Neil Carlson, U Mass Amherst

Professor Emeritus at the University of Massachusetts, Amherst, Neil Carlson's research has focused on the relationship between the brain and behavior—specifically, the role of the limbic system in learning and species-typical behavior. Dr. Carlson received his undergraduate degree and Ph.D. from the University of Illinois.  In addition to writing Foundations of Behavioral Neuroscience, Dr. Carlson is also the author of Physiology of Behavior and co-author of Psychology: The Science of Behavior.  Dr. Carlson's books have been translated into Spanish, Italian, Portuguese, German, Hebrew, Japanese, Chinese, and Korean.

Contents

Chapter 1 Origins of Behavioral Neuroscience 1

PROLOGUE ¿¿Rene’s Inspiration 2

Understanding Human Consciousness: A Physiological Approach 3

Split Brains 3

The Nature of Behavioral Neuroscience 6

The Goals of Research 6

Biological Roots of Behavioral Neuroscience 7

Natural Selection and Evolution 10

Functionalism and the Inheritance of Traits 10

Evolution of the Human Species 12

Evolution of Large Brains 14

Ethical Issues in Research with Animals 16

Careers in Neuroscience 19

Strategies for Learning 20

EPILOGUE ¿¿Models of Brain Functions 22

Key Concepts 23

Suggested Readings 23

Additional Resources 23

Chapter 2 Structure and Functions of Cells of the Nervous System 24

PROLOGUE ¿¿Unresponsive Muscles 25

Cells of the Nervous System 26

Neurons 26

Supporting Cells 30

The Blood—Brain Barrier 33

Communication Within a Neuron 34

Neural Communication: An Overview 34

Measuring Electrical Potentials of Axons 36

The Membrane Potential: Balance of Two Forces 38

The Action Potential 40

Conduction of the Action Potential 42

Communication Between Neurons 45

Structure of Synapses 45

Release of Neurotransmitter 46

Activation of Receptors 47

Postsynaptic Potentials 48

Termination of Postsynaptic Potentials 50

Effects of Postsynaptic Potentials: Neural Integration 51

Autoreceptors 52

Axoaxonic Synapses 52

Nonsynaptic Chemical Communication 52

EPILOGUE ¿¿Myasthenia Gravis 54

Key Concepts 55

Suggested Readings 55

Additional Resources 55

Chapter 3 Structure of the Nervous System 56

PROLOGUE ¿¿The Left Is Gone 57

Basic Features of the Nervous System 57

An Overview 60

Meninges 60

The Ventricular System and Production of Cerebrospinal Fluid 60

The Central Nervous System 63

Development of the Central Nervous System 63

The Forebrain 67

The Midbrain 76

The Hindbrain 78

The Spinal Cord 78

The Peripheral Nervous System 80

Spinal Nerves 80

Cranial Nerves 81

The Autonomic Nervous System 83

EPILOGUE ¿¿Unilateral Neglect 85

Key Concepts 87

Suggested Readings 87

Additional Resources 87

Chapter 4 Psychopharmacology 88

PROLOGUE ¿¿A Contaminated Drug 89

Principles of Psychopharmacology 89

Pharmacokinetics 90

Drug Effectiveness 92

Effects of Repeated Administration 93

Placebo Effects 94

Sites of Drug Action 95

Effects on Production of Neurotransmitters 96

Effects on Storage and Release of Neurotransmitters 97

Effects on Receptors 97

Effects on Reuptake or Destruction of Neurotransmitters 98

Neurotransmitters and Neuromodulators 99

Acetylcholine 99

The Monoamines 101

Amino Acids 106

Peptides 108

Lipids 109

Nucleosides 110

Soluble Gases 111

EPILOGUE ¿¿Helpful Hints from a Tragedy 113

Key Concepts 114

Suggested Readings 114

Additional Resources 114

Chapter 5 Methods and Strategies of Research 115

PROLOGUE ¿¿Heart Repaired, Brain Damaged 116

Experimental Ablation 117

Evaluating the Behavioral Effects of Brain Damage 117

Producing Brain Lesions 117

Stereotaxic Surgery 119

Histological Methods 121

Tracing Neural Connections 122

Studying the Structure of the Living Human Brain 125

