Brain and Behavior A Cognitive Neuroscience Perspective

Brain and Behavior: A Cognitive Neuroscience Perspective addresses the central aims of cognitive neuroscience, seeking to examine the brain not only by its components but also by their functions. It highlights the principles, discoveries, and remaining mysteries of modern cognitive neuroscience.

Brain and Behavior covers a wide swath of territory critical for understanding the brain, from the basics of the nervous system, to sensory and motor systems, sleep, language, memory, emotions and motivation, social cognition, and brain disorders. Throughout the narrative, the authors emphasize the dynamically changing nature of the brain, through the mechanisms of neuroplasticity. Wherever possible, they refer to elements of neuroscience that are encountered in everyday life. Key points and concepts are illustrated using case studies of rare but illuminating brain disorders. Brain and Behavior pulls together the best current knowledge about the brain while acknowledging current areas of ignorance and pointing students towards the most promising directions for future research.

David Eagleman is a neuroscientist, New York Times best-selling author, and Guggenheim Fellow who holds joint appointments in the Departments of Neuroscience and Psychiatry at Baylor College of Medicine in Houston, Texas. Dr. Eagleman's areas of research include time perception, vision, synesthesia, and the intersection of neuroscience with the legal system.

Jonathan Downar is the director of the MRI-Guided rTMS Clinic at the University Health Network Hospital in Toronto, Canada, and a scientist at the Toronto Western Research Institute. He currently holds appointments with the Department of Psychiatry and the Institute of Medical Science at the University of Toronto.

*Every chapter ends with the following: Conclusion, Key Principles, Key Terms, Review Questions, and Critical-Thinking Questions

Contents
Preface

Part I: The Basics

Chapter 1: Introduction
Learning Objectives
Starting Out: A Spark of Awe in the Darkness
Who Are We?
The Mission of Cognitive Neuroscience
Neuroscience Is a Relatively New Field
In Pursuit of Principles
The Functions behind the Form
Which Parts Matter?
What Is the Brain For?
How We Know What We Know
Connectional Methods
Correlational Methods
Research Methods: Magnetic Resonance Imaging
Lesion Methods
Stimulation Methods
A Toolbox of Complementary Methods
Thinking Critically about the Brain
Is the Brain Equipped to Understand Itself?
Biases and Pitfalls in Human Cognition
A Toolbox of Critical-Thinking Techniques
The Big Questions in Cognitive Neuroscience
Why Have a Brain at All? (Chapter 2)
How Is Information Coded in Neural Activity? (Chapter 3)
How Does the Brain Balance Stability against Change? (Chapter 4)
Why Does Vision Have So Little to Do with the Eyes? (Chapter 5)
How Does the Brain Stitch Together a Picture of the World from Different Senses? (Chapter 6)
How Does the Brain Control Our Actions? (Chapter 7)
What Is Consciousness? (Chapter 8)
How Are Memories Stored and Retrieved? (Chapter 9)
Why Do Brains Sleep and Dream? (Chapter 10)
How Does the Human Brain Acquire Its Unique Ability for Language? (Chapter 11)
How Do We Make Decisions? (Chapter 12)
What Are Emotions? (Chapter 13)
How Do We Set Our Priorities? (Chapter 14)
How Do I Know What You're Thinking? (Chapter 15)
What Causes Disorders of the Mind and the Brain? (Chapter 16)
The Payoffs of Cognitive Neuroscience
Healing the Disordered Brain
Enhancing Human Abilities
Blueprints for Artificial Cognition
Brain-Compatible Social Policies

