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Brief Contents

Preface xxi CHAPTER 7: Traumatic and Chronic Brain


Damage 191
About the Authors xxxiii
CHAPTER 8: Disorders of Anxiety:
Obsessive-Compulsive Disorder
P AR T I and Tourette Syndrome 218
Foundations of Clinical Neuroscience
CHAPTER 9: Mood Disorders 248
CHAPTER 1: Emergence of Clinical
Neuroscience 1 CHAPTER 10: Schizophrenia 290

CHAPTER 2: Research, Treatment, and CHAPTER 11: Drug Addiction 325


Points of View: Historical
Perspectives 31 PART IV
Maintaining Homeostasis
P AR T I I CHAPTER 12: Stress and Coping 357
Fundamentals: Establishing
CHAPTER 13: Psychoneuroimmunology 391
Homeostasis
CHAPTER 14: Eating Regulation and
CHAPTER 3: Macroanatomy and the
Associated Disorders 420
Dynamic Brain 63
CHAPTER 4: Microanatomy and
Neurotransmission 95 EPIL OGUE 454
CHAPTER 5: Neurochemistry and
Psychopharmacology 127 Glossary G-1

CHAPTER 6: Neurodevelopment Over the References R-1


Life Span 158 Name Index N-1
Subject Index S-1
P AR T I I I
Disruptions of Homeostasis:
Representative Clinical Disorders

vii
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Contents

Preface xxi

About the Authors xxxiii

PART I Foundations
of Clinical Neuroscience

CHAPTER 1: Emergence of Clinical Neuroscience 1


CONNECTIONS: Mara: Seeking Therapy 1
Jasmine: Seeking a Profession 2
Alex: Seeking Support 2

Status Reports on Psychiatry and Clinical Psychology 3


Research–Treatment Disconnect 5
Classifying Mental Disorders 8
Lessons Learned from the Multiple Personality Disorder Diagnosis 9

Exploring Various Windows of Mental Health 12


A CASE IN POINT: Diagnoses of Nadean Cool 13
Windows on Neurobiology 14
Neuroanatomy 15
Neurochemistry 16
Neurophysiology 17
Genetics Perspective 18
Developmental Perspective 20
Environmental Perspective 21
Evolutionary Perspective 22
Window Provided by Culture 24

What Lies Ahead 25


Multiple Perspectives 25

ix
x Contents

OLIVER SACKS: Educating the Public About Clinical Neuroscience 26


A Brain–Body Balance 28

Summary 28

CHAPTER 2: Research, Treatment, and Points of View:


Historical Perspectives 31
CONNECTIONS: Barbers and Neurosurgeons 31

Emergence of the Brain as the Controlling Center of


Behavior, Emotions, and Thought 33
Head, Heart, and Mind–Body Problem 33
Hippocrates: Heart of mental functions 33
Galen: Spirit of mental functions 34
René Descartes: Body meets mind 36
Localizing Functions in the Brain 37
BRAIN MATTERS: Mental Illness Loses Its Religion 38
Thomas Willis: Pioneer of brain functions 38
Franz Joseph Gall: Rise and fall of phrenology 39
Paul Broca: Localization of language 41
Pioneers of Cellular Investigations in the Brain: The Architects 42
Camillo Golgi: Revealing neuronal structure with the perfect stain 42
Santiago Ramón y Cajal: Providing evidence for the neuron doctrine 43
Charles Scott Sherrington: Describing the synapse 44
Pioneers of Cellular Investigations in the Brain: The Chemical Engineers 44
Otto Loewi: Dreaming of chemical communication 44
Edgar D. Adrian: Amplifying the electrical language of the neuron 45

Neurological Approaches to Mental Illness 46


Mental Illness Meets the Microscope 46
ERIC KANDEL: From Studying the History of Neuroscience
to Becoming the History of Neuroscience 47
Jean-Martin Charcot: Birth of Neurology 48

Biological Therapies for Mental Illness 50


Early Biological Treatments 50
Electroconvulsive Therapy 51
The Drug Revolution 52
Psychosurgery 52

Importance of a Therapeutic Environment 55


Philippe Pinel and the Fall of the Asylum 55
Behaviorism: A Different Type of Therapeutic Environment 55
A CASE IN POINT: The Couple Who Needed Each Other Like a
Hole in the Head 56
Beyond Behaviorism: Rethinking the Role of Mental Processes in
Mental Health 58
Contents xi

Profiling the Pioneers of Clinical Neuroscience 60

Summary 60

Fundamentals: Establishing Homeostasis PART II

CHAPTER 3: Macroanatomy and the Dynamic Brain 63


CONNECTIONS: Brain Plasticity in Evolutionary and in Real Time 63

Gross Structure and Function of the Nervous System 64


Organization of the Peripheral Nervous System 65
ANS and Homeostasis/Allostasis 66
Development of the CNS 67
Layers of protection 68
Ventricular system 69
Geography of the developing brain 69

CNS Structure and Function 70


Brain Stem 71
Medulla and basic survival 73
Cerebellum and motor control 73
Pons and information transfer 74
Midbrain 74
Forebrain 75
Diencephalon: Regulating social and survival behaviors 76
Telencephalon and higher-order behaviors 78
Structures and functions of the cortex 79
Subcortical circuits: Limbic system and basal ganglia 81
Rhinencephalon: Smell and the brain 83
PAUL MACLEAN: Neural Investigator Paul MacLean's Pursuit of the Brain's
Circuits 84
BRAIN MATTERS: Joking Around in the Laboratory 86
Spinal Cord 86

Cranial Nerves and Blood Supply 89


A CASE IN POINT: Katherine Sherwood 91

Summary 91

CHAPTER 4: Microanatomy and Neurotransmission 95


CONNECTIONS: Human and Mouse Genius 95

Microanatomy of the Neuron 97


External Structure and Function of the Neuron 97
Cell membrane and its gateways 98
xii Contents

Dendrites: Extending the neuron’s reach 100

Cell Body and Organelles: A Cellular Factory 101


Nucleus and executive function 102
Endoplasmic reticulum, ribosomes, and protein synthesis 103
Mitochondria and metabolism 103
Golgi apparatus and lysosomes: Packaging and recycling centers 104
Neuronal Cytoskeleton: Structure and Transport 104
Dynamic Elements of Neuronal Communication, Connection,
and the Synapse 106
Axon: Information conductor 106
Terminal buttons: Getting the message across 106

