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Nolte’s The Human Brain E-Book

Nolte’s The Human Brain E-Book

Todd Vanderah | Douglas J Gould


Additional Information


Popular for its highly visual and easy-to-follow approach, Nolte's The Human Brain helps demystify the complexities of the gross anatomy of the brain, spinal cord and brainstem. A clear writing style, interesting examples and visual cues bring this extremely complicated subject to life and more understandable.

  • Get the depth of coverage you need with discussions on all key topics in functional neuroanatomy and neuroscience, giving you well-rounded coverage of this complex subject.
  • Zero in on the key information you need to know with highly templated, concise chapters that reinforce and expand your knowledge.
  • Develop a thorough, clinically relevant understanding through clinical examples providing a real-life perspective.
  • Gain a greater understanding of every concept through a glossary of key terms that elucidates every part of the text; 3-dimensional brain.
  • Acquaint yourself with the very latest advancements in the field with many illustrations using the most current neuroimaging techniques, reflecting recent developments and changes in understanding. 
  • Keep up with the latest knowledge in neural plasticity including formation, modification, and repair of connections, with coverage of learning and memory, as well as the coming revolution in ways to fix damaged nervous systems, trophic factors, stem cells, and more. 
  • NEW! Gauge your mastery of the material and build confidence with over 100 multiple choice questions that provide effective chapter review and quick practice for your exams.

Table of Contents

Section Title Page Action Price
Front Cover cover
Inside Front Cover ifc1
Nolte's The Human Brain i
Copyright Page ii
Preface iii
In Memoriam v
Table Of Contents vii
Video Contents xi
1 Introduction to the Nervous System 1
Chapter Outline 1
The Nervous System Has Central and Peripheral Parts 1
The Principal Cellular Elements of the Nervous System Are Neurons and Glial Cells 2
Neurons Come in a Variety of Sizes and Shapes, yet All Are Variations on the Same Theme 2
Neuronal Cell Bodies and Axons Are Largely Segregated Within the Nervous System 6
Neuronal Organelles Are Distributed in a Pattern That Supports Neuronal Function 8
Neuronal Cell Bodies Synthesize Macromolecules 9
Dendrites Receive Synaptic Inputs 10
Axons Convey Electrical Signals Over Long Distances 13
Organelles and Macromolecules Are Transported in Both Directions Along Axons 13
Synapses Mediate Information Transfer Between Neurons 18
Schwann Cells Are Glial Cells of the PNS 20
PNS Axons Can Be Myelinated or Unmyelinated 20
CNS Glial Cells Include Oligodendrocytes, Astrocytes, Ependymal Cells, and Microglial Cells 22
Some CNS Axons Are Myelinated by Oligodendrocytes, but Others Are Unmyelinated 29
Astrocytes Provide Structural and Metabolic Support to Neurons 29
Ependymal Cells Line the Ventricles 29
Microglial Cells Respond to CNS Injury 29
Suggested Readings 38
2 Development of the Nervous System 39
Chapter Outline 39
The Neural Tube Gives Rise to the Central Nervous System 39
The Sulcus Limitans Separates Sensory and Motor Areas of the Spinal Cord and Brainstem 40
The Neural Tube Has a Series of Bulges and Flexures 41
There Are Three Primary Vesicles 41
There Are Five Secondary Vesicles 42
Growth of the Telencephalon Overshadows Other Parts of the Nervous System 42
The Cavity of the Neural Tube Persists as a System of Ventricles 46
The Neural Crest and Cranial Placodes Give Rise to the Peripheral Nervous System 48
Adverse Events During Development Can Cause Congenital Malformations of the Nervous System 49
Defective Closure of the Neural Tube Can Cause Spina Bifida or Anencephaly 51
Defective Secondary Neurulation Can Cause a Distinctive Set of Abnormalities 53
The Forebrain Can Develop Abnormally Even If Neural Tube Closure Is Complete 53
Suggested Readings 54
3 Gross Anatomy and General Organization of the Central Nervous System 56
Chapter Outline 56
The Long Axis of the CNS Bends at the Cephalic Flexure 57
Hemisecting a Brain Reveals Parts of the Diencephalon, Brainstem, and Ventricular System 58
Humans, Relative to Other Animals, Have Large Brains and Many Neurons 59
Named Sulci and Gyri Cover the Cerebral Surface 59
Each Cerebral Hemisphere Includes a Frontal, Parietal, Occipital, Temporal, and Limbic Lobe 61
The Frontal Lobe Contains Motor Areas 61
The Parietal Lobe Contains Somatosensory Areas 62
The Temporal Lobe Contains Auditory Areas 64
The Occipital Lobe Contains Visual Areas 65
The Limbic Lobe Is Interconnected with Other Limbic Structures, Some Buried in the Temporal Lobe 65
The Diencephalon Includes the Thalamus and Hypothalamus 66
The Thalamus Conveys Information to the Cerebral Cortex 66