Chapter 2: The Brain and Nervous System
Learning Objectives
Starting Out: The Brains of Creatures Great and Small
An Overview of the Nervous System
Why Put Your Neurons in a Brain at All?
The Common Features of Every Central Nervous System
Getting Oriented in the Brain
The Peripheral Nervous System
Separate Systems for the Inner and Outer Environments
A Nervous System with Segmental Organization
The Spinal Cord
Circuits within a Segment: Spinal Reflexes
Case Study: Christopher Reeve, 1952-2004
Complex Circuits across Segments: Central Pattern Generators
The Bigger Picture: In Search of a Cure for Spinal Cord Injury
The Brainstem
Medulla Oblongata and Pons
Neuroscience of Everyday Life: Why Do We Get the Hiccups?
Midbrain
Most Cranial Nerves Emerge from the Brainstem
The Cerebellum
Circuitry of the "Little Brain"
Functions of the Little Brain
The Diencephalon: Hypothalamus and Thalamus
Hypothalamus: A Keystone Structure in Homeostasis
Thalamus
Case Study: Waking the Brain
The Telencephalon: Cerebral Cortex and Basal Ganglia
Cerebral Cortex
Basal Ganglia
Research Methods: Cytoarchitecture of the Cortex
Uniting the Inside and Outside Worlds
The Limbic System
The Ventricular System and Brain Function
Conclusion
Key Principles
Key Terms
Review Questions
Critical-Thinking Questions

Chapter 3: Neurons and Synapses
Learning Objectives
Starting Out: The Kabuki Actor and the Pufferfish
The Cells of the Brain
Neurons: A Close-Up View
Many Different Types of Neurons
Glial Cells
Research Methods: Visualizing Neurons and Their Products
Synaptic Transmission: Chemical Signaling in the Brain
Release of Neurotransmitter at the Synapse
Types of Neurotransmitters
Receptors
Postsynaptic Potentials
The Bigger Picture: Psychoactive Drugs
Spikes: Electrical Signaling in the Brain
Adding Up the Signals
How an Action Potential Travels
Myelinating Axons to Make the Action Potential Travel Faster
Action Potentials Reach the Terminals and Cause Neurotransmitter Release
Case Study: Multiple Sclerosis
Neuroscience of Everyday Life: The Magic of a Local Anesthetic
What Do Spikes Mean? The Neural Code
Encoding Stimuli in Spikes
Decoding Spikes
Research Methods: Recording Action Potentials with Electrodes
Individuals and Populations
Populations of Neurons
Forming a Coalition: What Constitutes a Group?
Open Questions for Future Investigation

Chapter 4: Neuroplasticity
Learning Objectives
Starting Out: The Child with Half a Brain
The Brain Dynamically Reorganizes to Match Its Inputs
Changes to the Body Plan
Case Study: Phantom Sensation
Research Methods: Mapping Out the Brain
Changes to Sensory Input
The Brain Distributes Resources Based on Relevance
The Role of Behavior
The Role of Relevance: Gating Plasticity with Neuromodulation
Neuroscience of Everyday Life: Pianists and Violinists Have Different Brains
Case Study: The Government Worker with the Missing Brain
The Brain Uses the Available Tissue
Maps Adjust Themselves to the Available Brain Tissue
Cortical Reorganization after Brain Damage
A Sensitive Period for Plastic Changes
A Window of Time to Make Changes
Case Study: Danielle, the Feral Child in the Window
The Sensitive Period in Language
Neuromodulation in Young Brains
Hardwiring versus World Experience
Aspects of the Brain Are Preprogrammed
Experience Changes the Brain
Brains Rely on Experience to Unpack Their Programs Correctly
The Mechanisms of Reorganization
Neurons Compete for Limited Space
Competition for Neurotrophins
Rapid Changes: Unmasking Existing Connections
Slow Changes: Growth of New Connections
Changing the Input Channels
Case Study: The Man Who Climbs with his Tongue
The Bigger Picture: Adding New Peripherals


