Neuroscience

Cover
Front Endpaperss
Front Matter
	Copyright Page
	Contents in Brief
	Contributors and Unit Editors
	Contents
	Preface
	Acknowledgments
	Media and Supplements
Chapter 1 Studying the Nervous System
	Overview
	Genetics and Genomics
	Cellular Components of the Nervous System
	Neurons
	Glial Cells
	Cellular Diversity in the Nervous System
	Neural Circuits
	Other Ways to Study Neural Circuits
	Organization of the Human Nervous System
	Neural Systems
	Genetic Analysis of Neural Systems
	BOX 1A Model Organisms in Neuroscience
	Structural Analysis of Neural Systems
	Functional Analysis of Neural Systems
	Analyzing Complex Behavior
	Imaging the Living Human Brain
	Early Brain Imaging Using X-Rays
	Early Functional Mapping Using Cortical Surface Stimulation and Electroencephalography
	Computerized Tomography
	Magnetic Resonance Imaging
	Functional Brain Imaging
	Summary
Chapter 2 Electrical Signals of Nerve Cells
	Overview
	Electrical Signals of Nerve Cells
	Long-Distance Transmission of Electrical Signals
	How Ion Movements Produce Electrical Signals
	Forces That Create Membrane Potentials
	Electrochemical Equilibrium in an Environment with More Than One Permeant Ion
	The Ionic Basis of the Resting Membrane Potential
	BOX 2A The Remarkable Giant Nerve Cells of Squid
	The Ionic Basis of Action Potentials
	BOX 2B Action Potential Form and Nomenclature
	Summary
Chapter 3 Voltage-Dependent Membrane Permeability
	Overview
	Ion Currents across Nerve Cell Membranes
	BOX 3A The Voltage Clamp Method
	Two Types of Voltage-Dependent Ion Currents
	Two Voltage-Dependent Membrane Conductances
	Reconstruction of the Action Potential
	Long-Distance Signaling by Means of Action Potentials
	Increased Conduction Velocity as a Result of Myelination
	Summary
Chapter 4 Ion Channels and Transporters
	Overview
	Ion Channels Underlying Action Potentials
	BOX 4A The Patch Clamp Method
	How Ion Channels Work
	BOX 4B Toxins That Poison Ion Channels
	The Diversity of Ion Channels
	Voltage-Gated Ion Channels
	Ligand-Gated Ion Channels
	Thermosensitive and Mechanosensitive Channels
	Active Transporters Create and Maintain Ion Gradients
	ATPase Pumps
	Ion Exchangers
	Summary
Chapter 5 Synaptic Transmission
	Overview
	Two Classes of Synapses
	Signaling Transmission at Electrical Synapses
	Signaling Transmission at Chemical Synapses
	Properties of Neurotransmitters
	Quantal Release of Neurotransmitters
	Release of Transmitters from Synaptic Vesicles
	Local Recycling of Synaptic Vesicles
	The Role of Calcium in Transmitter Secretion
	Molecular Mechanisms of Synaptic Vesicle Cycling
	Neurotransmitter Receptors
	Postsynaptic Membrane Permeability Changes during Synaptic Transmission
	Relationship between Ion Fluxes and Postsynaptic Potential Changes
	Excitatory and Inhibitory Postsynaptic Potentials
	Summation of Synaptic Potentials
	BOX 5A The Tripartite Synapse
	Summary
Chapter 6 Neurotransmitters and Their Receptors
	Overview
	Categories of Neurotransmitters
	Acetylcholine
	BOX 6A Neurotoxins That Act on Neurotransmitter Receptors
	Glutamate
	GABA and Glycine
	BOX 6B Excitatory Actions of GABA in the Developing Brain
	Biogenic Amines
	ATP and Other Purines
	Peptide Neurotransmitters