The Hypothalamus Controls the Autonomic Nervous System 68
Most Cranial Nerves Are Attached to the Brainstem 68
The Cerebellum Includes a Vermis and Two Hemispheres 70
Sections of the Forebrain Reveal the Basal Nuclei and Limbic Structures 70
Many Parts of Each Cerebral Hemisphere Are Arranged in a C Shape 70
The Caudate Nucleus, Putamen, and Globus Pallidus Are Major Components of the Basal Nuclei 71
The Amygdala and Hippocampus Are Major Limbic Structures 71
Cerebral Structures Are Arranged Systematically 71
Parts of the Nervous System Are Interconnected in Systematic Ways (Generalizations) 72
Axons of Primary Afferents and Lower Motor Neurons Convey Information to and From the CNS 72
Axons of Primary Afferents Enter the CNS Without Crossing the Midline 72
Axons of Lower Motor Neurons Leave the CNS Without Crossing the Midline 73
Somatosensory Inputs Participate in Reflexes, Pathways to the Cerebellum, and Pathways to the Cerebral Cortex 73
Somatosensory Pathways to the Cerebral Cortex Cross the Midline and Pass Through the Thalamus 78
Somatosensory Cortex Contains a Distorted Map of the Body 79
Each Side of the Cerebellum Receives Information About the Ipsilateral Side of the Body 79
Other Sensory Systems Are Similar to the Somatosensory System 80
Higher Levels of the CNS Influence the Activity of Lower Motor Neurons 81
Corticospinal Axons Cross the Midline 81
Each Side of the Cerebellum Indirectly Affects Movements of the Ipsilateral Side of the Body 81
The Basal Nuclei of One Side Indirectly Affect Movements of the Contralateral Side of the Body 81
Suggested Readings 82
4 Meningeal Coverings of the Brain and Spinal Cord 84
Chapter Outline 84
There Are Three Meningeal Layers: The Dura Mater, Arachnoid, and Pia Mater 84
The Dura Mater Provides Mechanical Strength 86
Dural Folds Partially Separate Different Intracranial Compartments 86
The Dura Mater Contains Venous Sinuses That Drain the Brain 86
The Dura Mater Has Its Own Blood Supply 89
The Dura Mater Is Pain Sensitive 90
The Arachnoid Mater 90
The Arachnoid Bridges Over CNS Surface Irregularities, Forming Cisterns 91
CSF Enters the Venous Circulation Through Arachnoid Villi 93
The Arachnoid Has a Barrier Function 93
Pia Mater Covers the Surface of the CNS 93
The Vertebral Canal Contains a Spinal Epidural Space 94
Bleeding Can Open Up Potential Meningeal Spaces 96
Tearing of Meningeal Arteries Can Cause an Epidural Hematoma 97
Tearing of Veins Where They Enter Venous Sinuses Can Cause a Subdural Hematoma 97
Parts of the CNS Can Herniate from One Intracranial Compartment Into Another 97
Suggested Readings 101
5 Ventricles and Cerebrospinal Fluid 103
Chapter Outline 103
The Brain Contains Four Ventricles 103
A Lateral Ventricle Curves Through Each Cerebral Hemisphere 104
The Third Ventricle Is a Midline Cavity in the Diencephalon 105
The Fourth Ventricle Communicates with Subarachnoid Cisterns 106
The Ventricles Contain Only a Fraction of the CSF 106
Choroid Plexus Is the Source of Most CSF 107
The Ependymal Lining of Choroid Plexus Is Specialized as a Secretory Epithelium 107
CSF Is a Secretion of the Choroid Plexus 109
CSF Circulates Through and Around the CNS, Eventually Reaching the Venous System 110
CSF Has Multiple Functions 110
Imaging Techniques Allow Noninvasive Visualization of the CNS 111
Tomography Produces Images of Two-Dimensional “Slices” 114
CT Produces Maps of X-Ray Density 115
MRI Produces Maps of Water Concentration 116
Disruption of CSF Circulation Can Cause Hydrocephalus 121
Suggested Readings 124
6 Blood Supply of the Brain 126
Chapter Outline 126
The Internal Carotid Arteries and Vertebral Arteries Supply the Brain 126
The Internal Carotid Arteries Supply Most of the Cerebrum 127
Small Perforating Arteries Supply Deep Cerebral Structures 128
The Vertebral-Basilar System Supplies the Brainstem and Parts of the Cerebrum and Spinal Cord 130
The Cerebral Arterial Circle (Circle of Willis) Interconnects the Internal Carotid and Vertebral-Basilar Systems 134
Imaging Techniques Allow Arteries and Veins to Be Visualized 136
Blood Flow to the CNS Is Closely Controlled 136
The Overall Flow Rate Is Constant, but There Are Regional Changes in Blood Flow 138
Strokes Result From Disruption of the Vascular Supply 141
A System of Barriers Partially Separates the Nervous System From the Rest of the Body 144
Superficial and Deep Veins Drain the Brain 146
Most Superficial Veins Empty Into the Superior Sagittal Sinus 148
Deep Veins Ultimately Empty Into the Straight Sinus 149
Suggested Readings 151
7 Electrical Signaling by Neurons 154
Chapter Outline 154
A Lipid-Protein Membrane Separates Intracellular and Extracellular Fluids 155
The Lipid Component of the Membrane Is a Diffusion Barrier 155
Membrane Proteins Regulate the Movement of Solutes Across the Membrane 156
Ions Diffuse Across the Membrane Through Ion Channels—Protein Molecules With Pores 156