Recording and Stimulating Neural Activity 128

Recording Neural Activity 128

Recording the Brain’s Metabolic and

Synaptic Activity 131

Stimulating Neural Activity 133

Neurochemical Methods 136

Finding Neurons that Produce Particular Neurochemicals  136

Localizing Particular Receptors 137

Measuring Chemicals Secreted in the Brain 138

Genetic Methods 140

Twin Studies 140

Adoption Studies 141

Targeted Mutations 142

Antisense Oligonucleotides 142

EPILOGUE ¿¿Watch the Brain Waves 143

Key Concepts 144

Suggested Readings 145

Additional Resources 145

Chapter 6 Vision 146

PROLOGUE ¿¿Seeing Without Perceiving 147

The Stimulus 148

Anatomy of the Visual System 149

The Eyes 149

Photoreceptors 151

Connections Between Eye and Brain 152

Coding of Visual Information in the Retina 154

Coding of Light and Dark 154

Coding of Color 155

Analysis of Visual Information: Role of the Striate Cortex 158

Anatomy of the Striate Cortex 158

Orientation and Movement 158

Spatial Frequency 159

Retinal Disparity 160

Color 160

Modular Organization of the Striate Cortex 161

Analysis of Visual Information: Role of the Visual

Association Cortex 162

Two Streams of Visual Analysis 162

Perception of Color 164

Perception of Form 166

Perception of Movement 169

Perception of Spatial Location 172

EPILOGUE ¿¿Case Studies 175

Key Concepts 176

Suggested Readings 176

Additional Resources 176

Chapter 7 Audition, the Body Senses, and the Chemical Senses 177

PROLOGUE ¿¿All in Her Head? 178

Audition 178

The Stimulus 179

Anatomy of the Ear 179

Auditory Hair Cells and the Transduction of Auditory Information 181

The Auditory Pathway 182

Perception of Pitch 183

Perception of Timbre 186

Perception of Spatial Location 186

Perception of Complex Sounds 188

Vestibular System 191

Anatomy of the Vestibular Apparatus 192

The Vestibular Pathway 193

Somatosenses 194

The Stimuli 194

Anatomy of the Skin and Its Receptive Organs 195

Perception of Cutaneous Stimulation 195

The Somatosensory Pathways 198

Perception of Pain 199

Gustation 201

The Stimuli 201

Anatomy of the Taste Buds and Gustatory Cells 202

Perception of Gustatory Information 202

The Gustatory Pathway 203

Olfaction 204

The Stimulus 206

Anatomy of the Olfactory Apparatus 206

Transduction of Olfactory Information 207

Perception of Specific Odors 207

EPILOGUE ¿¿Natural Analgesia 209

Key Concepts 210

Suggested Readings 211

Additional Resources 211

Chapter 8 Sleep and Biological Rhythms 212

PROLOGUE ¿¿Waking Nightmares 213

A Physiological and Behavioral

Description of Sleep 213

Disorders of Sleep 217

Insomnia 217

Narcolepsy 218

REM Sleep Behavior Disorder 220

Problems Associated with Slow-Wave Sleep 220

Why Do We Sleep? 221

Functions of Slow-Wave Sleep 222

Functions of REM Sleep 223

Sleep and Learning 224

Physiological Mechanisms of Sleep and Waking 226

Chemical Control of Sleep 226

Neural Control of Arousal 227

Neural Control of Slow-Wave Sleep 230

Neural Control of REM Sleep 232

Biological Clocks 236

Circadian Rhythms and Zeitgebers 236

The Suprachiasmatic Nucleus 236

Changes in Circadian Rhythms: Shift Work and Jet Lag 239

EPILOGUE ¿¿Functions of Dreams 241

Key Concepts 242

Suggested Readings 243

Additional Resources 243

Chapter 9 Reproductive Behavior 244

PROLOGUE ¿¿From Boy to Girl 245

Sexual Development 245

Production of Gametes and Fertilization 245

Development of the Sex Organs 246

Sexual Maturation 249

Hormonal Control of Sexual Behavior 252

Hormonal Control of Female Reproductive Cycles 252

Hormonal Control of Sexual Behavior of Laboratory Animals 253

Organizational Effects of Androgens on Behavior: Masculinization and Defeminization 255

