Principles of Neural Science · 5th Edition by Kandel, Eric -- Read -- Imperial Library of Trantor

Index

Principles of Neural Science, Fifth Edition Copyright Page Brief Contents Contents Preface Acknowledgments Contributors Part I Overall Perspective

1 The Brain and Behavior

Two Opposing Views Have Been Advanced on the Relationship Between Brain and Behavior The Brain Has Distinct Functional Regions The First Strong Evidence for Localization of Cognitive Abilities Came from Studies of Language Disorders Affective States Are Also Mediated by Local, Specialized Systems in the Brain Mental Processes Are the End Product of the Interactions Between Elementary Processing Units in the Brain Selected Readings References

2 Nerve Cells, Neural Circuitry, and Behavior

The Nervous System Has Two Classes of Cells

Nerve Cells Are the Signaling Units of the Nervous System Glial Cells Support Nerve Cells

Each Nerve Cell Is Part of a Circuit That Has One or More Specific Behavioral Functions Signaling Is Organized in the Same Way in All Nerve Cells

The Input Component Produces Graded Local Signals The Trigger Zone Makes the Decision to Generate an Action Potential The Conductive Component Propagates an All-or-None Action Potential The Output Component Releases Neurotransmitter The Transformation of the Neural Signal from Sensory to Motor Is Illustrated by the Stretch-Reflex Pathway

Nerve Cells Differ Most at the Molecular Level Neural Network Models Simulate the Brain’s Parallel Processing of Information Neural Connections Can Be Modified by Experience Selected Readings References

3 Genes and Behavior

Genes, Genetic Analysis, and Heritability in Behavior The Nature of the Gene Genes Are Arranged on Chromosomes The Relationship Between Genotype and Phenotype Genes Are Conserved Through Evolution The Role of Genes in Behavior Can Be Studied in Animal Models

Circadian Rhythm Is Generated by a Transcriptional Oscillator in Flies, Mice, and Humans Natural Variation in a Protein Kinase Regulates Activity in Flies and Honeybees The Social Behaviors of Several Species Are Regulated by Neuropeptide Receptors

Genetic Studies of Human Behavior and Its Abnormalities

Neurological Disorders in Humans Suggest That Distinct Genes Affect Different Brain Functions Autism-Related Disorders Exemplify the Complex Genetic Basis of Behavioral Traits

Psychiatric Disorders and the Challenge of Understanding Multigenic Traits

Complex Inheritance and Genetic Imprinting in Human Genetics Multigenic Traits: Many Rare Diseases or a Few Common Variants?

An Overall View Glossary Selected Readings References

Part II Cell and Molecular Biology of the Neuron

4 The Cells of the Nervous System

Neurons and Glia Share Many Structural and Molecular Characteristics The Cytoskeleton Determines Cell Shape Protein Particles and Organelles Are Actively Transported Along the Axon and Dendrites

Fast Axonal Transport Carries Membranous Organelles Slow Axonal Transport Carries Cytosolic Proteins and Elements of the Cytoskeleton

Proteins Are Made in Neurons as in Other Secretory Cells

Secretory and Membrane Proteins Are Synthesized and Modified in the Endoplasmic Reticulum Secretory Proteins Are Modified in the Golgi Complex

Surface Membrane and Extracellular Substances Are Recycled in the Cell Glial Cells Play Diverse Roles in Neural Function

Glia Form the Insulating Sheaths for Axons Astrocytes Support Synaptic Signaling Choroid Plexus and Ependymal Cells Produce Cerebrospinal Fluid Microglia in the Brain Are Derived from Bone Marrow

An Overall View Selected Readings References

5 Ion Channels

Rapid Signaling in the Nervous System Depends on Ion Channels Ion Channels Are Proteins That Span the Cell Membrane Currents Through Single Ion Channels Can Be Recorded Ion Channels in All Cells Share Several Characteristics

The Flux of Ions Through a Channel Is Passive The Opening and Closing of a Channel Involve Conformational Changes

The Structure of Ion Channels Is Inferred from Biophysical, Biochemical, and Molecular Biological Studies

Ion Channels Can Be Grouped into Gene Families The Closed and Open Structures of Potassium Channels Have Been Resolved by X-Ray Crystallography The Structural Basis of Chloride Selectivity Reveals a Close Relation Between Ion Channels and Ion Transporters

An Overall View Selected Readings References

6 Membrane Potential and the Passive Electrical Properties of the Neuron

The Resting Membrane Potential Results from the Separation of Charge Across the Cell Membrane The Resting Membrane Potential Is Determined by Nongated and Gated Ion Channels

Open Channels in Glial Cells Are Permeable to Potassium Only Open Channels in Resting Nerve Cells Are Permeable to Several Ion Species The Electrochemical Gradients of Sodium, Potassium, and Calcium Are Established by Active Transport of the Ions Chloride Ions Are Also Actively Transported

The Balance of Ion Fluxes That Maintains the Resting Membrane Potential Is Abolished During the Action Potential The Contributions of Different Ions to the Resting Membrane Potential Can Be Quantified by the Goldman Equation The Functional Properties of the Neuron Can Be Represented as an Electrical Equivalent Circuit The Passive Electrical Properties of the Neuron Affect Electrical Signaling

Membrane Capacitance Slows the Time Course of Electrical Signals Membrane and Axoplasmic Resistance Affect the Efficiency of Signal Conduction Large Axons Are More Easily Excited Than Small Axons Passive Membrane Properties and Axon Diameter Affect the Velocity of Action Potential Propagation

An Overall View Selected Readings References

7 Propagated Signaling: The Action Potential

The Action Potential Is Generated by the Flow of Ions Through Voltage-Gated Channels

Sodium and Potassium Currents Through Voltage-Gated Channels Are Recorded with the Voltage Clamp Voltage-Gated Sodium and Potassium Conductances Are Calculated from Their Currents The Action Potential Can Be Reconstructed from the Properties of Sodium and Potassium Channels

Variations in the Properties of Voltage-Gated Ion Channels Expand the Signaling Capabilities of Neurons

The Nervous System Expresses a Rich Variety of Voltage-Gated Ion Channels Gating of Voltage-Sensitive Ion Channels Can Be Influenced by Various Cytoplasmic Factors Excitability Properties Vary Between Regions of the Neuron Excitability Properties Vary Between Types of Neurons

The Mechanisms of Voltage-Gating and Ion Permeation Have Been Inferred from Electrophysiological Measurements

Voltage-Gated Sodium Channels Open and Close in Response to Redistribution of Charges Within the Channel Voltage-Gated Sodium Channels Select for Sodium on the Basis of Size, Charge, and Energy of Hydration of the Ion

Voltage-Gated Potassium, Sodium, and Calcium Channels Stem from a Common Ancestor and Have Similar Structures

X-Ray Crystallographic Analysis of Voltage-Gated Channel Structures Provides Insight into Voltage-Gating The Diversity of Voltage-Gated Channel Types Is Generated by Several Genetic Mechanisms

An Overall View Selected Readings References

Part III Synaptic Transmission

8 Overview of Synaptic Transmission

Synapses Are Either Electrical or Chemical Electrical Synapses Provide Instantaneous Signal Transmission

Cells at an Electrical Synapse Are Connected by Gap-Junction Channels Electrical Transmission Allows the Rapid and Synchronous Firing of Interconnected Cells Gap Junctions Have a Role in Glial Function and Disease

Chemical Synapses Can Amplify Signals

Neurotransmitters Bind to Postsynaptic Receptors Postsynaptic Receptors Gate Ion Channels Either Directly or Indirectly

Selected Readings References

9 Signaling at the Nerve-Muscle Synapse: Directly Gated Transmission

The Neuromuscular Junction Is a Well-Studied Example of Directly Gated Synaptic Transmission The Motor Neuron Excites the Muscle by Opening Ligand-Gated Ion Channels at the End-Plate

The End-Plate Potential Is Produced by Ionic Current Through Acetylcholine Receptor-Channels The Ion Channel at the End-Plate Is Permeable to Both Sodium and Potassium

The Current Through Single Acetylcholine Receptor-Channels Can Be Measured Using the Patch Clamp

Individual Receptor-Channels Conduct All-or-None Unitary Currents Four Factors Determine the End-Plate Current

The Molecular Properties of the Acetylcholine Receptor-Channel Are Known An Overall View Postscript: The End-Plate Current Can Be Calculated from an Equivalent Circuit Selected Readings References

10 Synaptic Integration in the Central Nervous System

Central Neurons Receive Excitatory and Inhibitory Inputs Excitatory and Inhibitory Synapses Have Distinctive Ultrastructures Excitatory Synaptic Transmission Is Mediated by Ionotropic Glutamate Receptor-Channels That Are Permeable to Sodium and Potassium

The Excitatory Ionotropic Glutamate Receptors Are Encoded by a Distinct Gene Family Glutamate Receptors Are Constructed from a Set of Modules NMDA and AMPA Receptors Are Organized by a Network of Proteins at the Postsynaptic Density

Inhibitory Synaptic Action Is Usually Mediated by Ionotropic GABA and Glycine Receptor-Channels That Are Permeable to Chloride

Currents Through Single GABA and Glycine Receptor-Channels Can Be Recorded Chloride Currents Through Inhibitory GABAA and Glycine Receptor-Channels Normally Inhibit the Postsynaptic Cell

Ionotropic Glutamate, GABA, and Glycine Receptors Are Transmembrane Proteins Encoded by Two Distinct Gene Families

Ionotropic GABAA and Glycine Receptors Are Homologous to Nicotinic ACh Receptors Some Synaptic Actions Depend on Other Types of Ionotropic Receptors in the Central Nervous System

Excitatory and Inhibitory Synaptic Actions Are Integrated by the Cell into a Single Output