The Number and Selectivity of Ion Channels Determine the Membrane Potential 159
The Resting Membrane Potential of Typical Neurons Is Heavily Influenced, but Not Completely Determined, by the Potassium Concentration Gradient 160
Concentration Gradients Are Maintained by Membrane Proteins That Pump Ions 161
Inputs to Neurons Cause Slow, Local Potential Changes 161
Membrane Capacitance and Resistance Determine the Speed and Extent of the Response to a Current Pulse 162
Membranes Have a Time Constant, Allowing Temporal Summation 162
Larger-Diameter Neuronal Processes Have Longer Length Constants 163
Action Potentials Convey Information Over Long Distances 164
Opening and Closing of Voltage-Gated Sodium and Potassium Channels Underlie the Action Potential 164
Mammalian Neurons Contain Multiple Types of Voltage-Gated Na+ and K+ Channels 167
Action Potentials Are Followed by Brief Refractory Periods 168
Refractory Periods Limit the Repetition Rate of Action Potentials 168
Pathological Processes and Toxins Can Selectively Affect Voltage-Gated Channels 169
Action Potentials Are Propagated Without Decrement Along Axons 170
Propagation Is Continuous and Relatively Slow in Unmyelinated Axons 170
Refractory Periods Ensure That Action Potentials Are Propagated in Only One Direction 173
Action Potentials “Jump” Rapidly from Node to Node in Myelinated Axons 173
Demyelinating Diseases Can Slow or Block Conduction of Action Potentials 175
Resistors, Capacitors, and Neuronal Membranes 177
Calculating the Membrane Potential 179
Suggested Readings 180
8 Synaptic Transmission Between Neurons 182
Chapter Outline 182
There Are Five Steps in Conventional Chemical Synaptic Transmission 183
Neurotransmitters Are Synthesized in Presynaptic Endings and in Neuronal Cell Bodies 184
Neurotransmitters Are Packaged Into Synaptic Vesicles Before Release 184
Presynaptic Endings Release Neurotransmitters Into the Synaptic Cleft 185
Neurotransmitters Diffuse Across the Synaptic Cleft and Bind to Postsynaptic Receptors 185
Neurotransmitter Action Is Terminated by Uptake, Degradation, or Diffusion 186
Synaptic Transmission Can Be Rapid and Point-to-Point, or Slow and Often Diffuse 188
Rapid Synaptic Transmission Involves Transmitter-Gated Ion Channels 188
Slow Synaptic Transmission Usually Involves Postsynaptic Receptors Linked to Intracellular Proteins 189
The Postsynaptic Receptor Determines the Effect of a Neurotransmitter 190
The Size and Location of a Synaptic Ending Influence the Magnitude of Its Effects 191
Synapses With Many Active Zones Have a Greater Effect 191
Synapses Closer to the Action Potential Trigger Zone Have a Greater Effect 192
Presynaptic Endings Can Themselves Be Postsynaptic 193
Synaptic Strength Can Be Facilitated or Depressed 193
Messages Also Travel Across Synapses in a Retrograde Direction 195
Most Neurotransmitters Are Small Amine Molecules, Amino Acids, or Neuropeptides 196
Acetylcholine Mediates Rapid, Point-to-Point Transmission in the PNS 196
Amino Acids Mediate Rapid, Point-to-Point Transmission in the CNS 197
Excessive Levels of Glutamate Are Toxic 198
ATP Is Not Just an Energy Source but Also a Neurotransmitter 198
Amines and Neuropeptides Mediate Slow, Diffuse Transmission 198
Drugs, Diseases, and Toxins Can Selectively Affect Particular Parts of Individual Neurotransmitter Systems 200
Gap Junctions Mediate Direct Current Flow From One Neuron to Another 201
Suggested Readings 205
9 Sensory Receptors and the Peripheral Nervous System 207
Chapter Outline 207
Receptors Encode the Nature, Location, Intensity, and Duration of Stimuli 208
Each Sensory Receptor Has an Adequate Stimulus, Allowing It to Encode the Nature of a Stimulus 208
Many Sensory Receptors Have a Receptive Field, Allowing Them to Encode the Location of a Stimulus 208
Receptor Potentials Encode the Intensity and Duration of Stimuli 209
Most Sensory Receptors Adapt to Maintained Stimuli, Some More Rapidly Than Others 209
Sensory Receptors All Share Some Organizational Features 210
Sensory Receptors Use Ionotropic and Metabotropic Mechanisms to Produce Receptor Potentials 210
All Sensory Receptors Produce Receptor Potentials, but Some Do Not Produce Action Potentials 211
Somatosensory Receptors Detect Mechanical, Chemical, or Thermal Changes 212
Cutaneous Receptors Have Either Encapsulated or Nonencapsulated Endings 212
Capsules and Accessory Structures Influence the Response Properties of Cutaneous Mechanoreceptors 214
Nociceptors, Thermoreceptors, and Some Mechanoreceptors Have Free Nerve Endings 217
Nociceptors Have Both Afferent and Efferent Functions 219
Pain Serves Useful Functions 220
Cutaneous Receptors Are Not Distributed Uniformly 221
Receptors in Muscles and Joints Detect Muscle Status and Limb Position 221
Muscle Spindles Detect Muscle Length 222
Golgi Tendon Organs Detect Muscle Tension 222
