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 About the Authors

Bryan Kolb received his Ph.D. from The

Pennsylvania State University and con-
ducted postdoctoral work at the Univer-
sity of Western Ontario and the Montreal
Neuro­logical Institute. In 1976 he moved
to the University of Lethbridge, Alberta,
where he is a professor of neuroscience and
holds a Board of Governors Chair in Neuro-
science. His current research examines how
Deborah Muirhead

perinatal factors—including tactile stimula-

tion, psycho­active drugs, stress, and injury—
modify the developing cerebral cortex and how these changes are related to
behavior. Kolb is a fellow of the Royal Society of Canada, the Canadian Psy-
chological Association (CPA), the American Psychological Association, and the
Association of Psychological Science. Currently a senior fellow of the Child
Brain Development program of the Canadian Institute for Advanced Research,
he is a recipient of the Hebb Prize from the CPA and from the Canadian So-
ciety for Brain, Behaviour, and Cognitive Science (CSBBCS). He has received
honorary doctorates from the University of British Columbia, Thompson Riv-
ers University, and Concordia University. He is a recipient the Ingrid Speaker
Gold Medal for research, the distinguished teaching medal from the University
of Lethbridge, and the Key to the City of Lethbridge. He and his wife train and
show horses in Western riding performance events.

Ian Q. Whishaw received his Ph.D.

from Western University and is a professor
of neuroscience at the University of Leth-
bridge. He has held visiting appointments
at the University of Texas, the University
of Michigan, the University of Cambridge,
and the University of Strasbourg. He is a fel-
low of Clair Hall, Cambridge, the Canadian
Psychological Association, the American
David Benard

Psychological Association, and the Royal

Society of Canada. He is a recipient the Ca-
nadian Humane Society Bronze Medal for bravery, the Ingrid Speaker Gold
Medal for research, the distinguished teaching medal from the University of
Lethbridge, and the Donald O. Hebb Prize. He has received the Key to the
City of Lethbridge and has honorary doctorates from Thompson Rivers Uni-
versity and the University of Lethbridge. His research addresses the neural
basis of skilled movement and the neural basis of brain disease. The Institute
for Scientific Information includes him in its list of most-cited neuroscientists.
His hobby is training and showing horses for Western performance events.
Brief Contents

Preface xix chapter 14

The Parietal Lobes 374
Media and Supplements xxiii
chapter 15
The Temporal Lobes 400
Part I Background chapter 16
chapter 1 The Frontal Lobes 427
The Development of Neuropsychology 1 chapter 17
chapter 2 Cortical Networks and Disconnection
Research on the Origins of the Human Brain Syndromes 462
and Behavior 28
chapter 3 Part IV Higher Functions
Nervous System Organization 53 chapter 18
chapter 4 Learning and Memory 480
The Structure and Electrical Activity chapter 19
of Neurons 85 Language 515
chapter 5 chapter 20
Communication Between Neurons 115 Emotion and the Social Brain 548
chapter 21
chapter 6
The Influence of Drugs and Hormones Spatial Behavior 575
on Behavior 139 chapter 22
Attention and Consciousness 607
chapter 7
Imaging the Brain’s Activity 174
Part V Plasticity and Disorders
chapter 23
Part II Cortical Organization Brain Development and Plasticity 635
chapter 8 chapter 24
Organization of the Sensory Systems 202 Neurodevelopmental Disorders 670
chapter 9 chapter 25
Organization of the Motor System 232 Plasticity, Recovery, and Rehabilitation
chapter 10 of the Adult Brain 699
Principles of Neocortical Function 255 chapter 26
chapter 11 Neurological Disorders 730
Cerebral Asymmetry 283 chapter 27
Psychiatric and Related Disorders 761
chapter 12
Variations in Cerebral Asymmetry 316 chapter 28
Neuropsychological Assessment 793
Glossary G-1
Part III Cortical Functions
chapter 13 Name Index NI-1
The Occipital Lobes 350 Subject Index SI-1
v
Page intentionally left blank
Contents

Preface xix chapter 2

Media and Supplements xxiii Research on the Origins of the Human
Brain and Behavior
Part I Background Portrait Evolving a Capacity for
Language 28
chapter1
The Development of Neuropsychology 2.1 ​Human Origins and the Origins of Larger
Brains 29
Portrait Living with Traumatic Research on Hominid Evolution 29
Brain Injury 1 Evolution of the Human Brain and Behavior 31
1.1 ​The Brain Theory 2 Relating Brain Size and Behavior 32
What Is the Brain? 2 The Meaning of Human Brain-Size
How Does the Brain Relate to the Rest of the Comparisons 37
Nervous System? 4 The Acquisition of Culture 39
1.2 ​Perspectives on the Brain 2.2 ​Comparative Research
and Behavior 5 in Neuropsychology 39
Aristotle: Mentalism 5 Understanding Brain Mechanisms 40
Descartes: Dualism 5 Designing Animal Models of Disorders 40
Darwin: Materialism 7 Describing Evolutionary Adaptations 41
Contemporary Perspectives 8 2.3 ​Genes, Environment, and Behavior 41
1.3 ​Brain Function: Insights from Brain Snapshot A Genetic Diagnosis 42
Injury 8 Mendelian Genetics and the Genetic Code 43
Localization of Function 8 Applying Mendel’s Principles 44
Lateralization of Function 10 Genetic Engineering 47
Neuroplasticity 13 Phenotypic Plasticity and the Epigenetic Code 49
Hierarchical Organization 13

Snapshot The Dilemma in Relating Behavior chapter3

and Consciousness 14 Nervous System Organization
1.4 ​The Neuron Theory 17 Portrait Stroke 53
Nervous System Cells 17 3.1 ​Neuroanatomy: Finding Your Way
Identifying the Neuron 18 Around the Brain 54
Relating Electrical Activity in Neurons Describing Location in the Brain 54
to Behavior 19
A Wonderland of Nomenclature 56
Connections Between Neurons As the Basis
of Learning 20
3.2 ​Overview of Nervous System Structure
and Function 57
1.5 ​Contributions to Neuropsychology from
Support and Protection 58
Allied Fields 21
Blood Supply 59
Neurosurgery 22
Neurons and Glia 59
Psychometrics and Statistical Evaluation 23
Gray, White, and Reticular Matter 61
Brain Imaging 24
Layers, Nuclei, Nerves, and Tracts 62

