface xv
Acknowledgments xix
List of Boxes xxi
Part One
EVOLUTION AND THE ORGANIZATION OF THE CENTRAL NERVOUS SYSTEM
1 Evolution and Variation, 3
Introduction, 3
Diversity Over Time, 4
Evolutionary Mechanisms, 5
Genetic Factors, 5
Natural Selection, 5
Evolution of the Vertebrate Central Nervous System, 7
Sameness and Its Biological Significance, 8
Analogy, 8
Historical Homology, 8
Homoplasy, 9
Biological Homology, 11
Generative Homology or Syngeny, 12
Analysis of Variation, 13
Cladistic Analysis, 13
Parsimony, 14
Tests of Homology, 15
A Word of Caution, 15
Reconstructing Evolution, 16
2 Neurons and Sensory Receptors
Introduction, 19
The Nervous System, 19
Neurons and Sensory Receptors, 20
Transport Within Neurons, 21
Classification of Neurons, 21
Somata, 21
Dendrites, 21
Axons, 23
Synapses, 23
Chemical Synapses, 23
Neuroactive Substances, 24
Electrical Synapses, 26
Volume Transmission, 26
Neuronal Populations, 26
Golgi Type I and II Cells, 26
Nuclei and Planes of Section, 27
Techniques forTracing Connections Between Nuclei, 27
Receptors and Senses, 28
How Many Senses? 29
Receptors and Awareness, 29
Sensory Experience as a Private Mental Event, 30
Sensory Adaptation, 30
Receptor Types, 30
Mechanoreceptors, 3 I
Radiant-Energy Receptors, 34
Chemoreceptors, 37
Nervus Terminalis: An Unclassified Receptor, 41
Electroreceptors, 41
Nociceptors, 42
Magnetoreceptors, 43
Topographic Organization, 43
Receptive Fields, 46
The Senses and Evolution of the Central Nervous System, 46
3 The Vertebrate Central Nervous System, 49
Introduction, 49
Development of the Brain, 49
Segmental Development of the Vertebrate Brain, 50
Neurogenesis and Migration of Neurons, 54
Cortices and Nuclei, 55
Differing Patterns of Development, 57
Ontogeny and Recapitulation, 60
The Brain and Spinal Cord, 61
Cellular Organization of the Central Nervous System, 61
Regional Organization of the Nervous System, 63
The Spinal Cord, 63
The Brain, 63
The Meninges and the Ventricular System, 66
Major Systems of the Brain, 67
Sensory Systems, 68
Motor Systems, 68
Nomenclature of the Brain, 69
4 Vertebrate Phylogeny and Diversity in Brain Organization, 73
Introduction, 73
Vertebrate Phylogeny, 74
Chordate Relationships, 74
lawless Vertebrates, 74
Chrondrichthyes, 76
Actinopterygii, 76
Sarcopterygii, 76
The Big Picture of Vertebrate Evolution, 84
Two Types of Brain Organization, 84
Laminar Brains (Group 1), 86
Elaborated Brains (Group 11), 87
Glia and Brain Elaboration, 89
Laminar and Elaborated Brains across Evolution, 89
5 Evolution and Adaptation of the Brain, Behavior, and Intelligence, 93
Phylogeny and Adaptation, 93
Phyletic Studies, 93
Adaptation Studies, 94
Phylogenetic Scale, 95
The Phylogenetic Tree, 95
Complexity and Evolution, 96
Anagenesis, 97
Grades of Evolutionary Advancement, 99
Evolutionary Change, 99
Brain Evolution and Behavioral Adaptation, 100
Brain Size and Brain Allometry, 100
Brain Size and Behavioral Adaptation, 105
Brain Size and Intelligence, 106
What Is Intelligence? 108
Summary and Conclusions, 109
6 Theories of Brain Evolution
Introduction, 113
Some Common Assumptions, 113
Previous Theories of Vertebrate Brain Evolution: Addition of Structures or Areas, 114
Maclean, 114
Flechsig and Campbell, 114
Sanides, 115
Previous Theories of Vertebrate Brain Organization: New Formation and Reorganization of Circuits, 115