Neuroglia 107

Generating an Action Potential 110


Electrochemical Activity at the Cell Membrane 110
Membrane Activity as Trigger of the Action Potential 113
Propagating the action potential 114
Refractory periods: The neuron recharges 115
A CASE IN POINT: Lorenzo Odone’s Battle with Adrenoleukodystrophy 116
Integrating Information at the Cell Membrane 117
Activity at the Synapse 118
Postsynaptic Potentials 119
Activity at the Receptor 120
BRAIN MATTERS: Epilepsy: Continued Intrigue and Lessons Learned 122

Summary 124

CHAPTER 5: Neurochemistry and Psychopharmacology 127


CONNECTIONS: Coffee: Changing Our Culture and Our Nervous Systems 127

Neurochemicals 129
Small-Molecule Neurotransmitters 130
Acetylcholine 132
Catecholamines 134
Serotonin 137
BRAIN MATTERS: MDMA: Ecstasy or Agony for the Brain? 140
Amino Acid Transmitters 141
Excitatory amino acid 141
Inhibitory amino acids 142
Neuropeptides and Neurohormones 144
Endogenous opioids 145
Oxytocin 146
Vasopressin 147
Contents xiii

A CASE IN POINT: Controversial Dietary Intervention for Autism: Karyn


Seroussi’s Struggle to Help Her Son 148

Neurochemistry of the Stress Response 149


Autonomic Stress Response 150
Hypothalamic–Pituitary–Adrenal Axis Response 150
Other Brain Areas Involved in the Stress Response 151
The hippocampus 151
Amygdala 152
Locus coeruleus 153

Foundations of Psychopharmacology 153

Summary 155

CHAPTER 6: Neurodevelopment Over the Life Span 158


CONNECTIONS: Fate of the Aging Brain: Popular and Scientific
Perspectives 158

Prenatal Brain Development 159


Emergence and Journey of the Brain’s Neurons 160
Establishing and Fine-Tuning the Brain’s Circuits 164

Postnatal Brain Development 167


Infancy and Childhood 167
Adolescence and Attention-Deficit/Hyperactivity Disorder 169
A CASE IN POINT: Relieving Ashlee’s Seizure-Induced Wiggles 172

The Aging Brain 174


Normal Aging Processes 174
Neurodegenerative Decline: Alzheimer’s Disease 175

Development of Key Neurohormonal Circuits 176


Sexual Differentiation and Brain Development 177
BRAIN MATTERS: The Maternal Brain 178
Development of the Stress Response 180
Mom’s impact on the developing stress response 181
Stress and aging 182
Sex differences in the stress response 183

Scientific Interventions with Developmental Neurobiology 183


Stem Cells 184
Mammalian Cloning 186

Summary 188
xiv Contents

PART III Disruptions of Homeostasis:


Representative Clinical Disorders

CHAPTER 7: Traumatic and Chronic Brain Damage 191


CONNECTIONS: Educated Minds, Neural Reserves, and Recovery from
Brain Injury 191

Overview of Chapter 193

Traumatic Brain Injury 194


Epidemiological Factors 194
Hypothesized Events Associated With the Aftermath of TBI 195
Primary effects 196
After the impact: Secondary factors related to sustained brain injuries 197

BRAIN MATTERS: The Perils of Using Your Head in the National


Football League 199
Treatment and Recovery 199
A CASE IN POINT: Surviving a Life-Threatening Automobile Accident 200
Injuries necessitating surgical interventions 200
Pharmacological interventions 201
Discovering the impact of progesterone on TBI: A story of hard knocks 203
Cognitive rehabilitation 205
A Unique Behavioral Approach 206
BRAIN MATTERS: Gourmand Syndrome: TBI and Fine Dining 207
Environmental Enrichment and Recovery from TBI 207
Prevention of TBI: Impact of Safety Regulations and Legal Policies 210

Parkinson’s Disease 211


Identification of Relevant Brain Damage 211
Treatment Strategies 211
BRAIN MATTERS: Striatal Neurons Hit the Dance Floor 215

Summary 215

CHAPTER 8: Disorders of Anxiety: Obsessive–Compulsive


Disorder and Tourette Syndrome 218
CONNECTIONS: Common Denominators in Real-Life and Laboratory-Induced
Compulsions? 218

Anxiety Disorders: An Overview 219

Obsessive–Compulsive Disorder 220


BRAIN MATTERS: Panic Disorder: A Faulty Brain Stem Surveillance System? 222
Tracing the Neurobiological Roots of OCD 223
Contents xv

OCD neuroanatomical circuit 223

A CASE IN POINT: Joseph Dee’s Life with OCD 224


Additional evidence from neuroimaging studies 228
Neurochemistry of OCD 230
Evolutionary Perspectives: Fixed Action Patterns and
Ritualistic Behavior 232
Animal models 233
Human models: Baseball rituals 233
Therapies for OCD 234
Psychopharmacological therapy 234
Neurosurgical treatment 236
Deep brain stimulation 238
Cognitive-behavioral therapy 238

Tourette Syndrome 240


Tracing the Neurobiological Roots of Tourette Syndrome 240

A CASE IN POINT: Living with Tourette Syndrome 241


Role of the basal ganglia and associated neurochemistry 242
Possible environmental factors 243
Treatment for Tourette Syndrome 243
Psychopharmacological therapy 244
Behavioral and cognitive strategies 245

Summary 246

CHAPTER 9: Mood Disorders 248


CONNECTIONS: Foods and Moods 248

Mood Disorders: An Overview 250

Major Depression in Context 252


Sex Differences and Sociocultural Influences 252
Genetic Influences and Evolutionary Factors 253

Tracing the Neurobiological Roots of Depression 255


Depression’s Neuroanatomical Circuit 255
Prefrontal cortex 255
Hippocampus and amygdala 256
A CASE IN POINT: Andrew Solomon’s Noonday Demon 257
Cingulate cortex 258
Nucleus accumbens 259
Brain stem tracts 260
White matter 261
Neurochemistry of Depression 261
Serotonin (5-HT) 261
xvi Contents

Norepinephrine 265
Corticosteroids 266
Brain Plasticity 267
Brain structure and neurogenesis 267
BRAIN MATTERS: Animal Models of Depression 268
Role of BDNF in depression 269