Synaptic Inputs Are Integrated to Fire an Action Potential at the Axon Initial Segment Dendrites Are Electrically Excitable Structures That Can Fire Action Potentials Synapses on a Central Neuron Are Grouped According to Physiological Function

An Overall View Selected Readings References

11 Modulation of Synaptic Transmission: Second Messengers

The Cyclic AMP Pathway Is the Best Understood Second-Messenger Signaling Cascade Initiated by G Protein-Coupled Receptors The Second-Messenger Pathways Initiated by G Protein-Coupled Receptors Share a Common Molecular Logic

A Family of G Proteins Activates Distinct Second-Messenger Pathways Hydrolysis of Phospholipids by Phospholipase C Produces Two Important Second Messengers, IP3 and Diacylglycerol Hydrolysis of Phospholipids by Phospholipase A2 Liberates Arachidonic Acid to Produce Other Second Messengers

Transcellular Messengers Are Important for Regulating Presynaptic Function

Endocannabinoids Are Derived from Arachidonic Acid The Gaseous Second Messengers, Nitric Oxide and Carbon Monoxide, Stimulate Cyclic GMP Synthesis

A Family of Receptor Tyrosine Kinases Mediates Some Metabotropic Receptor Effects The Physiological Actions of Ionotropic and Metabotropic Receptors Differ

Second-Messenger Cascades Can Increase or Decrease the Opening of Many Types of Ion Channels G Proteins Can Modulate Ion Channels Directly Cyclic AMP-Dependent Protein Phosphorylation Can Close Potassium Channels

Synaptic Actions Mediated by Phosphorylation Are Terminated by Phosphoprotein Phosphatases Second Messengers Can Endow Synaptic Transmission with Long-Lasting Consequences An Overall View Selected Readings References

12 Transmitter Release

Transmitter Release Is Regulated by Depolarization of the Presynaptic Terminal Release Is Triggered by Calcium Influx

The Relation Between Presynaptic Calcium Concentration and Release Several Classes of Calcium Channels Mediate Transmitter Release

Transmitter Is Released in Quantal Units Transmitter Is Stored and Released by Synaptic Vesicles

Synaptic Vesicles Discharge Transmitter by Exocytosis and Are Recycled by Endocytosis Capacitance Measurements Provide Insight into the Kinetics of Exocytosis and Endocytosis Exocytosis Involves the Formation of a Temporary Fusion Pore The Synaptic Vesicle Cycle Involves Several Steps

Exocytosis of Synaptic Vesicles Relies on a Highly Conserved Protein Machinery

The Synapsins Are Important for Vesicle Restraint and Mobilization SNARE Proteins Catalyze Fusion of Vesicles with the Plasma Membrane Calcium Binding to Synaptotagmin Triggers Transmitter Release The Fusion Machinery Is Embedded in a Conserved Protein Scaffold at the Active Zone

Modulation of Transmitter Release Underlies Synaptic Plasticity

Activity-Dependent Changes in Intracellular Free Calcium Can Produce Long-Lasting Changes in Release Axo-axonic Synapses on Presynaptic Terminals Regulate Transmitter Release

An Overall View Selected Readings References

13 Neurotransmitters

A Chemical Messenger Must Meet Four Criteria to Be Considered a Neurotransmitter Only a Few Small-Molecule Substances Act as Transmitters

Acetylcholine Biogenic Amine Transmitters Catecholamine Transmitters Serotonin Histamine Amino Acid Transmitters ATP and Adenosine

Small-Molecule Transmitters Are Actively Taken Up into Vesicles Many Neuroactive Peptides Serve as Transmitters Peptides and Small-Molecule Transmitters Differ in Several Ways Peptides and Small-Molecule Transmitters Coexist and Can Be Co-released Removal of Transmitter from the Synaptic Cleft Terminates Synaptic Transmission An Overall View Selected Readings References

14 Diseases of the Nerve and Motor Unit

Disorders of the Peripheral Nerve, Neuromuscular Junction, and Muscle Can Be Distinguished Clinically A Variety of Diseases Target Motor Neurons and Peripheral Nerves

Motor Neuron Diseases Do Not Affect Sensory Neurons Diseases of Peripheral Nerves Affect Conduction of the Action Potential The Molecular Bases of Some Inherited Peripheral Neuropathies Have Been Defined

Diseases of the Neuromuscular Junction Have Multiple Causes

Myasthenia Gravis Is the Best Studied Example of a Neuromuscular Junction Disease Treatment of Myasthenia Targets the Physiological Effects and Autoimmune Pathogenesis of the Disease There Are Two Distinct Congenital Forms of Myasthenia Gravis Lambert-Eaton Syndrome and Botulism Are Two Other Disorders of Neuromuscular Transmission

Diseases of Skeletal Muscle Can Be Inherited or Acquired 320

Dermatomyositis Exemplifies Acquired Myopathy Muscular Dystrophies Are the Most Common Inherited Myopathies Some Inherited Diseases of Skeletal Muscle Arise from Genetic Defects in Voltage-Gated Ion Channels Periodic Paralysis Is Associated with Altered Muscle Excitability and Abnormal Levels of Serum Potassium

An Overall View Postscript: Diagnosis of Motor Unit Disorders Is Aided by Laboratory Criteria Selected Readings References

Part IV The Neural Basis of Cognition

15 The Organization of the Central Nervous System

The Central Nervous System Consists of the Spinal Cord and the Brain The Major Functional Systems Are Similarly Organized

Information Is Transformed at Each Synaptic Relay Neurons at Each Synaptic Relay Are Organized into a Neural Map of the Body Each Functional System Is Hierarchically Organized Functional Systems on One Side of the Brain Control the Other Side of the Body

The Cerebral Cortex Is Concerned with Cognition

Neurons in the Cerebral Cortex Are Organized in Layers and Columns The Cerebral Cortex Has a Large Variety of Neurons

Subcortical Regions of the Brain Are Functionally Organized into Nuclei Modulatory Systems in the Brain Influence Motivation, Emotion, and Memory The Peripheral Nervous System Is Anatomically Distinct from the Central Nervous System An Overall View Selected Readings References

16 The Functional Organization of Perception and Movement

Sensory Information Processing Is Illustrated in the Somatosensory System

Somatosensory Information from the Trunk and Limbs Is Conveyed to the Spinal Cord The Primary Sensory Neurons of the Trunk and Limbs Are Clustered in the Dorsal Root Ganglia The Central Axons of Dorsal Root Ganglion Neurons Are Arranged to Produce a Map of the Body Surface Each Somatic Submodality Is Processed in a Distinct Subsystem from the Periphery to the Brain

The Thalamus Is an Essential Link Between Sensory Receptors and the Cerebral Cortex for All Modalities Except Olfaction Sensory Information Processing Culminates in the Cerebral Cortex Voluntary Movement Is Mediated by Direct Connections Between the Cortex and Spinal Cord An Overall View Selected Readings References

17 From Nerve Cells to Cognition: The Internal Representations of Space and Action

The Major Goal of Cognitive Neural Science Is to Understand Neural Representations of Mental Processes The Brain Has an Orderly Representation of Personal Space

The Cortex Has a Map of the Sensory Receptive Surface for Each Sensory Modality Cortical Maps of the Body Are the Basis of Accurate Clinical Neurological Examinations

The Internal Representation of Personal Space Can Be Modified by Experience Extrapersonal Space Is Represented in the Posterior Parietal Association Cortex Much of Mental Processing Is Unconscious Is Consciousness Accessible to Neurobiological Analysis?

Consciousness Poses Fundamental Problems for a Biological Theory of Mind Neurobiological Research on Cognitive Processes Does Not Depend on a Specific Theory of Consciousness Studies of Binocular Rivalry Have Identified Circuits That May Switch Unconscious to Conscious Visual Perception Selective Attention to Visual Stimuli Can Be Studied on the Cellular Level in Nonhuman Primates How Is Self-Awareness Encoded in the Brain?

An Overall View Selected Readings References

18 The Organization of Cognition

Functionally Related Areas of Cortex Lie Close Together Sensory Information Is Processed in the Cortex in Serial Pathways Parallel Pathways in Each Sensory Modality Lead to Dorsal and Ventral Association Areas

The Dorsal Visual Pathway Carries Spatial Information and Leads to Parietal Association Cortex The Ventral Visual Pathway Processes Information About Form and Leads to Temporal Association Cortex

Goal-Directed Motor Behavior Is Controlled in the Frontal Lobe

Prefrontal Cortex Is Important for the Executive Control of Behavior Dorsolateral Prefrontal Cortex Contributes to Cognitive Control of Behavior Orbital-Ventromedial Prefrontal Cortex Contributes to Emotional Control of Behavior

Limbic Association Cortex Is a Gateway to the Hippocampal Memory System An Overall View Selected Readings References

19 Cognitive Functions of the Premotor Systems

Direct Connections Between the Cerebral Cortex and Spinal Cord Play a Fundamental Role in the Organization of Voluntary Movements The Four Premotor Areas of the Primate Brain Also Have Direct Connections in the Spinal Cord Motor Circuits Involved in Voluntary Actions Are Organized to Achieve Specific Goals The Hand Has a Critical Role in Primate Behavior The Joint Activity of Neurons in the Parietal and Premotor Cortex Encodes Potential Motor Acts

Some Neurons Encode the Possibilities for Interaction with an Object Mirror Neurons Respond to the Motor Actions of Others Potential Motor Acts Are Suppressed or Released by Motor Planning Centers

An Overall View Selected Readings References

20 Functional Imaging of Cognition

Functional Imaging Reflects the Metabolic Demand of Neural Activity

Functional Imaging Emerged from Studies of Blood Flow Functional Imaging Reflects Energy Metabolism

Functional Imaging Is Used to Probe Cognitive Processes

Imaging Perception with and Without Consciousness Imaging Memory with and Without Consciousness Imaging Attentional Modulation of Conscious Perception