Joints Have Receptors 225
Muscle Spindles Are Important Proprioceptors 225
Visceral Structures Contain a Variety of Receptive Endings 225
Particular Sensations Are Sometimes Related to Particular Receptor Types 226
Peripheral Nerves Convey Information To and From the CNS 227
Extensions of the Meninges Envelop Peripheral Nerves 227
The Diameter of a Nerve Fiber Is Correlated With Its Function 230
Suggested Readings 231
10 Spinal Cord 233
Chapter Outline 233
The Spinal Cord Is Segmented 234
Each Spinal Cord Segment Innervates a Dermatome 235
The Spinal Cord Is Shorter Than the Vertebral Canal 238
All Levels of the Spinal Cord Have a Similar Cross-Sectional Structure 238
The Spinal Cord Is Involved in Sensory Processing, Motor Outflow, and Reflexes 239
Spinal Gray Matter Is Regionally Specialized 240
The Posterior Horn Contains Sensory Interneurons and Projection Neurons 240
The Anterior Horn Contains Motor Neurons 241
The Intermediate Gray Matter Contains Autonomic Neurons 243
Spinal Cord Gray Matter Is Arranged in Layers 244
Reflex Circuitry Is Built Into the Spinal Cord 244
Muscle Stretch Leads to Excitation of Motor Neurons 244
Muscle Tension Can Lead to Inhibition of Motor Neurons 245
Painful Stimuli Elicit Coordinated Withdrawal Reflexes 246
Reflexes Are Accompanied by Reciprocal and Crossed Effects 246
Reflexes Are Modifiable 247
Ascending and Descending Pathways Have Defined Locations in the Spinal White Matter 248
The Posterior Column–Medial Lemniscus System Conveys Information About Touch and Limb Position 249
Information About the Location and Nature of a Stimulus Is Preserved in the Posterior Column–Medial Lemniscus System 250
Damage to the Posterior Column–Medial Lemniscus System Causes Impairment of Proprioception and Discriminative Tactile Functions 250
The Spinothalamic Tract Conveys Information About Pain and Temperature 252
Damage to the Anterolateral System Causes Diminution of Pain and Temperature Sensations 255
Additional Pathways Convey Somatosensory Information to the Thalamus 255
Spinal Information Reaches the Cerebellum Both Directly and Indirectly 256
The Posterior Spinocerebellar Tract and Cuneocerebellar Tract Convey Proprioceptive Information 256
The Anterior Spinocerebellar Tract Conveys More Complex Information 256
Descending Pathways Influence the Activity of Lower Motor Neurons 258
The Corticospinal Tracts Mediate Voluntary Movement 258
The Autonomic Nervous System Monitors and Controls Visceral Activity 260
Preganglionic Parasympathetic Neurons Are Located in the Brainstem and Sacral Spinal Cord 262
Preganglionic Sympathetic Neurons Are Located in Thoracic and Lumbar Spinal Segments 262
Visceral Distortion or Damage Causes Pain That Is Referred to Predictable Dermatomes 264
A Longitudinal Network of Arteries Supplies the Spinal Cord 264
Spinal Cord Damage Causes Predictable Deficits 267
Long-Term Effects of Spinal Cord Damage Are Preceded by a Period of Spinal Shock 267
The Side and Distribution of Deficits Reflect the Location of Spinal Cord Damage 267
Suggested Readings 269
11 Organization of the Brainstem 272
Chapter Outline 272
The Brainstem Has Conduit, Cranial Nerve, and Integrative Functions 273
The Medulla, Pons, and Midbrain Have Characteristic Gross Anatomical Features 274
The Medulla Includes Pyramids, Olives, and Part of the Fourth Ventricle 274
The Pons Includes the Basal Pons, Middle Cerebellar Peduncles, and Part of the Fourth Ventricle 276
The Midbrain Includes the Superior and Inferior Colliculi, the Cerebral Peduncles, and the Cerebral Aqueduct 277
The Internal Structure of the Brainstem Reflects Surface Features and the Position of Long Tracts 277
The Corticospinal Tract and Anterolateral System Have Consistent Locations Throughout the Brainstem 278
The Medial Lemniscus Forms in the Caudal Medulla 278
The Rostral Medulla Contains the Inferior Olivary Nucleus and Part of the Fourth Ventricle 281
The Caudal Pons Is Attached to the Cerebellum by the Middle Cerebellar Peduncle 281
The Superior Cerebellar Peduncle Joins the Brainstem in the Rostral Pons 283
The Superior Cerebellar Peduncles Decussate in the Caudal Midbrain 283
The Rostral Midbrain Contains the Red Nucleus and Substantia Nigra 285
The Reticular Core of the Brainstem Is Involved in Multiple Functions 286
The Reticular Formation Participates in the Control of Movement Through Connections With Both the Spinal Cord and the Cerebellum 287
The Reticular Formation Modulates the Transmission of Information in Pain Pathways 288
The Reticular Formation Contains Autonomic Reflex Circuitry 290
The Reticular Formation Is Involved in the Control of Arousal and Consciousness 290
Some Brainstem Nuclei Have Distinctive Neurochemical Signatures 290
Neurons of the Locus Ceruleus Contain Norepinephrine 291
Neurons of the Substantia Nigra and Ventral Tegmental Area Contain Dopamine 291