Stress and Depression 270


Life’s Losses 271
Sexual Abuse in Childhood 271
Seasonal Changes in Light 271

Treatment for Depression 272


Pharmacotherapy 272
BRAIN MATTERS: St. John’s Wort 273
Serotonergic manipulation 274
Noradrenergic manipulation 276
Inhibiting CRH 276
Electroconvulsive Therapy 277
Repetitive Transcranial Magnetic Stimulation 278
Deep Brain Stimulation (DBS) 279
Cognitive-Behavioral Therapy 280

Bipolar Disorder 281


Tracing the Neurobiological Roots of Bipolar Disorder 282
Treatment for Bipolar Disorder 283
A CASE IN POINT: Kay Redfield Jamison 284

Summary 287

CHAPTER 10: Schizophrenia 290


CONNECTIONS: Home Movies, Childhood Behavior, and Schizophrenia 290

Overview of Schizophrenia 291


History, Descriptions, and Definitions 293
Contemporary Views of Schizophrenia 294
Views of a Schizophrenic Mind 294

Neurobiology of Schizophrenia 296


Genetic Factors 296
A CASE IN POINT: The Transformations of John Nash’s Schizophrenic Mind 298
Neuroanatomy 301
Tuning in to the neural substrates of auditory hallucinations 301
Cellular disorganization and alterations 303
Structural alterations in schizophrenic brains 305
Contents xvii

Neurochemistry 307
Dopamine hypothesis of schizophrenia 307
Nondopaminergic neurochemical involvement in schizophrenia 310

Developmental Markers of Schizophrenia 311


Evidence from Minor Physical Anomalies 312
Sensory-Gating Deficits as a Common Denominator 312
Stress and Developmental Predisposition 313
BRAIN MATTERS: Animal Models of Schizophrenia 314
Viral Theory Controversy 315

Treatments for Schizophrenia 316


Somatic Therapies 316
Pharmacological Therapy 317
Neurocognitive Training 321
Psychosocial Therapy 322

Summary 322

CHAPTER 11: Drug Addiction 325


CONNECTIONS: Vulnerability to Addiction: Threatened Territories in
North America and in the Laboratory 325

Drug Addiction in Context 326


Prevalence of Addiction 328
Addiction Across Cultures 329
Hereditary Influences on Addiction 329
Considering Addiction in the Context of Evolution 329

Tracing the Neurobiological Roots of Addiction 331


Neural Circuits Involved in Establishing Addiction 331
A CASE IN POINT: Running from Addiction 332
Role of the nucleus accumbens 333
Role of the OFC 336
Neurochemistry of Addiction 337
Dopamine: Primary fuel of addiction 337
Potential role of acetylcholine 340
Craving Response 340

BRAIN MATTERS: Substance Abuse and Behavioral Compulsion:


Two Expressions of the Same Neurobiological System? 341
Craving as a conditioned response 342
Neural components of the craving response 342

Stress and Addiction 344

Therapy for Drug Addiction 346


Initial Step: Detoxification 347
xviii Contents

Maintaining Abstinence 347


Vaccination: Therapeutic Shot in the Dark? 349
Behavioral Therapies for Addiction 351
Ethical Questions About New Addiction Therapies 353

Summary 354

PART IV Maintaining Homeostasis

CHAPTER 12: Stress and Coping 357


CONNECTIONS: Pacing and Prozac 357

Revisiting the Stress Response 358


Acute Stress Response 359
Chronic Stress Response 361
Stress and personality traits 362
BRAIN MATTERS: Eating and Stress 363
Stress ulcers 364
The Brain’s Stress Circuit 365

Stress Responsivity Over the Life Span 367


Role of Mother–Infant Interactions 367
Aging-Related Effects of Stress 369
Effects of Sex Differences on Responses to Stress 370

Coping with Stress 371


Perception of Control 371
Research on control 372
Coping with lack of control 374
Social Support 374
Active Versus Passive Coping 379
Allostatic Load 382

Posttraumatic Stress Disorder 383


A CASE IN POINT: How Terry Anderson Survived as a Hostage 384
Tracing the Neurobiological Roots of PTSD 386
Treatment for PTSD 387

Summary 388

CHAPTER 13: Psychoneuroimmunology 391


CONNECTIONS: Species, Scents, and Sex 391

Fundamentals of the Immune System: The Body’s Internal


Security Agents 393
Contents xix

The Body’s Most Wanted Invaders 393


Innate Immunity: The Immune System’s Front-Line Defense 394
Acquired Immunity: Cells That Plan for the Future 396
Antibodies and B cells 397
T cells 397
Contrasting acquired immune defense strategies 400
Major Histocompatibility Complex and Information Dissemination 401
When Good Cells Turn Bad 402

Integrative Operations: Immune and Nervous System


Communication 404
Conditioning the Immune System: A Serendipitous Discovery 404
Additional Pieces of the Neuroimmune Puzzle 405
Stress: Mediator Between the Immune and Neural Agencies 406
Sickness Behavior and Its Role in Maintaining Health 407
Fever 407
Loss of appetite 408
Role of cytokines 408

Putting the “Psych” in Psychoneuroimmunology 409


Psychological Factors Leading to Compromised Immune Function 410
A CASE IN POINT: Evan Handler 411
Psychological Factors Leading to Enhanced Immune Function 413
BRAIN MATTERS: Humor’s Influence on Health: No Laughing Matter! 415

Summary 417

CHAPTER 14: Eating Regulation and Associated


Disorders 420
CONNECTIONS: Increased Rates in Eating Out Accompany Increased Inches
in Our Waistlines 420

Eating Regulation 422


Early Hunger Research Was Difficult to Swallow 422
Neurobiology of Hunger Regulation 424
Neurochemical Factors 425
Glucose 425
Cholecystokinin 426
Leptin 426
Neuropeptide Y 427
Environmental Factors 429

Evolutionary Theories of Hunger Regulation 430


Set-Point Theory 430
Preparatory Response Theory 431
Positive Incentive Theory 432
xx Contents

Sensory-Specific Satiety 433

Disorders of Eating Regulation 434


Obesity 434
Possible causes 435
Treatment 437
Children and obesity 438
BRAIN MATTERS: A Closer Look at Gastric Bypass Surgery 440
Anorexia Nervosa 440
Possible causes 441
Treatment 446
Bulimia Nervosa 447
Possible causes 447
A CASE IN POINT: Sara Hunnicutt’s Bout with Anorexia Nervosa 448
Treatment 450