Functional Imaging Has Limitations An Overall View Selected Readings References

Part V Perception

21 Sensory Coding

Psychophysics Relates the Physical Properties of Stimuli to Sensations

Psychophysical Laws Govern the Perception of Stimulus Intensity Psychophysical Measurements of Sensation Magnitude Employ Standardized Protocols Sensations Are Quantified Using Probabilistic Statistics Decision Times Are Correlated with Cognitive Processes

Physical Stimuli Are Represented in the Nervous System by Means of the Sensory Code

Sensory Receptors Are Responsive to a Single Type of Stimulus Energy Multiple Subclasses of Sensory Receptors Are Found in Each Sense Organ Neural Firing Patterns Transmit Sensory Information to the Brain The Receptive Field of a Sensory Neuron Conveys Spatial Information

Modality-Specific Pathways Extend to the Central Nervous System

The Receptor Surface Is Represented Topographically in Central Nuclei Feedback Regulates Sensory Coding Top-Down Learning Mechanisms Influence Sensory Processing

An Overall View Selected Readings References

22 The Somatosensory System: Receptors and Central Pathways

The Primary Sensory Neurons of the Somatosensory System Are Clustered in the Dorsal Root Ganglia Peripheral Somatosensory Nerve Fibers Conduct Action Potentials at Different Rates Many Specialized Receptors Are Employed by the Somatosensory System

Mechanoreceptors Mediate Touch and Proprioception Proprioceptors Measure Muscle Activity and Joint Positions Nociceptors Mediate Pain Thermal Receptors Detect Changes in Skin Temperature Itch Is a Distinctive Cutaneous Sensation Visceral Sensations Represent the Status of Various Internal Organs

Somatosensory Information Enters the Central Nervous System Through Cranial and Spinal Nerves Somatosensory Information Flows from the Spinal Cord to the Thalamus Through Parallel Pathways

The Dorsal Column–Medial Lemniscal System Relays Tactile and Proprioceptive Information The Spinothalamic System Conveys Noxious, Thermal, and Visceral Information

The Thalamus Has a Number of Specialized Somatosensory Regions

The Ventral Posterior Nucleus Relays Tactile and Proprioceptive Information Noxious, Thermal, and Visceral Information Is Processed in Several Thalamic Nuclei

An Overall View Selected Readings References

23 Touch

Active and Passive Touch Evoke Similar Responses in Mechanoreceptors The Hand Has Four Types of Mechanoreceptors

Receptive Fields Define the Zone of Tactile Sensitivity Two-Point Discrimination Tests Measure Texture Perception Slowly Adapting Fibers Detect Object Pressure and Form Rapidly Adapting Fibers Detect Motion and Vibration Both Slowly and Rapidly Adapting Fibers Are Important for Grip Control

Tactile Information Is Processed in the Central Touch System

Cortical Receptive Fields Integrate Information from Neighboring Receptors Neurons in the Somatosensory Cortex Are Organized into Functionally Specialized Columns Cortical Columns Are Organized Somatotopically

Touch Information Becomes Increasingly Abstract in Successive Central Synapses

Cognitive Touch Is Mediated by Neurons in the Secondary Somatosensory Cortex Active Touch Engages Sensorimotor Circuits in the Posterior Parietal Cortex

Lesions in Somatosensory Areas of the Brain Produce Specific Tactile Deficits An Overall View Selected Readings References

24 Pain

Noxious Insults Activate Nociceptors Signals from Nociceptors Are Conveyed to Neurons in the Dorsal Horn of the Spinal Cord Hyperalgesia Has Both Peripheral and Central Origins Nociceptive Information Is Transmitted from the Spinal Cord to the Thalamus

Five Major Ascending Pathways Convey Nociceptive Information Several Thalamic Nuclei Relay Nociceptive Information to the Cerebral Cortex

Pain Is Controlled by Cortical Mechanisms

Cingulate and Insular Areas Are Active During the Perception of Pain Pain Perception Is Regulated by a Balance of Activity in Nociceptive and Non-Nociceptive Afferent Fibers Electrical Stimulation of the Brain Produces Analgesia

Opioid Peptides Contribute to Endogenous Pain Control

Endogenous Opioid Peptides and Their Receptors Are Distributed in Pain-Modulatory Systems Morphine Controls Pain by Activating Opioid Receptors Tolerance and Addiction to Opioids Are Distinct Phenomena

An Overall View Selected Readings References

25 The Constructive Nature of Visual Processing

Visual Perception Is a Constructive Process Visual Perception Is Mediated by the Geniculostriate Pathway Form, Color, Motion, and Depth Are Processed in Discrete Areas of the Cerebral Cortex The Receptive Fields of Neurons at Successive Relays in an Afferent Pathway Provide Clues to How the Brain Analyzes Visual Form The Visual Cortex Is Organized into Columns of Specialized Neurons Intrinsic Cortical Circuits Transform Neural Information Visual Information Is Represented by a Variety of Neural Codes An Overall View Selected Readings References

26 Low-Level Visual Processing: The Retina

The Photoreceptor Layer Samples the Visual Image

Ocular Optics Limit the Quality of the Retinal Image There Are Two Types of Photoreceptors: Rods and Cones

Phototransduction Links the Absorption of a Photon to a Change in Membrane Conductance

Light Activates Pigment Molecules in the Photoreceptors Excited Rhodopsin Activates a Phosphodiesterase Through the G Protein Transducin Multiple Mechanisms Shut Off the Cascade Defects in Phototransduction Cause Disease

Ganglion Cells Transmit Neural Images to the Brain

The Two Major Types of Ganglion Cells Are ON Cells and OFF Cells Many Ganglion Cells Respond Strongly to Edges in the Image The Output of Ganglion Cells Emphasizes Temporal Changes in Stimuli Retinal Output Emphasizes Moving Objects Several Ganglion Cell Types Project to the Brain Through Parallel Pathways

A Network of Interneurons Shapes the Retinal Output

Parallel Pathways Originate in Bipolar Cells Spatial Filtering Is Accomplished by Lateral Inhibition Temporal Filtering Occurs in Synapses and Feedback Circuits Color Vision Begins in Cone-Selective Circuits Congenital Color Blindness Takes Several Forms Rod and Cone Circuits Merge in the Inner Retina

The Retina’s Sensitivity Adapts to Changes in Illumination

Light Adaptation Is Apparent in Retinal Processing and Visual Perception Multiple Gain Controls Occur Within the Retina Light Adaptation Alters Spatial Processing

An Overall View Selected Readings References

27 Intermediate-Level Visual Processing and Visual Primitives

Internal Models of Object Geometry Help the Brain Analyze Shapes Depth Perception Helps Segregate Objects from Background Local Movement Cues Define Object Trajectory and Shape Context Determines the Perception of Visual Stimuli

Brightness and Color Perception Depend on Context Receptive-Field Properties Depend on Context

Cortical Connections, Functional Architecture, and Perception Are Intimately Related

Perceptual Learning Requires Plasticity in Cortical Connections Visual Search Relies on the Cortical Representation of Visual Attributes and Shapes Cognitive Processes Influence Visual Perception

An Overall View Selected Readings References

28 High-Level Visual Processing: Cognitive Influences

High-Level Visual Processing Is Concerned with Object Identification The Inferior Temporal Cortex Is the Primary Center for Object Perception

Clinical Evidence Identifies the Inferior Temporal Cortex as Essential for Object Recognition Neurons in the Inferior Temporal Cortex Encode Complex Visual Stimuli Neurons in the Inferior Temporal Cortex Are Functionally Organized in Columns The Inferior Temporal Cortex Is Part of a Network of Cortical Areas Involved in Object Recognition

Object Recognition Relies on Perceptual Constancy Categorical Perception of Objects Simplifies Behavior Visual Memory Is a Component of High-Level Visual Processing 630

Implicit Visual Learning Leads to Changes in the Selectivity of Neuronal Responses Explicit Visual Learning Depends on Linkage of the Visual System and Declarative Memory Formation

Associative Recall of Visual Memories Depends on Top-Down Activation of the Cortical Neurons That Process Visual Stimuli An Overall View Selected Readings References

29 Visual Processing and Action

Successive Fixations Focus Our Attention in the Visual Field

Attention Selects Objects for Further Visual Examination Activity in the Parietal Lobe Correlates with Attention Paid to Objects

The Visual Scene Remains Stable Despite Continual Shifts in the Retinal Image Vision Lapses During Saccades The Parietal Cortex Provides Visual Information to the Motor System An Overall View Selected Readings References

30 The Inner Ear

The Ear Has Three Functional Parts Hearing Commences with the Capture of Sound Energy by the Ear The Hydrodynamic and Mechanical Apparatus of the Cochlea Delivers Mechanical Stimuli to the Receptor Cells

The Basilar Membrane Is a Mechanical Analyzer of Sound Frequency The Organ of Corti Is the Site of Mechanoelectrical Transduction in the Cochlea

Hair Cells Transform Mechanical Energy into Neural Signals

Deflection of the Hair Bundle Initiates Mechanoelectrical Transduction Mechanical Force Directly Opens Transduction Channels Direct Mechanoelectrical Transduction Is Rapid

The Temporal Responsiveness of Hair Cells Determines Their Sensitivity

Hair Cells Adapt to Sustained Stimulation Hair Cells Are Tuned to Specific Stimulus Frequencies Sound Energy Is Mechanically Amplified in the Cochlea

Hair Cells Use Specialized Ribbon Synapses Auditory Information Flows Initially Through the Cochlear Nerve

Bipolar Neurons in the Spiral Ganglion Innervate Cochlear Hair Cells Cochlear Nerve Fibers Encode Stimulus Frequency and Intensity

Sensorineural Hearing Loss Is Common but Treatable An Overall View Selected Readings References