Coverage links neuropsychological theory and assessment

vii
viii cont ent s

3.3 ​Origin and Development of the Central The Resting Potential 99

Nervous System 62 Graded Potentials 102
3.4 ​The Spinal Cord 64 The Action Potential 103
Spinal-Cord Structure and Spinal 4.3 ​Sending a Message Along
Nerve Anatomy 64 an Axon 106
Spinal-Cord Function and the Spinal Nerves 65 The Nerve Impulse 106
Cranial Nerve Connections 67 Saltatory Conduction and Myelin Sheaths 107
Autonomic Nervous System Connections 69
Snapshot Diagnosing MS 108
The Brainstem 70
3.5 ​
The Hindbrain 70 4.4 ​How Neurons Integrate
The Midbrain 71 Information 109
The Diencephalon 72 Excitatory and Inhibitory Postsynaptic
Potentials 109
The Forebrain 72
3.6 ​
Voltage-Sensitive Channels and the Action
The Basal Ganglia 73 Potential 110
The Limbic System 74 Summation of Inputs 110
The Neocortex 75 The Versatile Neuron 112
Fissures, Sulci, and Gyri 76
4.5 ​Stimulating and Recording
Cortical Organization in Relation to Inputs,
with Optogenetics 112
Outputs, and Function 77
Cellular Organization in the Cortex 78
Cortical Connections 80 5
chapter
Snapshot Brainbow and Clarity 81 Communication Between Neurons
The Crossed Brain 82
3.7 ​ Portrait Otto Loewi’s Dream
Breakthrough 115

chapter 4 5.1 ​Neurotransmitter Discovery 116

The Structure and Electrical Activity
of Neurons
Portrait The Halle Berry Neuron 85
The Neuron’s Structure 86
4.1 ​
Overview of a Neuron 86
The Neuron as a Factory 87
The Cell Membrane: Barrier and
Gatekeeper 88
The Nucleus: Blueprints for Proteins 90
Protein Synthesis: Transcription
and Translation 91
Applying Epigenetic Mechanisms 92
Proteins: The Cell’s Products 93
Golgi Bodies and Microtubules: Protein
Packaging and Shipment 93
Crossing the Cell Membrane: Channels, Gates,
and Pumps 94
4.2 ​The Neuron’s Electrical Activity 95
Recording from an Axon 96
How the Movement of Ions Creates
Electrical Charges 97
5.2 ​The Structure of Synapses 117
Chemical Synapses 117
Electrical Synapses 118
5.3 ​Neurotransmission in Four Steps 119
Step 1: Transmitter Synthesis and Storage 120
Step 2: Neurotransmitter Release 120
Step 3: Receptor-Site Activation 121
Step 4: Neurotransmitter Deactivation 121
5.4 ​Types of Synapses 122
Synaptic Variations 122
Excitatory and Inhibitory Messages 123
5.5 ​Varieties of Neurotransmitters 124
Four Criteria for Identifying
Neurotransmitters 124
Three Classes of Neurotransmitters 125
Peptide Transmitters 127
Transmitter Gases 128
5.6 ​Excitatory and Inhibitory Receptors 129
Ionotropic Receptors and Excitation 129
Metabotropic Receptors and Inhibition 129
Excitatory and Inhibitory Receptor Effects 131

5.7 ​Neurotransmitter Activating Systems
and Behavior 131
Neurotransmission in Peripheral Nervous System
Divisions 131
Activating Systems of the Central
Nervous System 132
Snapshot Neurochemical Links Between SIDS
and Sleep Apnea 136
chapter 6
The Influence of Drugs and Hormones on
Behavior
Portrait The Case of the Frozen Addict 139
6.1 ​Principles of Psychopharmacology 140
Routes of Drug Administration 140
Routes of Drug Removal 141
Revisiting the Blood–Brain Barrier 142
Drug Routes and Dosage 143
6.2 ​Drug Actions in Synapses 143
Steps in Synaptic Transmission 144
Examples of Drug Action: An Acetylcholine
Synapse 144
Tolerance 146
Sensitization 146
Can Drugs Cause Brain Damage? 148
6.3 ​Grouping Psychoactive Drugs 150
Group I: Antianxiety Agents and
Sedative Hypnotics 150
Group II: Antipsychotic Agents 152
Group III: Antidepressants and
Mood Stabilizers 153
Group IV: Opioid Analgesics 155
Group V: Psychotropics 156
Snapshot Cognitive Enhancement 158
General Stimulants 160
6.4 ​Individual Responses and Influences
on Addiction 160
Behavior on Drugs 160
Addiction and Dependence 161
Sex Differences in Addiction 162
Wanting-and-Liking Theory 162
Treating Drug Abuse 163
6.5 ​Hormones 164
Hierarchical Control of Hormones 165
Classes and Functions of Hormones 165
contents ix
Homeostatic Hormones 166
Gonadal Hormones 167
Anabolic–Androgenic Steroids 168
Glucocorticoids and Stress 168
Ending a Stress Response 169
chapter7
Imaging the Brain’s Activity
Portrait Angelo Mosso 174
7.1 ​Recording the Brain’s
Electrical Activity 175
Single-Cell Recording 175
Electroencephalographic Recording 177
Event-Related Potentials 181
Magnetoencephalography 183
7.2 ​Brain Stimulation 183
Deep Brain Stimulation 184
Transcranial Magnetic Stimulation 184
7.3 ​Static Imaging Techniques 185
Imaging by X-Ray 185
Computed Tomography 186
7.4 ​Dynamic Brain Imaging 187
Positron Emission Tomography 187
Magnetic Resonance Imaging 189
Magnetic Resonance Spectroscopy 191
Diffusion Tensor Imaging 192
Functional Magnetic Resonance Imaging 193
Resting-State fMRI 194
Optical Tomography 195
Snapshot Tuning in to Language 196
7.5 ​Comparing Brain-Imaging Techniques
and Uses 197
Imaging Techniques, Pro and Con 198
Toward Multimodal Brain Atlases 199
Part II Cortical Organization
chapter 8
Organization of the Sensory
Systems
Portrait Phantoms of the Brain 202
8.1 ​General Principles of Sensory-
System Function 203
Sensory Receptors and Neural Relays 203
x c ontents