Digesting the Conflicting Theories of Eating Regulation 450

Summary 451

EPILOGUE 454
CONNECTIONS: Superman, Stem Cells, and the Senate 454

Ethics Buffeted on the Sea of Neuroscience 455


Neuroscience and Criminal Behavior 456
Brain areas influencing criminal behavior: The usual suspects 456
Neuroscience and the burden of proof 459
Designer Brains, Designer Disorders 461

Genetics and Clinical Neuroscience: Basic Research, Basic


Applications 464
A CASE IN POINT: Richard Restak’s “Ted” 465

Looking to the Future Through the Lenses of Clinical


Neuroscience 466
Integrating Mental Health Perspectives 467
Cleansing Our Views with Empirical Evidence: Remembering Past
Lessons 470

Summary 472

Glossary G-1
References R-1
Name Index N-1
Subject Index S-1
Preface

THE NEED FOR CLINICAL NEUROSCIENCE


A major challenge for neuroscientists today is the translation of basic
research findings, such as those that begin as presentations at the Society for
Neuroscience meeting and then move through peer review to publication, into
applied clinical approaches to treating brain disorders and mental illness. At
the same time, however, mental-health practitioners—those on the front lines
of mental health delivery—must acquire broad-based knowledge of neurobi-
ology, behavior, and environment if they hope to understand the difference
between adaptive and maladaptive neural functions. Molecular outcomes of
drugs with equal clinical efficacy, for example, sometimes employ very dif-
ferent modes of action, depending on the underlying systems affected.
Another question practitioners struggle with is the value of the mental
health diagnostic categories that are presented in this text. For instance, it
may be more parsimonious to treat symptoms rather than mental illness
labels, especially, as you’ll learn in Chapter 1, when there is considerable
overlap in the symptoms for two completely different diagnostic categories.
Although considerable progress has been made in laboratories across the
world, significant challenges face practitioners and scientists working in
the field of clinical neuroscience. To address these challenges, the National
Institute of Mental Health (NIMH) awards translational research grants to
encourage collaboration between neuroscientists and mental health profes-
sionals. Such collaborations facilitate the construction of a “clinical neurosci-
ence” that converts data into treatments more rapidly for in-need patients.
By 2025, depression is forecast to become the second leading illness in
the United States (McEwen & Lashley, 2002). A meta-analysis published
in Psychological Science in the Public Interest explored the response rates of
depressed patients to various treatment modalities. Consider the treatment
implications of these results. Interpersonal therapy rates are equally as
effective as antidepressant drugs in reducing typical symptoms associated
with depression. Further, 30 percent of respondents who received a placebo
treatment reported improvement; see Figure 1.1 on page 5 (Hollon, Thase, &
Markowitz, 2002).
Research emphasizing methodological challenges in antidepressant
research had little impact on the public’s trust in these drugs as the number
of Americans taking antidepressants doubled from 1996 to 2005 (increasing
from 13.3 million to 27 million consumers). A recent article in Newsweek
magazine echoed the findings of several researchers suggesting that antide-
pressants are no more effective than placebos. Science writer Sharon Begley
xxi
xxii Preface

ended the article with the discouraging realization that the scientific evi-
dence supporting treatments such as antidepressants just isn’t that important
to most people (Begley, 2010).
Thus, in addition to the importance of clinical neuroscience research
to address the unmet needs of individuals suffering from various mental
illnesses, education about the importance of empirically-based mental
health therapies is necessary to educate the public about the most effective
treatment strategies. The clinical neuroscience course is a first-step toward
addressing these important mental health issues—motivating students to
conduct relevant research in the area of clinical neuroscience and reinforcing
the value of empirically-based approaches to future mental health practitio-
ners and consumers. Recent attention directed toward former Playboy model
and MTV celebrity Jenny McCarthy for authoritative advice on the causes
and treatments for autism certainly support the notion that empirical evi-
dence is less important than personal experiences when it comes to selecting
the most effective therapies for mental illnesses. As you’ll read in Chapter 1,
as treatment strategies stray from methodologically sound experimental
studies providing solid empirical evidence, the results can be disastrous.

THE GENESIS OF THIS BOOK


We designed a new course called Clinical Neuroscience for our students, before
we found a book for the course. We managed to find several impressive general
interest books and edited volumes, but we found no suitable single textbook
that integrated neurobiological mechanisms of mental health into the coverage
of disorders that was readable, affordable, and comprehensive. We decided to
write that textbook. We sincerely hope it satisfies the need.
We have conducted extensive reviews of all the subject matter in this book
and include novel interpretations and explanations. Clinical Neuroscience is a
resource for anyone interested in an integrative, empirically based approach
to mental health and for anyone who seeks to draw connections among
courses offered in departments of psychology, biology, and neuroscience—
the intersection of a modern psychology.
Initially, we envisioned the courses that we were designing within a psy-
chology curriculum, specifically for upper-level undergraduates who had a
prerequisite in basic biological psychology. Fittingly, practice informed our
theory: In teaching the course we discovered that students benefited from a
unit on neurobiology and neurochemistry, and we have added that unit as
Part II of the book. With this addition, a biological psychology prerequisite is
not necessary, but it may be complementary and certainly useful. Because of
the text’s inclusion of more basic neurobiological information, this book may
also be used in a nontraditional biology or psychology course.

AND THE AUDIENCE


Clinical Neuroscience will be of value to upper-level undergraduate courses in
clinical and counseling psychology, as well as elective courses focusing on
topics related to the neurobiology of mental illness in psychology and biol-
ogy major courses. Momentum gathers to prepare clinicians for the possibil-
ity of gaining prescription privileges (a practice already approved in New
Mexico and Louisiana and likely to spread to other states); here, Clinical
Preface xxiii

Neuroscience’s value is clear. Further, the chasm separating psychology from


biology—especially as it relates to brain regulation of mental activity—is
growing smaller. The current volume will introduce even the most diehard
dualist to evidence that the brain must be considered in all discussions of
clinical treatments.
This text represents the culmination of decades’-long conversations with
our neuroscientist colleagues at conferences, extensive reading of the neu-
robiology literature, and our own personal research. Since writing the fi rst
edition of this text, we are now more convinced than ever that clinical neuro-
science is emerging not just as a course, but as a burgeoning field. Reviewers
have elucidated the need for both separate courses and advanced seminars
in clinical neuroscience. We believe this book will benefit neuroscience stu-
dents as a reference as they navigate through their coursework and can also
help them to keep an eye on applications—how neuroscience can benefit its
subjects—thereby increasing the likelihood of translating the basic research
into positive real-world outcomes.
As another example of accelerating treatments from lab to clinic, the
medical student will also benefit from the book’s integrative, medical-model
approach to mental health. That avenue is appropriate for post-graduate audi-
ences (such as internship-level faculty) as they prepare to move into mental
health careers. By allowing instructors to draw connections between basic
neuroscience and mental health applications, Clinical Neuroscience is equally
appropriate for neuroscience students. Building bridges between and among
formerly disparate sub-disciplines will hasten beneficial applications.