Neurons of the Raphe Nuclei Contain Serotonin 292
Neurons of the Rostral Brainstem and Basal Forebrain Contain Acetylcholine 295
Neurochemical Imbalances Are Involved in Certain Forms of Mental Illness 295
The Brainstem Is Supplied by the Vertebral-Basilar System 295
Suggested Readings 299
12 Cranial Nerves and Their Nuclei 301
Chapter Outline 301
Cranial Nerve Nuclei Have a Generally Predictable Arrangement 301
The Sulcus Limitans Intervenes Between Motor and Sensory Nuclei of Cranial Nerves 302
Cranial Nerves III, IV, VI, XI, and XII Contain Somatic Motor Fibers 305
The Oculomotor Nerve (III) Innervates Four of the Six Extraocular Muscles 305
The Trochlear Nerve (IV) Innervates the Superior Oblique 307
The Abducens Nerve (VI) Innervates the Lateral Rectus 308
The Abducens Nucleus Also Contains Interneurons That Project to the Contralateral Oculomotor Nucleus 308
The Accessory Nerve (XI) Innervates Neck and Shoulder Muscles 310
The Hypoglossal Nerve (XII) Innervates Tongue Muscles 311
Branchiomeric Nerves Contain Axons From Multiple Categories 312
The Trigeminal Nerve (V) Is the General Sensory Nerve for the Head 312
The Main Sensory Nucleus Receives Information About Touch and Jaw Position 315
The Spinal Trigeminal Nucleus Receives Information About Pain and Temperature 315
The Trigeminal Motor Nucleus Innervates Muscles of Mastication 319
The Facial (VII), Glossopharyngeal (IX), and Vagus (X) Nerves All Contain Somatic and Visceral Sensory, Visceral Motor, and Pharyngeal Motor Fibers 319
The Facial Nerve (VII) Innervates Muscles of Facial Expression 320
Upper Motor Neuron Damage Affects the Upper and Lower Parts of the Face Differently 322
The Glossopharyngeal Nerve (IX) Conveys Information From Intraoral Receptors 323
The Vagus Nerve (X) Is the Principal Parasympathetic Nerve 324
Brainstem Damage Commonly Causes Deficits on One Side of the Head and the Opposite Side of the Body 325
Suggested Readings 327
13 The Chemical Senses of Taste and Smell 329
Chapter Outline 329
The Perception of Flavor Involves Gustatory, Olfactory, Trigeminal, and Other Inputs 330
Taste Is Mediated by Receptors in Taste Buds Innervated by Cranial Nerves VII, IX, and X 330
The Tongue Is Covered by a Series of Papillae, Some of Which Contain Taste Buds 330
Taste Receptor Cells Are Modified Epithelial Cells With Neuron-Like Properties 332
Taste Receptor Cells Utilize a Variety of Transduction Mechanisms to Detect Sweet, Salty, Sour, and Bitter Stimuli 332
Second-Order Gustatory Neurons Are Located in the Nucleus of the Solitary Tract 334
Information About Taste Is Coded, in Part, by the Pattern of Activity in Populations of Neurons 335
Olfaction Is Mediated by Receptors That Project Directly to the Telencephalon 336
The Axons of Olfactory Receptor Neurons Form Cranial Nerve I 336
Olfactory Receptor Neurons Utilize a Large Number of G Protein–Coupled Receptors to Detect a Wide Range of Odors 338
Olfactory Information Bypasses the Thalamus on Its Way to the Cerebral Cortex 338
The Olfactory Nerve Terminates in the Olfactory Bulb 338
The Olfactory Bulb Projects to Olfactory Cortex 342
Olfactory Information Reaches Other Cortical Areas Both Directly and Via the Thalamus 344
Conductive and Sensorineural Problems Can Affect Olfactory Function 344
Multiple Flavor-Related Signals Converge in Orbital Cortex 345
Suggested Readings 345
14 Hearing and Balance: 348
Chapter Outline 348
Auditory and Vestibular Receptor Cells Are Located in the Walls of the Membranous Labyrinth 349
The Membranous Labyrinth Is Suspended Within the Bony Labyrinth, a Cavity in the Temporal Bone 349
Endolymph Is Actively Secreted, Circulates Through the Membranous Labyrinth, and Is Reabsorbed 351
Auditory and Vestibular Receptors Are Hair Cells 351
Hair Cells Have Mechanosensitive Transduction Channels 352
Subtle Differences in the Physical Arrangements of Hair Cells Determine the Stimuli to Which They Are Most Sensitive 354
The Cochlear Division of the Eighth Nerve Conveys Information About Sound 354
The Outer and Middle Ears Convey Airborne Vibrations to the Fluid-Filled Inner Ear 355
The Cochlea Is the Auditory Part of the Labyrinth 355
Traveling Waves in the Basilar Membrane Stimulate Hair Cells in the Organ of Corti, in Locations That Depend on Sound Frequency 358
Inner Hair Cells Are Sensory Cells; Outer Hair Cells Are Amplifiers 361
Auditory Information Is Distributed Bilaterally in the CNS 363
Activity in the Ascending Auditory Pathway Generates Electrical Signals That Can Be Measured From the Scalp 365
Efferents Control the Sensitivity of the Cochlea 365
Middle Ear Muscles Contract in Response to Loud Sounds 366
Different Sets of Efferents Control Outer Hair Cells and the Afferent Endings on Inner Hair Cells 366
Conductive and Sensorineural Problems Can Affect Hearing 367
The Vestibular Division of the Eighth Nerve Conveys Information About Linear and Angular Acceleration of the Head 369