Herrick, 115
Bishop, 115
Ariens Kappers, 115
Bowsher, 115
Diamond and Hall, 116
Critique of Previous Theories of Vertebrate Brain Evolution, 116
Parcellation Theory, 117
Ebbesson, 117
Deacon, 117
Current Theories of Forebrain Evolution, 117
Forebrain Evolution: Experimental Foundations, 117
Karten: Equivalent Cell Hypothesis, 118
Other Theories of Pallial Evolution, 119
Perspective, 121
Part Two
THE SPINAL CORD AND HINDBRAIN
7 Overview of Spinal Cord and Hindbrain, 127
Overview of the Spinal Cord, 127
Segmentation Within the Spinal Cord, 127 Roots and Ganglia, 128
Columns of the Spinal Cord, 129
Pathways Within the Spinal Cord, 130 Reflexes, 131
Spinal Autonomy, 133
Rhythmic Movements and Central Pattern Generators, 133
Overview of the Hindbrain, 133
The Obex and the Fourth Ventricle, 135
The Pontine Nuclei, 135
Ganglia of the Cranial Nerves, 135
Organization of the Cranial Nerves, 135
Embryology of the Hindbrain and a New Classification of Cranial Nerves, 135
Efferent Axons in Afferent Nerves, 136
Evolutionary Perspectives on the Spinal Cord and Hindbrain, 136
The Transition to Land, 136
Tetrapod Locomotor Patterns, 137
8 The Spinal Cord, 139
The Spinal Cords of Nontetrapods, 139
Muscles and Locomotion, 139
Cell and Fiber Columns, 139
Giant Axons and Escape, 141
Electromotor Neurons, 144
The Curious Spinal Cords of Sharks, 144
Ascending and Descending Pathways, 144
Reissner's Fiber, 145
The Organization of the Tetrapod Spinal Cord, 145
Locomotor Patterns and Spinal Cord Organization, 145
The Curious Spinal Cords of Birds, 146
Segmental Organization, 147
Lamination, 147
Intrinsic Spinal Neurons, 148
Somatotopic Organization of the Ventral Horns, 148
Renshaw Cells, 149
Axon Columns and Cell Columns, 149
Marginal Cells, 150
Accessory Lobes, 150
Ascending Spinal Pathways, 150
Descending Spinal Pathways, 150
Tetrapod Central Pattern Generators, 152
Evolutionary Perspective, 152
9 Segmental Organization of the Head, Brain, and Cranial Nerves, 157
"Twelve" Cranial Nerves, 157
The Vertebrate Head: Segmental Organization, 158
Head Skeleton, 159
The Striated Musculature of the Head, 159
Neural Crest and Placodes, 162
Segmentation of the Head, 164
Theoretical Head Segments, 165
Segmental Organization of the Individual Cranial Nerves, 166
The Forebrain, 168
The First Head Segment, 168
The Second Head Segment, 169
The Third Head Segment, 169
The Fourth Head Segment, 169
The Fifth Head Segment, 170
10 Functional Organization of the Cranial Nerves, 173
Introduction, 173
The Cranial Nerves and the Spinal Cord, 173
The Organization of Sensory and Motor Columns of the Caudal Brainstem, 176
Afferent Columns of the Brainstem, 177
Efferent Columns of the Brainstem, 179
Five Cranial Nerves Rostral to the Brainstem, 180
General Considerations, 181
11 Sensory Cranial Nerves of the Brainstem, 183
Introduction, 183
Dorsal Cranial Nerves: Sensory Components for General Somatosensory Sensation, 183
Somatosensory Innervation of the Head, 184
Central Terminations of the Trigeminal Nerve, 185
The Mesencephalic Division of the Trigeminal System, 185
Secondary Connections of the Trigeminal Nuclei, 186
Ventrolateral Placodal Cranial Nerves: Taste, 189
The Gustatory System, 190
The Gustatory Nerves and the Nucleus Solitarius, 190
Secondary Connections of the Gustatory Nucleus and Nucleus Solitarius, 190