Part II: How the Brain Interacts with the World

Chapter 5: Vision
Learning Objectives
Starting Out: Vision Is More Than the Eyes
Visual Perception
What Is It Like to See?
Signal Transduction
Anatomy of the Visual System
Sensory Transduction: The Eye and Its Retina
Case Study: The Bionic Retina
Path to the Visual Cortex: The Lateral Geniculate Nucleus
The Visual Cortex
Two Eyes Are Better Than One: Stereo Vision
Neuroscience of Everyday Life: Random-Dot Stereograms
Higher Visual Areas
Secondary and Tertiary Visual Cortex: Processing Becomes More Complex
Ventral Stream: What an Object Is
The Bigger Picture: Reading the Movies in Our Minds
Dorsal Stream: How to Interact with the World
Case Study: The World in Snapshots
Attention and the Dorsal Stream
Comparing the Ventral and Dorsal Processing Streams
The Bigger Picture of the Visual Brain
Case Study: The Blind Woman Who Could See, Sort Of
Perception Is Active, Not Passive
Interrogating the Scene with Our Eyes
The Blind Spot
Seeing the Same Object Different Ways: Multistability
Binocular Rivalry: Different Images in the Two Eyes
We Don't See Most of What Hits Our Eyes: Fetching Information on a Need-to-Know Basis
Vision Relies on Expectations
Change Blindness
Saving Resources by Embedding Prior Experience
Unconscious Inference
Activity from Within
Feedback Allows an Internal Model

Chapter 6: Other Senses
Learning Objectives
Starting Out: The Man with the Bionic Ear
Detecting Data from the World
Hearing
Research Methods: Psychophysics
The Outer and Middle Ear
Converting Mechanical Information into Electrical Signals: The Inner Ear
Neuroscience of Everyday Life: The Undetectable Cell Phone
The Auditory Nerve and Primary Auditory Cortex
The Hierarchy of Sound Processing
Sound Localization
Balance
The Somatosensory System
Touch
Temperature
Pain
Case Study: The Pain of a Painless Existence
Proprioception
Interoception
The Somatosensory Pathway
Chemical Senses
Taste
Smell
The Sense of Flavor
Pheromones
The Brain Is Multisensory
Synesthesia
Combining Sensory Information
The Binding Problem
The Internal Model of the World
Case Study: The Paralyzed Supreme Court Justice Who Claimed He Could Play Football
Time Perception

Chapter 7: The Motor System
Learning Objectives
Starting Out: "'Locked-In Syndrome"'
Muscles
Skeletal Muscle: Structure and Function
The Neuromuscular Junction
The Spinal Cord
Lower Motor Neurons
Spinal Motor Circuits: Reflexes
Spinal Motor Circuits: Central Pattern Generators
Descending Pathways of Motor Control
The Cerebellum
The Circuitry of the Cerebellum
Motor Functions of the Cerebellum
Nonmotor Functions of the Cerebellum
The Motor Cortex
Motor Cortex: Neural Coding of Movements
Motor Cortex: Recent Controversies
The Bigger Picture: Neural Implants for Motor Control
The Prefrontal Cortex: Goals to Strategies to Tactics to Actions
The Functional Organization of the Prefrontal Cortex in Motor Control
Sensory Feedback
Mirror Neurons in Premotor Cortex
Control Stages of the Motor Hierarchy
Basal Ganglia
Components of the Basal Ganglia
Circuitry of the Basal Ganglia
Diseases of the Basal Ganglia
Medial and Lateral Motor Systems: Internally and Externally Guided Movement Control
Organization of Medial Motor Areas
Functions of Medial and Lateral Motor Systems
Neuroscience of Everyday Life: Why Can't I Multitask?
Did I Really Do That? The Neuroscience of Free Will
Research Methods: Neurosurgical Stimulation
Case Study: Alien Hand Syndrome