CHANGES TO THE NEW EDITION


■ A new chapter on traumatic and chronic brain injury that empha-
sizes traditionally viewed neurobiological disorders such as TBI and
Parkinson’s Disease
■ Existing content has been updated with hundreds of new references
■ New color insert gives students a better sense of the visual complexity of
the brain and related issues in the text

FORMAT OF TEXT
Because clinical approaches are an emerging area within the broad field of
neuroscience, we put considerable thought into the identification, organization,
and categorization of the topics covered in the text. Although diagnostic criteria
from the Diagnostic and Statistical Manual (DSM) IV-TR (and its newer variant)
were included as a reference for students for some conditions, we do not always
follow the DSM categorizations (in part, due to their inherent vagueness and
hand-waving imprecision). The structure of the text is as follows:

PART I: Foundations of Clinical Neuroscience


Chapter 1 Emergence of Clinical Neuroscience
Chapter 2 Research, Treatment, and Points of View: Historical Perspectives
The introductory chapters include historical and contemporary overviews
of theories and therapies in the mental health field. They establish a need for
xxiv Preface

the more integrative, empirically based discipline of clinical neuroscience,


emphasizing the scientific method. Various disciplines (e.g., psychology, psy-
chiatry, neurology, chemistry, basic research) and their unique perspectives
are also addressed; it becomes increasingly apparent that their once unique
boundaries are becoming less distinct. Finally, a description of these many
perspectives, or levels of analysis, in Clinical Neuroscience sets the stage for
the multifaceted approach in the text.

PART II: Fundamentals: Establishing Homeostasis


Chapter 3 Macroanatomy and the Dynamic Brain
Chapter 4 Microanatomy and Neurotransmission
Chapter 5 Neurochemistry and Psychopharmacology
Chapter 6 Neurodevelopment Over the Life Span

Many students may already have some experience with the basics of neu-
ral structure and function. These early chapters, then, will allow for a brief
review for those students and for initial exposure to the basics of nervous
function for the uninitiated. The inclusion of pedagogical components that
relate directly to the field of clinical neuroscience will enable students to
see direct connections between the basic information about the underlying
“wet” tissue, and its regulation and dysregulation in the manifestation of the
human’s so-called “mental health and life.” For example, neurodegenerative
and cognitive processing disorders (such as Alzheimer’s and attention deficit
hyperactivity disorder) are included in the neurodevelopment chapter to
show a direct link between basic mechanisms and the overt conditions they
produce.

PART III: Disruptions of Homeostasis:


Representative Clinical Disorders
Chapter 7 Traumatic and Chronic Brain Damage
Chapter 8 Disorders of Anxiety: Obsessive-Compulsive Disorder and
Tourette Syndrome
Chapter 9 Mood Disorders
Chapter 10 Schizophrenia
Chapter 11 Drug Addiction

In this text, mental disorders are viewed as disruptions of homeostasis/


allostasis, departures from endogenous averages and predisposed physi-
ological coping strategies. As the internal milieu is disrupted, the teeter
totters when it should not, the nervous system’s attempt to compensate
sometimes leads to extreme conditions that are classified as mental disor-
ders. We have included traumatic brain injury and Parkinson’s disease in
Chapter 7 (a new chapter in this edition!), obsessive-compulsive disorder
and Tourette syndrome in Chapter 8 (Disorders of Anxiety); major depres-
sion and bipolar disorder in Chapter 9 (Mood Disorders); and, as implied,
schizophrenia and addiction are the sole disorders for Chapters 10 and
11, respectively. Although other disorders fit in these categories, a deci-
sion was made to include in-depth coverage of representative disorders
Another random document with
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the anal fin; and one confined to the extremity of the tail—the caudal
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The caudal fin is rarely symmetrical, so that its upper half would
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subsequently, Figs. 31, 41). In fishes in which it is nearly symmetrical
it is frequently prolonged into an upper and lower lobe, its hind
margin being concave or more or less deeply excised; in others the
hind margin is rounded, and when the middle rays greatly exceed in
length the outer ones the fin assumes a pointed form.
Fig. 4.—Labrax lupus (Bass), an Acanthopterygian with anterior spinous, and
posterior soft dorsal fin.
Many and systematically important differences are observed in
the dorsal fin, which is either spiny-rayed (spinous)
(Acanthopterygian), or soft-rayed (Malacopterygian). In the former, a
smaller or greater number of the rays are simple and without
transverse joints; they may be flexible, or so much osseous matter is
deposited in them that they appear hard and truly spinous (Fig. 3);
these spines form always the anterior portion of the fin, which is
detached from, or continuous with, the remaining jointed rays. The
spines can be erected or depressed at the will of the fish; if in the
depressed position the spines cover one another completely, their
points lying in the same line, the fish is called homacanth; but if the
spines are asymmetrical, alternately broader on one side than on the
other, the fish is called heteracanth. The spinous division, as well as
the one consisting of jointed rays, may again be subdivided. In the
Malacopterygian type all the rays remain jointed; indeed, sometimes
the foremost ray, with its preceding short supports, is likewise
ossified, and a hard spine, but the articulations can nearly always be
distinctly traced. Sometimes the dorsal fin of Malacopterygian fishes
is very long, extending from the head to the end of the tail,
sometimes it is reduced to a few rays only, and in a few cases it is
entirely absent. In addition to the rayed dorsal fin, many
Malacopterygian fishes (as the Salmonoids, many Siluroids,
Scopeloids, etc.) have another of greater or lesser extent, without
any rays; and as always fat is deposited within this fold, it is called a
fatty fin (pinna adiposa).