Cyprinid and Silurid Gustatory Specializations, 192
Dorsolateral Cranial Nerves: Lateral Line and Octaval Systems, 194
The Lateral Line System, 195
The Octaval System, 196
12 Motor Cranial Nerves, 205
Introduction, 205
Feeding and Swallowing, 207
The Neural Control of Feeding and Swallowing, 209
The Communication Systems of Fishes, 211
The Acoustic Reflex, 213
Motor Control of Eye Muscles, 214
The Extraocular Muscles in lawless Vertebrates, 214
The Extraocular Muscles in ]awled Vertebrates, 214 The Intraocular Muscles, 215
Central Control of the Eye Muscles, 215
The Oculomotor Complex, 217
Coordination of Eye Muscle Action, 218
Evolutionary Perspective on the Hindbrain and Midbrain Cranial Nerves, 218
13 The Reticular Formation, 221
Introduction, 221
The Organization of the Reticular Formation, 222
Neurons of the Reticular Formation, 222
Giant Reticulospinal Neurons, 223
Nomenclature of the Reticular Formation, 224
The Reticular Formation of the Medulla, Pons, and Midbrain, 225
The Reticular Formation of the Diencephalon, 228
Pathways of the Reticular Formation, 230
Chemical Pathways of the Reticular Formation, 232
The Reticular Formation and Sleep, 234
Evolutionary Perspective on the Reticular Formation, 236
14 The Cerebellum, 241
Introduction, 241
Overview of the Cerebellum, 241
Cerebellar Size, 242
The Various Forms of the Cerebellum, 243
Corpus Cerebelli, 243
Electroreception and the Cerebellum, 244
The Cerebellar Auricle, 245
Phyletic Development of the Form of the Cerebellum, 245
The Cerebella of Tetrapods, 246
The Cerebella of Nontetrapods, 246
Agnathans and Cartilaginous Fishes, 246
Ray-Finned Fishes, 246
The Cerebellar Cortex, 247
The Purkinje Cell Layer, 247
The Granule Cell Layer, 249
The Molecular Layer, 253
Afferent Inputs to the Cerebellar Cortex, 253
Interconnections Within the Cerebellar Cortex, 255
The Precerebellar Nuclei, 256
Cerebelloid Structures Associated With the Cerebellum in
Nontretapods, 257
The Exceptional Cerebella of Weakly Electric Fishes, 259
Cerebellar Efferents and the Deep Cerebellar Nuclei, 260
Evolutionary Perspective, 262
Functions of the Cerebellum, 262
Part Three
THE MIDBRAIN
15 Overview of the Midbrain, 267
Introduction, 267
The Isthmus, 268
The Tegmentum, 273
The Tectum, 273
16 Isthmus, 275
Introduction, 275
Nuclei of the Raphe, 275
Group I, 275
Group II, 275
Evolutionary Perspective, 277
Locus Coeruleus, 277
Group I, 277
Group II, 277
Evolutionary Perspective, 278
Nucleus Isthmi, 281
Group I, 281
Group II, 281
Evolutionary Perspective, 283
Isthmo-Optic Nucleus, 283
Group I, 283
Group II, 283
Evolutionary Perspective, 284
Midbrain Locomotor Region and Pedunculopontine Tegmental Nucleus, 284
Group I, 284
Group II, 284
Evolutionary Perspective, 285
\Interpeduncular Nucleus, 285
Group I, 285
Group II, 285
Evolutionary Perspective, 285
17 Tegmentum and Tori, 289
Introduction, 289
Mesencephalic Nucleus of the Trigeminal Nerve, 289
Group I, 289
Group II, 290
Evolutionary Perspective, 290
Red Nucleus and Related Nuclei, 290
Group I, 290
Group II, 290
Evolutionary Perspective, 292
Substantia Nigra and Ventral Tegmental Area, 292
Group I, 293
Group II, 294
Evolutionary Perspective, 303
Torus Lateralis, 304
Group I, 304
Group II, 304
Evolutionary Perspective, 304
Torus Semicircularis, 304
Group I, 304
Group II, 305
Evolutionary Perspective, 306
18 Optic Tectum, 311
