xiv Content Chapter 24 Plasticity of Mature Synapses Long-Term Synaptic Depression 592 nd Circuits 575 Box C Silent sy ses 594 Overview 575 Changes in Gene Expression Cause Enduring ic Plasticity Underlies Behavioral Modification Changes in Synaptic Function during LTP and in Invertebrates 575 LTD 597 Box A Genetics of Learning and Memory in the Plasticity in the Adult Cerebral Cortex 599 Fruit Fly 581 Box D Epilepsy: The Effect of Pathological Short-Term Synaptic Plasticity in the Mammalian Activity on Neural Circuitry 600 Nervous System 582 Recovery from Neural Injury 602 Long-Term Synaptic Plasticity in the Mammalian Generation of Neurons in the adult brain 605 Nervous System 583 Box E Why Aren t We More Like Fish and Long-Term Potentiation of Hippocampal Synapses 584 Frogs? 606 Molecular Mechanisms Underlying LTP 587 umm Box B Dendritic Spines 590 Unity COMPLEX BRAIN FUNCTIONS Chapter 25 The Association Cortices 613 Chapter 26 Language and Speech 637 Overview 613 Overview 637 The Association Cortices 613 Language Is Both Localized and Lateralized 637 An Overview of Cortical Structure 614 Aphasias 638 Specific Features of the Association Cortices 615 Box A Speech 640 BOx AA More detailed look at cortical Box B Do Other Animals Have Language? 642 Lamination 617 Box C Words and Meaning 645 Lesions of the Parietal Association Cortex: Deficits of A Dramatic Confirmation of Language Lateralization Attention 619 Lesions of the Temporal Association Cortex: Anatomical Differences between the Right and Left Deficits of Recognition 622 Lesions of the Frontal Association Cortex: Deficits of Mapping Language Functions 649 Box D Language and Handedness 650 Box B Psychosurgery 625 The Role of the Right Hemisphere in Language 654 "Attention Neurons" in the monkey parietal Cortex 626 Sign Language 655 Recognition Neurons"in the Monkey Temporal Summary 656 Cortex 627 Planning Neurons"in the Monkey Frontal Cortex bsu Chapter 27 Sleep and Wakefulness 659 Box C Neuropsychological Testing 632 Box D Brain Size and Intelligence 634 umma Why Do Humans(and Many Other Animals) Sleep 659 Box A Styles of Sleep in Different Species 661
xiv Contents Chapter 25 The Association Cortices 613 Overview 613 The Association Cortices 613 An Overview of Cortical Structure 614 Specific Features of the Association Cortices 615 BOX A A More Detailed Look at Cortical Lamination 617 Lesions of the Parietal Association Cortex: Deficits of Attention 619 Lesions of the Temporal Association Cortex: Deficits of Recognition 622 Lesions of the Frontal Association Cortex: Deficits of Planning 623 BOX B Psychosurgery 625 “Attention Neurons” in the Monkey Parietal Cortex 626 “Recognition Neurons” in the Monkey Temporal Cortex 627 “Planning Neurons” in the Monkey Frontal Cortex 630 BOX C Neuropsychological Testing 632 BOX D Brain Size and Intelligence 634 Summary 635 Chapter 26 Language and Speech 637 Overview 637 Language Is Both Localized and Lateralized 637 Aphasias 638 BOX A Speech 640 BOX B Do Other Animals Have Language? 642 BOX C Words and Meaning 645 A Dramatic Confirmation of Language Lateralization 646 Anatomical Differences between the Right and Left Hemispheres 648 Mapping Language Functions 649 BOX D Language and Handedness 650 The Role of the Right Hemisphere in Language 654 Sign Language 655 Summary 656 Chapter 27 Sleep and Wakefulness 659 Overview 659 Why Do Humans (and Many Other Animals) Sleep? 659 BOX A Styles of Sleep in Different Species 661 Unit V COMPLEX BRAIN FUNCTIONS Chapter 24 Plasticity of Mature Synapses and Circuits 575 Overview 575 Synaptic Plasticity Underlies Behavioral Modification in Invertebrates 575 BOX A Genetics of Learning and Memory in the Fruit Fly 581 Short-Term Synaptic Plasticity in the Mammalian Nervous System 582 Long-Term Synaptic Plasticity in the Mammalian Nervous System 583 Long-Term Potentiation of Hippocampal Synapses 584 Molecular Mechanisms Underlying LTP 587 BOX B Dendritic Spines 590 Long-Term Synaptic Depression 592 BOX C Silent Synapses 594 Changes in Gene Expression Cause Enduring Changes in Synaptic Function during LTP and LTD 597 Plasticity in the Adult Cerebral Cortex 599 BOX D Epilepsy: The Effect of Pathological Activity on Neural Circuitry 600 Recovery from Neural Injury 602 Generation of Neurons in the Adult Brain 605 BOX E Why Aren’t We More Like Fish and Frogs? 606 Summary 609 Purves3/eFM 5/13/04 12:59 PM Page xiv
