This text is a unique inquiry-based introduction to behavioral neuroscience. Each chapter focuses on a central question (i.e., How Does the Nervous System Function?) guiding students through the key concepts. The authors emphasize a distinctive clinical perspective, with examples showing students what happens when common neuronal processes malfunction with real life applications. The book reflects the latest findings at the time of publication on specific disorders including Parkinson disease, Alzheimer disease, depression and drug dependency, sleep disorders, schizophrenia, glaucoma, and abnormal development related to prenatal experience.

Chapter 1
What Are the Origins of Brain and Behavior?
CLINICAL FOCUS 1-1 Living with Traumatic Brain Injury
1-1 The Brain in the Twenty-First Century
Why Study Brain and Behavior?
What Is the Brain?
What Is Behavior?
1-2 Theories of Brain and Behavior
Aristotle and Mentalism
Descartes and Dualism
COMPARATIVE FOCUS 1-2 The Speaking Brain
Darwin and Materialism
EXPERIMENT 1-1 Question: How do parents transmit heritable factors to offspring?
Toward Contemporary Perspectives on Consciousness
1-3 Evolution of Brains and of Behavior
Origin of Brain Cells and Brains
The Basics: Classification of Life
Evolution of Nervous Systems in Animals
Chordate Brain
1-4 Evolution of the Human Brain and Behavior
Humans: Members of the Primate Order
Australopithecus: Our Distant Ancestor
The First Humans
Relating Brain Complexity and Behavior
COMPARATIVE FOCUS 1-3 The Elephant’s Brain
Why the Hominin Brain Became More Complex
1-5 Modern Human Brain Size, Intelligence, and Culture
The Significance of Human Brain Size Comparisons
The Significance of Human Intelligence
The Significance of Human Culture

Chapter 2
What Is the Nervous System’s Functional Anatomy?
RESEARCH FOCUS 2-1 Agenesis of the Cerebellum
2-1 Overview of Brain Function and Structure
Plastic Patterns of Neural Organization
Functional Organization of the Nervous System
The Brain’s Surface Features
The Basics: Finding Your Way Around the Brain
CLINICAL FOCUS 2-2 Meningitis and Encephalitis
The Brain’s Internal Features
CLINICAL FOCUS 2-3 Stroke
2-2 The Conserved Pattern of Nervous System Development
Comparative Brain Evolution
The Nervous System and Intelligent Behavior
EXPERIMENT 2-1 Question: Does intelligent behavior require a vertebrate
nervous system organization?
2-3 The Central Nervous System: Mediating Behavior
Spinal Cord
Brainstem
Forebrain
Cerebral Cortex
Basal Ganglia
2-4 Somatic Nervous System: Transmitting Information
Cranial Nerves
Spinal Nerves
Somatic Nervous System Connections
Integrating Spinal Function
CLINICAL FOCUS 2-4 Bell Palsy
2-5 Autonomic and Enteric Nervous Systems: Visceral Relations
ANS: Regulating Internal Functions
ENS: Controlling the Gut
2-6 Ten Principles of Nervous System Function
Principle 1: Neuronal Circuits Are the Functional Units of the Nervous System
Principle 2: Sensory and Motor Divisions Pervade the Nervous System
Principle 3: The CNS Functions on Multiple Levels and Is Organized Hierarchically and in Parallel
Principle 4: Many Brain Circuits Are Crossed
Principle 5: Brain Functions Are Localized and Distributed
Principle 6: The Brain Is Symmetrical and Asymmetrical
Principle 7: The Nervous System Works by Juxtaposing Excitation and Inhibition
Principle 8: The Brain Divides Sensory Input for Object Recognition and Movement
Principle 9: The Nervous System Produces Movement in a Perceptual World the Brain Constructs
Principle 10: Neuroplasticity Is the Hallmark of Nervous System Functioning