	Unconventional Neurotransmitters
	BOX 6C Marijuana and the Brain
	Summary
Chapter 7 Molecular Signaling within Neurons
	Overview
	Strategies of Molecular Signaling
	Activation of Signaling Pathways
	Receptor Types
	G-Proteins and Their Molecular Targets
	Second Messengers
	BOX 7A Dynamic Imaging of Intracellular Signaling
	Second-Messenger Signaling via Protein Phosphorylation
	Protein Kinases
	Protein Phosphatases
	BOX 7B Dendritic Spines
	Nuclear Signaling
	Examples of Neuronal Signal Transduction
	Summary
Chapter 8 Synaptic Plasticity
	Overview
	Short-Term Synaptic Plasticity
	Long-Term Synaptic Plasticity and Behavioral Modification in
	BOX 8A Genetics of Learning and Memory in the Fruit Fly
	Long-Term Potentiation at a Hippocampal Synapse
	Mechanisms Underlying Long-Term Potentiation
	BOX 8B Silent Synapses
	Mechanisms Underlying Long-Term Depression
	Spike Timing-Dependent Plasticity
	Summary
Chapter 9 The Somatosensory System: Touch and Proprioception
	Overview
	Afferent Fibers Convey Somatosensory Information to the Central Nervous System
	Somatosensory Afferents Convey Different Functional Information
	Mechanoreceptors Specialized to Receive Tactile Information
	Mechanoreceptors Specialized for Proprioception
	Central Pathways Conveying Tactile Information from the Body: The Dorsal Column–Medial Lemniscal System
	Central Pathways Conveying Tactile Information from the Face: The Trigeminothalamic System
	Central Pathways Conveying Proprioceptive Information from the Body
	Central Pathways Conveying Proprioceptive Information from the Face
	Somatosensory Components of the Thalamus
	Primary Somatosensory Cortex
	BOX 9A Patterns of Organization within the Sensory Cortices: Brain Modules
	Beyond SI: Corticocortical and Descending Pathways
	Plasticity in the Adult Cerebral Cortex
	Summary
Chapter 10 Pain
	Overview
	Nociceptors
	Transduction and Transmission of Nociceptive Signals
	BOX 10A Capsaicin
	Central Pain Pathways Are Distinct from Mechanosensory Pathways
	BOX 10B Referred Pain
	Parallel Pain Pathways
	BOX 10C A Dorsal Column Pathway for Visceral Pain
	Pain and Temperature Pathways for the Face
	Other Modalities Mediated by the Anterolateral System
	Sensitization
	Descending Control of Pain Perception
	The Placebo Effect
	The Physiological Basis of Pain Modulation
	Summary
Chapter 11 Vision: The Eye
	Overview
	Anatomy of the Eye
	Image Formation on the Retina
	BOX 11A Myopia and Other Refractive Errors
	The Retinal Surface
	Retinal Circuitry
	The Pigment Epithelium
	Phototransduction
	BOX 11B Retinitis Pigmentosa
	Functional Specialization of the Rod and Cone Systems
	Anatomical Distribution of Rods and Cones
	Cones and Color Vision
	BOX 11C The Importance of Context in Color Perception
	Retinal Circuits for Light and Dark
	BOX 11D The Perception of Light Intensity
	The Adjustable Operating Range of Retinal Ganglion Cells
	Luminance Contrast and Receptive Field Surrounds
	Summary
Chapter 12 Central Visual Pathways
	Overview
	Central Projections of Retinal Ganglion Cells
	Functionally Distinct Types of Retinal Ganglion Cells
	Retinotopic Representation of the Visual Field
	Spatiotemporal Tuning Properties of Neurons in Primary Visual Cortex