Neural Relays Determine the Hierarchy The Cortex: Intention 263

of Motor Responses 207 10.2 ​The Structure of the Cortex 264
Central Organization of Sensory Systems 208 Cortical Cells 264
Sensory Receptors and Pathways 211
8.2 ​
Snapshot Mapping the Human Cortex 265
Vision 211
Hearing 214 Cortical Layers, Efferents, and Afferents 266
Body Senses 217 Cortical Columns, Spots, and Stripes 268
The Chemical Senses: Taste and Smell 223 Multiple Representations: Mapping Reality 270
Cortical Systems: Frontal Lobe, Paralimbic
Snapshot Watching the Brain Make Cortex, and Subcortical Loops 272
Flavor 225 Cortical Connections, Reentry, and
8.3 ​Perception 228 the Binding Problem 273
Illusions 228 10.3 ​Functional Organization of
Synesthesia 229 the Cortex 275
Sensory Synergies 229 A Hierarchical Model of Cortical Function 275
Evaluating the Hierarchical Model 276
A Contemporary Model of Cortical
chapter9 Function 278
Organization of the Motor System 10.4 ​Do Human Brains Possess
Portrait Mind in Motion 232 Unique Properties? 279
The Neocortex: Initiating Movement 233
9.1 ​
Mapping the Motor Cortex Using chapter11
Electrical Stimulation 235 Cerebral Asymmetry
Multiple Representations in the Motor Cortex 236
Portrait Words and Music 283
The Movement Lexicon 239
Mirroring Movement 242
Anatomical Asymmetries
11.1 ​
in the Human Brain 284
Snapshot Recording Mirror Neuron Cerebral Asymmetry 284
Activity 244 Neuronal Asymmetry 288
9.2 ​The Brainstem: Motor Control 245 Genetic Asymmetry 288
The Basal Ganglia and Movement Force 245 Asymmetries in
11.2 ​
The Cerebellum and Motor Learning 247 Neurological Patients 289
9.3 ​Communicating with the Spinal Patients with Lateralized Lesions 289
Cord 250 Commissurotomy Patients 291
Spinal-Cord Pathways 250 Brain Stimulation 294
Spinal Motor Neurons 251 Carotid Sodium Amobarbital Injection 296
Behavioral Asymmetries
11.3 ​
chapter 10 in the Intact Brain 298
Asymmetry in the Visual System 298
Principles of Neocortical Function
Asymmetry in the Auditory System 299
Portrait Hemispherectomy 255 Asymmetry in the Somatosensory System 300
A Hierarchy of Function from Spinal Cord
10.1 ​ Asymmetry in the Motor System 301
to Cortex 256 What Do Laterality Studies Tell Us about
The Spinal Cord: Reflexes 258 Brain Function? 303
The Hindbrain: Postural Support 258 Neuroimaging and Asymmetry 304
11.4 ​
The Midbrain: Spontaneous Movement 260 11.5 ​Theoretical Arguments:
The Diencephalon: Affect and Motivation 261 What Is Lateralized? 306
The Basal Ganglia: Self-Maintenance 262 Specialization Models 306

Interaction Models 308
Preferred Cognitive Mode 309
Snapshot Imaging the Brain’s Plasticity 310
Measuring Behavior in
Neuropsychology 311
chapter 12
Variations in Cerebral Asymmetry
Portrait Individual Responses to Injury 316
Handedness and
12.1 ​
Functional Asymmetry 316
Anatomical Studies 317
Functional Cerebral Organization in Left-
Handers 318
Theories of Hand Preference 319
Snapshot Genetic Influences
on Brain Structure 321
Sex Differences in
12.2 ​
Cerebral Organization 323
Sex Differences in Children’s Behavior 323
Sex Differences in Adult Behavior 324
Sex Differences in Brain Structure 328
The Homosexual Brain 330
Sex Differences Revealed in Functional
Imaging Studies 331
Research with Neurological Patients 332
Explanations for Sex Differences 333
12.3 ​Environmental Effects
on Asymmetry 338
Language and Culture 338
Sensory or Environmental Deficits 340
Asymmetry in Nonhuman Animals 343
12.4 ​
Asymmetry in Birds 344
Asymmetry in Nonhuman Primates 344
Part III Cortical Functions
chapter13
The Occipital Lobes
Portrait An Injured Soldier’s Visual
World 350
13.1 ​Occipital Lobe Anatomy 350
Subdivisions of the Occipital Cortex 351
Connections of the Visual Cortex 353
contents xi
A Theory of Occipital-Lobe Function 353
13.2 ​
Visual Functions Beyond the Occipital Lobe 354
Visual Pathways Beyond the Occipital Lobe 357
Imaging Studies of Dorsal and Ventral
Streams 359
Top-Down Predictions in Vision 360
13.3 ​Disorders of Visual Pathways 360
13.4 ​Disorders of Cortical Function 362
Case B.K.: V1 Damage and a Scotoma 362
Case D.B.: V1 Damage and Blindsight 364
Case G.Y. and Related Cases: V1 Damage
and Conscious Vision 364
Case J.I.: V4 Damage and Loss of Color
Vision 364
Case P.B.: Conscious Color Perception
in a Blind Patient 365
Case L.M.: V5 (MT) Damage and
the Perception of Movement 365
Case D.F.: Occipital Damage and Visual
Agnosia 366
Case V.K.: Parietal Damage and
Visuomotor Guidance 367
Cases D. and T.: Higher-Level
Visual Processes 367
Conclusions from the Case Studies 368
13.5 ​Visual Agnosia 368
Object Agnosias 368
Other Visual Agnosias 369
Visual Imagery 370
13.6 ​
Snapshot Generating Mental Images 371
chapter 14
The Parietal Lobes
Portrait Varieties of Spatial Information 374
Parietal Lobe Anatomy 374
14.1 ​
Subdivisions of the Parietal Cortex 375
Connections of the Parietal Cortex 376
Anatomy of the Dorsal Stream 377
A Theory of Parietal-Lobe Function 378
14.2 ​
Behavioral Uses of Spatial Information 379
The Complexity of Spatial Information 382
Other Parietal-Lobe Functions 382
Snapshot Spatial Cognition and
White-Matter Organization 383
14.3 ​Somatosensory Symptoms of Parietal
Lesions 384
xii cont ent s
Somatosensory Thresholds 384
Somatoperceptual Disorders 385
Numb Touch 385
Somatosensory Agnosias 386
14.4 ​Symptoms of Posterior
Parietal Damage 387
Bálint’s Syndrome 387
Contralateral Neglect and Other Symptoms
of Right Parietal Lesions 388
The Gerstmann Syndrome and Other Left
Parietal Symptoms 390
Apraxia and the Parietal Lobe 391
Drawing 392
Spatial Attention 392
Disorders of Spatial Cognition 393
Left and Right Parietal Lobes Compared 394
14.5 ​Major Symptoms and
Their Assessment 394
Clinical Neuropsychological Assessment 395
chapter15
The Temporal Lobes
Portrait Living with Temporal-Lobe
Damage 400
Temporal-Lobe Anatomy 400
15.1 ​
Subdivisions of the Temporal Cortex 401
Connections of the Temporal Cortex 402
Anatomy of the Ventral Stream 403
A Theory of Temporal-Lobe Function 404
15.2 ​
The Superior Temporal Sulcus and
Biological Motion 406
Visual Processing in the Temporal Lobe 406
Are Faces Special? 409
Auditory Processing in the Temporal Lobe 411
Asymmetry of Temporal-Lobe Function 415
15.3 ​Symptoms of Temporal-
Lobe Lesions 416
Disorders of Auditory and Speech
Perception 417
Disorders of Music Perception 417
Snapshot Imaging Auditory
Hallucinations 418
Disorders of Visual Perception 419
Disturbance of Visual- and Auditory-
Input Selection 420
Impaired Organization and Categorization 420
Inability to Use Contextual Information 421
Memory Impairment 421
Altered Affect and Personality 422
Changes in Sexual Behavior 423
15.4 ​Clinical Neuropsychological
Assessment of Temporal-Lobe
Damage 423
chapter 16
The Frontal Lobes
Portrait Losing Frontal-Lobe Functions 427
Frontal-Lobe Anatomy 427
16.1 ​
Subdivisions of the Frontal Cortex 428
The Connectome and the Frontal Cortex 430
A Theory of Frontal-Lobe Function 431
16.2 ​
Functions of the Premotor Cortex 432
Functions of the Prefrontal Cortex 433
Asymmetry of Frontal-Lobe Function 435
Heterogeneity of Frontal-Lobe Function 435
Snapshot Heterogeneity of Function in
the Orbitofrontal Cortex 436
16.3 ​Symptoms of Frontal-Lobe Lesions 437
Disturbances of Motor Function 437
Loss of Divergent Thinking 440
Environmental Control of Behavior 443
Poor Temporal Memory 446
Impaired Social and Sexual Behavior 449
Does a Spatial Deficit Exist? 452
Clinical Neuropsychological Assessment
of Frontal-Lobe Damage 452
Intelligence and the Frontal Lobes 454
16.4 ​
Imaging Frontal-Lobe Function 455
16.5 ​
16.6 ​Disorders Affecting
the Frontal Lobe 457
chapter 17
Cortical Networks and Disconnection
Syndromes
Portrait At Cross Purposes 462
Disconnecting Cognitive Functions 463
17.1 ​
Anatomy of Cerebral Connections 464
17.2 ​
17.3 ​Cortical Networks and Hubs 466
17.4 ​Behavioral Effects
of Disconnection 468
 contents xiii