Fig. 5.—Saurus undosquamis, a Malacopterygian with anterior soft dorsal, and


additional adipose fin.

The anal fin is built on the same plan as the dorsal, and may be
single or plural, long or short, or entirely absent; in
Acanthopterygians its foremost rays are frequently simple and
spinous.
The horizontal or paired fins consist of two pairs: the pectorals
and ventrals.
The pectoral fins (with their osseous supports) are the
homologues of the anterior limbs of the higher Vertebrata. They are
always inserted immediately behind the gill-opening; either
symmetrical with a rounded posterior margin, or asymmetrical, with
the upper rays longest and strongest; in Malacopterygians with a
dorsal spine the upper pectoral ray is frequently developed into a
similar defensive weapon.
The ventral fins are the homologues of the hind-limbs, and
inserted on the abdominal surface, either behind the pectorals
(Pisces s. Pinnæ abdominales), or below them (Pisces s. Pinnæ
thoracicæ), or in advance of them (Pisces s. Pinnæ jugulares). They
are generally narrow, composed of a small number of rays, the outer
of which is frequently osseous. In some small groups of fishes, like
the Gobies, the fins coalesce and form a suctorial disk.

Fig. 6.—Salmo salar (Salmon), with abdominal ventral fins.

Fig. 7.—Mullus barbatus (Red Mullet), with thoracic ventral fins.


Fig. 8.—Burbot (Lota vulgaris), with jugular ventral fins.

For the definition of the smaller systematic groups, and the


determination of species, the numbers of the spines and rays are
generally of the greatest importance. This holds good, especially for
the ventral rays, by the number of which the Acanthopterygian
affinities of a fish can nearly always be determined. The numbers of
the dorsal and anal rays generally correspond to the number of
vertebræ in a certain portion of the spine, and are therefore constant
specific, generic, or even family characters; but when their number is
very great, a proportionally wide margin must be allowed for
variation, and the taxinomic value of this character becomes
uncertain. The numbers of the pectoral and caudal rays are rarely of
any account.
The fins are organs of motion; but it is chiefly the tail
Function of the and the caudal fin by which the fish impels itself
Fins. forward. To execute energetic locomotion the tail and
caudal fin are strongly bent, with rapidity, alternately
towards the right and left; whilst a gentle motion forwards is effected
by a simple undulating action of the caudal fin, the lobes of which act
like the blades of a screw. Retrograde motions can be made by fish
in an imperfect manner only, by forward-strokes of the pectoral fins.
When the fish wants to turn towards the left, he gives a stroke of the
tail towards the right, the right pectoral acting simultaneously, whilst
the left remains ad-pressed to the body. Thus the pectoral fins assist
in the progressive motions of the fish, but rather directing its course
than acting as powerful propellers. The chief function of the paired
fins is to maintain the balance of the fish in the water, which is
always the most unsteady where there is no weight to sink it: when
the pectoral of one side, or the pectoral and ventral of the same side
are removed, the fish loses its balance and falls on the side
opposite; when both pectorals are removed, the fish’s head sinks; on
removal of the dorsal and anal fins the motion of the fish assumes a
zig-zag course. A fish deprived of all fins, as well as a dead fish,
floats with the belly upwards, the back being the heavier part of the
body.
In numerous groups of fishes which live in mud, or are enabled to
pass a longer or shorter time in soil periodically dried and hardened
during the hot season, forms occur entirely devoid of, or with only
rudimentary, ventral fins (Cyprinodon, Ophiocephalidæ, Galaxiidæ,
Siluridæ). The chief function of these fins being to balance the body
of the fish whilst swimming, it is evident that in fishes moving during
a great part of their life over swampy ground, or through more or less
consistent mud, this function of the ventral fins ceases, and that
nature can readily dispense with these organs altogether.

Fig. 9.—Ventrals of Gobius.


In certain fishes the shape and function of the fins are
considerably modified: thus, in the Rays, locomotion is almost
entirely effected and regulated by the broad and expanded pectoral
fins acting with an undulatory motion of their margins, similar to the
undulations of the long vertical fins of the Flat-fishes; in many
Blennies the ventral fins are adapted for walking on the sea-bottom;
in some Gobioids (Periophthalmus), Trigloids, Scorpænioids, and
Pediculati, the pectoral fins are perfect organs of walking; in the
Gobies, Cyclopteri, and Discoboli the ventral fins are transformed
into an adhesive disk, and finally in the Flying-fish, in which the
pectorals act as a parachute. In the Eels and other snake-like fishes,
the swimming as well as the gliding motions are effected by several
curvatures of the body, alternate towards the right and left,
resembling the locomotion of Snakes. In the Syngnathi (Pipe-fishes)
and Hippocampi, whose body admits of but a slight degree of lateral
curvature, and whose caudal fin is generally small, if present at all,
locomotion is very limited, and almost wholly dependent on the
action of the dorsal fin, which consists of a rapid undulating
movement.

Fig. 10.—Cycloid scale of Gadopsis


marmoratus (magn.)
Fig. 11.—Cycloid scale of Scopelus
resplendens (magn.)
The skin of fishes is either covered with scales, or
Skin and naked, or provided with more or less numerous
Scales. scutes of various forms and sizes. Some parts, like
the head and fins, are more frequently naked than
scaly. All fishes provided with electric organs, the majority of Eels,
and the Lampreys, are naked. Scales of fishes are very different
from those of Reptiles; the latter being merely folds of the cutis,
whilst the scales of fishes are distinct horny elements, developed in
grooves or pockets of the skin, like hairs, nails, or feathers. Very
small or rudimentary scales are extremely thin, homogeneous in
structure, and more or less imbedded in the skin, and do not cover
each other. When more developed, they are imbricated (arranged in
the manner of tiles), with the posterior part extruded and free, the
surface of the anterior portion being usually covered by the skin to a
greater or less extent. On their surface (Figs. 10 and 11) may be
observed a very fine striation concentric and parallel to the margin,
and coarser striæ radiating from a central point towards the hind
margin. Scales without a covering of enamel, with an entire (not
denticulated) posterior margin, and with a concentric striation, are
called Cycloid scales. Ctenoid scales (Figs. 12–15) are generally
thicker, and provided with spinous teeth on the posterior edges of the
layers of which the scale consists. In some species only the layer
nearest to the margin is provided with denticulations (Fig. 14).
Scales, the free surface of which is spiny, and which have no
denticulation on the margin, have been termed Sparoid scales; but
their distinction from ctenoid scales is by no means sharp, and there
are even intermediate forms between the cycloid and ctenoid types.
Both kinds of scales may occur not only in species of the same
genus of fishes, but in the same fish.