Receptors in the Semicircular Ducts Detect Angular Acceleration of the Head 370
Receptors in the Utricle and Saccule Detect Linear Acceleration and Position of the Head 371
Vestibular Primary Afferents Project to the Vestibular Nuclei and the Cerebellum 372
The Vestibular Nuclei Project Primarily to the Spinal Cord, Cerebellum, and Nuclei of Cranial Nerves III, IV, and VI 373
Vestibulospinal Fibers Influence Antigravity Muscles and Neck Muscles 374
The Vestibular Nuclei Participate in the Vestibuloocular Reflex 375
Nystagmus Can Be Physiological or Pathological 376
Conditions That Make the Cupula Sensitive to Gravity Cause Nystagmus and Illusions of Movement 379
Efferents Control the Sensitivity of Vestibular Hair Cells 379
Position Sense Is Mediated by the Vestibular, Proprioceptive, and Visual Systems Acting Together 380
Suggested Readings 381
15 Atlas of the Human Brainstem 383
16 The Thalamus and Internal Capsule: 394
Chapter Outline 394
The Diencephalon Includes the Epithalamus, Subthalamus, Hypothalamus, and Thalamus 395
The Epithalamus Includes the Pineal Gland and the Habenular Nuclei 395
The Subthalamus Includes the Subthalamic Nucleus and the Zona Incerta 397
The Thalamus Is the Gateway to the Cerebral Cortex 398
The Thalamus Has Anterior, Medial, and Lateral Divisions, Defined by the Internal Medullary Lamina 398
Intralaminar Nuclei Are Embedded in the Internal Medullary Lamina 401
The Thalamic Reticular Nucleus Partially Surrounds the Thalamus 401
Midline Nuclei Cover the Ventricular Surface of Each Thalamus 404
Patterns of Input and Output Connections Define Functional Categories of Thalamic Nuclei 404
All Thalamic Nuclei (Except the Reticular Nucleus) Are Variations on a Common Theme 405
Thalamic Projection Neurons Have Two Physiological States 405
There Are Relay Nuclei for Sensory, Motor, and Limbic Systems 407
The Dorsomedial Nucleus and Pulvinar Are the Principal Association Nuclei 408
Intralaminar Nuclei Project to Both the Cerebral Cortex and the Basal Nuclei 409
The Thalamic Reticular Nucleus Projects to Other Thalamic Nuclei and Not to the Cerebral Cortex 410
Small Branches of the Posterior Cerebral Artery Provide Most of the Blood Supply to the Thalamus 410
Interconnections Between the Cerebral Cortex and Subcortical Structures Travel Through the Internal Capsule 411
The Internal Capsule Has Five Parts 411
Small Branches of the Middle Cerebral Artery Provide Most of the Blood Supply to the Internal Capsule 413
Suggested Readings 417
17 The Visual System 419
Chapter Outline 419
The Eye Has Three Concentric Tissue Layers and a Lens 420
Intraocular Pressure Maintains the Shape of the Eye 421
The Cornea and Lens Focus Images on the Retina 422
The Iris Affects the Brightness and Quality of the Image Focused on the Retina 423
A System of Barriers Partially Separates the Retina From the Rest of the Body 423
The Retina Contains Five Major Neuronal Cell Types 424
Retinal Neurons and Synapses Are Arranged in Layers 424
The Retina Is Regionally Specialized 428
Retinal Neurons Translate Patterns of Light Into Patterns of Contrast 429
Photopigments Are G Protein–Coupled Receptors That Cause Hyperpolarizing Receptor Potentials 430
Rods Function in Dim Light 433
Populations of Cones Signal Spatial Detail and Color 435
Ganglion Cells Have Center-Surround Receptive Fields 435
Center-Surround Receptive Fields Are Formed in the Outer Plexiform Layer 437
Rod and Cone Signals Reach the Same Ganglion Cells 441
Half of the Visual Field of Each Eye Is Mapped Systematically in the Contralateral Cerebral Hemisphere 441
Fibers From the Nasal Half of Each Retina Cross in the Optic Chiasm 442
Most Fibers of the Optic Tract Terminate in the Lateral Geniculate Nucleus 442
The Lateral Geniculate Nucleus Projects to Primary Visual Cortex 444
Damage at Different Points in the Visual Pathway Results in Predictable Deficits 445
Some Fibers of the Optic Tract Terminate in the Superior Colliculus, Accessory Optic Nuclei, and Hypothalamus 447
Primary Visual Cortex Sorts Visual Information and Distributes It to Other Cortical Areas 450
Visual Cortex Has a Columnar Organization 450
Visual Information Is Distributed in Dorsal and Ventral Streams 450
Early Experience Has Permanent Effects on the Visual System 454
Reflex Circuits Adjust the Size of the Pupil and the Focal Length of the Lens 454
Illumination of Either Retina Causes Both Pupils to Constrict 454
Both Eyes Accommodate for Near Vision 456
Suggested Readings 457
18 Overview of Motor Systems 459
Chapter Outline 459
Each Lower Motor Neuron Innervates a Group of Muscle Fibers, Forming a Motor Unit 459
Lower Motor Neurons Are Arranged Systematically 460
There Are Three Kinds of Muscle Fibers and Three Kinds of Motor Units 460
Motor Units Are Recruited in Order of Size 461