Introduction, 311
Overview of Tectal Organization, 311
Overview ofTectal Connections, 312
The Optic Tectum in Group I Vertebrates, 315
Lampreys, 315
Squalomorph Sharks and Ratfishes, 316
Nonteleost Ray-Finned Fishes, 316
Amphibians, 319
The Optic Tectum in Group II Vertebrates, 321
Hagfishes, 321
Galeomorph Sharks, Skates, and Rays, 322
Teleosts, 322
Amniotes, 325
Evolutionary Perspective, 334
Part Four
THE FOREBRAIN: DIENCEPHALON
19 Overview of the Forebrain, 343
Introduction, 343
Nomenclature of the Forebrain in Amniotes, 344
The Diencephalon, 344
Pretectum, 344
Posterior Tuberculum, 344
Epithalamus, 345
Dorsal Thalamus, 346
Ventral Thalamus, 347
Hypothalamus and Preoptic Area, 352
The Telencephalon: Pallium, 352
The Telencephalic Pallium of Mammals, 353
The Telencephalic Pallium of Nonmammalian Amniotes, 361
The Telencephalon: Subpallium, 364
The Ventrolateral Telencephalon of Anamniotes, 364
The Ventrolateral Telencephalon of Mammals, 364
The Ventrolateral Telencephalon of Nonmammalian Vertebrates, 368
The Septum, 369
20 Pretectum, Accessory Optic System, and Migrated Posterior
Tuberculum, 373
Introduction, 373
Pretectum, 373
Group I, 374
Group II, 375
Accessory Optic System, 389
Group I, 391
Group II, 392
Evolutionary Perspective, 394
Migrated Posterior Tuberculum, 396
Group I, 396
Group II, 396
Evolutionary Perspective, 399
21 Epithalamus, 407
Introduction, 407
Epiphysis, 407
Habenula, 409
Evolutionary Perspective, 414
22 Dorsal Thalamus, 417
Introduction, 417
Collothalamic Auditory System, 418
Group I, 418
Group IIA, 418
Group IIB, 420
Collothalamic Visual and Somatosensory Systems, 426
Group I, 427
Group IIA, 430
Group IIB, 430
Lemnothalamus, 432
Group I, 432
Group IIA, 434
Group IIB, 434
Evolutionary Perspective, 437
Collothalamus,437
Lemnothalamus,437
A New Definition of the Dorsal Thalamus in Vertebrates, 439
23 The Visceral Brain: The Hypothalamus and the Autonomic Nervous
System, 445
Introduction, 445
The Hypothalamus, 445
The Hypothalamus and the Endocrine System, 446
Circumventricular Organs, 449
Biological Rhythms, the Epiphysis, and the Hypothalamus, 449
The Hypothalamus and the Limbic System, 450
The Preoptic Area, 450
The Hypothalamus in Anamniotes, 451
Jawless Fishes, 451
Cartilaginous Fishes, 451
Actinopterygians, 451
Sarcopterygians, 455
The Hypothalamus in Amniotes, 455
Connections of the Hypothalamus in Reptiles and Birds, 456
Connections of the Hypothalamus in Mammals, 457
Functions of the Hypothalamus, 460
The Autonomic Nervous System, 460
Autonomic Neurochemistry, 462
Amniotes, 462
Anamniotes, 462
Evolutionary Perspective, 462
Part Five
THE FOREBRAIN: TELENCEPHALON
24 Basal Telencephalon, 471
Introduction, 471
The Striatopallidal Complexes, 472
Group I, 472
Group IIA, 476
Group lIB, 477
The Striatal Amygdala, 487
Cholinergic Neuronal Populations of the Basal Telencephalon, 488
Evolutionary Perspective, 489
25 Nonlimbic Pallium, 495
Introduction, 495
The Nonlimbic Pallium in Group I Vertebrates, 496
The Nonlimbic Pallium in Group IIA Vertebrates, 498
Neuroanatomical Organization, 498
Behavioral Issues, 501
The Nonlimbic Pallium in Amniotes, 501
Mammals: Neocortex, 501
Mammals: Claustrum-Endopiriform Formation and Frontotemporal Amygdala, 504
Reptiles and Birds, 504
Ascending Sensory Pathways to the Pallium in Amniotes, 507