Chapter 3
What Are the Nervous System’s Functional Units?
RESEARCH FOCUS 3-1 A Genetic Diagnosis
3-1 Cells of the Nervous System
Neurons: The Basis of Information Processing
EXPERIMENT 3-1 Question: Can the principles of neural excitation and inhibition control the activity of a simple robot that behaves like a cricket?
Classes of Glial Cells
CLINICAL FOCUS 3-2 Brain Tumors
3-2 Internal Structure of a Cell
The Basics: Chemistry Review
The Cell as a Factory
Cell Membrane: Barrier and Gatekeeper
The Nucleus and Protein Synthesis
The Endoplasmic Reticulum and Protein Manufacture
Proteins and RNA: The Cell’s Products
Golgi Bodies and Microtubules: Protein Packaging and Shipment
Crossing the Cell Membrane: Channels, Gates, and Pumps
3-3 Genes, Cells, and Behavior
Mendelian Genetics and the Genetic Code
Applying Mendel’s Principles
CLINICAL FOCUS 3-3 Huntington Disease
Genetic Engineering
The Epigenetic Code

Chapter 4
How Do Neurons Use Electrical Signals to Transmit Information?
4-1 Searching for Electrical Activity in the Nervous System
CLINICAL FOCUS 4-1 Epilepsy
Early Clues That Linked Electricity and Neuronal Activity
The Basics: Electricity and Electrical Stimulation
Tools for Measuring a Neuron’s Electrical Activity
How Ion Movement Produces Electrical Charges
4-2 Electrical Activity at the Dendritic and Cell Body Membrane
Resting Potential
Maintaining the Resting Potential
How Neurons Integrate Information
Excitatory and Inhibitory Postsynaptic Potentials
EXPERIMENT 4-1 Question: How does stimulating a neuron influence its excitability?
Summation of Inputs
4-3 Generation and Propagation of the Action Potential
Triggering an Action Potential
Role of Voltage-Activated Ion Channels
The Versatile Neuron
RESEARCH FOCUS 4-3 Optogenetics and Light-Sensitive Ion Channels
Action Potentials and Refractory Periods
Nerve Impulse
Refractory Periods and Nerve Action
Saltatory Conduction and the Myelin Sheath
CLINICAL FOCUS 4-2 Multiple Sclerosis
4-4 Into the Nervous System and Back Out
How Sensory Stimuli Produce Action Potentials
How Nerve Impulses Produce Movement
CLINICAL FOCUS 4-4 ALS: Amyotrophic Lateral Sclerosis

Chapter 5
How Do Neurons Communicate and Adapt?
5-1 A Chemical Message
RESEARCH FOCUS 5-1 The Basis of Neural Communication in a Heartbeat
EXPERIMENT 5-1 Question: How does a neuron pass on a message?
CLINICAL FOCUS 5-2 Parkinson Disease
Structure of Synapses
Neurotransmission in Five Steps
Varieties of Synapses
Excitatory and Inhibitory Messages
RESEARCH FOCUS 5-3 Dendritic Spines: Small but Mighty
Evolution of Complex Neurotransmission Systems
5-2 Varieties of Neurotransmitters and Receptors
Four Criteria for Identifying Neurotransmitters
Classes of Neurotransmitters
CLINICAL FOCUS 5-4 Awakening with L-Dopa
Varieties of Receptors
5-3 Neurotransmitter Systems and Behavior
Neurotransmission in the Somatic Nervous System (SNS)
Dual Activating Systems of the Autonomic Nervous System (ANS)
Enteric Nervous System (ENS) Autonomy
Four Activating Systems in the Central Nervous System
CLINICAL FOCUS 5-5 The Case of the Frozen Addict
5-4 Hormones
Hierarchical Control of Hormones
Classes and Functions of Hormones
Homeostatic Hormones
Anabolic–Androgenic Steroids
Glucocorticoids and Stress