Part III: Higher Levels of Interaction

Chapter 8: Attention and Consciousness
Learning Objectives
Starting Out: The Stream of Consciousness
Awareness Requires Attention
Change Blindness
Inattentional Blindness
Neuroscience of Everyday Life: Stage Magic
Approaches to Studying Attention and Awareness
Attentional Orienting Paradigms: Aiming the "Spotlight" of Attention
The Oddball Paradigm: Monitoring a Physiological Measure of Attention
Uncoupling Sensory Input from Perception: Sensory Rivalry
Neural Mechanisms of Attention and Awareness
Seeking the Correlates of Consciousness
Hemineglect: A Disorder of Attention and Awareness
Case Study: Unaware of Half of the World
Neural Correlates of Attention: A Single Network, or Many?
Case Study: Whose Arm Is This, Anyway?
Sites of Attentional Modulation: Neurons and Neural Populations
The Biased-Competition Model of Attention
Attention and Single Neurons: Enhancing the Signal
Attention and Local Groups of Neurons
Synchronization, Attention, and Awareness
Coma and Vegetative State: Anatomy of the Conscious State
Why Should Synchronization Matter?
Unconsciousness: Coma and Vegetative State
Case Study: Waking the Brain
Midbrain and Thalamus: Key Players in the Conscious State
Anesthesia and Sleep: Rhythms of Consciousness
Sleep: Unraveling the Rhythm of Consciousness
Anesthesia: Reversible, Artificial Unconsciousness
Theories of Consciousness
Dualism: The Mind-Body Problem
Functionalist Theories of Consciousness
Consciousness and the Integration of Information

Chapter 9: Memory
Learning Objectives
Starting Out: "The Woman Who Cannot Forget"
The Many Kinds of Memory
Working and Long-Term Memory
Implicit Memory
Explicit Memory
Travels in Space and Time: The Hippocampus and Temporal Lobe
Case Study: Gone but Not Forgotten: Henry Molaison, 1926-2008
A Map of the Medial Temporal Lobe
Episodic Memory
Spatial Memory
Theories of Hippocampal Function
Unifying the Functions of the Hippocampus
Remembering the Future: Prospection and Imagination
How We Imagine Future Experiences
Research Methods: Localizing Human Brain Function
The Circuitry of Prospection and Recollection
Neuroscience of Everyday Life: Simonides and the Champions of Memory
Prospection in Other Species
Models of Prospection
The Confabulation of Reality
Confabulation in the Injured Brain
Case Study: The Woman with a Thirty-Year-Old Baby
The Anatomy of Spontaneous Confabulation
Confabulation in the Normal Brain
The Anatomy of a False Memory
The Bigger Picture: Scanning for the Truth
The Mechanisms of Memory
General Mechanisms of Learning and Memory
Memory as Synaptic Change
Long-Term Potentiation and Depression of Synaptic Connections
The NMDA Receptor
Consolidation and Reconsolidation
Associative Neural Networks
Beyond Synaptic Plasticity: The Frontiers of Memory Mechanisms
Whole Neurons as a Substrate for Memory?
New Neurons for New Memories
Spines: Another Structural Basis for Memory?
Looking inside the Cell: Memory in Chemical Reactions
Case Study: The Flies with Photographic Memory
Epigenetics: Making a Single Genome Play Different Tunes
The Mysteries of Memory
Are the Roles Of LTP and LTD Overstated?
The Timing of Spikes
The Limitations of Neural Networks
Neural Networks: Solving the Wrong Problem?
Remembering Relationships, Not Features
The Future of Memory Research

Chapter 10: Sleep
Learning Objectives
Starting Out: Caught between Sleeping and Waking
Sleep and the Brain
The Brain Is Active during Sleep
Research Methods: Electroencephalography
The Neural Networks of Sleep
The Brain during REM Sleep
The Circadian Rhythm
Entrainment of the Circadian Rhythm by Light Cues
The Circadian Rhythm Is Not Fixed
Case Study: The Shifted Circadian Rhythm
The Circadian Rhythm and Napping
The Bigger Picture: Schools and Circadian Rhythms
Why Do Brains Sleep?
Four Theories of Sleeping: Restoration, Survival, Simulation, Learning
Rehearsal
Forgetting
Insight and the Restructuring of Information
Dreaming
Dream Content
The Neuroscience of Everyday Life: Lucid Dreaming
Can Dreams Shed Light on Consciousness?
Dreams of the Future and How to Study Them
Sleep Deprivation and Disorders
Sleep Deprivation
Case Study: Staying Awake
Insomnia
Hypersomnia
Case Study: The Family Who Couldn't Sleep
Parasomnias