Pallial Evolution and Persistent Questions of Homologies, 510
26 Visual Forebrain in Amniotes, 523
Introduction, 523
Ipsilateral Retinal Pathways and Stereoscopic
Vision, 524
Visual Pathways to the Telencephalon in Mammals, 524
Lemnothalamic Visual Forebrain, 524
Collothalamic Visual Forebrain, 536
Pathways to the Visual Telencephalon in Reptiles and Birds, 537
Lemnothalamic Visual Pathways, 538
Collothalamic Visual Pathways, 540
Evolutionary Trends in the Visual System of
Amniotes, 540
27 Somatosensory and Motor Forebrain in Amniotes, 547
Introduction, 547
The Somatosensory and Motor Forebrain of Mammals, 547
The Ventral Tier Nuclei of the Dorsal Thalamus, 548
Somatosensory Lemnothalamus, 548
Somatosensory Collothalamus, 548
Motor Lemnothalamus, 548
Afferents to Somatosensory Cortex, 549
Efferents of Somatosensory Cortex, 549
Pain Pathways, 549
Somatotopic Organization, 552
Motor Cortex, 557
Multiple Motor Representations of the Body, 558
The Cortical Eye Fields, 558
Afferents and Efferents of the Motor Cortex, 558
The Somatosensory and Motor Forebrain of Nonmammalian Amniotes, 559
Somatosensory System, 559
Motor System, 564
Evolutionary Perspective, 566
28 Auditory and Vocal Forebrain in Amniotes, 571
Introduction, 571
Location of Sound Sources, 571
Echolocation, 572
Auditory Channels for Time and Intensity, 573
Design Features of the Auditory System, 574
Topographic Organization, 574
Bilateral Interaction in the Auditory Pathway, 574
Descending Auditory Pathways, 574
Auditory Pathways in Tetrapods, 574
Auditory Telencephalon, 577
Columnar Organization, 577
Mammals, 577
Reptiles and Birds, 579
Vocal Telencephalon, 580
Vocalization and Hearing, 581
Anurans, 582
Reptiles and Birds, 583
Mammals, 587
Evolutionary Perspective, 589
29 Terminal Nerve and Olfactory Forebrain, 593
Introduction, 593
Olfactory System, 593
Group I, 594
Group II, 595
Vomeronasal System, 601
Terminal Nerve, 605
Evolutionary Perspective, 606
30 Limbic Telencephalon, 611
Introduction, 611
The Limbic Pallium in Anamniotes, 612
Group I, 612
Group IIA, 6 I3
The Limbic Pallium in Amniotes (Group lIB), 617
Limbic Pallium of Mammals, 619
Limbic Pallium in Nonmammalian Amniotes, 623
Limbic Subpallium: Septum, 628
Evolutionary Perspective, 629
Part Six
CONCLUSION
31 Evolution of Brains: A Bilaterian View, 637
Introduction, 637
Invertebrate Brains and the Inversion Hypothesis, 638
Insect Brain Organization, 639
Urbilateria and the Ancestral Condition of Bilaterian
Brains, 641
Deuterostomes and Dorsoventral Inversion, 641
Brain Evolution within Chordates, 644
The Origin of Vertebrates, 649
Haikouella, 650
Sensory System Evolution in the Vertebrate Lineage, 652
Organization of the Vertebrate Brain, 653
The Advent of Jaws, 655
Onto the Land and Into the Air, 656
Theories of Vertebrate Brain Evolution, 657
How Vertebrate Brains Evolve, 657
Appendix: Terms Used in Neuroanatomy, 665
Introduction, 665
Direction and Location Terms, 665
Planes of Section, 666
Neuroanatomical Names, 668
Derivation of Terms, 668
Glossary, 671
Index, 679 WHERE TO ORDER ISBN number 0471210056 ($125.00 hardcover) John Wiley & Sons, Inc. 111 River Street Hoboken, NJ 07030 Website: http://www.wiley.com/WileyCDA/ Direct link to catalog entry: http://www.wiley.com/WileyCDA/WileyTitle/productCd-0471210056.html Posted Date: 10/26/05
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