31 The Auditory Central Nervous System

Multiple Types of Information Are Present in Sounds The Neural Representation of Sound Begins in the Cochlear Nuclei

The Cochlear Nerve Imposes a Tonotopic Organization on the Cochlear Nuclei and Distributes Acoustic Information into Parallel Pathways The Ventral Cochlear Nucleus Extracts Information About the Temporal and Spectral Structure of Sounds The Dorsal Cochlear Nucleus Integrates Acoustic with Somatosensory Information in Making Use of Spectral Cues for Localizing Sounds

The Superior Olivary Complex of Mammals Contains Separate Circuits for Detecting Interaural Time and Intensity Differences

The Medial Superior Olive Generates a Map of Interaural Time Differences The Lateral Superior Olive Detects Interaural Intensity Differences

Efferent Signals from the Superior Olivary Complex Provide Feedback to the Cochlea

Brain Stem Pathways Converge in the Inferior Colliculus Sound Location Information from the Inferior Colliculus Creates a Spatial Map of Sound in the Superior Colliculus Midbrain Sound-Localization Pathways Are Sensitive to Experience in Early Life

The Inferior Colliculus Transmits Auditory Information to the Cerebral Cortex

The Auditory Cortex Maps Numerous Aspects of Sound Auditory Information Is Processed in Multiple Cortical Areas Insectivorous Bats Have Cortical Areas Specialized for Behaviorally Relevant Features of Sound A Second Sound-Localization Pathway from the Inferior Colliculus Involves the Cerebral Cortex in Gaze Control Auditory Circuits in the Cerebral Cortex Are Segregated into Separate Processing Streams The Cerebral Cortex Modulates Processing in Subcortical Auditory Areas

Hearing Is Crucial for Vocal Learning and Production in Both Humans and Songbirds

Normal Vocal Behavior Cannot Be Learned in Isolation Vocal Learning Is Optimal During a Sensitive Period Both Humans and Songbirds Possess Specialized Neural Networks for Vocalization Songbirds Have Feature Detectors for Learned Vocalizations

An Overall View Selected Readings References

32 Smell and Taste: The Chemical Senses

A Large Number of Olfactory Receptor Proteins Initiate the Sense of Smell

Mammals Share a Large Family of Odorant Receptors Different Combinations of Receptors Encode Different Odorants

Olfactory Information Is Transformed Along the Pathway to the Brain

Odorants Are Encoded in the Nose by Dispersed Neurons Sensory Inputs in the Olfactory Bulb Are Arranged by Receptor Type The Olfactory Bulb Transmits Information to the Olfactory Cortex Output from the Olfactory Cortex Reaches Higher Cortical and Limbic Areas Olfactory Acuity Varies in Humans

Odors Elicit Characteristic Innate Behaviors

Pheromones Are Detected in Two Olfactory Structures Invertebrate Olfactory Systems Can Be Used to Study Odor Coding and Behavior The Anatomy of the Insect Olfactory System Resembles That of Vertebrates Olfactory Cues Elicit Stereotyped Behaviors and Physiological Responses in the Nematode Strategies for Olfaction Have Evolved Rapidly

The Gustatory System Controls the Sense of Taste

Taste Has Five Submodalities or Qualities Taste Detection Occurs in Taste Buds Each Taste Is Detected by a Distinct Sensory Transduction Mechanism and Distinct Population of Taste Cells Sensory Neurons Carry Taste Information from the Taste Buds to the Brain Taste Information Is Transmitted from the Thalamus to the Gustatory Cortex Perception of Flavor Depends on Gustatory, Olfactory, and Somatosensory Inputs Insect Taste Organs Are Distributed Widely on the Body

An Overall View Selected Readings References

Part VI Movement

33 The Organization and Planning of Movement

Motor Commands Arise Through Sensorimotor Transformations

The Central Nervous System Forms Internal Models of Sensorimotor Transformations Movement Inaccuracies Arise from Errors and Variability in the Transformations Different Coordinate Systems May Be Employed at Different Stages of Sensorimotor Transformations Stereotypical Patterns Are Employed in Many Movements

Motor Signals Are Subject to Feedforward and Feedback Control

Feedforward Control Does Not Use Sensory Feedback Feedback Control Uses Sensory Signals to Correct Movements Prediction Compensates for Sensorimotor Delays Sensory Processing Is Different for Action and Perception

Motor Systems Must Adapt to Development and Experience

Motor Learning Involves Adapting Internal Models for Novel Kinematic and Dynamic Conditions Kinematic and Dynamic Motor Learning Rely on Different Sensory Modalities

An Overall View Selected Readings References

34 The Motor Unit and Muscle Action

The Motor Unit Is the Elementary Unit of Motor Control

A Motor Unit Consists of a Motor Neuron and Multiple Muscle Fibers The Properties of Motor Units Vary Physical Activity Can Alter Motor Unit Properties Muscle Force Is Controlled by the Recruitment and Discharge Rate of Motor Units The Input–Output Properties of Motor Neurons Are Modified by Input from the Brain Stem

Muscle Force Depends on the Structure of Muscle

The Sarcomere Contains the Contractile Proteins Noncontractile Elements Provide Essential Structural Support Contractile Force Depends on Muscle Fiber Activation, Length, and Velocity Muscle Torque Depends on Musculoskeletal Geometry

Different Movements Require Different Activation Strategies

Contraction Velocity Can Vary in Magnitude and Direction Movements Involve the Coordination of Many Muscles Muscle Work Depends on the Pattern of Activation

An Overall View Selected Readings References

35 Spinal Reflexes

Reflexes Are Adaptable to Particular Motor Tasks Spinal Reflexes Produce Coordinated Patterns of Muscle Contraction

Cutaneous Reflexes Produce Complex Movements That Serve Protective and Postural Functions The Stretch Reflex Resists the Lengthening of a Muscle

Local Spinal Circuits Contribute to the Coordination of Reflex Responses

The Stretch Reflex Involves a Monosynaptic Pathway Ia Inhibitory Interneurons Coordinate the Muscles Surrounding a Joint Divergence in Reflex Pathways Amplifies Sensory Inputs and Coordinates Muscle Contractions Convergence of Inputs on Ib Interneurons Increases the Flexibility of Reflex Responses

Central Motor Commands and Cognitive Processes Can Alter Synaptic Transmission in Spinal Reflex Pathways

Central Neurons Can Regulate the Strength of Spinal Reflexes at Three Sites in the Reflex Pathway Gamma Motor Neurons Adjust the Sensitivity of Muscle Spindles

Proprioceptive Reflexes Play an Important Role in Regulating Both Voluntary and Automatic Movements

Reflexes Involving Limb Muscles Are Mediated Through Spinal and Supraspinal Pathways Stretch Reflexes Reinforce Central Commands for Movements

Damage to the Central Nervous System Produces Characteristic Alterations in Reflex Response and Muscle Tone

Interruption of Descending Pathways to the Spinal Cord Frequently Produces Spasticity Transection of the Spinal Cord in Humans Leads to a Period of Spinal Shock Followed by Hyperreflexia

An Overall View Selected Readings References

36 Locomotion

A Complex Sequence of Muscle Contractions Is Required for Stepping The Motor Pattern for Stepping Is Organized at the Spinal Level

Contraction in Flexor and Extensor Muscles of the Hind Legs Is Controlled by Mutually Inhibiting Networks Central Pattern Generators Are Not Driven by Sensory Input Spinal Networks Can Generate Complex Locomotor Patterns

Sensory Input from Moving Limbs Regulates Stepping

Proprioception Regulates the Timing and Amplitude of Stepping Sensory Input from the Skin Allows Stepping to Adjust to Unexpected Obstacles

Descending Pathways Are Necessary for Initiation and Adaptive Control of Stepping

Pathways from the Brain Stem Initiate Walking and Control Its Speed The Cerebellum Fine-Tunes Locomotor Patterns by Regulating the Timing and Intensity of Descending Signals The Motor Cortex Uses Visual Information to Control Precise Stepping Movements Planning and Coordination of Visually Guided Movements Involves the Posterior Parietal Cortex

Human Walking May Involve Spinal Pattern Generators An Overall View Selected Readings References

37 Voluntary Movement: The Primary Motor Cortex

Motor Functions Are Localized within the Cerebral Cortex Many Cortical Areas Contribute to the Control of Voluntary Movements

Voluntary Motor Control Appears to Require Serial Processing The Functional Anatomy of Precentral Motor Areas is Complex The Anatomical Connections of the Precentral Motor Areas Do Not Validate a Strictly Serial Organization

The Primary Motor Cortex Plays an Important Role in the Generation of Motor Commands

Motor Commands Are Population Codes The Motor Cortex Encodes Both the Kinematics and Kinetics of Movement Hand and Finger Movements Are Directly Controlled by the Motor Cortex Sensory Inputs from Somatic Mechanoreceptors Have Feedback, Feed-Forward, and Adaptive Learning Roles The Motor Map Is Dynamic and Adaptable The Motor Cortex Contributes to Motor Skill Learning

An Overall View Selected Readings References

38 Voluntary Movement: The Parietal and Premotor Cortex

Voluntary Movement Expresses an Intention to Act Voluntary Movement Requires Sensory Information About the World and the Body Reaching for an Object Requires Sensory Information About the Object’s Location in Space

Space Is Represented in Several Cortical Areas with Different Sensory and Motor Properties The Inferior Parietal and Ventral Premotor Cortex Contain Representations of Peripersonal Space The Superior Parietal Cortex Uses Sensory Information to Guide Arm Movements Toward Objects in Peripersonal Space Premotor and Primary Motor Cortex Formulate More Specific Motor Plans About Intended Reaching Movements