Fig. 12.—Ctenoid scale of


Scatophagus multifasciatus (magn.)
Fig. 13.—Ctenoid scale of Platycephalus
cirrhonasus (magn.)

Fig. 14.—Ctenoid scale of Gobius


ommaturus (magn.)
Fig. 15.—Ctenoid scale of Lethrinus
(magn.)

Fig. 16. Ganoid


Scales.
Ganoid scales are hard and bony, covered with a layer of
enamel; they are generally rhombic or quadrangular, rarely rounded
and imbricate; and arranged in oblique rows, those of one row being
linked together by an articulary process. This type of scales,
common in fossil Ganoid fishes, occurs among recent fishes in
Lepidosteus and Polypterus only.
Finally, in Sharks, the Balistidæ, and others, true scales are
absent and replaced by the ossified papillæ of the cutis, which give
the surface the appearance of fine-grained chagreen. These
generally small bodies, as well as the large osseous scutes of the
Rays, Sturgeons, etc., have been comprised under the common
name Placoid scales; a term which deservedly is being abandoned.

Fig. 17.—Dermal papillæ of Monacanthus trossulus.

Fig. 18.—Dermal papillæ of Monacanthus hippocrepis (magn.)


Fig. 19.—Cycloid scale from the lateral line of Odax lineatus
(magn.)
Along the side of the body of osseous fishes runs a series of
perforated scales, which is called the lateral line (Fig. 21). The
perforating duct is simple at its base, and may be also simple at its
outer opening (Fig. 19), or (and this is frequently the case) the
portion on the free surface of the scale is ramified (Fig. 20). The
lateral line runs from the head to the tail, sometimes reaching the
caudal fin, sometimes stopping in front of it, sometimes advancing
over its rays. It is nearer to the dorsal profile in some fishes than in
others. Some species have several lateral lines, the upper one
coasting the dorsal, the lower the abdominal outline, one running
along the middle as usual. The scales of the lateral line are
sometimes larger than the others, sometimes smaller, sometimes
modified into scutes, sometimes there are no other scales beside
them, the rest of the body being naked. The foramina of the lateral
line are the outlets of a muciferous duct which is continued on to the
head, running along the infraorbital bones, and sending off a branch
into the præopercular margin and mandible. In many fishes, as in
many Sciænoids, Gadoids, and in numerous deep-sea fishes, the
ducts of this muciferous system are extraordinarily wide, and
generally filled with mucus, which is congealed or contracted in
specimens preserved in spirits, but swells again when the specimens
are immersed in water. This system is abundantly provided with
nerves, and, therefore, has been considered to be the seat of a
sense peculiar to fishes, but there cannot be any doubt that its
function is the excretion of mucus, although probably mucus is
excreted also from the entire surface of the fish.

Fig. 20.—Cycloid scale from the lateral line of Labrichthys


laticlavius (magn.)
The scales, their structure, number and arrangement, are an
important character for the determination of fishes; in most scaly
fishes they are arranged in oblique transverse series; and as the
number of scales in the lateral line generally corresponds to the
number of transverse series, it is usual to count the scales in that
line. To ascertain the number of longitudinal series of scales, the
scales are counted in one of the transverse series, generally in that
running from the commencement of the dorsal fin, or the middle of
the back to the lateral line, and from the lateral line down to the vent
or ventral fin, or middle of the abdomen.[4]
Fig. 21.—Arrangement of scales in the Roach (Leuciscus ratilus): Ll = Lateral line;
tr = Transverse line. a, Transverse line from lateral line to ventral fin.

The scales of many fishes are modified for special purposes,


especially to form weapons of defence or a protective armour, but
the details of such modifications are better mentioned under the
several families in which they occur. All scales are continually
growing and wasting away on the surface, and it seems that some
fish, at least,—for instance, Salmonoids—“shed” them periodically;
during the progress of this shedding the outlines of the scales are
singularly irregular.
CHAPTER III.

TERMINOLOGY AND TOPOGRAPHY OF THE SKELETON.