The Circadian Cycle of Sleep and Wakefulness 662 Box C The Actions of sex hormones 718 Stages of Sleep 665 Other Central Nervous System Dimorphisms Box B Molecular Mechanisms of Biological Specifically Related to Reproductive Behaviors 720 Clocks 666 Brain Dimorphisms Related to Cognitive Function 728 Box C Electroencephalograph 668 Hormone-Sensitive Brain Circuits in Adult Animals 729 Physiological Changes in Sleep States 671 Summary 731 The Possible Functions of REM Sleep and Dreaming Chapter 30 Memory 733 Neural Circuits Governing Sleep 674 Overview 733 Box D Consciousness 675 Qualitative Categories of Human Memory 733 Thalamocortical Interactions 679 Temporal Categories of Memory 734 Sleep disorders 681 Box A Phylogenetic Memory 735 Box E Drugs and Sleep 682 The Importance of Association in Information S orage Forgetting 738 Chapter 28 Emotions 687 Box B Savant Syndrome 739 Brain Systems Underlying Declarative Memory Physiological Changes Associated with Emotion 687 Formation 741 The Integration of Emotional Behavior 688 Box C Clinical Cases that Reveal the anatomical Box A Facial Expressions: Pyramidal and Substrate for declarative memories 742 Extrapyramidal Contributions 690 Brain Systems Underlying Long- Term Storage of The Limbic System 693 Declarative Memory 746 Box B The Anatomy of the Amygdala 696 Brain Systems Underlying Nondeclarative Learnin The Importance of the Amygdala 697 Box C The Reasoning Behind an Important Memory and Aging 749 Discovery 698 Box D Alzheimer' s Disease 750 The Relationship between Neocortex and amygdala Summary 753 701 Box D Fear and the human Amygdala Appendix A The Brainstem and Cranial A Case Study 702 Nerves 755 BOx E Affective Disorders 704 Cortical Lateralization of Emotional Functions 705 Appendix B Vascular Supply, the Meninges, Emotion Reason, and Social Behavior 707 and the ventricular System 763 Summary 708 The Blood Supply of the Brain and Spinal Cord 763 The Blood-Brain barrier 766 Chapter 29 Sex, Sexuality, and the Brain 711 Box A Stroke 767 Overview 711 Sexually Dimorphic Behavior 711 The Meninges 76 What Is Sex? 712 The Ventricular System 770 Box A The Development of Male and Female Phenotypes 714 Glossary Hormonal Influences on Sexual Dimorphism 715 Illustration Source References Box B The Case of Bruce/Brenda 716 Index The Effect of Sex Hormones on Neural Circuitry 718
The Circadian Cycle of Sleep and Wakefulness 662 Stages of Sleep 665 BOX B Molecular Mechanisms of Biological Clocks 666 BOX C Electroencephalography 668 Physiological Changes in Sleep States 671 The Possible Functions of REM Sleep and Dreaming 671 Neural Circuits Governing Sleep 674 BOX D Consciousness 675 Thalamocortical Interactions 679 Sleep Disorders 681 BOX E Drugs and Sleep 682 Summary 684 Chapter 28 Emotions 687 Overview 687 Physiological Changes Associated with Emotion 687 The Integration of Emotional Behavior 688 BOX A Facial Expressions: Pyramidal and Extrapyramidal Contributions 690 The Limbic System 693 BOX B The Anatomy of the Amygdala 696 The Importance of the Amygdala 697 BOX C The Reasoning Behind an Important Discovery 698 The Relationship between Neocortex and Amygdala 701 BOX D Fear and the Human Amygdala: A Case Study 702 BOX E Affective Disorders 704 Cortical Lateralization of Emotional Functions 705 Emotion, Reason, and Social Behavior 707 Summary 708 Chapter 29 Sex, Sexuality, and the Brain 711 Overview 711 Sexually Dimorphic Behavior 711 What Is Sex? 712 BOX A The Development of Male and Female Phenotypes 714 Hormonal Influences on Sexual Dimorphism 715 BOX B The Case of Bruce/Brenda 716 The Effect of Sex Hormones on Neural Circuitry 718 BOX C The Actions of Sex Hormones 718 Other Central Nervous System Dimorphisms Specifically Related to Reproductive Behaviors 720 Brain Dimorphisms Related to Cognitive Function 728 Hormone-Sensitive Brain Circuits in Adult Animals 729 Summary 731 Chapter 30 Memory 733 Overview 733 Qualitative Categories of Human Memory 733 Temporal Categories of Memory 734 BOX A Phylogenetic Memory 735 The Importance of Association in Information Storage 736 Forgetting 738 BOX B Savant Syndrome 739 Brain Systems Underlying Declarative Memory Formation 741 BOX C Clinical Cases That Reveal the Anatomical Substrate for Declarative Memories 742 Brain Systems Underlying Long-Term Storage of Declarative Memory 746 Brain Systems Underlying Nondeclarative Learning and Memory 748 Memory and Aging 749 BOX D Alzheimer’s Disease 750 Summary 753 Appendix A The Brainstem and Cranial Nerves 755 Appendix B Vascular Supply, the Meninges, and the Ventricular System 763 The Blood Supply of the Brain and Spinal Cord 763 The Blood-Brain Barrier 766 BOX A Stroke 767 The Meninges 768 The Ventricular System 770 Glossary Illustration Source References Index Contents xv Purves3/eFM 5/13/04 12:59 PM Page xv