Grasping an Object Requires Sensory Information About Its Physical Properties

Neurons in the Inferior Parietal Cortex Associate the Physical Properties of an Object with Specific Motor Acts The Activity of Neurons of the Inferior Parietal Cortex Is Influenced by the Purpose of an Action The Activity of Neurons in the Ventral Premotor Cortex Correlates with Motor Acts The Primary Motor Cortex Transforms a Grasping Action Plan into Appropriate Finger Movements

The Supplementary Motor Complex Plays a Crucial Role in Selecting and Executing Appropriate Voluntary Actions The Cortical Motor System Is Involved in Planning Action

Cortical Motor Areas Apply the Rules That Govern Behavior The Premotor Cortex Contributes to Perceptual Decisions That Guide Motor Behavior The Premotor Cortex Is Involved in Learning Motor Skills

Cortical Motor Areas Contribute to Understanding the Observed Actions of Others The Relationship between Motor Acts, the Sense of Volition, and Free Will Is Uncertain An Overall View Selected Readings References

39 The Control of Gaze

Six Neuronal Control Systems Keep the Eyes on Target

An Active Fixation System Keeps the Fovea on a Stationary Target The Saccadic System Points the Fovea Toward Objects of Interest The Smooth-Pursuit System Keeps Moving Targets on the Fovea The Vergence System Aligns the Eyes to Look at Targets at Different Depths

The Eye Is Moved by the Six Extraocular Muscles

Eye Movements Rotate the Eye in the Orbit The Six Extraocular Muscles Form Three Agonist–Antagonist Pairs Movements of the Two Eyes Are Coordinated The Extraocular Muscles Are Controlled by Three Cranial Nerves Extraocular Motor Neurons Encode Eye Position and Velocity

The Motor Circuits for Saccades Lie in the Brain Stem

Horizontal Saccades Are Generated in the Pontine Reticular Formation Vertical Saccades Are Generated in the Mesencephalic Reticular Formation Brain Stem Lesions Result in Characteristic Deficits in Eye Movements

Saccades Are Controlled by the Cerebral Cortex Through the Superior Colliculus

The Superior Colliculus Integrates Visual and Motor Information into Oculomotor Signals to the Brain Stem The Rostral Superior Colliculus Facilitates Visual Fixation The Basal Ganglia Inhibit the Superior Colliculus Two Regions of Cerebral Cortex Control the Superior Colliculus The Control of Saccades Can Be Modified by Experience

Smooth Pursuit Involves the Cerebral Cortex, Cerebellum, and Pons Some Gaze Shifts Require Coordinated Head and Eye Movements An Overall View Selected Readings References

40 The Vestibular System

The Vestibular Apparatus in the Inner Ear Contains Five Receptor Organs

Hair Cells Transduce Mechanical Stimuli into Receptor Potentials The Semicircular Canals Sense Head Rotation The Otolith Organs Sense Linear Accelerations Most Movements Elicit Complex Patterns of Vestibular Stimulation

Vestibulo-Ocular Reflexes Stabilize the Eyes and Body When the Head Moves

The Rotational Vestibulo-Ocular Reflex Compensates for Head Rotation The Otolithic Reflexes Compensate for Linear Motion and Head Deviations Vestibulo-Ocular Reflexes Are Supplemented by Optokinetic Responses

Central Connections of the Vestibular Apparatus Integrate Vestibular, Visual, and Motor Signals

The Vestibular Nerve Carries Information on Head Velocity to the Vestibular Nuclei A Brain Stem Network Connects the Vestibular System with the Oculomotor System Two Visual Pathways Drive the Optokinetic Reflexes The Cerebral Cortex Integrates Vestibular, Visual, and Somatosensory Inputs The Cerebellum Adjusts the Vestibulo-Ocular Reflex

Clinical Syndromes Elucidate Normal Vestibular Function

Unilateral Vestibular Hypofunction Causes Pathological Nystagmus Bilateral Vestibular Hypofunction Interferes with Normal Vision

An Overall View 933 Selected Readings 933 References

41 Posture

Postural Equilibrium and Orientation Are Distinct Sensorimotor Processes Postural Equilibrium Requires Control of the Body’s Center of Mass

Balance During Stance Requires Muscle Activation Automatic Postural Responses Counteract Unexpected Disturbances Automatic Postural Responses Adapt to Changes in the Requirements for Support Anticipatory Postural Adjustments Compensate for Voluntary Movements

Postural Orientation Is Important for Optimizing Execution of Tasks, Interpreting Sensations, and Anticipating Disturbances to Balance Sensory Information from Several Modalities Must Be Integrated to Maintain Equilibrium and Orientation

Somatosensory Afferents Are Important for Timing and Direction of Automatic Postural Responses Vestibular Information Is Important for Balance on Unstable Surfaces and During Head Movements Visual Information Provides Advance Knowledge of Potentially Destabilizing Situations and Assists in Orienting to the Environment Information from a Single Sensory Modality Can Be Ambiguous The Postural Control System Uses a Body Schema that Incorporates Internal Models for Balance The Influence of Each Sensory Modality on Balance and Orientation Changes According to Task Requirements

Control of Posture Is Distributed in the Nervous System

Spinal Cord Circuits Are Sufficient for Maintaining Antigravity Support but Not Balance The Brain Stem and Cerebellum Integrate Sensory Signals for Posture The Spinocerebellum and Basal Ganglia Are Important in Adaptation of Posture Cerebral Cortex Centers Contribute to Postural Control

An Overall View Suggested Readings References

42 The Cerebellum

Cerebellar Diseases Have Distinctive Symptoms and Signs The Cerebellum Has Several Functionally Distinct Regions The Cerebellar Microcircuit Has a Distinct and Regular Organization

Neurons in the Cerebellar Cortex Are Organized into Three Layers Two Afferent Fiber Systems Encode Information Differently Parallel Pathways Compare Excitatory and Inhibitory Signals Recurrent Loops Occur at Several Levels

The Vestibulocerebellum Regulates Balance and Eye Movements The Spinocerebellum Regulates Body and Limb Movements

Somatosensory Information Reaches the Spinocerebellum Through Direct and Indirect Mossy Fiber Pathways The Spinocerebellum Modulates the Descending Motor Systems The Vermis Controls Saccadic and Smooth-Pursuit Eye Movements Spinocerebellar Regulation of Movement Follows Three Organizational Principles Are the Parallel Fibers a Mechanism for Motor Coordination?

The Cerebrocerebellum Is Involved in Planning Movement

The Cerebrocerebellum Is Part of a High-Level Internal Feedback Circuit That Plans Movement and Regulates Cortical Motor Programs Lesions of the Cerebrocerebellum Disrupt Motor Planning and Prolong Reaction Time The Cerebrocerebellum May Have Cognitive Functions Unconnected with Motor Control

The Cerebellum Participates in Motor Learning

Climbing-Fiber Activity Produces Long-Lasting Effects on the Synaptic Efficacy of Parallel Fibers Learning Occurs at Multiple Sites in the Cerebellar Microcircuit

An Overall View Selected Readings References

43 The Basal Ganglia

The Basal Ganglia Consist of Several Interconnected Nuclei A Family of Cortico–Basal Ganglia–Thalamocortical Circuits Subserves Skeletomotor, Oculomotor, Associative, and Limbic Functions The Cortico–Basal Ganglia–Thalamocortical Motor Circuit Originates and Terminates in Cortical Areas Related to Movement

The Motor Circuit Plays a Role in Multiple Aspects of Movement Dopaminergic and Cholinergic Inputs to the Striatum Are Implicated in Reinforcement Motor Learning

Other Basal Ganglia Circuits Are Involved in the Regulation of Eye Movements, Mood, Reward, and Executive Functions Diseases of the Basal Ganglia Are Associated with Disturbances of Movement, Executive Function, Behavior, and Mood

Abnormalities in the Basal Ganglia Motor Circuit Result in a Wide Spectrum of Movement Disorders A Deficiency of Dopamine in the Basal Ganglia Leads to Parkinsonism Reduced and Abnormally Patterned Basal Ganglia Output Results in Hyperkinetic Disorders Abnormal Neuronal Activity in Nonmotor Circuits Is Associated with Several Neuropsychiatric Disorders

An Overall View Selected Readings References

44 Genetic Mechanisms in Degenerative Diseases of the Nervous System

Expanded Trinucleotide Repeats Characterize Several Neurodegenerative Diseases

Huntington Disease Involves Degeneration of the Striatum Spinobulbar Muscular Atrophy Is Due to Abnormal Function of the Androgen Receptor Hereditary Spinocerebellar Ataxias Include Several Diseases with Similar Symptoms but Distinct Etiologies

Parkinson Disease Is a Common Degenerative Disorder of the Elderly Selective Neuronal Loss Occurs After Damage to Ubiquitously Expressed Genes Animal Models Are Powerful Tools for Studying Neurodegenerative Diseases

Mouse Models Reproduce Many Features of Neurodegenerative Diseases Invertebrate Models Manifest Progressive Neurodegeneration

Several Pathways Underlie the Pathogenesis of Neurodegenerative Diseases

Protein Misfolding and Degradation Contribute to Parkinson Disease Protein Misfolding Triggers Pathological Alterations in Gene Expression Mitochondrial Dysfunction Exacerbates Neurodegenerative Disease Apoptosis and Caspase Modify the Severity of Neurodegeneration

Advances in Understanding the Molecular Basis of Neurodegenerative Diseases Are Opening Possibilities for Approaches to Therapeutic Intervention An Overall View Selected Readings References

Part VII The Unconscious and Conscious Processing of Neural Information

45 The Sensory, Motor, and Reflex Functions of the Brain Stem

The Cranial Nerves Are Homologous to the Spinal Nerves

Cranial Nerves Mediate the Sensory and Motor Functions of the Face and Head and the Autonomic Functions of the Body Cranial Nerves Leave the Skull in Groups and Often Are Injured Together