	Primary Visual Cortex Architecture
	Combining Inputs from Two Eyes
	BOX 12A Random Dot Stereograms and Related Amusements
	Division of Labor within the Primary Visual Pathway
	Functional Organization of Extrastriate Visual Areas
	Summary
Chapter 13 The Auditory System
	Overview
	Sound
	The Audible Spectrum
	A Synopsis of Auditory Function
	The External Ear
	The Middle Ear
	The Inner Ear
	Hair Cells and the Mechanoelectrical Transduction of Sound Waves
	BOX 13A The Sweet Sound of Distortion
	The Ionic Basis of Mechanotransduction in Hair Cells
	The Hair Cell Mechanoelectrical Transduction Channel
	The Cochlear Amplifier
	Tuning and Timing in the Auditory Nerve
	How Information from the Cochlea Reaches Targets in the Brainstem
	Integrating Information from the Two Ears
	Monaural Pathways from the Cochlear Nucleus to the Nuclei of the Lateral Lemniscus
	Integration in the Inferior Colliculus
	The Auditory Thalamus
	The Auditory Cortex
	BOX 13B Representing Complex Sounds in the Brains of Bats and Humans
	Summary
Chapter 14 The Vestibular System
	Overview
	The Vestibular Labyrinth
	Vestibular Hair Cells
	Otolith Organs: The Utricle and Saccule
	How Otolith Neurons Sense Head Tilts and Translational Head Movements
	Semicircular Canals
	How Semicircular Canal Neurons Sense Head Rotations
	Central Pathways for Stabilizing Gaze, Head, and Posture
	BOX 14A Mauthner Cells in Fish
	Vestibular–Cerebellar Pathways
	Vestibular Pathways to the Thalamus and Cortex
	Spatial Orientation Perception and Multisensory Integration
	Summary
Chapter 15 The Chemical Senses
	Overview
	Organization of the Olfactory System
	Olfactory Perception in Humans
	Assessing Olfactory Function in the Laboratory or Clinic
	Physiological and Behavioral Responses to Olfactory Stimuli
	Olfactory Epithelium and Olfactory Receptor Neurons
	Odor Transduction and Odorant Receptor Proteins
	BOX 15A The “Dogtor” Is In
	Molecular and Physiological Mechanisms of Olfactory Odor Transduction
	The Olfactory Bulb
	Pyriform Cortical Processing of Information Relayed from the Olfactory Bulb
	The Vomeronasal System: Predators, Prey, and Mates
	The Vomeronasal System: Molecular Mechanisms of Sensory Transduction
	Organization of the Taste System
	Taste Perception in Humans
	Taste Receptor Proteins and Transduction
	Neural Coding in the Taste System
	Summary
Chapter 16 Lower Motor Neuron Circuits and Motor Control
	Overview
	Neural Centers Responsible for Movement
	Motor Neuron–Muscle Relationships
	The Motor Unit
	BOX 16A Motor Unit Plasticity
	Regulation of Muscle Force
	The Spinal Cord Circuitry Underlying Muscle Stretch Reflexes
	Modifying the Gain of Muscle Stretch Reflexes
	The Spinal Cord Circuitry Underlying the Regulation of Muscle Force
	Complementary Functions of Muscle Spindles and Golgi Tendon Organs
	Flexion Reflex Pathways
	Spinal Cord Circuitry and Locomotion
	BOX 16B Locomotion in the Leech and the Lamprey
	The Lower Motor Neuron Syndrome
	Summary
Chapter 17 Upper Motor Neuron Control of the Brainstem and Spinal Cord
	Overview
	Organization of Descending Motor Control
	The Corticospinal and Corticobulbar Tracts
	Functional Organization of the Primary Motor Cortex
	BOX 17A What Do Motor Maps Represent?