Motor Control Systems Involve Both Hierarchical and Parallel Connections 463
Reflex and Motor Program Connections Provide Some of the Inputs to Lower Motor Neurons 463
Upper Motor Neurons Control Lower Motor Neurons Both Directly and Indirectly 464
Association Cortex, the Cerebellum, and the Basal Nuclei Modulate Motor Cortex 465
The Corticospinal Tract Has Multiple Origins and Terminations 465
Corticospinal Axons Arise in Multiple Cortical Areas 466
Motor Cortex Projects to Both the Spinal Cord and the Brainstem 469
Corticospinal Input Is Essential for Only Some Movements 469
Upper Motor Neuron Damage Causes a Distinctive Syndrome 470
There Are Upper Motor Neurons for Cranial Nerve Motor Nuclei 471
Suggested Readings 473
19 Basal Nuclei 475
Chapter Outline 475
The Basal Nuclei Include Five Major Nuclei 476
The Striatum and Globus Pallidus Are the Major Forebrain Components of the Basal Nuclei 478
The Subthalamic Nucleus and Substantia Nigra Are Interconnected With the Striatum and Globus Pallidus 479
Basal Nuclei Circuitry Involves Multiple Parallel Loops That Modulate Cortical Output 480
Afferents From the Cortex Reach the Striatum and Subthalamic Nucleus; Efferents Leave From the Globus Pallidus and Substantia Nigra 480
Interconnections of the Basal Nuclei Determine the Pattern of Their Outputs 481
The Cerebral Cortex, Substantia Nigra, and Thalamus Project to the Striatum 482
Different Parts of the Striatum Are Involved in Movement, Cognition, and Affect 482
The Striatum Projects to the Globus Pallidus and Substantia Nigra 483
The External Segment of the Globus Pallidus Distributes Inhibitory Signals Within the Basal Nuclei 483
The Internal Segment of the Globus Pallidus and the Reticular Part of the Substantia Nigra Provide the Output From the Basal Nuclei 483
The Subthalamic Nucleus Is Part of Additional Pathways Through the Basal Nuclei 485
Part of the Substantia Nigra Modulates the Output of the Striatum and Other Parts of the Basal Nuclei 485
Perforating Branches From the Cerebral Arterial Circle (of Willis) Supply the Basal Nuclei 487
Many Basal Nuclei Disorders Result in Abnormalities of Movement 488
Anatomical and Neurochemical Properties of the Basal Nuclei Suggest Effective Treatments for Disorders 490
Suggested Readings 493
20 Cerebellum 495
Chapter Outline 495
The Cerebellum Can Be Divided Into Both Transverse and Longitudinal Zones 496
Transverse Fissures Divide the Cerebellum Into Lobes 496
Functional Connections Divide the Cerebellum Into Longitudinal Zones 498
Three Peduncles Convey the Inputs and Outputs of Each Half of the Cerebellum 498
Deep Nuclei Are Embedded in the Cerebellar White Matter 500
Inputs Reach the Cerebellar Cortex as Mossy and Climbing Fibers 500
Purkinje Cells of the Cerebellar Cortex Project to the Deep Nuclei 503
Each Side of the Cerebellum Affects the Ipsilateral Side of the Body 506
Details of Connections Differ Among Zones 506
Cerebellar Cortex Receives Multiple Inputs 507
Vestibular Inputs Reach the Flocculus and Vermis 508
The Spinal Cord Projects to the Vermis and Medial Hemisphere 508
Cerebral Cortex Projects to the Cerebellum by Way of Pontine Nuclei 508
Climbing Fibers Arise in the Contralateral Inferior Olivary Nucleus 511
Visual and Auditory Information Reaches the Cerebellum 512
Each Longitudinal Zone Has a Distinctive Output 513
The Vermis Projects to the Fastigial Nucleus 513
The Medial and Lateral Parts of Each Hemisphere Project to the Interposed and Dentate Nuclei 513
The Lateral Hemispheres Are Involved in Planning and Skilled Movements 514
The Medial Hemispheres Are Involved in Adjusting Limb Movements 515
The Vermis Is Involved in Postural Adjustments 516
The Flocculus and Vermis Are Involved in Eye Movements 516
The Cerebellum Is Involved in Motor Learning 516
The Cerebellum Is Also Involved in Cognitive Functions 518
Clinical Syndromes Correspond to Functional Zones 518
Midline Damage Causes Postural Instability 518
Lateral Damage Causes Limb Ataxia 518
Damage to the Flocculus Affects Eye Movements 521
Suggested Readings 521
21 Eye Movements 524
Chapter Outline 524
Six Extraocular Muscles Move the Eye in the Orbit 526
The Medial and Lateral Recti Adduct and Abduct the Eye 527
The Superior and Inferior Recti and the Obliques Have More Complex Actions 527
There Are Fast and Slow Conjugate Eye Movements 529
Fast, Ballistic Eye Movements Get Images Onto the Fovea 530
Motor Programs for Saccades Are Located in the Pons and Midbrain 530
The Frontal Eye Field and Superior Colliculus Trigger Saccades to the Contralateral Side 531
Slow, Guided Eye Movements Keep Images on the Fovea 533
Vestibuloocular and Optokinetic Movements Compensate for Head Movement 533
Smooth Pursuit Movements Compensate for Target Movement 534
Changes in Object Distance Require Vergence Movements 535
The Basal Nuclei and Cerebellum Participate in Eye Movement Control 536
Suggested Readings 539
