Section I Orientation, Development, Gross Anatomy, Blood Supply and Meninges... 1 1 Orientation... 3 References... 3 2 Development... 7 Early Morphogenesis... 7 FormationoftheBrainRegions... 9 Histogenesis... 16 Fundamental Morphological Pattern... 24 SpinalCord... 26 Hindbrain... 29 Cerebellum... 32 Midbrain... 39 Forebrain... 41 Diencephalon... 44 Telencephalon... 48 New Insights into the Development of Forebrain..... 53 References... 59 3 Gross Anatomy... 67 References... 71 4 Blood Supply, Meninges and Cerebrospinal Fluid Circulation... 95 Introduction... 95 Arteries of the Brain... 95 Meninges, Cisterns and Cerebrospinal Fluid Circulation..... 97 Circumventricular Organs... 99 Veins of the Brain... 99 Vessels and Meninges of the Spinal Cord..... 100 References... 101
VIII 5 Brain Slices... 137 Introduction... 137 CoronalSections... 138 Sections Perpendicular to the Axis of the Brain Stem... 153 Sagittal Sections... 158 HorizontalSections... 165 Section II Structure of Spinal Cord and Brain Parts... 175 6 Topography of Spinal Cord, Brain Stem and Cerebellum... 177 SpinalCord... 177 Introduction... 177 The Grey Matter of the Cord... 178 Primary Afferents... 180 TheWhiteMatteroftheCord... 182 Topography of the Brain Stem and the Cerebellum... 190 Introduction... 190 Cranial Nerve Nuclei in the Brain Stem... 190 Medulla Oblongata... 191 Metencephalon and the Cerebellum... 198 Mesencephalon... 204 Addendum: Diencephalon and the Corpus Striatum... 209 References... 210 7 Diencephalon: Introduction and Epithalamus... 247 Introduction... 247 Epithalamus... 247 References... 250 8 Diencephalon: Dorsal Thalamus... 253 IntroductoryNotes... 253 Parcellation of the Thalamus... 255 Thalamic and Thalamocortical Circuits... 259 Ventral Nuclear Group... 262 Anterior Nuclear Group and Lateral Dorsal Nucleus... 263 Mediodorsal Nucleus... 264 Medial Geniculate Body... 264 Lateral Geniculate Body... 265 Lateral Nuclear Group... 265 Posterior Nuclear Complex... 266 Intralaminar and Midline Nuclei... 266 FunctionalAspects... 268 References... 271
IX 9 Diencephalon: Ventral Thalamus or Subthalamus... 281 IntroductoryNotes... 281 Subthalamic Cell Masses... 281 Subthalamic Fibre Systems... 284 References... 286 10 Diencephalon: Hypothalamus... 289 IntroductoryNotes... 289 Topography and Subdivision... 289 Major Conduction Channels... 290 FunctionalAspects1:IntroductoryNotes... 293 Functional Aspects 2: Organization of Specific Hypothalamic FunctionalSystems... 298 Circadian Rhythms... 298 Wakefulness and Sleep... 301 StressResponse... 305 Thermoregulation... 307 FoodIntake... 308 Thirst... 311 Sexual Behaviour... 314 Defensive Behaviour... 320 Functional Aspects 3: Summary and Concluding Remarks... 322 References... 323 11 Telencephalon: Introduction and Olfactory System... 337 Introduction... 337 OlfactorySystem... 341 GrossFeatures... 341 OlfactoryReceptors... 341 OlfactoryBulb... 343 Secondary Olfactory Projections and Their Sites of Termination..... 347 Further Spread of Olfactory Information... 353 Accessory Olfactory System and Terminal Nerve... 355 References... 357 12 Telencephalon: Hippocampus and Related Structures... 361 Introduction... 361 Topography and Structure of Hippocampus... 366 Microcircuitry of Ammon s Horn... 372 IntrinsicConnectionsoftheHippocampus... 375 Major Fibre Systems Connecting the Hippocampal Formation and Its Dependencies with Other Parts of the Brain... 378 Afferents to the Hippocampus... 379 EfferentsfromtheHippocampus... 383 The Circuit of Papez... 386