Cranial Nerve Nuclei in the Brain Stem Are Organized on the Same Basic Plan As Are Sensory and Motor Regions of the Spinal Cord

Adult Cranial Nerve Nuclei Have a Columnar Organization Embryonic Cranial Nerve Nuclei Have a Segmental Organization The Organization of the Brain Stem and Spinal Cord Differs in Three Important Ways

Neuronal Ensembles in the Brain Stem Reticular Formation Coordinate Reflexes and Simple Behaviors Necessary for Homeostasis and Survival

Cranial Nerve Reflexes Involve Mono- and Polysynaptic Brain Stem Relays Pattern Generator Neurons Coordinate Stereotypic and Autonomic Behaviors A Complex Pattern Generator Regulates Breathing

An Overall View Selected Readings References

46 The Modulatory Functions of the Brain Stem

Ascending Monoaminergic and Cholinergic Projections from the Brain Stem Maintain Arousal Monoaminergic and Cholinergic Neurons Share Many Properties and Functions

Many Monoaminergic and Cholinergic Neurons Are Linked to the Sleep-Wake Cycle Monoaminergic and Cholinergic Neurons Maintain Arousal by Modulating Neurons in the Thalamus and Cortex

Monoamines Regulate Many Brain Functions Other Than Arousal

Cognitive Performance Is Optimized by Ascending Projections from Monoaminergic Neurons Monoamines Are Involved in Autonomic Regulation and Breathing Pain and Anti-nociceptive Pathways Are Modulated by Monoamines Monoamines Facilitate Motor Activity

An Overall View Postscript: Evaluation of the Comatose Patient Selected Readings References

47 The Autonomic Motor System and the Hypothalamus

The Autonomic Motor System Mediates Homeostasis The Autonomic System Contains Visceral Motor Neurons That Are Organized into Ganglia

Preganglionic Neurons Are Localized in Three Regions Along the Brain Stem and Spinal Cord Sympathetic Ganglia Project to Many Targets Throughout the Body Parasympathetic Ganglia Innervate Single Organs The Enteric Ganglia Regulate the Gastrointestinal Tract

Both the Pre- and Postsynaptic Neurons of the Autonomic Motor System Use Co-Transmission at Their Synaptic Connections Autonomic Behavior Is the Product of Cooperation Between All Three Autonomic Divisions Autonomic and Endocrine Function Is Coordinated by a Central Autonomic Network Centered in the Hypothalamus The Hypothalamus Integrates Autonomic, Endocrine, and Behavioral Responses

Magnocellular Neuroendocrine Neurons Control the Pituitary Gland Directly 1072 Parvicellular Neuroendocrine Neurons Control the Pituitary Gland Indirectly 1072 An Overall View 1076 Selected Readings 1076 References

48 Emotions and Feelings

The Modern Search for the Emotional Brain Began in the Late 19th Century The Amygdala Emerged as a Critical Regulatory Site in Circuits of Emotions

Studies of Avoidance Conditioning First Implicated the Amygdala in Fear Responses Pavlovian Conditioning Is Used Extensively to Study the Contribution of the Amygdala to Learned Fear The Amygdala Has Been Implicated in Unconditioned (Innate) Fear in Animals The Amygdala Is Also Important for Fear in Humans The Amygdala Is Involved in Positive Emotions in Animals and Humans

Other Brain Areas Contribute to Emotional Processing The Neural Correlates of Feeling Are Beginning to Be Understood An Overall View Selected Readings References

49 Homeostasis, Motivation, and Addictive States

Drinking Occurs Both in Response to and in Anticipation of Dehydration

Body Fluids in the Intracellular and Extracellular Compartments Are Regulated Differentially The Intravascular Compartment Is Monitored by Parallel Endocrine and Neural Sensors The Intracellular Compartment Is Monitored by Osmoreceptors Motivational Systems Anticipate the Appearance and Disappearance of Error Signals

Energy Stores Are Precisely Regulated

Leptin and Insulin Contribute to Long-Term Energy Balance Long-Term and Short-Term Signals Interact to Control Feeding

Motivational States Influence Goal-Directed Behavior

Both Internal and External Stimuli Contribute to Motivational States Motivational States Serve Both Regulatory and Nonregulatory Needs Brain Reward Circuitry May Provide a Common Logic for Goal Selection

Drug Abuse and Addiction Are Goal-Directed Behaviors

Addictive Drugs Recruit the Brain’s Reward Circuitry Addictive Drugs Alter the Long-Term Functioning of the Nervous System Dopamine May Act As a Learning Signal

An Overall View Selected Readings References

50 Seizures and Epilepsy

Classification of Seizures and the Epilepsies Is Important for Pathogenesis and Treatment

Seizures Are Temporary Disruptions of Brain Function Epilepsy Is the Chronic Condition of Recurrent Seizures

The Electroencephalogram Represents the Collective Behavior of Cortical Neurons Focal Seizures Originate Within a Small Group of Neurons Known as a Seizure Focus

Neurons in a Seizure Focus Have Characteristic Activity The Breakdown of Surround Inhibition Leads to Synchronization The Spread of Focal Seizures Involves Normal Cortical Circuitry

Primary Generalized Seizures Are Driven by Thalamocortical Circuits Locating the Seizure Focus Is Critical to the Surgical Treatment of Epilepsy Prolonged Seizures Can Cause Brain Damage

Repeated Convulsive Seizures Are a Medical Emergency Excitotoxicity Underlies Seizure-Related Brain Damage

The Factors Leading to Development of Epilepsy Are an Unfolding Mystery

Among the Genetic Causes of Epilepsy Are Ion Channel Mutations Epilepsies Involving Focal Seizures May Be a Maladaptive Response to Injury

An Overall View Selected Readings References

51 Sleep and Dreaming

Sleep Consists of Alternating REM and Non-REM Periods

Non-REM Sleep Has Four Stages REM and Non-REM Dreams Are Different

Sleep Obeys Circadian and Ultradian Rhythms

The Circadian Rhythm Clock Is Based on a Cyclic Production of Nuclear Transcription Factors The Ultradian Rhythm of Sleep Is Controlled by the Brain Stem Sleep-Related Activity in the EEG Is Generated Through Local and Long-Range Circuits

Sleep Changes with Age The Characteristics of Sleep Vary Greatly Between Species Sleep Disorders Have Behavioral, Psychological, and Neurological Causes

Insomnia Is the Most Common Form of Sleep Disruption Excessive Daytime Sleepiness Is Indicative of Disrupted Sleep The Disruption of Breathing During Sleep Apnea Results in Fragmentation of Sleep Narcolepsy Is Characterized by Abnormal Activation of Sleep Mechanisms Restless Leg Syndrome and Periodic Leg Movements Disrupt Sleep Parasomnias Include Sleep Walking, Sleep Talking, and Night Terrors Circadian Rhythm Sleep Disorders Are Characterized by an Activity Cycle That Is Out of Phase with the World

An Overall View Selected Readings References

Part VIII Development and the Emergence of Behavior

52 Patterning the Nervous System

The Neural Tube Becomes Regionalized Early in Embryogenesis Secreted Signals Promote Neural Cell Fate

Development of the Neural Plate Is Induced by Signals from the Organizer Region Neural Induction Is Mediated by Peptide Growth Factors and Their Inhibitors

Rostrocaudal Patterning of the Neural Tube Involves Signaling Gradients and Secondary Organizing Centers

Signals from the Mesoderm and Endoderm Define the Rostrocaudal Pattern of the Neural Plate Signals from Organizing Centers within the Neural Tube Pattern the Forebrain, Midbrain, and Hindbrain

Dorsoventral Patterning of the Neural Tube Involves Similar Mechanisms at Different Rostrocaudal Levels

The Ventral Neural Tube Is Patterned by Sonic Hedgehog Protein Secreted from the Notochord and Floor Plate The Dorsal Neural Tube Is Patterned by Bone Morphogenetic Proteins Dorsoventral Patterning Mechanisms Are Conserved Along the Rostrocaudal Extent of the Neural Tube

Local Signals Determine Functional Subclasses of Neurons

Rostrocaudal Position Is a Major Determinant of Motor Neuron Subtype Local Signals and Transcriptional Circuits Further Diversify Motor Neuron Subtypes

The Developing Forebrain Is Patterned by Intrinsic and Extrinsic Influences

Inductive Signals and Transcription Factor Gradients Establish Regional Differentiation Afferent Inputs Also Contribute to Regionalization

An Overall View Selected Readings References

53 Differentiation and Survival of Nerve Cells

The Proliferation of Neural Progenitor Cells Involves Symmetric and Asymmetric Modes of Cell Division Radial Glial Cells Serve As Neural Progenitors and Structural Scaffolds The Generation of Neurons or Glial Cells Is Regulated by Delta-Notch Signaling and Basic Helix-Loop-Helix Transcription Factors Neuronal Migration Establishes the Layered Organization of the Cerebral Cortex Central Neurons Migrate Along Glial Cells and Axons to Reach Their Final Settling Position

Glial Cells Serve As a Scaffold in Radial Migration Axon Tracts Serve As a Scaffold for Tangential Migration Neural Crest Cell Migration in the Peripheral Nervous System Does Not Rely on Scaffolding

The Neurotransmitter Phenotype of a Neuron Is Plastic

The Transmitter Phenotype of a Peripheral Neuron Is Influenced by Signals from the Neuronal Target The Transmitter Phenotype of a Central Neuron Is Controlled by Transcription Factors

The Survival of a Neuron Is Regulated by Neurotrophic Signals from the Neuron’s Target

The Neurotrophic Factor Hypothesis Was Confirmed by the Discovery of Nerve Growth Factor Neurotrophins Are the Best Studied Neurotrophic Factors Neurotrophic Factors Suppress a Latent Death Program in Cells

An Overall View Selected Readings References

54 The Growth and Guidance of Axons

Differences in the Molecular Properties of Axons and Dendrites Emerge Early in Development