Preface Whether judged in molecular, cellular, systemic, behavioral, or cogni tive terms, the human nervous system is a stupendous piece of bio- logical machinery. Given its accomplishments--all the artifacts of human culture, for instance-there is good reason for wanting to understand how the brain and the rest of the nervous system works The debilitating and costly effects of neurological and psychiatric dis ease add a further sense of urgency to this quest. The aim of this book is to highlight the intellectual challenges and excitement--as well as the uncertainties--of what many see as the last great frontier of bio- logical science. The information presented should serve as a starting point for undergraduates, medical students, graduate students in the neurosciences, and others who want to understand how the human nervous system operates. Like any other great challenge, neuro- ence should be, and is full of debate dissension and considerable fun. All these ingredients have gone into the construction of the third edition of this book; we hope they will be conveyed in equal measure to readers at all levels
Whether judged in molecular, cellular, systemic, behavioral, or cognitive terms, the human nervous system is a stupendous piece of biological machinery. Given its accomplishments—all the artifacts of human culture, for instance—there is good reason for wanting to understand how the brain and the rest of the nervous system works. The debilitating and costly effects of neurological and psychiatric disease add a further sense of urgency to this quest. The aim of this book is to highlight the intellectual challenges and excitement—as well as the uncertainties—of what many see as the last great frontier of biological science. The information presented should serve as a starting point for undergraduates, medical students, graduate students in the neurosciences, and others who want to understand how the human nervous system operates. Like any other great challenge, neuroscience should be, and is, full of debate, dissension, and considerable fun. All these ingredients have gone into the construction of the third edition of this book; we hope they will be conveyed in equal measure to readers at all levels. Preface Purves3/eFM 5/13/04 12:59 PM Page xvi
Acknowledgments We are grateful to numerous colleagues who provided helpful contri- butions, criticisms and suggestions to this and previous editions. We particularly wish to thank Ralph Adolphs, David Amaral, Eva Anton, Gary Banker, Bob Barlow, Marlene Behrmann, Ursula Bellugi, Dan Blazer, Bob Burke, Roberto Cabeza, Nell Cant, Jim Cavanaugh, John Chapin, Milt Charlton, Michael Davis, Rob Deaner, Bob Desimone Allison Doupe, Sasha du Lac, Jen Eilers, Anne Fausto-Sterling, Howard Fields, Elizabeth Finch, Nancy Forger, Jannon Fuchs, Michela Gallagher, Dana Garcia, Steve George, the late Patricia Gold man-Rakic, Mike Haglund, Zach Hall, Kristen Harris, Bill Henson, John Heuser, Jonathan Horton, Ron Hoy, Alan Humphrey, Jon Kaas Jagmeet Kanwal, Herb Killackey, Len Kitzes, Arthur Lander, Story Landis, Simon LeVay, Darrell Lewis, Jeff Lichtman, Alan Light, Steve Lisberger, Donald Lo, Arthur Loewy, Ron Mangun, Eve Marder, Robert McCarley, Greg McCarthy, Jim Mcllwain, Chris Muly, Vic Nadler, Ron Oppenheim, Larysa Pevny, Michael Platt, Franck Polleux, Scott Pomeroy, Rodney Radtke, Louis Reichardt, Marnie Rid dle, Jamie Roitman, Steve Roper, John Rubenstein, Ben Rubin, David Rubin, Josh Sanes, Cliff Saper, Lynn Selemon, Carla Shatz, Bill Snider, Larry Squire, John Staddon, Peter Strick, Warren Strittmatter, Joe Takahashi, Richard Weinberg, Jonathan Weiner, Christina Williams, Joel Winston, and Fulton Wong. It is understood, of course, that any errors are in no way attributable to our critics and advisors We also thank the students at Duke University Medical School as rell as many other students and colleagues who provided sugges tions for improvement of the last edition. Finally, we owe special thanks to Robert Reynolds and Nate O'Keefe, who labored long and hard to put the third edition together, and to Andy Sinauer, Graig Donini, Carol Wigg, Christopher Small, Janice Holabird, and the rest of the staff at Sinauer Associates for their outstanding work and high standards