17.5 ​
Hemispheric Disconnection 470 Unique Aspects of Emotional Memory 502
Commissurotomy 470 18.5 ​Short-Term Memory 502
Callosal Agenesis and Early Transections 471 Short-Term Memory and the Temporal and
17.6 ​Disconnecting Parietal Lobes 503
Sensorimotor Systems 472 Short-Term Memory and the Frontal Lobes 503
Olfaction 472 Neuropsychological Testing for Short-Term
Memory Function 503
Vision 473
Somatosensory Functions 474 Snapshot Disrupting Memory
Audition 474 Formation 504
Movement 475 18.6 ​Neurological Diseases and
Effects of Partial Disconnection 476 Long‑Term Memory 507
Snapshot An fMRI Study of Transient Global Amnesia 507
Disconnection 476 Herpes Simplex Encephalitis 507
Alzheimer’s Disease 508
17.7 ​Lesion Effects Reinterpreted As
Korsakoff’s Syndrome 508
Disconnection Syndromes 477
Neurotransmitter Activating Systems
Apraxia 477 and Memory 509
Agnosia and Alexia 477
18.7 ​Special Memory Abilities 510
Contralateral Neglect 478
Savant Syndrome 510
Hubs and Connectivity in Brain
Superior Autobiographical Memory 511
Dysfunction 478

chapter19
Part IV Higher Functions Language
Portrait Multilingual Meltdown 515
chapter 18
Learning and Memory What Is Language? 516
19.1 ​
Language Structure 516
Portrait The Mystery of Memory 480
Producing Sound 517
18.1 ​Learning, Memory, Core Language Skills 518
and Amnesia 481 Searching for the Origins
19.2 ​
Varieties of Amnesia 482 of Language 519
Anterograde and Retrograde Amnesia 484
Snapshot Genetic Basis for an Inherited
Time-Dependent Retrograde Amnesia 484
Speech and Language Disorder 520
Three Theories of Amnesia 485
Continuity Theory 520
18.2 ​Long-Term Explicit Memory 486
Discontinuity Theory 523
Episodic Memory 486
Experimental Approaches to Language
Autonoetic Awareness of Time 487
Origins 524
Semantic Memory 488
Localization of Language 526
19.3 ​
Neural Substrates of Explicit Memory 489
Anatomical Areas Associated with
Hemispheric Specialization for Explicit
Language 527
Memory 495
Speech Zones Mapped by Brain Stimulation
18.3 ​Long-Term Implicit Memory 497 and Surgical Lesions 529
Sparing of Implicit Memory in Amnesia 497 Speech Zones Mapped by Brain-
Neural Substrates of Implicit Memory 498 Imaging Techniques 531
18.4 ​Long-Term Emotional Memory 501 Neural Networks for Language 533
Evoking Negative Emotions 501 19.4 ​Language Disorders 536
Neural Substrates of Emotional Memory 501 Fluent Aphasias 536
xiv contents
Nonfluent Aphasias 538
Pure Aphasias 538
19.5 ​Localization of Lesions in
Aphasia 538
Cortical Language Components 539
Subcortical Language Components 540
Right-Hemisphere Contributions to
Language 540
19.6 ​Neuropsychological Assessment of
Aphasia 541
Assessing Developmental Language
Disorders 542
chapter20
Emotion and the Social Brain
Portrait Agenesis of the Frontal Lobe 548
The Nature of Emotion 549
20.1 ​
What Are Emotions? 549
Components of Emotion 549
Historical Views 550
20.2 ​
Investigating the Anatomy of Emotion 550
The Emotional Brain 551
Cortical Connections of Emotion 551
Candidate Structures in
20.3 ​
Emotional Behavior 553
Processing Emotional Stimuli 553
Brain Circuits for Emotion 554
20.4 ​Neuropsychological Theories
of Emotion 556
Appraisal Theories of Emotion 556
Snapshot Brain Activation in Social
Cognition 558
Cognitive–Emotional Interactions 559
Cognitive Asymmetry and Emotion 561
20.5 ​Asymmetry in
Emotional Processing 562
Producing Emotional Behavior 562
Interpreting Emotional Behavior 564
Temporal-Lobe Personality 566
20.6 ​The Social Brain and
Social Cognition 567
Frontal Lesions in Monkeys 567
Cerebral Lesions in Humans 568
Social Neural Networks 569
The Self and Social Cognition 570
Cognitive Control of Emotion 571
chapter 21
Spatial Behavior
Portrait Lost in Space 575
Spatial Behavior and
21.1 ​
Spatial Impairments 576
Explaining Spatial Behavior 577
Clinical Descriptions of Spatial
Impairments 577
Topographic Disorientation 578
21.2 ​Dorsal- and Ventral-Stream
Contributions to Spatial Behavior 581
The Dorsal Stream in Parietal Cortex 581
The Dorsal Stream in Frontal Cortex 584
The Dorsal and Ventral Streams in
Temporal Cortex 585
SNAPSHOT Imaging the Hippocampi of London
Taxi Drivers 586
Experimental Models of
21.3 ​
Spatial Behavior 587
Route Following 588
Piloting 588
Caching Behavior 590
Dead Reckoning 591
Neuropsychological Tests of Spatial
Behavior 593
Single-Cell Recording and Spatial Behavior 594
Location of Spatial Cells 597
21.4 ​Individual Differences in
Spatial Abilities 598
Sex-Related Differences 598
Handedness and Spatial Ability 601
21.5 ​Episodic Memory, Scene Construction,
and Theory of Mind 601
Spatial Activity in Episodic Memory 601
Spatial Memory as Distinct from
Episodic Memory 602
Spatial and Episodic Memory as
Hippocampal Functions 602
Theory of Mind 603
chapter 22
Attention and Consciousness
Portrait A Curious Case of Neglect 607
22.1 ​Defining Attention and
Consciousness 608
 contents xv