Effects of Pheromones 255

Human Sexual Behavior 258

Sexual Orientation 259

Neural Control of Sexual Behavior 264

Males 264

Females 266

Formation of Pair Bonds 267

Parental Behavior 269

Maternal Behavior of Rodents 269

Hormonal Control of Maternal Behavior 270

Neural Control of Maternal Behavior 271

Neural Control of Paternal Behavior 271

EPILOGUE ¿¿From Boy to Girl and Back Again 272

Key Concepts 273

Suggested Readings 274

Additional Resources 274

Chapter 10 Emotion 275

PROLOGUE ¿¿Intellect and Emotion 276

Emotions as Response Patterns 276

Fear 277

Anger, Aggression, and Impulse Control 281

Communication of Emotions 288

Facial Expression of Emotions: Innate Responses 288

Neural Basis of the Communication of Emotions: Recognition 290

Neural Basis of the Communication of Emotions: Expression 292

Feelings of Emotions 295

The James-Lange Theory 295

Feedback from Simulated Emotions 296

EPILOGUE ¿¿Mr. V. Revisited 298

Key Concepts 299

Suggested Readings 300

Additional Resources 300

Chapter 11 Ingestive Behavior 301

PROLOGUE ¿¿Out of Control 302

Physiological Regulatory Mechanisms 302

Drinking 304

Some Facts About Fluid Balance 304

Two Types of Thirst 305

Eating: Some Facts About Metabolism 309

What Starts a Meal? 312

Signals from the Environment 312

Signals from the Stomach 312

Metabolic Signals 313

What Stops a Meal? 315

Gastric Factors 315

Intestinal Factors 315

Liver Factors 316

Insulin 317

Long-Term Satiety: Signals from Adipose Tissue 317

Brain Mechanisms 318

Brain Stem 318

Hypothalamus 319

Obesity 323

Possible Causes 323

Treatment 326

Anorexia Nervosa/Bulimia Nervosa 330

Possible Causes 330

Treatment 332

EPILOGUE ¿¿An Insatiable Appetite 334

Key Concepts 334

Suggested Readings 335

Additional Resources 335

Chapter 12 Learning and Memory 336

PROLOGUE ¿¿Every Day Is Alone 337

The Nature of Learning 337

Synaptic Plasticity: Long-Term Potentiation and Long-Term Depression 341

Induction of Long-Term Potentiation 341

Role of NMDA Receptors 343

Mechanisms of Synaptic Plasticity 345

Long-Term Depression 348

Perceptual Learning 349

Classical Conditioning 351

Instrumental Conditioning 353

Role of the Basal Ganglia 353

Reinforcement 355

Relational Learning 358

Human Anterograde Amnesia 358

Spared Learning Abilities 360

Declarative and Nondeclarative Memories 361

Anatomy of Anterograde Amnesia 363

Role of the Hippocampal Formation in Memory Consolidation 363

Episodic and Semantic Memories 365

Spatial Memory 366

Relational Learning in Laboratory Animals 367

Role of Hippocampal Neurogenesis in Learning 372

EPILOGUE ¿¿What Causes Confabulation? 374

Key Concepts 375

Suggested Readings 375

Additional Resources 376

Chapter 13 Human Communication 377

Speech Production and Comprehension: Brain Mechanisms 378

Lateralization 379

Speech Production 380

Speech Comprehension 382

Aphasia in Deaf People 392

Prosody: Rhythm, Tone, and Emphasis in Speech 393

Stuttering 394

Disorders of Reading and Writing 397

Pure Alexia 397

Toward an Understanding of Reading 399

Toward an Understanding of Writing 404

Developmental Dyslexias 405

EPILOGUE ¿¿Speech Sounds and the Left Hemisphere 408

Key Concepts 408

Suggested Readings 409

Additional Resources 409

Chapter 14 Neurological Disorders 410

PROLOGUE ¿¿It Started with Her Foot 411

Tumors 411

Seizure Disorders 413

Cerebrovascular Accidents 416

Disorders of Development 420

Toxic Chemicals 420

Inherited Metabolic Disorders 421

Down Syndrome 422

Degenerative Disorders 423

Transmissible Spongiform Encephalopathies 423

Parkinson’s Disease 425

Huntington’s Disease 429

Alzheimer’s Disease 430

Amyotrophic Lateral Sclerosis 434

Multiple Sclerosis 435

Disorders Caused by Infectious Diseases 437

EPILOGUE ¿¿Seizure Surgery 439

Key Concepts 440

Suggested Readings 440

Additional Resources 441

Chapter 15 Schizophrenia, Affective Disorders, and Anxiety Disorders 442

PROLOGUE ¿¿Anxiety Surgery 443

Schizophrenia 443

Description 443

Heritability 445

Pharmacology of Schizophrenia:

The Dopamine Hypothesis 445

Schizophrenia as a Neurological Disorder 446

Major Affective Disorders 455

Description 455

Heritability 456

Biological Treatments 457

The Monoamine Hypothesis 460

Evidence for Brain Abnormalities 461

Role of Neurogenesis 463

Role of Circadian Rhythms 464

Anxiety Disorders 467

Panic Disorder, Generalized Anxiety Disorder,

and Social Anxiety Disorder 467

Obsessive-Compulsive Disorder 469

EPILOGUE ¿¿Prefrontal Lobotomy 474

Key Concepts 474

Suggested Readings 475

Additional Resources 475

Chapter 16 Autistic, Attention-Deficit/Hyperactivity, Stress, and Substance Abuse Disorders 476

PROLOGUE ¿¿A Sudden Craving 477

Autistic Disorder 477

Description 477

Possible Causes 478

Attention-Deficit/Hyperactivity Disorder 481

Description 481

Possible Causes 482

Stress Disorders 483

Physiology of the Stress Response 483

Health Effects of Long-Term Stress 485

Effects of Stress on the Brain 485

Posttraumatic Stress Disorder 487

Stress and Infectious Diseases 488

Substance Abuse Disorders 490

What Is Addiction? 490

Commonly Abused Drugs 496

Heredity and Drug Abuse 503

Therapy for Drug Abuse 504

EPILOGUE ¿¿Classically Conditioned Craving 508

Key Concepts 509

Suggested Readings 509

Additional Resources 510

References 511

Name Index 537

Subject Index

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