In order to readily comprehend the subsequent account of the modifications of


the skeleton in the various sub-classes and groups of Fishes, the student has to
acquaint himself with the terms used for the numerous bones of the fish skeleton,
as well as with their relative position. The skeleton of any of the more common
kinds of osseous fish may serve for this purpose; that of the Perch is chosen
here.
The series of bones constituting the axis of the body, and destined to protect
the spinal chord and some large longitudinal blood-vessels, is called the vertebral
or spinal column; the single bones are the vertebræ. The skull consists of the
bones surrounding the brain and organs of sense, and of a number of arches
suspended from it, to support the commencement of the alimentary canal and the
respiratory organs.
The vertebra (Fig. 22) consists of a body or centrum (c), with a concave
anterior and posterior surface, and generally of several processes or apophyses,
as—1. Two neurapophyses (na), which, on the dorsal side, rising upwards, form
the neural arch over the canal, in which the spinal chord is lodged. 2. Two
parapophyses (pa) usually projecting from the lower part of the sides of the body,
or two hæmapophyses (ha) which actually coalesce to form on the ventral side
the hæmal canal for a large trunk of the vascular system. 3. A neural spine (ns),
which crowns the neurapophyses, or is interposed between their tips. 4. A hæmal
spine (hs), having the same relation to the hæmapophyses. 5. Two
pleurapophyses or floating ribs, suspended from, or from the base of, the
parapophyses. 6. In most fishes the neural arches are connected together by
articular or oblique processes, zygapophyses (za), which are developed from the
base of each neurapophysis.
Fig. 22.—Side and Front view of Fish-vertebra.
The vertebræ are either abdominal or caudal vertebræ, the coalescence of the
parapophyses into a complete hæmal ring, and the suspension of the anal fin
generally forming a sufficiently well-marked boundary between abdominal and
caudal regions (Fig. 23). In the Perch there are twenty-one abdominal and as
many caudal vertebræ. The centrum of the first vertebra or atlas is very short,
with the apophyses scarcely indicated, and lacking ribs like the succeeding
vertebra. All the other abdominal vertebræ, with the exception of the last or two
last, are provided with ribs, many of which are bifid (72). A series of flat spines
(74), called interneurals, to which the spines and rays of the dorsal fins are
articulated, are supported by the neural spines, the strength of the neurals and
interneurals corresponding to that of the dermal spines (75). The caudal vertebræ
differ from the abdominal in having the hæmapophyseal elements converted into
spines similar to the neurals, the anterior being likewise destined to support a
series of interhæmals (79), to which the anal rays are articulated. The last and
smallest caudal vertebra articulates with the hypural (70), a fan-like bone, which,
together with the dilated hindmost neural and hæmal elements, supports the
caudal rays.
Looking at a perch’s skull from the side (Fig. 24), the most superficial bones
will be found to be those of the jaws, a chain of thin bones round the lower half of
the eye, and the opercles.
The anterior margin of the upper jaw is formed by the intermaxillary or
premaxillary (17) which bears teeth, terminates in a pedicle above, to allow of a
forward sliding motion of the jaw, and is dilated into a flat triangular process
behind, on which leans the second bone of the upper jaw, the maxillary (18). This
bone is toothless, articulates with the vomer and palatine bone, and is greatly
dilated towards its distal extremity. Both the maxillary and intermaxillary lie and
move parallel to each other, being connected by a narrow membrane; in many
other fishes their relative position is very different.
The mandible or lower jaw consists of a right and left ramus; their union by a
ligament in front is called symphysis. Each ramus is formed of several pieces;
that which, by a sigmoid concavity articulates with the quadrate, is the articulary
bone (35); it sends upwards a coronoid process, to which a ligament from the
maxillary and the masticatory muscles are attached; and forwards a long-pointed
process, to be sheathed in the deep notch of the anterior piece. A small separate
piece (36) at the lower posterior angle of the mandible is termed angular. The
largest piece (34) is tooth-bearing, and hence termed dentary; at its inner surface
it is always deeply excavated, to receive a cylindrical cartilage, called Meckel’s
cartilage, the remains of an embryonic condition of the jaw, the articulary and
angular being but ossified parts of it. In other Teleostei this number is still more
increased by a splenial and other bones.
The infraorbital ring of bones (Fig. 23, 19) consists of several (four) pieces, of
which the anterior is the largest, and distinguished as præorbital.
The so-called præoperculum (30) belongs rather to the bones of the
suspensorium of the mandible, presently to be described, than to the opercles
proper. It is narrow, strong, angularly bent, so as to consist of a vertical and
horizontal limb, with an incompletely closed canal running along both limbs. As it
is quite a superficial bone, and frequently armed with various spines, its form and
configuration form an important item in the descriptive details of many fishes.
The principal piece of the gill-cover is the operculum (28), triangular in shape,
situated behind, and movably united with, the vertical limb of the præoperculum.
There is an articulary cavity at its upper anterior angle for its junction with the
hyomandibular. The oblong lamella below the operculum is the sub-operculum
(32), and the one in front of this latter, below the horizontal limb of the
præoperculum, is the interoperculum (33), which is connected by ligament with
the angular piece of the lower jaw, and is also attached to the outer face of the
hyoid, so that the gill-covers cannot open or shut without the hyoid apparatus
executing a corresponding movement.
The chain of flat bones which, after the removal of the temporal muscles,
appear arranged within the inner concavity of the præoperculum (Fig. 24), are
comprised with the latter under the common name of mandibulary suspensorium.
They connect the mandible with the cranium. The uppermost, the epitympanic or
hyomandibular (23), is articulated by a double articulary head with the mastoid
and posterior frontal. Another articulary head is destined for the opercular joint.
The mesotympanic or symplectic (31) appears as a styliform prolongation of the
lower part of the hyomandibular; is entirely cartilaginous in the young, but nearly
entirely ossified in the adult. The position of this bone is noteworthy, because,
directly inwards of its cartilaginous junction with the hyomandibular, there is
situated the uppermost piece of the hyoid arch, the stylohyal. The next bone of
the series is the pretympanic or metapterygoid (27), a flat bone forming a bridge
towards the pterygoid, and not rarely absent in the teleosteous sub-class. Finally,
the large triangular hypo-tympanic or quadrate (26) has a large condyle for the
mandibulary joint.
The palatine arch (Fig. 26) connects the suspensorium with the anterior
extremity of the skull, and is formed by three bones: the entopterygoid (25), an
oblong and thin bone attached to the inner border of the palatine and pterygoid,
and increasing the surface of the bony roof of the mouth towards the median line;
it constitutes also the floor of the orbit. The pterygoid (24) (or os transversum)
starts from the quadrate, and is joined by suture to the palatine, which is toothed,
and reaches to the vomer and anterior frontal.
In the occipital region there are distinguished the basi-occipital (5), readily
recognised by the conical excavation corresponding and similar to that of the
atlas, with which it is articulated through the intervention of a capsule filled with a
gelatinous substance (the remains of the notochord); the exoccipitals (10),
articulated, one on each side, to the basi-occipital, and expanding on the upper
surface of that bone, so as to meet and support the spinal column; a superficial
thin lamella (13), suturally connected with the exoccipitals, not constant in fishes,
and erroneously believed by Cuvier to be the petrosal (os petrosum) of higher
animals; further, the paroccipitals (9), which are wedged in between the
exoccipitals and supraoccipital. This last bone (8) forms the key of the arch over
the occipital foramen, and raises a strong high crest from the whole length of its
mesial line; a transverse supraoccipital ridge, coming from each side of the base
of this spine runs outwards laterally to the external angles of the bone. The
supraoccipital separates the parietals, and forms a suture with the frontals.
In front of the basi-occipital the base of the skull is formed by the
basisphenoid (parasphenoid of Huxley) (6). This very long and narrow bone
extends from the basi-occipital beyond the brain-capsule to between the orbits,
where it forms the support of the fibro-membranous interorbital septum. Anteriorly
it is connate with another long hammer-shaped bone (16), the vomer, the head of
which marks the anterior end of the palate, and is beset with teeth. The
alisphenoids (11) are short broad bones, rising from the basisphenoid; their
posterior margins are suturally connected with the anterior of the basi- and
exoccipitals.

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