22.2 ​
Attention 609 23.6 ​Brain Injury and Plasticity 656
Automatic and Conscious Effects of Age 657
Processing Compared 609 Effects of Brain Damage on Language 657
Neurophysiological Evidence of Attention 612
SNAPSHOT Distinct Cortical Areas for Second
Parallel Processing of Sensory Input 615
Languages 658
Functional Imaging and Attention 615
Networks of Attention 618 Reorganization of Language 660
Mechanisms of Attention 621 Absence of Language After Bilateral
Lesions 662
22.3 ​
Inattention 622
Absence of Visual Attention 622
23.7 ​Studying Plasticity After Early Brain
Injury 662
Sensory Neglect 624
Effects of Early Brain Lesions on Behaviors
22.4 ​
Consciousness 625 Later in Life 663
The Neural Basis of Consciousness 627 Effects of Early Brain Lesions on Brain Structure
Cerebral Substrates of Consciousness 629 Later in Life 665
SNAPSHOT Stimulating Nonconscious Factors Influencing Plasticity After Early
Cortical Injury 665
Emotion 630
Emotion and Consciousness 630
24
chapter
Nonconscious Processing 632
Neurodevelopmental Disorders
Portrait Life Without Reading 670
Part V Plasticity and Disorders
24.1 ​Neurodevelopmental Disorders 671
chapter 23 Historical Background and Evolution
Brain Development and Plasticity of Understanding 671
Portrait Plasticity and Language 635 Incidence of Neurodevelopmental
Disorders 673
23.1 ​Approaches to Studying Types of Neurodevelopmental Disorders 673
Brain Development 635 Learning Disorders That
24.2 ​
23.2 ​Development of the Human Brain 636 Affect Reading 674
Neuron Generation 638 Types of Reading 674
Cell Migration and Differentiation 639 Causes of Reading Disorders 675
Neural Maturation 640
Synapse Formation and Pruning 641
Snapshot Imaging Sound Perception in
Glial Development 643
Controls and Subjects with Dyslexia 678
The Adolescent Brain 643 Neuropsychological Evaluation 679
Imaging Studies of Brain
23.3 ​ 24.3 ​Nonlanguage Neurodevelopmental
Development 644 Disorders 681
23.4 ​Development of Problem-​ Attention-Deficit/Hyperactivity Disorder 681
Solving Ability 647 Cerebral Palsy 683
23.5 ​Environmental Effects on Hydrocephalus 685
Brain Development 650 Autism Spectrum Disorders 686
Developmental Effects of Aversive Fragile-X Syndrome 689
Environments 651 Fetal Alcohol Spectrum Disorder 690
Environmental Influences on Brain 24.4 ​Developmental Influences on
Organization 652 Neurodevelopmental Disorders 692
Experience and Neural Connectivity 654 Structural Damage and Toxic Effects 692
Plasticity of Representational Zones in Hormonal Effects: The Geschwind–
the Developing Brain 655 Galaburda Theory 693
xvi contents

Environmental Deprivation 694 25.5 ​Variables Affecting Recovery 720

The Birthday Effect 694 25.6 ​Therapeutic Approaches to Recovery
24.5 ​Adult Outcome of After Brain Damage 721
Neurodevelopmental Disorders 695 Rehabilitation 722
Pharmacological Therapies 724
chapter 25 Electrical Stimulation 725
Plasticity, Recovery, and Rehabilitation Brain-Tissue Transplants and Stem-Cell
of the Adult Brain Induction 725
Diet 726
Portrait Concussion 699
25.1 ​Principles of Brain Plasticity 700 chapter26
Principle 1: Plasticity is common to all Neurological Disorders
nervous systems, and the principles are
conserved. 700 Portrait Posttraumatic Stress Disorder 730
Principle 2: Plasticity can be analyzed at 26.1 ​The Neurological Examination 731
many levels. 700 The Patient’s History 731
Principle 3: The two general types of plasticity The Physical Examination 731
derive from experience. 707 Cerebral Vascular Disorders 733
26.2 ​
Principle 4: Similar behavioral changes can Types of Cerebral Vascular Disease 733
correlate with different plastic changes. 707
Treating Cerebral Vascular Disorders 735
Principle 5: Experience-dependent
changes interact. 707 26.3 ​Traumatic Brain Injuries 736
Principle 6: Plasticity is age-dependent. 708 Open Head Injuries 737
Principle 7: Plastic changes are time- Closed Head Injuries 737
dependent. 708 Behavioral Assessment of Head Injury 739
Principle 8: Plasticity is related to an Recovering from and Preventing Head Injury 741
experience’s relevance to the animal. 708 Epilepsy 741
26.4 ​
Principle 9: Plasticity is related to the intensity Classifying Seizures 742
or frequency of experiences. 709 Treating Epilepsy 743
Principle 10: Plasticity can be Tumors 743
26.5 ​
maladaptive. 709
Headache 745
26.6 ​
25.2 ​Can Plasticity Support Functional Types of Headache 745
Recovery After Injury? 709 Treating Headache 747
Compensation Compared with Recovery 710
Infections 747
26.7 ​
What Happens When a Brain Is Injured? 711
Types of CNS Infection 748
25.3 ​Examples of Functional Treating CNS Infection 749
Restitution 712 26.8 ​Disorders of Motor Neurons
Recovery from Motor-Cortex Damage 712 and the Spinal Cord 750
Recovery from Aphasia 712 Myasthenia Gravis 750
Recovery from Traumatic Lesions 713 Poliomyelitis 751
Recovery from Surgical Lesions 714 Multiple Sclerosis 751
Return to Daily Life 715 Paraplegia 752
25.4 ​Research on Plasticity Brown-Séquard Syndrome 752
in the Injured Brain 717 Hemiplegia 753
Functional Imaging After Cerebral Injury 717 26.9 ​Sleep Disorders 753
SNAPSHOT Using Imaging Narcolepsy 755
to Study Recovery 718 Insomnia 756

Physiological Mapping After Cerebral Injury 719 Snapshot Restless Legs Syndrome 757

chapter 27
Psychiatric and Related Disorders
Portrait Losing Touch with Reality 761
27.1 ​The Brain and Behavior 761
27.2 ​Schizophrenia 762
Structural Abnormalities in Schizophrenic
Brains 763
Biochemical Abnormalities in the Brains
of People with Schizophrenia 764
Schizophrenia as a
Neurodevelopmental Disorder 765
Cognitive Symptoms in Schizophrenia 765
Mood Disorders 767
27.3 ​
Neurochemical Aspects of Depression 767
Neuropathological and Blood-Flow
Abnormalities in Depression 768
SNAPSHOT Cortical Metabolic and Anatomical
Abnormalities in Mood Disorders 769
Neurobiological Aspects of Bipolar
Disorder 770
27.4 ​Anxiety Disorders 771
27.5 ​Psychiatric Symptoms of Cerebral
Vascular Disease 772
27.6 ​Psychosurgery 773
27.7 ​Motor Disorders 774
Hyperkinetic Disorders 774
Hypokinetic Disorders 778
Causes of Parkinsonism 780
Treating Parkinson’s Disease 781
Psychological Aspects of Parkinson’s
Disease 782
Dementias 783
27.8 ​
Anatomical Correlates of Alzheimer’s
Disease 784
Putative Causes of Alzheimer’s Disease 786
contents xvii
Clinical Symptoms and the Progression
of Alzheimer’s Disease 787
27.9 ​Micronutrients and Behavior 788
chapter28
Neuropsychological Assessment
Portrait Lingering Effects of Brain
Trauma 793
The Changing Face of
28.1 ​
Neuropsychological Assessment 794
Functional Brain Imaging 794
Cognitive Neuroscience 795
Managed Care 796
Rationale Behind Neuropsychological
28.2 ​
Assessment 797
Factors Affecting Test Choice 798
Goals of Neuropsychological Assessment 798
Intelligence Testing in Neuropsychological
Assessment 799
Categories of Neuropsychological
Assessment 801
Neuropsychological Tests and
28.3 ​
Brain Activity 802
The Problem of Effort 803
28.4 ​
28.5 ​Case Histories 804
Case 1: Epilepsy Caused by Left-
Hemisphere Tumor 804
Case 2: Epilepsy Caused by Right-
Hemisphere Infection 805
Case 3: Rehabilitation 805
Glossary G-1
Name Index NI-1
Subject Index SI-1
Page intentionally left blank
Preface

