Sensation Perception THIRD EDITION Jeremy M. Wolfe Harvard University Medical School Keith R. Kluender Purdue University Dennis M. Levi University of California, Berkeley Linda M. Bartoshuk University of Florida Rachel S. Herz Brown University Roberta Klatzky Carnegie Mellon University Susan J. Lederman Queen s University Daniel M. Merfeld Harvard University Medical School Sinauer Associates, Inc. Publishers Sunderland, Massachusetts U.S.A.
Brief Contents CHAPTER 1 Introduction 3 CHAPTER 2 The First Steps in Vision: From Light to Neural Signals 31 CHAPTER 3 Spatial Vision: From Spots to Stripes 55 CHAPTER 4 Perceiving and Recognizing Objects 85 CHAPTER 5 The Perception of Color 117 CHAPTER 6 Space Perception and Binocular Vision 149 CHAPTER 7 Attention and Scene Perception 187 CHAPTER 8 Motion Perception 221 CHAPTER 9 Hearing: Physiology and Psychoacoustics 243 CHAPTER 10 Hearing in the Environment 275 CHAPTER 11 Music and Speech Perception 303 CHAPTER 12 Spatial Orientation and the Vestibular System 329 CHAPTER 13 Touch 363 CHAPTER 14 Olfaction 399 CHAPTER 15 Taste 433
Contents CHAPTER 1 Introduction 3 Welcome to Our World 3 Sensation and Perception 3 Method 1: Thresholds 4 Method 2: Scaling Measuring Private Experience 4 Method 3: Signal Detection Theory Measuring Difficult Decisions 4 Method 4: Sensory Neuroscience 4 Method 5: Neuroimaging An Image of the Mind 5 Thresholds and the Dawn of Psychophysics 5 Psychophysical Methods 7 Scaling Methods and Supertasters 9 Signal Detection Theory 11 Fourier Analysis 15 Sensory Neuroscience and the Biology of Perception 17 Neuronal Connections 21 Neural Firing: The Action Potential 22 Neural Firing: Populations of Neurons 24 CHAPTER 2 The First Steps in Vision: From Light to Neural Signals 31 A Little Light Physics 31 Eyes That See Light 32 Focusing Light onto the Retina 34 The Retina 35 Retinal Information Processing 38 Light Transduction by Rod and Cone Photoreceptors 38 The Man Who Could Not Read 42 Lateral Inhibition through Horizontal and Amacrine Cells 43 Convergence and Divergence of Information via Bipolar Cells 43 Communicating to the Brain via Ganglion Cells 44 Whistling in the Dark: Dark and Light Adaptation 48 The Man Who Could Not See Stars 50 CHAPTER 3 Spatial Vision: From Spots to Stripes 55 Visual Acuity: Oh Say, Can You See? 55 A Visit to the Eye Doctor 58 Acuity for Low-Contrast Stripes 59 Why Sine Wave Gratings? 61 Retinal Ganglion Cells and Stripes 61 The Lateral Geniculate Nucleus 62 Striate Cortex 64 Topography of the Human Cortex 66 Some Perceptual Consequences of Cortical Magnification 67 Receptive Fields in Striate Cortex 68 Orientation Selectivity 68 Other Receptive-Field Properties 69 Simple and Complex Cells 70 Further Complications 71
x CONTENTS Columns and Hypercolumns 72 Selective Adaptation: The Psychologist s Electrode 75 The Site of Selective Adaptation Effects 78 Spatial Frequency Tuned Pattern Analyzers in Human Vision 78 The Development of Spatial Vision 80 Development of the Contrast Sensitivity Function 81 The Girl Who Almost Couldn t See Stripes 82 CHAPTER 4 Perceiving and Recognizing Objects 85 What and Where Pathways 85 The Problems of Perceiving and Recognizing Objects 91 Middle Vision 93 Finding Edges 93 Texture Segmentation and Grouping 96 Perceptual Committees Revisited 99 Figure and Ground 101 Dealing with Occlusion 103 CHAPTER 5 The Perception of Color 117 Basic Principles of Color Perception 117 Three Steps to Color Perception 117 Step 1: Color Detection 118 Step 2: Color Discrimination 118 The Problem of Univariance 118 The Trichromatic Solution 120 Metamers 120 The History of Trichromatic Theory 122 A Brief Digression into Lights, Filters, and Finger Paints 123 From Retina to Brain: Repackaging the Information 124 Cone-Opponent Cells in the Retina and LGN 125 Step 3: Color Appearance 125 Three Numbers, Many Colors 125 Parts and Wholes 104 Summarizing Middle Vision 104 From Metaphor to Formal Model 105 Object Recognition 106 Templates versus Structural Descriptions 108 Problems with Structural-Description Theories 110 Multiple Recognition Committees? 