22 Cerebral Cortex 541
Chapter Outline 541
Most Cerebral Cortex Is Neocortex 542
Pyramidal Cells Are the Most Numerous Neocortical Neurons 543
Neocortex Has Six Layers 543
Different Neocortical Layers Have Distinctive Connections 544
The Corpus Callosum and Anterior Commissure Interconnect the Two Cerebral Hemispheres 547
Association Bundles Interconnect Areas Within Each Cerebral Hemisphere 548
Neocortex Also Has a Columnar Organization 549
Neocortical Areas Are Specialized for Different Functions 549
Different Neocortical Areas Have Subtly Different Structures 549
There Are Sensory, Motor, Association, and Limbic Areas 551
Primary Somatosensory Cortex Is in the Parietal Lobe 553
Primary Visual Cortex Is in the Occipital Lobe 555
Primary Auditory Cortex Is in the Temporal Lobe 555
There Are Primary Vestibular, Gustatory, and Olfactory Areas 556
Most Motor Areas Are in the Frontal Lobe 556
The Right and Left Cerebral Hemispheres Are Specialized for Different Functions 557
Language Areas Border on the Lateral Sulcus, Usually on the Left Hemisphere 558
Parietal Association Cortex Mediates Spatial Orientation 562
Prefrontal Cortex Mediates Working Memory and Decision Making 565
The Corpus Callosum Unites the Two Cerebral Hemispheres 568
Disconnection Syndromes Can Result From White Matter Damage 570
Consciousness and Sleep Are Active Processes 570
There Are Two Forms of Sleep 572
Both Brainstem and Forebrain Mechanisms Regulate Sleep-Wake Transitions 573
Control Circuits for REM Sleep Are Located in the Brainstem 574
Suggested Readings 575
23 Drives and Emotions: 579
Chapter Outline 579
The Hypothalamus Coordinates Drive-Related Behaviors 580
The Hypothalamus Can Be Subdivided in Both Longitudinal and Medial-Lateral Directions 581
Hypothalamic Inputs Arise in Widespread Neural Sites 584
Most Inputs From the Forebrain Arise in Limbic Structures 584
Inputs From the Brainstem and Spinal Cord Traverse the Medial Forebrain Bundle and Dorsal Longitudinal Fasciculus 585
The Hypothalamus Contains Intrinsic Sensory Neurons 586
Hypothalamic Outputs Largely Reciprocate Inputs 586
The Hypothalamus Controls Both Lobes of the Pituitary Gland 586
Perforating Branches From the Circle of Willis Supply the Hypothalamus 588
The Hypothalamus Collaborates With a Network of Brainstem and Spinal Cord Neurons 588
Normal Micturition Involves a Central Pattern Generator in the Pons 590
The Hypothalamus and Associated Central Pattern Generators Keep Physiological Variables Within Narrow Limits 592
Limbic Structures Are Interposed Between the Hypothalamus and Neocortex 593
The Hippocampus and Amygdala Are the Central Components of the Two Major Limbic Subsystems 594
The Amygdala Is Centrally Involved in Emotional Responses 594
The Amygdala Receives a Wide Variety of Sensory Inputs 595
The Amygdala Projects to the Cerebral Cortex and Hypothalamus 599
The Amygdala Is Involved in Emotion-Related Aspects of Learning 599
Bilateral Temporal Lobe Damage Causes a Complex, Devastating Syndrome 603
Suggested Readings 603
24 Formation, Modification, and Repair of Neuronal Connections 605
Chapter Outline 605
Both Neurons and Connections Are Produced in Excess During Development 606
Neurotrophic Factors Ensure That Adequate Numbers of Neurons Survive 606
Axonal Branches Are Pruned to Match Functional Requirements 608
Pruning of Neuronal Connections Occurs During Critical Periods 609
Synaptic Connections Are Adjusted Throughout Life 612
There Are Short-Term and Long-Term Adjustments of Synaptic Strength 612
Multiple Memory Systems Depend on Adjustments of Synaptic Strength 614
Cortical Maps Are Adjusted Throughout Life 614
The Hippocampus and Nearby Cortical Regions Are Critical for Declarative Memory 616
The Hippocampus Is a Cortical Structure That Borders the Inferior Horn of the Lateral Ventricle 617
The Fornix Is a Prominent Output Pathway From the Hippocampus 620
Entorhinal Cortex Is the Principal Source of Inputs to the Hippocampus 620
Hippocampal Outputs Reach Entorhinal Cortex, the Mammillary Body, and the Septal Nuclei 621
Bilateral Damage to the Hippocampus or Medial Diencephalon Impairs Declarative Memory 621
The Amygdala Is Centrally Involved in Emotional Memories 623
The Basal Ganglia Are Important for Some Forms of Nondeclarative Memory 623
The Cerebellum Is Important for Some Forms of Nondeclarative Memory 624
PNS Repair Is More Effective Than CNS Repair 625
Peripheral Nerve Fibers Can Regrow After Injury 625
CNS Glial Cells Impede Repair After Injury 627
Limited Numbers of New Neurons Are Added to the CNS Throughout Life 627
Suggested Readings 630
25 Atlas of the Human Forebrain 632
Glossary 652
Index 679
A 679
B 680
C 682
D 685
E 686
F 686
G 687
H 688
I 689
J 690
K 690
L 690
M 691
N 692
O 694
P 695
Q 697
R 697
S 698
T 700
U 701
V 702
W 703
X 703
Z 703
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