Looking back to 1980, when Fundamentals of Human Neuropsychology’s first edi-

tion appeared, reminds us that in the 1970s, human neuropsychology did not
yet exist as a unified body of knowledge about the human brain. The field had
coalesced around hunches and inferences based on laboratory studies of mon-
keys, cats, and rats as well as on scattered studies of humans with assorted brain
injuries. Over the past 40 years, as neuropsychology expanded, cognitive and
social neuroscience have emerged as disciplines. Advances in and ever-more
incisive use of noninvasive neuroimaging and abundant research innovations
all have improved our understanding of brain anatomy.
Studies of nonhuman species remain central to human neuropsychology’s
core principles, especially in understanding the structure and connectivity of
the primate brain, but are more focused on mechanisms than behavioral phe-
nomena. Many researchers may share a bias that functional neuroimaging can
replace studying brain-injured humans and laboratory animals. To others, this
seems unlikely given the complexity of brain processes and the nature of sub-
traction methods used in imaging. The two approaches have become comple-
mentary, and this seventh edition reflects their intellectual evolution:
• Neuroimaging has led the renaissance in understanding neural networks and ap-
preciating the brain’s connectome. In this edition, we have expanded Chapter 7,
Imaging the Brain’s Activity, both to include new methods and to consider
the pros and cons of different techniques in light of their relevant uses
and costs (see Section 7.5). Coverage of dynamic neural networks appears
throughout the book, especially in Chapters 10, 16 to 22, and 27.
• Epigenetics explains how our behaviors change our brains. We introduce basic
genetics and epigenetic principles in Section 2.3 and highlight both factors
throughout the book to reflect the expanding emphasis on epigenetics as a
factor in cerebral organization.
• Neuropsychological assessment is vital for evaluating patients with focal brain injuries.
One unexpected consequence of the cognitive neuroscience revolution is a de-
clining appreciation for neuropsychological theory and clinical focus. In this
new edition, we employ the venerable maze as a graphic icon (shown at right)
to identify for the reader particular discussions, cases, tables, and figures that
link theory and assessment throughout the book.

Content and Structure

Fundamentals differs from other textbooks of psychology, cognitive neuro­
science, or neuroscience. In our experience, students find it helpful to see the
brain from two organizational perspectives, anatomical and behavioral.
• We continue to provide the requisite basic background—about history, evo-
lution, genetics and epigenetics, anatomy, physiology, pharmacology, and
methodology—in Part I, Chapters 1 to 7.

xix
xx p ref ace

• Equally fundamental to understanding subsequent material, Part II, Chap-

ters 8 to 12, outlines the general organization and functions of the cerebral
cortex.
• Part III, Chapters 13 to 17, focuses on the anatomically defined cortical
regions. Understanding the organization of the cerebral cortex is central to
appreciating how the brain functions to produce the complex processes that
underlie complex behaviors.
• The psychological constructs presented in Part IV, Chapters 18 to 22, in-
cluding language, memory, social behavior and affect, spatial behavior, and
attention and consciousness, emerge from the neuronal networks explored
in Part III. Shifting from anatomy to psychological processes naturally
means revisiting material from earlier parts, but this time in the context of
psychological theory rather than anatomy.
• Part V, Chapters 23 to 28, considers brain development and plasticity and
includes more-detailed discussions of brain disorders introduced earlier in
the book. Chapters on neurological and psychiatric disorders and on neuro­
psychological assessment continue the book’s emphasis on approaching
human brain functions from an interdisciplinary perspective.
We have updated all of the chapters and the glossary that follows, both to
correspond to new material that reflects the changing face of neuro­psychology
and to include some unexpected topics—neuroeconomics in Section 22.4 and
micronutrients in Section 27.9 are two. Maintaining a manageable length
meant sacrificing some detail that may have been prominent in previous edi-
tions, sometimes reaching back to the first edition.
To address the challenge inherent in using a comprehensive text and to fa-
cilitate access to information, we added section numbers to each chapter’s main
headings. Readers can easily locate interrelated material relevant across several
topics, refresh their knowledge, or jump ahead to learn more.

Acknowledgments
As in the past, we must sincerely thank many people who have contributed to
the development of this edition. We are particularly indebted to colleagues
from around the world who have been so supportive and have strongly encour-
aged us to include their favorite topics. We have done so wherever possible.
We also thank the reviewers solicited by our editors on the sixth edition
of Fundamentals. Their anonymous comments contributed varied perspectives
and valuable points of consensus that helped us shape the new edition.

Julie Alvarez Peter Donovick

Tulane University The State University of New York at
Marlene Behrmann Binghamton
Carnegie Mellon University Amanda Higley
Edward Castañeda Point Loma Nazarene University
The University of Texas at El Paso Jamie Lillie
Pauline Dibbets Argosy University, Schaumburg
Maastricht University
 PREFACE xxi

Salvatore Massa Tony Robertson

Marist College Vancouver Island University
Taryn Myers Joe Wayand
Virginia Wesleyan College Walsh University
Martin Paczynski Robin Wellington
George Mason University St. John’s University
Rosie Reid
Dublin Business School