Neuronal Polarity Is Established Through Rearrangements of the Cytoskeleton Dendrites Are Patterned by Intrinsic and Extrinsic Factors

The Growth Cone Is a Sensory Transducer and a Motor Structure Molecular Cues Guide Axons to Their Targets The Growth of Retinal Ganglion Axons Is Oriented in a Series of Discrete Steps

Growth Cones Diverge at the Optic Chiasm Ephrins Provide Gradients of Inhibitory Signals in the Brain

Axons from Some Spinal Neurons Cross the Midline

Netrins Direct Developing Commissural Axons Across the Midline Chemoattractant and Chemorepellent Factors Pattern the Midline

An Overall View Selected Readings References

55 Formation and Elimination of Synapses

Recognition of Synaptic Targets Is Specific

Recognition Molecules Promote Selective Synapse Formation Different Synaptic Inputs Are Directed to Discrete Domains of the Postsynaptic Cell Neural Activity Sharpens Synaptic Specificity

Principles of Synaptic Differentiation Are Revealed at the Neuromuscular Junction

Differentiation of Motor Nerve Terminals Is Organized by Muscle Fibers Differentiation of the Postsynaptic Muscle Membrane Is Organized by the Motor Nerve The Nerve Regulates Transcription of Acetylcholine Receptor Genes The Neuromuscular Junction Matures in a Series of Steps

Central Synapses Develop in Ways Similar to Neuromuscular Junctions

Neurotransmitter Receptors Become Localized at Central Synapses Synaptic Organizing Molecules Pattern Central Nerve Terminals Glial Cells Promote Synapse Formation

Some Synapses Are Eliminated After Birth An Overall View Selected Readings References

56 Experience and the Refinement of Synaptic Connections

Development of Human Mental Function Is Influenced by Early Experience

Early Experience Has Lifelong Effects on Social Behaviors Development of Visual Perception Requires Visual Experience

Development of Binocular Circuits in the Visual Cortex Depends on Postnatal Activity

Visual Experience Affects the Structure and Function of the Visual Cortex Patterns of Electrical Activity Organize Binocular Circuits in the Visual Cortex

Reorganization of Visual Circuits During a Critical Period Involves Alterations in Synaptic Connections

Reorganization Depends on a Change in the Balance of Excitatory and Inhibitory Inputs Postsynaptic Structures Are Rearranged During the Critical Period Thalamic Inputs Are Also Remodeled Synaptic Stabilization Contributes to Closing the Critical Period

Segregation of Retinal Inputs in the Lateral Geniculate Nucleus Is Driven by Spontaneous Neural Activity In Utero Activity-Dependent Refinement of Connections Is a General Feature of Circuits in the Central Nervous System

Many Aspects of Visual System Development Are Activity-Dependent Auditory Maps Are Refined During a Critical Period Distinct Regions of the Brain Have Different Critical Periods of Development

Critical Periods Can Be Reopened in Adulthood An Overall View Selected Readings References

57 Repairing the Damaged Brain

Damage to Axons Affects Neurons and Neighboring Cells

Axon Degeneration Is an Active Process Axotomy Leads to Reactive Responses in Nearby Cells

Central Axons Regenerate Poorly After Injury Therapeutic Interventions May Promote Regeneration of Injured Central Neurons

Environmental Factors Support the Regeneration of Injured Axons Components of Myelin Inhibit Neurite Outgrowth Injury-Induced Scarring Hinders Axonal Regeneration An Intrinsic Growth Program Promotes Regeneration Formation of New Connections by Intact Axons Can Lead to Functional Recovery

Neurons in the Injured Brain Die but New Ones Can Be Born

Therapeutic Interventions May Retain or Replace Injured Central Neurons Transplantation of Neurons or Their Progenitors Can Replace Lost Neurons Stimulation of Neurogenesis in Regions of Injury May Contribute to Restoring Function Transplantation of Nonneuronal Cells or Their Progenitors Can Improve Neuronal Function Restoration of Function Is the Aim of Regenerative Therapies

An Overall View Selected Readings References

58 Sexual Differentiation of the Nervous System

Genes and Hormones Determine Physical Differences Between Males and Females

Chromosomal Sex Directs the Gonadal Differentiation of the Embryo Gonads Synthesize Hormones That Promote Sexual Differentiation Steroid Hormones Act by Binding to Specific Receptors

Sexual Differentiation of the Nervous System Generates Sexually Dimorphic Behaviors

A Sexually Dimorphic Neural Circuit Controls Erectile Function A Sexually Dimorphic Neural Circuit Controls Song Production in Birds A Sexually Dimorphic Neural Circuit in the Hypothalamus Controls Mating Behavior

Environmental Cues Control Some Sexually Dimorphic Behaviors

Pheromones Control Partner Choice in Mice Early Experience Modifies Later Maternal Behavior

Sexual Dimorphism in the Human Brain May Correlate with Gender Identity and Sexual Orientation An Overall View Selected Readings References

59 The Aging Brain

The Structure and Function of the Brain Change with Age Cognitive Decline Is Dramatic in a Small Percentage of the Elderly Alzheimer Disease Is the Most Common Senile Dementia The Brain in Alzheimer Disease Is Altered by Atrophy, Amyloid Plaques, and Neurofibrillary Tangles

Amyloid Plaques Contain Toxic Peptides That Contribute to Alzheimer Pathology Neurofibrillary Tangles Contain Microtubule-Associated Proteins Risk Factors for Alzheimer Disease Have Been Identified

Alzheimer Disease Can Be Diagnosed Well but Available Treatments Are Poor An Overall View Selected Readings References

Part IX Language, Thought, Affect, and Learning

60 Language

Language Has Many Functional Levels: Phonemes, Morphemes, Words, and Sentences Language Acquisition in Children Follows a Universal Pattern

The “Universalist” Infant Becomes Linguistically Specialized by Age 1 Year Language Uses the Visual System Prosodic Cues Assist Learning of Words and Sentences Infants Use Transitional Probabilities to Identify Words in Continuous Speech There Is a Critical Period for Language Learning “Motherese” Enhances Language Learning

Several Cortical Regions Are Involved in Language Processing

Language Circuits in the Brain Were First Identified in Studies of Aphasia The Left Hemisphere Is Specialized for Phonetic, Word, and Sentence Processing Prosody Engages Both Right and Left Hemispheres Depending on the Information Conveyed Language Processing in Bilinguals Depends on Age of Acquisition and Language Use The Model for the Neural Basis of Language Is Changing

Brain Injuries Responsible for the Aphasias Provide Important Insights into Language Processing

Broca Aphasia Results from a Large Lesion in the Left Frontal Lobe Wernicke Aphasia Results from Damage to Left Posterior Temporal Lobe Structures Conduction Aphasia Results from Damage to a Specific Sector of Posterior Language Areas Global Aphasia Results from Widespread Damage to Several Language Centers Transcortical Aphasias Result from Damage to Areas Near Broca’s and Wernicke’s Areas The Classical Aphasias Have Not Implicated All Brain Areas Important for Language

An Overall View Selected Readings References

61 Disorders of Conscious and Unconscious Mental Processes

Conscious and Unconscious Cognitive Processes Have Distinctive Neural Correlates Differences Between Conscious Processes in Perception Can Be Seen in Exaggerated Form after Brain Damage The Control of Action Is Largely Unconscious The Conscious Recall of Memory Is a Creative Process Behavioral Observation Needs to Be Supplemented with Subjective Reports

Brain Imaging Can Corroborate Subjective Reports Malingering and Hysteria Can Lead to Unreliable Subjective Reports

An Overall View Selected Readings References

62 Disorders of Thought and Volition: Schizophrenia

Diagnosis of Schizophrenia Is Based on Standardized Clinical Criteria

The Symptoms of Schizophrenia Can Be Grouped into Positive, Negative, and Cognitive Schizophrenia Is Characterized by Psychotic Episodes

Both Genetic and Nongenetic Risk Factors Contribute to Schizophrenia Neuroanatomic Abnormalities May Be a Causative Factor in Schizophrenia

Loss of Gray Matter in the Cerebral Cortex Appears to Result from Loss of Synaptic Contacts Rather Than Loss of Cells Abnormalities in Brain Development During Adolescence May Contribute to Schizophrenia

Antipsychotic Drugs Act on Dopaminergic Systems in the Brain An Overall View Selected Readings References

63 Disorders of Mood and Anxiety

The Most Common Disorders of Mood Are Unipolar Depression and Bipolar Disorder

Unipolar Depression Often Begins Early in Life Bipolar Disorder Includes Episodes of Mania Mood Disorders Are Common and Disabling

Both Genetic and Nongenetic Risk Factors Play an Important Role in Mood Disorders Specific Brain Regions and Circuits Are Involved in Mood Disorders Depression and Stress Are Interrelated Major Depression Can Be Treated Effectively

Antidepressant Drugs Target Monoaminergic Neural Systems Psychotherapy Is Effective in the Treatment of Major Depression Electroconvulsive Therapy Is Highly Effective Against Depression Bipolar Disorder Can Be Treated with Lithium and Several Drugs Initially Developed a Anticonvulsants

Anxiety Disorders Stem from Abnormal Regulation of Fear

Anxiety Disorders Have a Genetic Component Animal Models of Fear May Shed Light on Human Anxiety Disorders Neuro-imaging Implicates Amygdala-Based Circuits in Human Fear and Anxiety Anxiety Disorders Can Be Treated Effectively with Medications and Psychotherapy

An Overall View Selected Readings References

64 Autism and Other Neurodevelopmental Disorders Affecting Cognition

Autism Has Characteristic Behavioral Features There Is a Strong Genetic Component in Autism Autism Has Characteristic Neurological Abnormalities There Are Distinctive Cognitive Abnormalities in Autism