	The Premotor Cortex
	BOX 17B Minds and Machines
	Motor Control Centers in the Brainstem: Upper Motor Neurons That Maintain Balance, Govern Posture, Initiate Locomotion, and Orie
	BOX 17C The Reticular Formation
	Damage to Descending Motor Pathways: The Upper Motor Neuron Syndrome
	BOX 17D Muscle Tone
	Summary
Chapter 18 Modulation of Movement by the Basal Ganglia
	Overview
	Projections to the Basal Ganglia
	BOX 18A Making and Breaking Habits
	Projections from the Basal Ganglia to Other Brain Regions
	Evidence from Studies of Eye Movements
	Circuits within the Basal Ganglia System
	Dopamine Modulates Basal Ganglia Circuits
	Hypokinetic Movement Disorders
	Hyperkinetic Movement Disorders
	BOX 18B Basal Ganglia Loops and Non-Motor Brain Functions
	Summary
Chapter 19 Modulation of Movement by the Cerebellum
	Overview
	Organization of the Cerebellum
	Projections to the Cerebellum
	Projections from the Cerebellum
	Circuits within the Cerebellum
	Cerebellar Circuitry and the Coordination of Ongoing Movement
	Further Consequences of Cerebellar Lesions
	BOX 19A Genetic Analysis of Cerebellar Function
	Summary
Chapter 20 Eye Movements and Sensorimotor Integration
	Overview
	What Eye Movements Accomplish
	BOX 20A The Perception of Stabilized Retinal Images
	Actions and Innervation of Extraocular Muscles
	Types of Eye Movements and Their Functions
	Neural Control of Saccadic Eye Movements
	BOX 20B Sensorimotor Integration in the Superior Colliculus
	BOX 20C From Place Codes to Rate Codes
	Neural Control of Smooth Pursuit Movements
	Neural Control of Vergence Movements
	Summary
Chapter 21 The Visceral Motor System
	Overview
	Early Studies of the Visceral Motor System
	Distinctive Features of the Visceral Motor System
	The Sympathetic Division of the Visceral Motor System
	BOX 21A The Hypothalamus
	The Parasympathetic Division of the Visceral Motor System
	The Enteric Nervous System
	Sensory Components of the Visceral Motor System
	Central Control of Visceral Motor Functions
	BOX 21B Obesity and the Brain
	Neurotransmission in the Visceral Motor System
	Autonomic Regulation of Cardiovascular Function
	Autonomic Regulation of the Bladder
	Autonomic Regulation of Sexual Function
	Summary
Chapter 22 Early Brain Development
	Overview
	Formation of the Nervous System: Gastrulation and Neurulation
	BOX 22A Stem Cells: Promise and Peril
	Formation of the Major Brain Subdivisions
	The Molecular Basis of Neural Induction
	Stem Cells
	Integrated Inductive Signals Establish Neuron Identity
	Initial Differentiation of Neurons and Glia
	Molecular Regulation of Neurogenesis
	Generation of Neuronal Diversity
	Molecular and Genetic Disruptions of Early Neural Development
	Neuronal Migration in the Peripheral Nervous System
	Neuronal Migration in the Central Nervous System
	Molecular Mechanisms of Neuronal Migration and Cortical Migration Disorders
	Cranial Placodes: The Critical Connection between the Brain and the Outside World
	Summary
Chapter 23 Construction of Neural Circuits
	Overview
	Neuronal Polarization: The First Step in Neural Circuit Formation
	The Axon Growth Cone
	BOX 23A Choosing Sides: Axon Guidance at the Optic Chiasm
	The Molecular Basis of Growth Cone Motility
	Non-Diffusible Signals for Axon Guidance
	Chemoattraction and Chemorepulsion
	Directed Dendritic Growth: Ensuring Polarity
	Dendritic Tiling: Defining Synaptic Space
	Formation of Topographic Maps
	Selective Synapse Formation
	Regulation of Neuronal Connections by Trophic Interactions
	Competitive Interactions and the Formation of Neuronal Connections
	BOX 23B Why Do Neurons Have Dendrites?