Chapter 6
How Do Drugs Influence Brain and Behavior?
6-1 Principles of Psychopharmacology
CLINICAL FOCUS 6-1 Cognitive Enhancement?
Drug Routes into the Nervous System
Drug Action at Synapses: Agonists and Antagonists
An Acetylcholine Synapse: Examples of Drug Action
Tolerance
EXPERIMENT 6-1 Question: Will the constant consumption of alcohol produce tolerance?
Sensitization
EXPERIMENT 6-2 Question: Does the injection of a drug always produce the same behavior?
6-2 Psychoactive Drugs
Adenosinergic
Cholinergic
GABAergic
CLINICAL FOCUS 6-2 Fetal Alcohol Spectrum Disorder
Glutamatergic
Dopaminergic
RESEARCH FOCUS 6-3 Attention-Deficit/Hyperactivity Disorder
Serotonergic
CLINICAL FOCUS 6-4 Major Depression
Opioidergic
CLINICAL FOCUS 6-5 The Opioid Overdose Death Epidemic
Cannabinergic
6-3 Factors Influencing Individual Responses to Drugs
Behavior on Drugs
Substance Use Disorder, Withdrawal, and Addiction
Risk Factors for Substance Use Disorder
Individual and Sex Differences in Substance Use Disorder
6-4 Explaining Substance Use and Misuse
Wanting-and-Liking Theory
Why Doesn’t Everyone Develop Substance Use Disorder?
Issues Related to Treating Substance Use Disorder
Can Drugs Cause Brain Damage?
CLINICAL FOCUS 6-6 Drug-Induced Psychosis

Chapter 7
How Do We Study the Brain’s Structures and Functions?
7-1 Measuring and Manipulating Brain and Behavior
RESEARCH FOCUS 7-1 Tuning in to Language
Early Origins of Behavioral Neuroscience
RESEARCH FOCUS 7-2 Brainbow: Rainbow Neurons
EXPERIMENT 7-1 Question: Do hippocampal neurons contribute to memory formation?
Methods of Behavioral Neuroscience
Manipulating Brain–Behavior Interactions
CLINICAL FOCUS 7-3 Brain Organoids and Personalized Medicine
7-2 Measuring the Brain’s Electrical Activity
Recording Action Potentials from Single Cells
EEG: Recording Graded Potentials from Thousands of Cells
Mapping Brain Function with Event-Related Potentials
Magnetoencephalography
7-3 Anatomical Imaging Techniques: CT and MRI
7-4 Functional Brain Imaging
Functional Magnetic Resonance Imaging
Optical Tomography
Positron Emission Tomography
7-5 Chemical and Genetic Measures of Brain and Behavior
Measuring Brain Chemistry
Measuring Genes in Brain and Behavior
Epigenetics: Measuring Gene Expression
7-6 Comparing Neuroscience Research Methods
7-7 Computational Neuroscience and Deep Learning
7-8 Ethical Considerations
Neuroethics
Using Nonhuman Species in Brain-Behavior Research

Chapter 8
How Does the Nervous System Develop and Adapt?
8-1 Three Perspectives on Brain Development
RESEARCH FOCUS 8-1 Linking Socioeconomic Status to Cortical Development
Correlating Emerging Brain Structures with Emerging Behaviors
Correlating Emerging Behaviors with Neural Maturation
Identifying Influences on Brain and Behavior
8-2 Neurobiology of Development
Gross Development of the Human Nervous System
Origins of Neurons and Glia
Neuronal Growth and Development
CLINICAL FOCUS 8-2 Autism Spectrum Disorder
Glial Development
Unique Aspects of Frontal Lobe Development
8-3 Using Emerging Behaviors to Infer Neural Maturation
Motor Behaviors
Language Development
Development of Problem-Solving Ability
EXPERIMENT 8-1 Question: In what sequence do the forebrain structures required for learning and memory mature?
A Caution about Linking Correlation to Causation
8-4 Brain Development and the Environment
Experience and Cortical Organization
RESEARCH FOCUS 8-3 Keeping Brains Young by Making Music
Environmental Influences on the Pace of Brain Development
Experience and Neural Connectivity
Critical Periods for Experience and Brain Development
Gut Bacteria and Brain Development
8-5 Abnormal Experience and Brain Development
Aversive Experience and Brain Development
CLINICAL FOCUS 8-4 Romanian Orphans
Injury and Brain Development
Other Sources of Abnormal Brain Development
CLINICAL FOCUS 8-5 Schizophrenia
Developmental Disability
How Do Any of Us Develop a Normal Brain?