The staff at Worth Publishers and W. H. Freeman and Company are amaz-
ing and have made this task far more enjoyable than it would be without them.
These folks include our sponsoring editor, Daniel DeBonis, assistant editor
Nadina Persaud, and editorial assistant Katie Pachnos; our project editors, En-
rico Bruno, Andrew Roney, and Janice Stangel; and our production manager,
Sarah Segal. Our thanks to art director Diana Blume and interior designer
Tamara Newnam for a fresh, inviting, accessible new book design, and to Kevin
Kall for a striking cover.
Once again, Cecilia Varas coordinated photo research, ably assisted by re-
searcher Richard Fox. They found photographs and other illustrative materials
that we would not have found on our own. We remain indebted to art manager
Matt McAdams and the artists at Dragonfly Media for their excellent work in
expanding the illustration program and to Kate Scully and her team at North-
eastern Graphic for their talents in translating the manuscript onto the page.
Our manuscript editor, Martha Solonche, has contributed to the book’s
clarity and consistency and proofreader Kate Daly to its accuracy. And as in
the past, our gratitude to Barbara Brooks, our development editor, knows no
bounds. She has provided a strong guiding hand to our thinking and organiza-
tion and has done so with humor and a commitment to excellence that shows
its stamp all over the book. Thank you, Barbara, for reminding us that the book
is for students, not senior investigators, and thus requires us to write simply
and clearly, and for keeping us abreast of topical news items that we might
other­wise not encounter in our reading of the field’s diverse literature.
Once again, errors remain solely attributable to us. In a field that has ex-
panded so dramatically since our first edition, we hope that readers continue to
acquire a breadth of knowledge in the ever-expanding world of human neuro-
psychology. Finally, we thank our students, who have motivated us to continue
the journey of Fundamentals of Human Neuropsychology for nearly 40 years. See-
ing the faces of students light up when they begin to understand how the mar-
velous brain can produce cognition and behavior continues to be rewarding
and is what this endeavor is all about. Once again, we must thank our wives for
putting up with us when we were distracted by deadlines and may not always
have been our “usual” selves.

Bryan Kolb and Ian Q. Whishaw

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him who had been father, mother, friend, and companion to
her. She had indeed other friends, and a choice of homes
for the future, but none could ever fill his place in her heart.

Yet, in the worst of her woe, Annie could not but be

thankful. She could not but know how very tenderly her
Heavenly Father had dealt with them both. When she
thought of the terrible death which might have been his
portion, this most childlike falling "asleep in the Arms of
Jesus" did indeed seem only mercy.

Almost all Littleburgh went to the funeral. So great a

throng had never before been seen in the cemetery. Few
dry eyes could be perceived throughout the concourse of
people, and more than once a widespread burst of weeping
drowned the voice of the clergyman who read the service.

For Arthur Wilmot had given up his life for others; had
spent lavishly time and powers, money and strength, upon
those who needed help. No marvel that years of outpoured
sympathies should have brought a wealth of love in return.

Mrs. Stuart like others had been to the cemetery, and

like others she had wept there freely for the friend whose
face she could never see again on earth. Since returning
home, she had sat long in thought, giving vent to no
remarks.

Archie had noted his mother's gravity and absence of

mind since the day of Mr. Wilmot's death—not grimness,
which was usual, but a softened gravity which was unusual.
He noted too that her manner of speaking was gentler,
more humble, less stiff and self-asserting. He felt sure that
she was grieving deeply over the loss they had all
sustained.
 "It was a wonderful sight this afternoon, mother, wasn't
it?" he said, to break in upon a silence which lasted long.
Archie had been to the funeral as well as his mother: and so
had many scores of working-men, set free from work for
the purpose by their employers.

"Ay," she said, with a deep sigh.

"There wasn't a soul stayed away that could manage to

go," pursued Archie.

"Ay," repeated Mrs. Stuart. "'Twould have been a deal

more wonderful if they hadn't gone, after all he's been and
done."

"I'm afraid we'll never get another Rector like him,

mother."

Mrs. Stuart sighed again.

"And to think of his being bitten after all by that dog,—

and nobody knowing all this while what might come of it,"
said Archie; for the fact had become known in the town.
There was no longer any reason for concealing what had
occurred. It was well that the people should know to the full
the brave and self-sacrificing spirit of the man who so long
had lived among them. If any more were needed to make
them cling to the memory of their beloved Rector, that
"more" was now supplied.

He had not, indeed, in one sense, died from the results

of the bite or scratch; but in another sense, he perhaps
had, since without it, the break-down of his health might
have been long postponed. There could be no question as to
the tremendous and terrible peril which he had willingly
incurred for the sake of helpless little ones.
 "It isn't many would do what he did," said Mrs. Stuart.

"It isn't many would have thought of such a thing,

either," added Archie. "I'm sure I shouldn't. But I liked that
what the preacher said to-day, mother, about Mr. Wilmot
being willing to give up his life for the children, and about
God's pity being greater, and the Son of God giving up His
life for us. It seemed to bring the thought out so clear. I
don't know as I ever saw it so plain before. And I can
remember Mr. Wilmot telling us how pretty near all the
kindness and pity we do see round us was learnt from
Christ."

Mrs. Stuart said "Yes" thoughtfully. After a pause, she

added, "You put your life in danger too, Archie, for the sake
of Nancy Dunn."

Archie was very much astonished at such a remark from

his mother. He did not, however, show his astonishment,
but said only—"It wasn't for Nancy, mother. I didn't see her
to be Nancy till after. I hope I'd do all I could for any girl in
danger—though what I did wasn't what Mr. Wilmot did."

"But you'd sooner help Nancy than any other girl?" said
Mrs. Stuart.

"Yes—I would," said Archie, his face kindling. "Mother, I

love Nannie so much—I do think I could die for her."

"That's easy enough said," rejoined Mrs. Stuart. "It

wouldn't be so easy if you'd got to do it. But you're a brave
lad, I know. And as the preacher told us to-day—it's one
thing to be willing to die for one that loves us, and another
thing to die for them that hates us. There wouldn't be many
who'd do the last."
 "Only Christ," said Archie, wondering anew at his
mother's mood.

"Only—Christ," she repeated. "Yes—He did. I don't know

as I've hated Him, like some do—but I haven't cared. I
haven't thought about Him; and that's bad enough, after all
He's done. And I hope I shan't go on so—not any longer.
You needn't tell what I'm saying—not to anybody, Archie.
But it did seem as if I must say it to somebody. And Mr.
Wilmot 'll never come again. He'd have helped me—and
there's nobody now."

"Mother, there's One," said Archie shyly; "there's One

who'll help, if anybody asks Him—and He's best of all. Mr.
Wilmot would tell you to go to Him."

"But if He shouldn't want me?" she said.

"I shouldn't think there's any chance of that, mother,"

said Archie. "I should think, if He cared enough about you
to die for you, when you weren't even born—why, it isn't
likely He'd stop caring, and turn away, just the minute
you're beginning to want to know Him."

"No—I think you're about right," said Mrs. Stuart. And

she stood up slowly, left him alone, and went upstairs to her
bedroom for a while. There are times when one needs to be
all alone with God.

CHAPTER XXVII.
HOW THINGS CAME ABOUT.
 THE next evening Mrs. Stuart asked abruptly of her son
—

"Seen any of the Dunns to-day?"

"Just for a minute." Archie questioned with himself what

might be coming next.

"Nancy Dunn?"

"Yes, mother. I'd hardly a word with her, though."

"And you're hankering after her just as much as ever?"

"Mother, there isn't another girl like Nancy in all the

whole world! I give you my word for it," said Archie
earnestly.

Mrs. Stuart began to sob.

"It isn't my way to give in," she said. "I'm not one o'
those folks that's for ever chopping and changing. When I
says a thing, I mean it, and I keep to it. And I did say I'd
never have you marry Nancy Dunn. I didn't think her good
enough. Nor I don't. Leastways, her father isn't your
father's equal."