	The Molecular Basis of Trophic Interactions
	Neurotrophin Signaling
	Summary
Chapter 24 Circuit Development: Intrinsic Factors and Sex Differences
	Overview
	Sexual Dimorphisms and Sexually Dimorphic Behaviors
	Sex, Gonads, Bodies, and Brains
	BOX 24A The Science of Love (or, Love As a Drug)
	Hormonal Influences on Sexually Dimorphic Development
	Development of Dimorphisms in the Central Nervous System
	Brain Dimorphisms and the Establishment of Reproductive Behaviors
	Cellular and Molecular Basis of Sexually Dimorphic Development
	Steroid Receptors and Responses in the Adult Brain
	Human Genetic Disorders of Genotypic and Phenotypic Sex
	Sexual Orientation and Human Brain Structure
	Sex-Based Differences in Cognitive Functions
	Summary
Chapter 25 Experience-Dependent Plasticity in the Developing Brain
	Overview
	Neural Activity and Brain Development
	Critical Periods
	BOX 25A Built-In Behaviors
	The Role of Oscillations in Establishing Critical Periods
	Critical Periods in Visual System Development
	Effects of Visual Deprivation on Ocular Dominance
	Manipulating Competition
	Binocular Competition and Orientation Tuning for Binocular Vision
	Amblyopia, Strabismus, and Critical Periods for Human Vision
	Cellular and Molecular Regulation of Critical Periods
	Evidence for Critical Periods in Other Sensory Systems
	Language Development: A Critical Period for a Distinctly Human Behavior
	Human Brain Development, Activity-Dependent Plasticity, and Critical Periods
	Summary
Chapter 26 Repair and Regeneration in the Nervous System
	Overview
	The Damaged Brain
	Functional Reorganization without Repair
	Three Types of Neuronal Repair
	Peripheral Nerve Regeneration
	The Cellular and Molecular Basis of Peripheral Nerve Repair
	Regeneration of Peripheral Synapses
	BOX 26A Specific Regeneration of Synaptic Connections in Autonomic Ganglia
	Regeneration in the Central Nervous System
	Cellular and Molecular Responses to Injury in the Central Nervous System
	Glial Scar Formation in the Injured Brain
	Immune Activation and Inflammation Following Brain Injury
	Axon Growth after Brain Injury
	Neurogenesis in the Mature Central Nervous System
	Adult Neurogenesis in Non-Mammalian Vertebrates
	Neurogenesis in the Adult Mammalian Brain
	Cellular and Molecular Mechanisms of Adult Neurogenesis
	Adult Neurogenesis, Stem Cells, and Brain Repair in Humans
	BOX 26B Nuclear Weapons and Neurogenesis
	Summary
Chapter 27 Cognitive Functions and the Organization of the Cerebral Cortex
	Overview
	A Primer on Cortical Structure
	BOX 27A Cortical Lamination
	Unique Features of the Association Cortices
	The Parietal Association Cortex
	BOX 27B Synesthesia
	The Temporal Association Cortex
	The Frontal Association Cortex
	BOX 27C Neuropsychological Testing
	Summary
Chapter 28 Cortical States
	Overview
	The Circadian Cycle
	Molecular Mechanisms of Biological Clocks
	BOX 28A Electroencephalography
	Sleep
	The Purpose of Sleep
	BOX 28B Sleep and Memory
	Sleep in Different Species
	The Stages of Sleep
	BOX 28C Dreaming
	The Physiological Changes in Sleep States
	The Neural Circuits Governing Sleep
	Thalamocortical Interactions in Sleep
	Consciousness
	The Neural Correlates of Consciousness
	Comatose States
	The “Default State” of the Brain
	Summary
Chapter 29 Attention
	Overview
	Attention as “Selective” Processing
	Endogenous versus Exogenous Attention
	Covert Attention
	Attention across Sensory Modalities
	Evidence for a Brain System That Controls Attention
	Evidence from Neurological Patients: Hemispatial Neglect Syndrome
	Other Brain Regions That Affect Attention
	Evidence from Normal Subjects
	BOX 29A Attention and the Frontal Eye Fields
	Studies in Non-Human Primates
	Problems with the Concept of Attention as Executive Control
	Alternative Interpretations of Attention
	Summary
Chapter 30 Memory
	Overview
	Qualitative Categories of Human Memory
	BOX 30A Phylogenetic Memory
	Temporal Categories of Memory
	Priming
	The Importance of Association in Information Storage
	Conditioned Learning
	BOX 30B Savant Syndrome
	Forgetting
	Amnesia
	Brain Systems Underlying Declarative Memory Acquisition and Storage
	Sites of Long-Term Memory Storage
	BOX 30C Alzheimer’s Disease
	BOX 30D lace Cells and Grid Cells
	Brain Systems Underlying Nondeclarative Memory Acquisition and Storage
	Memory and Aging
	Summary
Chapter 31 Emotion
	Overview
	Defining Emotion
	Physiological Changes Associated with Emotion
	The Integration of Emotional Behavior
	Is the Motor Activity Underlying Emotion Cause or Effect?