X Synopsis of Anatomical Data: Functional and Clinical Aspect... 386 References... 391 13 Telencephalon: Amygdala and Claustrum... 401 Amygdala... 401 Topography, Subdivision and Structure... 401 Development... 403 Fibre Systems Related to the Amygdala... 404 Afferents to the Amygdala... 404 Intra-Amygdaloid Connections... 409 Efferents from the Amygdala... 409 The Bed Nucleus of the Stria Terminalis and the Extended Amygdala Concept... 414 FibreConnectionsoftheBST... 415 Functional and Clinical Aspects... 415 Claustrum... 417 Topography, Subdivision and Structure... 417 FibreConnections... 417 Circuitry and Function... 419 References... 420 14 Telencephalon: Basal Ganglia... 427 Introduction... 427 CellMasses... 429 Fibreconnections... 439 IntroductoryNotes... 439 The Direct Circuit: Cerebral Cortex Striatum Globus Pallidus Thalamus Cerebral Cortex; or: Cerebral Cortex Striatum Substantia Nigra, pars reticulata Thalamus Cortex... 439 Subdivision of Direct Striatal Circuit... 444 ABriefExcursiontotheCerebellum... 447 Thalamostriate Projections... 448 The Indirect Circuit: Cerebral Cortex Striatum External Segment of Globus Pallidus Subthalamic Nucleus Internal Segment of Globus Pallidus and Substantia Nigra, pars reticulata Thalamus CerebralCortex... 449 The Striato-Nigro-Striatal Circuit... 451 Additional Striatal Connections... 454 InputSystems... 454 Output Channels... 454 ConnectionsoftheVentralStriatum... 456 Connections of the Substantia Innominata, Basal Nucleus ofmeynertandrelatedcellgroups... 460 Clinical Aspects... 468 IntroductoryNote... 468 Résumé of the Circuitry of the Basal Ganglia... 468 Disorders of the Basal Ganglia... 471 References... 474
XI 15 Telencephalon: Neocortex... 491 Introduction... 491 Sulcal Pattern... 498 Structural and Functional Subdivision of Neocortex... 498 Structural Subdivision 1: Cytoarchitecture... 498 Structural Subdivision 2: Myeloarchitecture... 506 Structural Subdivision 3: Myelogenesis.... 510 Structural Subdivision 4: Connectivity..... 510 Functional Subdivision... 516 Structural and Functional Subdivision: Overview... 528 Structural and Functional Localization in the Neocortex: Current Research and Perspectives... 530 Neocortical Afferents... 536 Neocortical Neurons and Their Synaptic Relationships...... 544 IntroductoryNote... 544 Typical Pyramidal Cells... 544 Atypical Pyramidal Cells... 559 Local Circuit Neurons... 560 Microcircuitry of Neocortex... 569 Introduction... 569 Networks of Pyramidal Neuron... 570 Interneuronal Systems... 571 Neocortical Columns and Modules... 575 Introduction... 575 The Investigations of Lorente de Nó: Elementary Units and Glomérulos... 576 The Columnar Organization of the Somatosensory Cortex... 576 The Columnar Organization of the Visual Cortex... 578 The Auditory Cortex... 579 TheMotorCortex... 579 Columnar Patterns Shown by the Cells of Origin and the Terminal Ramifications of Cortico-cortical Connections..... 579 Minicolumns and the Radial Unit Hypothesis of Cortical Development... 581 Dendritic Clusters, Axonal Bundles and Radial Cell Cords As (Possible) Constituents of Neocortical Minicolumns.... 582 Microcircuitry of Neocortical Columns.... 586 Neocortical Columns and Modules: A Critical Commentary... 586 ComparativeAspects... 591 Synopsis of Main Neocortical Regions... 592 Introduction... 592 Association and Commissural Connections... 592 Functional and Structural Asymmetry of the Two Hemispheres..... 599 Occipital Lobe... 600 Parietal Lobe... 605 TemporalLobe... 611 Limbic Lobe and Paralimbic Belt... 617
XII FrontalLobe... 620 Insula... 649 References... 653 Section III Functional Systems... 681 16 General Sensory Systems and Taste... 683 Introduction... 683 Peripheral Receptors and Sensory Afferents... 684 ArchitectureoftheDorsalHornoftheSpinalCord... 687 Dorsal Column-Medial Lemniscus System... 693 The Sensory Relay Nuclei of the Thalamus andthesomatosensorycortex... 694 Descending Control by the Somatosensory Cortex... 697 Ascending Pathways from the Spinal Cord: Anterolateral Fasciculus and Spinothalamic Tract. Conduction of Pain, Itch and Temperature... 697 Control Mechanisms of Nociceptive Transmission... 701 The Trigeminal System... 702 The Visceral Afferent System... 705 References... 709 17 Vestibular System... 715 Introduction... 715 The Vestibular Labyrinth, the Vestibular Nerve and the Vestibular Nuclei... 