"Oh, bother!" broke out Archie.

"Yes, it's easy to say 'bother!'" she retorted, rather

disposed to flare up. She had been somewhat irritable all
day—no unusual state of things in the earlier stages of
trouble, or of any marked change in heart and life. "But if
you'd have a bit of patience and hear me out, you'd maybe
speak in a different sort of way."
 "Yes, mother," Archie said meekly, a sudden hope
springing up.

"I did say I wouldn't have it," repeated Mrs. Stuart;

"and I meant it too, and I'd have kept to my word. But he—
he came in to see me, just the very same day he died—he
did, Archie, that very afternoon. And I made him a cup o'
tea, and he said—he said—I was the best tea-maker—"

"Yes, yes, mother, I know," said Archie, as she broke

out crying. "You told me all that."

"And if I did," she retorted, "there was something else I

didn't tell you nor nobody, and didn't mean to neither, till I'd
made up my mind what to do. And I couldn't make it up till
yesterday, when I was standing in the crowd, and all of 'em
in black, and everybody crying round me, and Nancy Dunn
with her pretty face all blistered—for there's no denying
she's got a pretty face."

"Yes," assented Archie, with much warmth.

"It gave me a sort of a fellow-feeling with her, and I

won't deny it," said Mrs. Stuart. "And it sort of made me
think of his talking to me like that, about you and she, and
how he hoped she'd be a good wife to you one day."

"Did he?" cried Archie.

"Yes, he did," said Mrs. Stuart, heaving another sigh.

"And I won't deny as I was a bit grumpy, not thinking as it
was the last time I'd ever see him again. O dear, dear me!
To be sure! And the very last words he says, as he was
going out of the door—looking so bad, for all he was better
for the tea, as I'm sure I'd a foreboding in my mind it
couldn't be long—and the very last words he says to me
was how he wished I'd give in and let it be."
 "And you will, won't you?" begged Archie.

"I'm not one to give in easy," repeated Mrs. Stuart. "But

seeing it was, as one may say, his last wish, it do make a
difference. And I won't deny neither that I've maybe been
too stiff; and got to learn to be different. And understanding
you don't mean to think of marrying for many a year to
come—"

"Marrying! No," cried Archie. "Not yet, mother. Not till

I'm earning enough to keep you and she too in comfort. But
I do want to have her promise. I want to know she's safe to
be my wife by-and-by."

"Folks chop and change," said Mrs. Stuart. "I'd sooner

you'd keep yourself free. Maybe you'll get tired of it, or
she'll change her mind. But there, you're set on the girl,
and he wanted me to give in, and I'm not going to stand
out against you no longer. So you just go and see Nancy
Dunn, and say whatever you like, and have it settled."

Perhaps the consent might have been more graciously

worded; but Archie was far too glad to be critical. He felt
almost ashamed of his own gladness, at a time when
everybody else was sad; yet doubtless it was only natural,
and it did not prevent his sharing in the sorrow of others for
the loss they had all endured.

That same evening, Archie made his way to Woodbine

Cottage, and told the Dunns about his mother's newly-given
permission. Nannie looked hardly so pretty as usual, for her
face was still blistered with many tears, and they
threatened to flow fast again when she heard of Mr.
Wilmot's last call on Mrs. Stuart.

"It don't seem as if we'd ought to think about ourselves

yet," she faltered. But Archie could not take that view of the
question.

"I've waited ever so long, and mother's willing at last,"

he said. No doubt the few months that he had been
acquainted with Nancy did really seem long at his age. "You
won't put me off, will you, Nancy? Say she won't, Mrs.
Dunn. I do want to have it a settled thing. And you've as
good as told me already you'd say 'Yes' if my mother didn't
make difficulties. I don't mean that I'm thinking of marrying
yet awhile. I've got my mother to keep, you know. But by-
and-by—"

"It'll have to be a good long by-and-by, I shouldn't

wonder," observed Dunn. "Mind you, Archie, I'm willing to
have it all fair and straight between you and Nannie. I do
believe it'll be for her happiness and yours too, please God.
There was a time when I wouldn't have been so willing. But
I do believe it's your wish now to be a servant of God."

"Yes," Archie answered heartily. "And if I may have

Nannie, why, I hope she and I'll be able to serve God
together, Dunn."

"That's it, lad. If I didn't think so, I'd be loth enough to

give her to you. But look you here, there's something else
I've got to say. You've got your mother to keep, and it's
right you should do it. She cared for you, and you must
care for her."

"But I hope I'll soon be making enough to keep her and

a wife too," said Archie.

"Maybe," responded Dunn. "You're a capable young

fellow, and you're steady, and I hope you'll do well. Seems
no reason why you shouldn't. Only mind this, Archie, you
don't marry our Nannie, till you've got a right good sum laid
by in the Savings Bank against a rainy day. It is all very fine
to be making enough to live on in comfort, and to spend
every penny of what you've got, never giving a thought to
the future. And if illness comes, or an accident lays you by,
or trade grows slack and work fails, what's to be done then?
No, no, I'll never give in to that for Nannie. I've seen
enough of the misery of it for wives and children, let alone a
man himself. If you're bent on marrying her, you may do it;
but you'll marry with a good provision laid by; and, what's
more, you'll not squander it all away on a fine wedding, nor
on a lot of smart furniture."

Archie was well content to bind himself by these

conditions. Perhaps, like many young men, he thought them
just a little needless. In his young vigour, he could not yet
believe that bodily strength would ever fail him. But after
all, he knew that Dunn was right, and he knew that his
mother would put no difficulties in the way. Before winter,
he would be receiving the wages of a full-blown artisan, and
then, as she had often told him, would be the time to lay
by. Archie had not quite seen the necessity till now.

Laying by for the future must of course mean some

measure of self-denial. It meant this often in Archie's case.
What if it did? His love for Nannie would have been a very
poor sort of article, If it could not have endured the least
touch of self-denial for her sake.

As time went on, months following months till they grew

into years, while the amount down to Archie's name at the
Savings Bank grew also, Archie became very grateful for
the wise advice of Richard Dunn.

For, after all, though he and Nannie waited years for

their wedding day, they did not wait so long as must have
been the case, if Archie had not persistently from the first
put aside every little sum he could spare from present
needs. Sometimes it was a few shillings, sometimes only a
few pence. But "many a little makes a mickle," says the old
proverb. Archie proved the truth of this proverb.

Little more need be said, except that when the wedding

did take place, Annie Wilmot was present, besides many
other kind friends. The wedding cake was a present from
Annie, as well as a beautiful family Bible, bound in morocco.
Nannie made a sweet-looking young bride, in her neat
brown dress and bonnet; and even Nannie's own parents
scarcely saw her with more of fond pride than did Archie's
mother. For Mrs. Stuart had long ceased to regret the
thought of Nannie becoming Archie's wife; and on that
wedding day, she gained a "daughter indeed" to be the
comfort of her old age.

THE END.

Printed by BALLANTYNE, HANSON & CO.

Edinburgh & London

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