	BOX 31A Determination of Facial Expressions
	The Limbic System
	BOX 31B The Amygdala
	The Importance of the Amygdala
	BOX 31C Fear and the Human Amygdala
	The Relationship between Neocortex and Amygdala
	Cortical Lateralization of Emotional Functions
	Emotion, Reason, and Social Behavior
	Emotion and Addiction
	Posttraumatic Stress Disorder
	Summary
Chapter 32 Thinking, Planning, and Deciding
	Overview
	A Sketch of the Relevant Circuitry
	Orbitofrontal Cortex and the Evaluation of Options
	BOX 32A Dopamine and Reward Prediction Errors
	Dorsolateral Prefrontal Cortex and the Planning and Organization of Behavior
	Cingulate Cortex and Learning from the Consequences of Behavior
	Ventrolateral Prefrontal Cortex and Self-Control
	Anterior Insula and the Internal Milieu
	Posterior Cingulate Cortex and Self-Awareness
	BOX 32B What Does Neuroscience Have to Say about Free Will?
	Summary
Chapter 33 Speech and Language
	Overview
	Representation of Language in the Brain
	Aphasias
	BOX 33A The Generation of Speech
	An Ingenious Confirmation of Language Lateralization
	BOX 33B Language and Handedness
	The Search for Anatomical Differences between the Hemispheres
	Mapping Language Functions
	The Role of the Right Hemisphere in Language
	Genes and Language
	Sign Language
	A Critical Period for Language Acquisition
	Reading and Dyslexia
	Do Other Animals Have Language?
	BOX 33C Learned Vocal Communication in Birds
	Summary
Chapter 34 Development and Evolution of Cognitive Functions
	Overview
	Foundations of Developmental and Evolutionary Cognitive Neurobiology
	Development of Cognitive Abilities
	BOX 34A Darwin and the Brain
	Brain Size and Cognitive Development
	Brain Size and the Evolution of Cognition
	BOX 34B Brain Differences among Modern Humans
	Relative Brain Size and Cerebral Complexity
	Evolution of Brain Development
	Evolutionary Specializations of the Brain and Behavior
	BOX 34C The Neurobiology of Intelligence
	Fossil Evidence on the Evolution of Brain and Cognition
	Evolution of Learning and Memory
	Evolution of Social Cognition
	Summary
Appendix: Survey of Human Neuroanatomy
	Overview
	Neuroanatomical Terminology
	Basic Subdivisions of the Central Nervous System
	External Anatomy of the Spinal Cord
	Internal Anatomy of the Spinal Cord
	Brainstem and Cranial Nerves
	Lateral Surface of the Brain
	Dorsal and Ventral Surfaces of the Brain
	Midsagittal Surface of the Brain
	Internal Anatomy of the Forebrain
	BOX A Thalamus and Thalamocortical Relations
	Blood Supply of the Brain and Spinal Cord
	The Blood–Brain Barrier
	The Meninges
	The Ventricular System
Atlas: The Human Central Nervous System
Glossary
Box References
Illustration Credits
Index
About the Book