111 Faces: An Illustrative Special Case 112 The Pathway Runs in Both Directions: Feedback and Reentrant Processing 113 Opponent Colors 128 Color in the Visual Cortex 131 Adaptation and Afterimages 133 Does Everyone See Colors the Same Way? 134 Does Everyone See Colors the Same Way? Yes 134 Does Everyone See Colors the Same Way? No 135 Does Everyone See Colors the Same Way? Maybe 136 From the Color of Lights to a World of Color 138 CHAPTER 6 Space Perception and Binocular Vision 149 Color Constancy 139 The Problem with the Illuminant 139 Physical Constraints Make Constancy Possible 141 Color Vision in Animals 143 Monocular Cues to Three-Dimensional Binocular Vision and Stereopsis 162 Space 152 Stereoscopes and Stereograms 166 Occlusion 152 Random Dot Stereograms 168 Size and Position Cues 153 Stereo Movies, TV, and Video Games 169 Aerial Perspective 157 Using Stereopsis 170 Linear Perspective 158 Stereoscopic Correspondence 171 Pictorial Depth Cues and Pictures 158 The Physiological Basis of Stereopsis 173 Motion Cues 160 Accommodation and Convergence 162
CONTENTS xi Combining Depth Cues 174 The Bayesian Approach Revisited 175 Illusions and the Construction of Space 176 Binocular Rivalry and Suppression 178 Development of Binocular Vision and Stereopsis 180 Abnormal Visual Experience Can Disrupt Binocular Vision 183 CHAPTER 7 Attention and Scene Perception 187 Selection in Space 189 The Spotlight of Attention 190 Visual Search 191 Feature Searches Are Efficient 192 Many Searches Are Inefficient 193 In Real-World Searches, Basic Features Guide Visual Search 194 In Real-World Searches, the Real World Guides Visual Search 194 The Binding Problem in Visual Search 196 Attending in Time: RSVP and the Attentional Blink 197 The Physiological Basis of Attention 200 Attention Could Enhance Neural Activity 200 Attention Could Enhance the Processing of a Specific Type of Stimulus 201 Attention and Single Cells 202 Disorders of Visual Attention 204 Neglect 204 Extinction 205 Balint Syndrome 206 Perceiving and Understanding Scenes 206 Two Pathways to Scene Perception 206 The Nonselective Pathway Computes Ensemble Statistics 207 The Nonselective Pathway Computes Scene Gist and Layout Very Quickly 208 Memory for Objects and Scenes Is Amazingly Good 210 Memory for Objects and Scenes Is Amazingly Bad: Change Blindness 210 What Do We Actually See? 214 CHAPTER 8 Motion Perception 221 Computation of Visual Motion 222 Apparent Motion 224 The Correspondence Problem 225 The Aperture Problem 226 Detection of Global Motion in Area MT 227 Motion Aftereffects Revisited 229 Second-Order Motion 230 Using Motion Information 231 Going with the Flow: Using Motion Information to Navigate 232 Something in the Way You Move: Using Motion Information to Identify Objects 233 Avoiding Imminent Collision: The Tao of Tau 234 Eye Movements 235 Physiology and Types of Eye Movements 236 Eye Movements and Reading 238 Saccadic Suppression and the Comparator 238 Development of Motion Perception 239 The Man Who Couldn t See Motion 240 CHAPTER 9 Hearing: Physiology and Psychoacoustics 243 The Function of Hearing 243 What Is Sound? 244 Basic Qualities of Sound Waves: Frequency and Amplitude 244 Sine Waves and Complex Sounds 246 Basic Structure of the Mammalian Auditory System 249 Outer Ear 249 Middle Ear 249 Inner Ear 250 The Auditory Nerve 256 Auditory Brain Structures 261 Basic Operating Characteristics of the Auditory System 264 Intensity and Loudness 264 Frequency and Pitch 266 Hearing Loss 268