We are grateful to numerous colleagues who provided helpful contributions, criticisms and suggestions to this and previous editions. We particularly wish to thank Ralph Adolphs, David Amaral, Eva Anton, Gary Banker, Bob Barlow, Marlene Behrmann, Ursula Bellugi, Dan Blazer, Bob Burke, Roberto Cabeza, Nell Cant, Jim Cavanaugh, John Chapin, Milt Charlton, Michael Davis, Rob Deaner, Bob Desimone, Allison Doupe, Sasha du Lac, Jen Eilers, Anne Fausto-Sterling, Howard Fields, Elizabeth Finch, Nancy Forger, Jannon Fuchs, Michela Gallagher, Dana Garcia, Steve George, the late Patricia Goldman-Rakic, Mike Haglund, Zach Hall, Kristen Harris, Bill Henson, John Heuser, Jonathan Horton, Ron Hoy, Alan Humphrey, Jon Kaas, Jagmeet Kanwal, Herb Killackey, Len Kitzes, Arthur Lander, Story Landis, Simon LeVay, Darrell Lewis, Jeff Lichtman, Alan Light, Steve Lisberger, Donald Lo, Arthur Loewy, Ron Mangun, Eve Marder, Robert McCarley, Greg McCarthy, Jim McIlwain, Chris Muly, Vic Nadler, Ron Oppenheim, Larysa Pevny, Michael Platt, Franck Polleux, Scott Pomeroy, Rodney Radtke, Louis Reichardt, Marnie Riddle, Jamie Roitman, Steve Roper, John Rubenstein, Ben Rubin, David Rubin, Josh Sanes, Cliff Saper, Lynn Selemon, Carla Shatz, Bill Snider, Larry Squire, John Staddon, Peter Strick, Warren Strittmatter, Joe Takahashi, Richard Weinberg, Jonathan Weiner, Christina Williams, Joel Winston, and Fulton Wong. It is understood, of course, that any errors are in no way attributable to our critics and advisors. We also thank the students at Duke University Medical School as well as many other students and colleagues who provided suggestions for improvement of the last edition. Finally, we owe special thanks to Robert Reynolds and Nate O’Keefe, who labored long and hard to put the third edition together, and to Andy Sinauer, Graig Donini, Carol Wigg, Christopher Small, Janice Holabird, and the rest of the staff at Sinauer Associates for their outstanding work and high standards. Acknowledgments Purves3/eFM 5/13/04 12:59 PM Page xvii
Supplements to Accompany NEUrOSCIENCE Third Edition For the student Sylvius for neuroscience sylvius A visual Glossary of Human Neuroanatomy(CD-ROM) S Mark Williams, Leonard E. White, and Andrew C. Mace Sylvius for Neuroscience: A Visual Glossary of Human Neuroanatomy, included in every copy of the textbook, is an interactive CD reference guide to the structure of the human nervous system. By entering a corresponding page number from the textbook, students can quickly search the CD for any neuroanatomical structure or term and view corresponding images and animations. Descriptive information is provided with all images and animations. Additionally, students can take notes on the content and share these with other Sylvius users Sylvius is an essential study aid for learning basic human neuro- anatomy. Sylvius for Neuroscience features Over 400 neuroanatomical structures and terms High-resolution images Animations of pathways and 3-D reconstructions Definitions and descriptions Audio pronunciations. A searchable glo Categories of anatomical structures and terms (e. g cranial nerves,spinal cord tracts, lobes, cortical areas, etc.), th easily browsed. In addition, structures can be browsed by text
For the Student Sylvius for Neuroscience: A Visual Glossary of Human Neuroanatomy (CD-ROM) S. Mark Williams, Leonard E. White, and Andrew C. Mace Sylvius for Neuroscience: A Visual Glossary of Human Neuroanatomy, included in every copy of the textbook, is an interactive CD reference guide to the structure of the human nervous system. By entering a corresponding page number from the textbook, students can quickly search the CD for any neuroanatomical structure or term and view corresponding images and animations. Descriptive information is provided with all images and animations. Additionally, students can take notes on the content and share these with other Sylvius users. Sylvius is an essential study aid for learning basic human neuroanatomy. Sylvius for Neuroscience features: • Over 400 neuroanatomical structures and terms. • High-resolution images. • Animations of pathways and 3-D reconstructions. • Definitions and descriptions. • Audio pronunciations. • A searchable glossary. • Categories of anatomical structures and terms (e.g., cranial nerves, spinal cord tracts, lobes, cortical areas, etc.), that can be easily browsed. In addition, structures can be browsed by textbook chapter. Supplements to Accompany NEUROSCIENCE Third Edition Purves3/eFM 5/13/04 1:00 PM Page xviii