715 Nonvestibular Afferents, Intrinsic and Commissural Connections of the Vestibular Nuclei... 720 Efferents From the Vestibular Nuclei... 721 Vestibular Projections to the Cerebellum... 721 ProjectionstotheSpinalCordandtheOculomotorCentres... 725 Vestibulo-ocular Reflex Pathways... 728 Vestibulo-collic Reflex Pathways... 729 Projections From the Vestibular Nuclei to the Thalamus andthecerebralcortex... 729 References... 730 18 Auditory System... 733 Auditory Centres... 733 Auditory Pathways... 736 Ascending Auditory Projection... 737 Nuclei of the Lateral Lemniscus and the Inferior Colliculus..... 741 Medial Geniculate Body and Auditory Cortex... 742 Descending Auditory Projection... 744 Auditory Reflex Pathways... 745 References... 747
XIII 19 Visual System... 751 Retina... 751 Optic Nerve, Chiasm and Tract... 757 Lateral Geniculate Body and the Visual Cortex: Retinotopical Projections to the Lateral Geniculate Body and the Primary Visual Cortex (V1)... 757 Visual Areas: Localization... 760 Visual Areas: Thalamocortical Projections of the Lateral Geniculate Body... 765 Visual Areas: Functional Streams. Intrinsic and Interareal ConnectionsofV1andV2... 766 Visual Areas: Dorsal and Ventral Processing Streams..... 767 VisuomotorPathways... 769 The Saccade Generating System... 770 Superior Colliculus... 772 Afferent Connections of the Superficial Layers of the Superior Colliculus... 772 Afferent Connections of the Intermediate and Deep Layers of the Superior Colliculus... 777 Efferent Connections of the Superior Colliculus... 778 Accessory Optic System and the Nucleus of the Optic Tract... 781 The Frontal and Parietal Eye Fields... 784 Pretectum and the Visual Reflexes... 787 VisualSystemandtheCerebellum... 790 References... 796 20 Cerebellum... 807 Introduction... 807 GrossAnatomy... 808 Lattice Structure of the Cerebellar Cortex and the Microcircuitry ofthecerebellum... 812 The Cerebellar Nuclei, the Modular Organization of the Cerebellar Output and the Efferent Connections of the Cerebellum..... 820 Localization in the Cerebellum... 826 FunctionandMalfunctionoftheCerebellum... 833 References... 834 21 Motor Systems... 841 Introduction... 841 Kuypers Concept of the Motor System: Medial and Lateral Brain Stem SystemsandtheProjectionoftheMotorCortex... 841 TheMotorCortex... 849 The Primary Motor Cortex and the Premotor Areas...... 849 The Origin of the Corticospinal and Corticobulbar Tracts... 851 Corticocortical, Cerebellar and Pallidal Connections of the Primary Motor Cortex and the Premotor Areas..... 855
XIV Properties of the Primary Motor and the Premotor Areas.... 859 The Primary Motor Area... 859 Premotor Cortical Areas... 859 The Cranial Motor Nuclei... 862 The Motor Nucleus of the Trigeminal Nerve... 863 The Nucleus of the Facial Nerve... 865 The Nucleus Ambiguus and the Nucleus of the Hypoglossal Nerve: Swallowing and Vocalization... 867 Brain Stem Control of Spinal Motor Systems... 872 Locomotion... 872 Brain Stem Control of Micturition and Ejaculation... 878 References... 880 22 Reticular Formation and the Monoaminergic and Cholinergic Cell Groups... 889 Introduction... 889 Subdivisions of the Reticular Formation... 889 Monoaminergic Cell Groups and Pathways... 893 SerotonergicCellGroups... 893 Adrenergic and Noradrenergic Cell groups... 898 Dopaminergic Cell Groups... 901 Cholinergic Cell Groups... 902 Connections of the Reticular Formation and Associated Cell Groups... 903 References... 908 23 Greater Limbic System... 917 Historical Introduction: From Limbic Lobe to Limbic System... 917 The Concept of a Greater Limbic System (GLS)... 923 Functional, Structural and Chemical Features of GLS... 925 A Brief Guide to Discussions of Limbic Structures in Previous Chapters 930 Inputs to Limbic Domains... 930 TheLimbicMotorSystem... 934 Concluding Remarks... 941 References... 942 Subject Index... 947
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