xii CONTENTS CHAPTER 10 Hearing in the Environment 275 Sound Localization 275 Interaural Time Difference 276 Interaural Level Difference 279 Cones of Confusion 281 Pinna and Head Cues 282 Auditory Distance Perception 285 Complex Sounds 286 Harmonics 287 Timbre 288 CHAPTER 11 Music and Speech Perception 303 Music 303 Music and Emotion 303 Musical Notes 304 Making Music 307 Modalities and Qualities of Spatial Orientation 331 Sensing Angular Motion, Linear Motion, and Tilt 331 Basic Qualities of Spatial Orientation: Amplitude and Direction 332 The Mammalian Vestibular System 334 Hair Cells: Mechanical Transducers 334 Semicircular Canals 336 Otolith Organs 340 Spatial Orientation Perception 344 Rotation Perception 345 Translation Perception 346 Tilt Perception 347 Auditory Color Constancy 289 Attack and Decay 291 Auditory Scene Analysis 292 Spatial, Spectral, and Temporal Segregation 292 Grouping by Timbre 294 Grouping by Onset 296 When Sounds Become Familiar 297 Continuity and Restoration Effects 298 Restoration of Complex Sounds 299 Speech 309 Speech Production 310 Speech Perception 314 Learning to Listen 319 Speech in the Brain 323 CHAPTER 12 Spatial Orientation and the Vestibular System 329 CHAPTER 13 Touch 363 Touch Physiology 364 The Skin and Its Tactile Receptors 364 From Skin to Brain 369 Pain 373 Pleasant Touch 376 Tactile Sensitivity and Acuity 377 How Sensitive Are We to Mechanical Pressure? 377 How Finely Can We Resolve Spatial Details? 379 How Finely Can We Resolve Temporal Details? 380 Sensory Integration 348 Visual-Vestibular Integration 348 Reflexive Vestibular Responses 350 Vestibulo-Ocular Responses 350 Vestibulo-Autonomic Responses 354 Vestibulo-Spinal Responses 354 Spatial Orientation Cortex 357 Vestibular Thalamocortical Pathways 358 Cortical Influences 358 When the Vestibular System Goes Bad 359 Mal de Debarquement Syndrome 360 Ménière s Syndrome 360 Haptic Perception 380 Perception for Action 381 Action for Perception 381 Role of Fingerprints in Perception and Action 383 The What System of Touch: Perceiving Objects and Their Properties 384 The Where System of Touch: Locating Objects 387 Tactile Spatial Attention 390 Social Touch 391 Interactions between Touch and Other Modalities 392 Haptic Virtual Environments 394
CONTENTS xiii CHAPTER 14 Olfaction 399 Olfactory Physiology 399 Odors and Odorants 399 The Human Olfactory Apparatus 400 The Genetic Basis of Olfactory Receptors 406 The Feel of Scent 407 From Chemicals to Smells 407 Theories of Olfactory Perception 407 The Importance of Patterns 410 Odor Mixtures 411 The Power of Sniffing 413 Odor Imagery 413 Olfactory Psychophysics, Identification, and Adaptation 414 Detection, Discrimination, and Recognition 414 Psychophysical Methods for Detection and Discrimination 415 Identification 416 Adaptation 417 Cognitive Habituation 419 Olfactory Hedonics 420 Familiarity and Intensity 420 Nature or Nurture? 420 An Evolutionary Argument 422 Caveats 423 Olfaction, Memory, and Emotion 423 Neuroanatomical and Evolutionary Connections between Odor and Emotion 425 The Vomeronasal Organ, Human Pheromones, and Chemosignals 426 CHAPTER 15 Taste 433 Taste versus Flavor 433 Localizing Flavor Sensations 434 Anatomy and Physiology of the Gustatory System 435 Papillae 436 Taste Buds and Taste Receptor Cells 437 Taste Processing in the Central Nervous System 438 The Four Basic Tastes 439 Salty 439 Sour 440 Bitter 441 Sweet 442 The Survival Value of Taste 444 The Pleasures of Taste 445 Specific Hungers 446 The Special Case of Umami 447 The Special Case of Fat 448 Coding of Taste Quality 449 Taste Adaptation and Cross-Adaptation 450 Genetic Variation in Taste Experience 450 Supertasters 451 Health Consequences of Taste Sensation 453 Pleasure and Retronasal versus Orthonasal Olfaction 453 Chili Peppers 453 Glossary 457 References 473 Photo Credits 489 Index 491