Social Communication Is Impaired: The Mind Blindness Hypothesis Other Social Mechanisms Contribute to Autism People with Autism Show a Lack of Behavioral Flexibility Some People with Autism Have Special Talents

Some Neurodevelopmental Disorders Have a Known Genetic Basis

Fragile X Syndrome Rett Syndrome Down Syndrome Prader-Willi and Angelman Syndrome and Other Disorders

An Overall View Selected Readings References

65 Learning and Memory

Short-Term and Long-Term Memory Involve Different Neural Systems

Short-Term Memory Maintains Transient Representations of Information Relevant to Immediate Goals Short-Term Memory Is Selectively Transferred to Long-Term Memory

Long-Term Memory Can Be Classified As Explicit or Implicit Explicit Memory Has Episodic and Semantic Forms

Explicit Memory Processing Involves at Least Four Distinct Operations Episodic Knowledge Depends on Interaction Between the Medial Temporal Lobe and Association Cortices Semantic Knowledge Is Stored in Distinct Association Cortices and Retrieval Depends on the Prefrontal Cortex

Implicit Memory Supports Perceptual Priming

Implicit Memory Can Be Associative or Nonassociative Classical Conditioning Involves Associating Two Stimuli Operant Conditioning Involves Associating a Specific Behavior with a Reinforcing Event Associative Learning Is Constrained by the Biology of the Organism

Errors and Imperfections in Memory Shed Light on Normal Memory Processes An Overall View Selected Readings References

66 Cellular Mechanisms of Implicit Memory Storage and the Biological Basis of Individuality

Storage of Implicit Memory Involves Changes in the Effectiveness of Synaptic Transmission

Habituation Results from an Activity-Dependent Presynaptic Depression of Synaptic Transmission Sensitization Involves Presynaptic Facilitation of Synaptic Transmission Classical Conditioning of Fear Involves Coordinated Pre- and Postsynaptic Facilitation of Synaptic Transmission

Long-Term Storage of Implicit Memory Involves Changes in Chromatin Structure and Gene Expression Mediated by the cAMP-PKA-CREB Pathway

Cyclic AMP Signaling Has a Role in Long-Term Sensitization Long-Term Synaptic Facilitation Is Synapse Specific Long-Term Facilitation Requires a Prion-Like Protein Regulator of Local Protein Synthesis for Maintenance

Classical Fear Conditioning in Flies Uses the cAMP-PKACREB Pathway Memory for Learned Fear in Mammals Involves the Amygdala Habit Learning and Memory Require the Striatum Learning-Induced Changes in the Structure of the Brain Contribute to the Biological Basis of Individuality An Overall View Selected Readings References

67 Prefrontal Cortex, Hippocampus, and the Biology of Explicit Memory Storage

Working Memory Depends on Persistent Neural Activity in the Prefrontal Cortex

Intrinsic Membrane Properties Can Generate Persistent Activity Network Connections Can Sustain Activity Working Memory Depends on the Modulatory Transmitter Dopamine

Explicit Memory in Mammals Involves Different Forms of Long-Term Potentiation in the Hippocampus

Long-Term Potentiation in the Mossy Fiber Pathway Is Nonassociative Long-Term Potentiation in the Schaffer Collateral Pathway Is Associative Long-Term Potentiation in the Schaffer Collateral Pathway Follows Hebbian Learning Rules Long-Term Potentiation Has Early and Late Phases

Spatial Memory Depends on Long-Term Potentiation in the Hippocampus A Spatial Map of the External World Is Formed in the Hippocampus Different Subregions of the Hippocampus Are Required for Pattern Separation and for Pattern Completion Memory Also Depends on Long-Term Depression of Synaptic Transmission Epigenetic Changes in Chromatin Structure Are Important for Long-Term Synaptic Plasticity and Learning and Memory Are There Molecular Building Blocks for Learning? An Overall View Selected Readings References

Appendices

A Review of Basic Circuit Theory

Basic Electrical Parameters

Potential Difference (V or E) Current (I) Conductance (g) Capacitance (C)

Rules for Circuit Analysis

Conductance Current Capacitance Potential Difference

Current in Circuits with Capacitance

Circuit with Capacitor Circuit with Resistor and Capacitor in Series Circuit with Resistor and Capacitor in Parallel

B The Neurological Examination of the Patient

Mental Status

Alertness and Attentiveness Behavior, Mood, and Thought Orientation and Memory Cognitive Abilities Language Disorders

Cranial Nerve Function

Olfactory Nerve (Cranial N. I) Optic Nerve (Cranial N. II) Oculomotor, Trochlear, and Abducens Nerves (Cranial N. III, IV, VI) Trigeminal Nerve (Cranial N. V) Facial Nerve (Cranial N. VII) Vestibulocochlear Nerve (Cranial N. VIII) Glossopharyngeal and Vagus Nerves (Cranial N. IX, X) Spinal Accessory Nerve Hypoglossal Nerve (Cranial N. XII)

Musculoskeletal System Sensory Systems Motor Coordination Gait and Stance Balance Deep Tendon Reflexes

C Circulation of the Brain

The Blood Supply of the Brain Can Be Divided into Arterial Territories The Cerebral Vessels Have Unique Physiological Responses A Stroke Is the Result of Disease Involving Blood Vessels Clinical Vascular Syndromes May Follow Vessel Occlusion, Hypoperfusion, or Hemorrhage

Infarction Can Occur in the Middle Cerebral Artery Territory Infarction Can Occur in the Anterior Cerebral Artery Territory Infarction Can Occur in the Posterior Cerebral Artery Territory The Anterior Choroidal and Penetrating Arteries Can Become Occluded The Carotid Artery Can Become Occluded The Brain Stem and Cerebellum Are Supplied by Branches of the Vertebral and Basilar Arteries Infarcts Affecting Predominantly Medial or Lateral Brain Stem Structures Produce Characteristic Syndromes Infarction Can Be Restricted to the Cerebellum Infarction Can Affect the Spinal Cord Diffuse Hypoperfusion Can Cause Ischemia or Infarction Cerebrovascular Disease Can Cause Dementia The Rupture of Microaneurysms Causes Intraparenchymal Stroke The Rupture of Saccular Aneurysms Causes Subarachnoid Hemorrhage

Stroke Alters the Vascular Physiology of the Brain Selected Readings

D The Blood–Brain Barrier, Choroid Plexus, and Cerebrospinal Fluid

The Blood–Brain Barrier Regulates the Interstitial Fluid in the Brain

Distinctive Properties of the Endothelial Cells of Brain Capillaries Account for the Blood–Brain Barrier Tight Junctions Are a Major Feature of the Anatomical Blood–Brain Barrier Composition and Structure The Blood–Brain Barrier Is Permeable in Three Ways Endothelial Enzyme Systems Form a Metabolic Blood–Brain Barrier Some Areas of the Brain Lack a Blood–Brain Barrier Brain-Derived Signals Induce Endothelial Cells to Express a Blood–Brain Barrier Diseases Can Alter the Blood–Brain Barrier

Cerebrospinal Fluid Is Secreted by the Choroid Plexuses

Cerebrospinal Fluid Has Several Functions Epithelial Cells of the Choroid Plexuses Account for the Blood–Cerebral Spinal Fluid Barrier Choroid Plexuses Nurture the Developing Brain Increased Intracranial Pressure May Harm the Brain

Brain Edema Is an Increase in Brain Volume Because of Increased Water Content Hydrocephalus Is an Increase in the Volume of the Cerebral Ventricles Selected Readings References

E Neural Networks

Early Neural Network Modeling Neurons Are Computational Devices

A Neuron Can Compute Conjunctions and Disjunctions A Network of Neurons Can Compute Any Boolean Logical Function

Perceptrons Model Sequential and Parallel Computation in the Visual System

Simple and Complex Cells Could Compute Conjunctions and Disjunctions The Primary Visual Cortex Has Been Modeled As a Multilayer Perceptron Selectivity and Invariance Must Be Explained by Any Model of Vision Visual Object Recognition Could Be Accomplished by Iteration of Conjunctions and Disjunctions

Associative Memory Networks Use Hebbian Plasticity to Store and Recall Neural Activity Patterns

Hebbian Plasticity May Store Activity Patterns by Creating Cell Assemblies Cell Assemblies Can Complete Activity Patterns Cell Assemblies Can Maintain Persistent Activity Patterns Interference Between Memories Limits Capacity Synaptic Loops Can Lead to Multiple Stable States Symmetric Networks Minimize Energy-Like Functions Hebbian Plasticity May Create Sequential Synaptic Pathways

An Overall View Selected Readings References

F Theoretical Approaches to Neuroscience: Examples from Single Neurons to Networks

Single-Neuron Models Allow Study of the Integration of Synaptic Inputs and Intrinsic Conductances

Neurons Show Sharp Threshold Sensitivity to the Number and Synchrony of Synaptic Inputs in Quiet Conditions Resembling In Vivo Neurons Show Graded Sensitivity to the Number and Synchrony of Synaptic Inputs in Noisy Conditions Resembling In Vitro Neuronal Messages Depend on Intrinsic Activity and Extrinsic Signals

Network Models Provide Insight into the Collective Dynamics of Neurons

Balanced Networks of Active Neurons Can Generate the Ongoing Noisy Activity Seen In Vivo Feed-forward and Recurrent Networks Can Amplify or Integrate Inputs with Distinct Dynamics Balanced Recurrent Networks Can Behave Like Feed-forward Networks Paradoxical Effects in Balanced Recurrent Networks May Underlie Surround Suppression in the Visual Cortex Recurrent Networks Can Model Decision-Making

Selected Reading References

Index Footnotes

Contribution Chapter 3 Chapter 6 Chapter 7 Chapter 9 Chapter 10 Chapter 47 part05fn part06fn appafn appefn

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