Chapter 9
How Do We Sense, Perceive, and See the World?
9-1 Nature of Sensation and Perception
CLINICAL FOCUS 9-1 Migraines and a Case of Blindsight
Sensory Receptors
Neural Relays
Sensory Coding and Representation
Perception
9-2 The Visual System’s Functional Anatomy
Structure of the Retina
The Basics: Visible Light and the Structure of the Eye
Photoreceptors
CLINICAL FOCUS 9-2 Visual Illuminance
Types of Retinal Neurons
CLINICAL FOCUS 9-3 Glaucoma
Visual Pathways
Dorsal and Ventral Visual Streams
9-3 Location in the Visual World
Coding Location in the Retina
Location in the Lateral Geniculate Nucleus and Region V1
Visual Corpus Callosum
9-4 Neuronal Activity
Seeing Shape
Seeing Color
RESEARCH FOCUS 9-4 Color-Deficient Vision
Neuronal Activity in the Dorsal Stream
9-5 The Visual Brain in Action
Injury to the Visual Pathway Leading to the Cortex
Injury to the What Pathway
Injury to the How Pathway
9-6 Plasticity in the Visual Pathways

Chapter 10
How Do We Hear, Speak, and Make Music?
10-1 Sound Waves: Stimulus for Audition
RESEARCH FOCUS 10-1 Evolution of Music and Language
Physical Properties of Sound Waves
CLINICAL FOCUS 10-2 Tinnitus
Perception of Sound
Properties of Spoken Language and Music as Sounds
10-2 Functional Anatomy of the Auditory System
Structure of the Ear
Auditory Receptors
RESEARCH FOCUS 10-3 Otoacoustic Emissions
Pathways to the Auditory Cortex
RESEARCH FOCUS 10-4 Seeing with Sound
Auditory Cortex
10-3 Neural Activity and Hearing
Hearing Pitch
CLINICAL FOCUS 10-5 Cochlear Implant in a Deaf Child
Detecting Loudness
Detecting Location
Detecting Patterns in Sound
10-4 Anatomy of Language and Music
Processing Language
Processing Music
RESEARCH FOCUS 10-6 The Brain’s Music System
10-5 Auditory Communication in Nonhuman Species
Birdsong
Whale Songs

Chapter 11
How Does the Nervous System Respond to Stimulation and Produce Movement?
11-1 Hierarchical and Parallel Movement Control
RESEARCH FOCUS 11-1 Neuroprosthetics
The Basics: Relating the Somatosensory and Motor Systems
Forebrain: Initiating Movement
Brainstem: Species-Typical Movement
EXPERIMENT 11-1 Question: What are the effects of brainstem stimulation under different conditions?
CLINICAL FOCUS 11-2 Cerebral Palsy
Spinal Cord: Executing Movement
11-2 Motor System Organization
Motor Cortex
Mapping the Motor Cortex
Motor Cortex and Skilled Movement
EXPERIMENT 11-2 Question: How does the motor cortex take part in the control of movement?
Plasticity in the Motor Cortex
EXPERIMENT 11-3 Question: What is the effect of rehabilitation on the cortical representation of the forelimb after brain damage?
Corticospinal Tracts
Motor Neurons
Control of Muscles
11-3 Basal Ganglia, Cerebellum, and Movement
Basal Ganglia and the Force of Movement
CLINICAL FOCUS 11-3 Tourette Syndrome
Cerebellum and Movement Skill
EXPERIMENT 11-4 Question: Does the cerebellum help make adjustments required to keep movements accurate?
11-4 Somatosensory System Receptors and Pathways
Somatosensory Receptors and Perception
Posterior Root Ganglion Neurons
Somatosensory Pathways to the Brain
Spinal Reflexes
Feeling and Treating Pain
RESEARCH FOCUS 11-4 Phantom Limb Pain
Vestibular System and Balance
11-5 Exploring the Somatosensory Cortex
Somatosensory Body Map
RESEARCH FOCUS 11-5 Tickling
Secondary Somatosensory Cortex
Effects of Somatosensory Cortex Damage
Somatosensory Cortex and the Hierarchy and Parallel Control of Movement