Chapter 11: Language and Lateralization
Learning Objectives
Starting Out: The Stuttering King
Speech, Language, and Communication
Aphasia: The Loss of Language
Case Study: The Woman Who Couldn't Find Her Words
Broca's Aphasia
Wernicke's Aphasia
Case Study: The Woman Who Makes Up Words
A Language Network
The Larger Picture of Language-Specific Regions
Dyslexia
Stuttering
Lateralization: The Two Hemispheres Are Not Identical
Tests for Dominance
Apraxia
Hemispheric Differences
Two Brains in One? The Case of the Split-Brain Patients
Thinking about Cerebral Asymmetry
Development of Language
Learning Language from Experience
Innate Language Tendencies
Socially and Emotionally Directed Learning
Research Methods: The Baby with No Privacy

Part IV: Motivated Behaviors

Chapter 12: Decision Making
Learning Objectives
Starting Out: A Fatal Mistake, at the Highest Place on Earth
How Do We Decide What to Do?
The Scorpion and the Frog
The Search for a "Physics" of Human Decisions
Homo economicus and Rational Choice Theory
The Predictably Irrational Homo sapiens
Homo sapiens versus Homo economicus
Confused by Uncertainty
The Framing Effect and the Endowment Effect
The Illusory Value of Procrastination
Where Do Our Irrational Decisions Come From?
Decision Making in Other Species
Do Irrational Decisions Come from Irrational People?
One Brain, Two Systems
How the Brain Decides
The Neural Mechanisms of Delay Discounting
Neural Mechanisms of Decisions under Risk
The Neural Basis of the Endowment Effect
The Neural Basis of the Framing Effect
The Common Currency of Subjective Value
Comparing Apples to Oranges
Research Methods: Charting the Landscape of Subjective Value
A Consistent Neural Basis for Subjective Value
Evaluation and the Orbitofrontal Cortex
One Currency, But Many Markets
The Neuroscience of Everyday Life: Snack Food or Brussels Sprouts?
A Hierarchy of Internally Guided Decision Making
Internally and Externally Guided Decision Making
Values into Goals
Goals into Plans
Plans into Behavior and Action
Modulators of Decision Making
Strategic Use of Decision-Making Systems
Neurotransmitter Effects on Decision Making
The Bigger Picture: How to Avoid the Scorpion's Sting

Chapter 13: Emotions
Learning Objectives
Starting Out: Sadness, at the Flip of a Switch
Early Theories of Emotion
Emotional Expressions: Signposts on a Landscape of Inner States
The James-Lange Theory of Emotion: A Bottom-Up Theory
The Cannon-Bard Theory: A Top-Down Theory
Case Study: Pathological Laughter and Crying
Two-Factor Theories: Reconciling Central and Peripheral Influences on Emotion
Core Limbic Structures: Amygdala and Hypothalamus
Hypothalamus: Internal States, Homeostatic Drives
Case Study: An Internal Growth of Rage
Do Hypothalamic Circuits Generate Inner Emotional Experiences?
Amygdala: Externally Generated States and Drives
The Amygdala and Emotional Experience
Case Study: The Woman Who Knows No Fear
Hippocampus: Emotional Memories
Ventral Striatum: Pleasure and Reward
Bringing It All Together: The Circuit of Papez and the Ring of Limbic Cortex
The Bigger Picture: The Ethics of Brain Stimulation in Human Beings
The Limbic Cortex and Emotions
The Interoceptive Insula: The "Feeling" Side of Emotions
Cingulate Cortex: A Motor Cortex for the Limbic System
Neuroscience of Everyday Life: Mental Effort
Ventromedial Prefrontal Cortex: A Generator of Gut Feelings
Limbic Association Cortex: Modulation of Emotion
The Mechanisms of Emotional Reappraisal
Brain Injury, Brain Stimulation, and Emotion Regulation
Neurochemical Influences on Emotion
Case Study: A Cure Born of Desperation
Serotonin and Mood
Norepinephrine and Mood
GABA and Anxiety