Chapter 12
What Causes Emotional and Motivated Behavior?
12-1 Identifying the Causes of Behavior
RESEARCH FOCUS 12-1 The Pain of Rejection
Behavior for Brain Maintenance
Neural Circuits and Behavior
Evolutionary Influences on Behavior
Environmental Influences on Behavior
12-2 Neuroanatomy of Motivated Behavior
Regulatory and Nonregulatory Behavior
Activities of the Hypothalamic Circuit
12-3 The Role of the Chemical Senses in Motivated Behavior
Olfaction
Gustation
Impairments in the Chemical Sense and Behavior
12-4 Control of Regulatory Behavior
Controlling Eating
EXPERIMENT 12-1 Question: Does the hypothalamus play a role in eating?
CLINICAL FOCUS 12-2 Diets and Rhythms
Controlling Drinking
12-5 Sexual Differences and Sexual Behavior
Sexual Differentiation of the Brain
RESEARCH FOCUS 12-3 The Brain Gender Continuum
Effects of Sex Hormones on the Brain
CLINICAL FOCUS 12-4 Androgen Insensitivity Syndrome and the Androgenital Syndrome
Neural Control of Sexual Behavior
Sexual Orientation, Sexual Identity, and Brain Organization
Cognitive Influences on Sexual Behavior
12-6 The Neural Control of Emotion
Theories of Emotion
Emotion and the Limbic Circuit
CLINICAL FOCUS 12-5 Agenesis of the Frontal Lobes
12-7 Reward
The Reward System
Mapping Pleasure in the Brain
Pleasure Electrodes?

Chapter 13
Why Do We Sleep and Dream?
13-1 A Clock for All Seasons
CLINICAL FOCUS 13-1 Doing the Right Thing at the Right Time
Biological Rhythms
The Origin of Biorhythms
EXPERIMENT 13-1 Question: Is plant movement exogenous or endogenous?
Free-Running Rhythms
Zeitgebers
CLINICAL FOCUS 13-2 Seasonal Affective Disorder
13-2 The Suprachiasmatic Biological Clock
Suprachiasmatic Rhythms
Keeping Time
RESEARCH FOCUS 13-3 Synchronizing Biorhythms at the Molecular Level
Pacemaking Circadian Rhythms
Pacemaking Circannual Rhythms
Chronotypes
Rhythms of Cognitive and Emotional Behavior
13-3 Sleep Stages and Dreaming
Measuring How Long We Sleep
Measuring Sleep
Stages of Waking and Sleeping
A Typical Night’s Sleep
Contrasting N-Sleep and R-Sleep
Dreaming
What We Dream About
13-4 What Does Sleep Accomplish?
Sleep as a Biological Adaptation
Sleep as a Restorative Process
Sleep for Memory Storage
Brain Events During Memory Storage
13-5 The Neural Bases of Sleep
Reticular Activating System and Sleep
Neural Basis of R-Sleep
13-6 Disorders of Sleep
Insomnia: Inability to Sleep
Hypersomnia: Inability to Stay Awake
Breathing Disorders
Parasomnias
Cataplexy
CLINICAL FOCUS 13-4 Orexin
Sleep-Related Movement Disorder
13-7 What Does Sleep Tell Us about the Brain?