Chapter 14: Motivation and Reward
Learning Objectives
Starting Out: "More Important Than Survival Itself"
Motivation and Survival
Addiction: An Illness of Motivation
Why Motivation Matters
Feelings: The Sensory Side of Motivation
The Circuitry of Motivation: Basic Drives
Hypothalamus and Homeostatic Drives
Amygdala and External-World Drives
Midbrain Dopamine Neurons and the Common Currency of Motivation
Reward, Learning, and the Brain
Defining Reward
Learning from Reward Using Prediction Error
"Liking" Is Different from "Wanting"
Opioids and the Sensation of Pleasure
Opioids, Opioid Receptors, and Opioid Functions
Opioids and Reward
Dopamine, Learning, Motivation, and Reward
Dopamine Functions in Motivation and Reward
Unifying the Functions of Dopamine
Research Methods: Measuring Neurotransmitter Levels in the Brain
Neurotransmitters Are Messengers, Not Functions
Addiction: Pathological Learning and Motivation
Addictive Substances Have Distorted Reward Value
Neuroscience of Everyday Life: The Pursuit of Happiness
Addiction Is a Result of Pathological Learning
The Circuitry and Chemistry of Addiction
Unlearning Addiction
The Challenge of Treatment
Case Study: Pathological Gambling in a Patient with Parkinson's Disease
Existing Approaches to Treatment
Future Approaches to Treatment
The Bigger Picture: Finding the Motivation to Change

Chapter 15: Social Cognition
Learning Objectives
Starting Out: Why Risk Your Life for a Yellow T-shirt?
Social Perception
What's in a Face?
Do I Look Like a Liar to You?
Neuroscience of Everyday Life: A Poker Face
Social Knowledge and the Temporal Pole
Social Signals and the Superior Temporal Sulcus
Social Thinking: Theory of Mind
What Is Theory of Mind?
Neural Mechanisms of Theory of Mind
Mirror Neurons and Theory of Mind
Disorders of Theory of Mind
Social Feelings: Empathy and Its Many Components
An Emotional Theory of Mind
Empathy, Sympathy, and Compassion
Neural Mechanisms of Emotional Mimicry and Contagion
Neural Mechanisms of Empathy, Sympathy, and Antipathy
Disorders of Empathy
Social Emotions, Motivations, and Behavior
Social Emotions from Theory of Mind
Case Study: Acquired Sociopathy
Social Emotions from Social Values
Social Reward and Social Aversion
The Anatomy of a Lie
Neurotransmitters and Social Behavior
Research Methods: Transcranial Direct Current Stimulation
An Ancient and Fundamental System
Oxytocin
Vasopressin
The Bigger Picture: The Brave New World of the "Cuddle Hormone"?
The Social Self
The Wondrous Self-Awareness of the Human Brain
Forms of Self-Awareness
Why Bother with Self-Awareness?
Neural Correlates of Self-Awareness
Disorders of Self-Awareness
Self-Awareness and Social Cognition
Case Study: The Man in the Mirror

Part V: Disorders of Brain and Behavior

Chapter 16: Neurological and Psychiatric Disorders
Learning Objectives
Starting Out: Epilepsy: "The Sacred Disease"
Alzheimer's Disease: Burning Out with Age?
Frontotemporal Dementia: Like a Cancer of the Soul
Case Study: Ravel and "Bolero"
Huntington's Disease: A Genetic Rarity, in Two Senses
Tourette Syndrome: A Case of Involuntary Volition?
Obsessive-Compulsive Disorder: Neurological or Psychiatric?
Research Methods: Voxel-Based Morphometry
Schizophrenia: A Dementia of the Young
Bipolar Disorder
Depression: A Global Burden
Impact of Depression
Case Study: A Lifetime Studying, and Living with, Bipolar Disorder
Causes of Depression
Neurochemical Effects of Depression on the Brain
Functional Effects of Depression on the Brain
Treatment of Depression