Chapter 14
How Do We Learn and Remember?
14-1 Connecting Learning and Memory
CLINICAL FOCUS 14-1 Remediating Dyslexia
Studying Learning and Memory in the Laboratory
EXPERIMENT 14-1 Question: Does an animal learn the association between emotional experience and environmental stimuli?
Two Categories of Memory
What Makes Explicit and Implicit Memory Different?
What Is Special about Personal Memories?
14-2 Dissociating Memory Circuits
Disrupting Explicit Memory
CLINICAL FOCUS 14-2 Patient Boswell’s Amnesia
Disrupting Implicit Memory
14-3 Neural Systems Underlying Explicit and Implicit Memories
Neural Circuit for Explicit Memories
CLINICAL FOCUS 14-3 Alzheimer Disease
CLINICAL FOCUS 14-4 Korsakoff Syndrome
Consolidation of Explicit Memories
Neural Circuit for Implicit Memories
Neural Circuit for Emotional Memories
Evolution of Memory Systems
14-4 Structural Basis of Brain Plasticity
Habituation and Sensitization in Aplysia
EXPERIMENT 14-2 Question: What happens to the gill response after repeated stimulation?
EXPERIMENT 14-3 Question: What happens to the gill response in sensitization?
Long-Term Potentiation
Measuring Synaptic Change
Enriched Experience and Plasticity
Sensory or Motor Training and Plasticity
EXPERIMENT 14-4 Question: Does the learning of a fine motor skill alter the cortical motor map?
RESEARCH FOCUS 14-5 Movement, Learning, and Neuroplasticity
Epigenetics of Memory
Plasticity, Hormones, Trophic Factors, and Drugs
EXPERIMENT 14-5 Question: What effect do repeated doses of amphetamine, a psychomotor stimulant, have on neurons?
Some Guiding Principles of Brain Plasticity
14-5 Recovery from Brain Injury
Donna’s Experience with Traumatic Brain Injury

Chapter 15
How Does the Brain Think?
15-1 The Nature of Thought
RESEARCH FOCUS 15-1 Split Brain
Characteristics of Human Thought
Neural Units of Thought
COMPARATIVE FOCUS 15-2 Animal Intelligence
EXPERIMENT 15-1 Question: How do individual neurons mediate cognitive activity?
15-2 Cognition and the Association Cortex
Knowledge about Objects
Multisensory Integration
Spatial Cognition
Attention
Planning and Executing Functioning
Imitation and Understanding
15-3 Expanding Frontiers of Cognitive Neuroscience
Mapping the Brain
CLINICAL FOCUS 15-3 Neuropsychological Assessment
Mapping the Brain
Cognition and the Cerebellum
Social Neuroscience
Neuroeconomics
15-4 Cerebral Asymmetry in Thinking
Anatomical Asymmetry
Functional Asymmetry in Neurological Patients
Functional Asymmetry in the Healthy Brain
Functional Asymmetry in the Split Brain
EXPERIMENT 15-2 Question: Will severing the corpus callosum affect the way in which the brain responds?
EXPERIMENT 15-3 (A) Question: How can the right hemisphere of a split-brain patient show that it knows information? (B) Question: What happens if both hemispheres are asked to respond to competing information?
Explaining Cerebral Asymmetry
Left Hemisphere, Language, and Thought
15-5 Variations in Cognitive Organization
Sex Differences in Cognitive Organization
Handedness and Cognitive Organization
CLINICAL FOCUS 15-4 Sodium Amobarbital Test
Synesthesia
15-6 Intelligence
Concept of General Intelligence
Divergent and Convergent Intelligence
Intelligence, Heredity, Epigenetics, and the Synapse
How Smart Brains Differ
15-7 Consciousness
Why Are We Conscious?
EXPERIMENT 15-4 Question: Can people alter their movements without conscious awareness?
What Is the Neural Basis of Consciousness?

Chapter 16
What Happens When the Brain Misbehaves?
16-1 Multidisciplinary Contributions to Brain and Behavior
RESEARCH FOCUS 16-1 Posttraumatic Stress Disorder
Clinical Neuroscience
Behavioral Disorders
16-2 Psychiatric Disorders
Schizophrenia Spectrum and Other Psychotic Disorders
Mood Disorders
RESEARCH FOCUS 16-2 Antidepressant Action and Brain Repair
16-3 Neurological Disorders
Traumatic Brain Injury
CLINICAL FOCUS 16-3 Concussion
Stroke
CLINICAL FOCUS 16-4 Cerebral Aneurysms
Epilepsy
Multiple Sclerosis
Neurocognitive Disorders
Treatments for Neurocognitive Disorders
RESEARCH FOCUS 16-5 Treating Behavioral Disorders with Transcranial Magnetic Stimulation
16-4 Research Challenges
Organizational Complexity
Systemic Complexity
Neuronal Plasticity
Compensatory Plasticity
Technological Resolution
Modeling Simplicity
Modeling Limitations
16-5 Posttraumatic Growth and Apathy