Sensation & Perception 6th Edition

Jeremy M. Wolfe
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Sensation &
Perception
SIXTH EDITION
 Sensation &
Perception SIXTH EDITION

Jeremy M. Wolfe Linda M. Bartoshuk

Brigham & Women’s Hospital University of Florida
Harvard Medical School
Rachel S. Herz
Keith R. Kluender Brown University
Purdue University
Roberta L. Klatzky
Dennis M. Levi Carnegie Mellon University
University of California, Berkeley
Daniel M. Merfeld
The Ohio State University

SINAUER ASSOCIATES

NEW YORK OXFORD

OXFORD UNIVERSITY PRESS
 Sensation & Perception, Sixth Edition
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Not only did we get an allusion to vestibular
Figure 15.10 Data sources: Sucrose, glucose, sucralose (Y. Nie et al. Curr Biol 15: 1948-1952); Aspartame,
sensation, but I think you will agree, you can
neotame (H. Xu et al. Proc Natl Acad Sci 101: 14258-14263); Neoculin (A. Koizumi et al. Biochem Bio-
imagine the sights, sounds, smell, tastes, and
phys Res Commun 358: 585-589); Cyclamate (P. Jiang et al. J Biol Chem 280: 34296e34305 and H. Xu et
touch of the fair as well.
al. 2004. Proc Natl Acad Sci USA 101: 14258e14263); Neohesperidin (M. Winnig et al. 2007. BMC Struct
Biol 7: 66); Lactisole (P. Jiang et al. 2005. J Biol Chem 280: 15238e15246 and H. Xu et al. 2004. Proc Natl
Acad Sci USA 101: 14258e14263); Saccharin, inhibitor @ high conc (V. Galindo-Cuspinera et al. 2006.
Nature 441: 354); Brazzein (F. M. Assadi-Porter et al. 2010. J Mol Biol 398: 584-599 and P. Jiang et al.
2004. J Biol Chem 279: 45068e45075); S819 (S. K. Kim et al. 2017. Proc Natl Acad Sci 114: 2568-2573).

Library of Congress Cataloging-in-Publication Data

Names: Wolfe, Jeremy M., author. | Kluender, Keith R., author. | Levi, Dennis M., author.
Title: Sensation & perception / Jeremy M. Wolfe, Brigham & Women’s Hospital/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 L.
Klatzky, Carnegie Mellon University, Daniel M. Merfeld, Ohio State University.
Other titles: Sensation and perception
Description: Sixth Edition. | New York: Sinauer Associates: Oxford University Press, 2020. | Revised
edition of Sensation & perception, [2017] | Includes bibliographical references and index.
Identifiers: LCCN 2020018364 (print) | LCCN 2020018365 (ebook) | ISBN 9781605359724 (hardcov-
er) | ISBN 9780197542682 | ISBN 9780197542705 (epub)
Subjects: LCSH: Senses and sensation. | Perception. |
Classification: LCC QP431.S445 2020 (print) | LCC QP431 (ebook) | DDC 612.8--dc23
LC record available at https://lccn.loc.gov/2020018364
LC ebook record available at https://lccn.loc.gov/2020018365

987654321
Printed in the United States of America
Brief Contents
CHAPTER 1 Introduction 3
CHAPTER 2 The First Steps in Vision:
From Light to Neural Signals 33
CHAPTER 3 Spatial Vision: From Spots to Stripes 57
CHAPTER 4 Perceiving and Recognizing Objects 93
CHAPTER 5 The Perception of Color 131
CHAPTER 6 Space Perception and Binocular Vision 167
CHAPTER 7 Attention and Scene Perception 211
CHAPTER 8 Visual Motion Perception 249
CHAPTER 9 Hearing: Physiology and Psychoacoustics 273
CHAPTER 10 Hearing in the Environment 305
CHAPTER 11 Music and Speech Perception 335
CHAPTER 12 Vestibular Sensation 365
CHAPTER 13 Touch 407
CHAPTER 14 Olfaction 451
CHAPTER 15 Taste 495
About the Authors
JEREMY M. WOLFE is Professor of Ophthalmology and Radiology at Harvard Med-
ical School. Dr. Wolfe was trained as a vision researcher/experimental psychologist
and remains one today. His early work includes papers on binocular vision, adapta-
tion, and accommodation. The bulk of his recent work has dealt with visual search
and visual attention in the lab and in real-world settings such as airport security and
cancer screening. He taught Introductory Psychology for over twenty-five years at
the Massachusetts Institute of Technology, where he won the Baker Memorial Prize
for undergraduate teaching in 1989. He directs the Visual Attention Lab of Brigham
and Women’s Hospital

DENNIS M. LEVI is Professor of Optometry, Vision Science and Neuroscience at the

University of California, Berkeley. He served as Dean of the School of Optometry
from 2001–2014. In the lab, Dr. Levi and colleagues use psychophysics, computational
modeling, and brain imaging (fMRI) to study the neural mechanisms of normal
pattern vision in humans, and to learn how they are degraded by abnormal visual
experience (amblyopia).

KEITH R. KLUENDER is Professor of Speech, Language, and Hearing Sciences, and

Professor of Psychological Sciences at Purdue University. His research encompasses:
how people hear complex sounds such as speech; how experience shapes the way we
hear; how what we hear guides our actions and communication; clinical problems of
hearing impairment or language delay; and practical concerns about computer speech
recognition and hearing aid design. Dr. Kluender is deeply committed to teaching, and
has taught a wide array of courses—philosophical, psychological, and physiological.

LINDA M. BARTOSHUK is Bushnell Professor, Department of Food Science and

Human Nutrition at the University of Florida. Her research on taste has opened up
broad new avenues for further study, establishing the impact of both genetic and
pathological variation in taste on food preferences, diet, and health. She discovered
that taste normally inhibits other oral sensations such that damage to taste leads to
unexpected consequences like weight gain and intensified oral pain. Most recently,
working with colleagues in Horticulture, her group found that a considerable amount
of the sweetness in fruit is actually produced by interactions between taste and olfac-
tion in the brain. This may lead to a new way to reduce sugar in foods and beverages.
RACHEL S. HERZ is an Adjunct Assistant Professor in the Department of Psychiatry
and Human Behavior at Brown University’s Warren Alpert Medical School and Part-
time Faculty in the Department of Psychology and Neuroscience at Boston College.
Her research focuses on a number of facets of olfactory cognition and perception
and on emotion, memory, and motivated behavior. Using an experimental approach
grounded in evolutionary theory and incorporating both cognitive–behavioral and
neuropsychological techniques, Dr. Herz aims to understand how biological mecha-
nisms and cognitive processes interact to influence perception, cognition, and behavior.

ROBERTA L. KLATZKY is the Charles J. Queenan, Jr. University Professor of Psy-

chology at Carnegie Mellon University, where she also holds faculty appointments in
the Human–Computer Interaction Institute and the Neuroscience Institute. She has
done extensive research on haptic and visual object recognition, space perception and
spatial thinking, and perceptually guided action. Her work has application to haptic
interfaces, navigation aids for the blind, image-guided surgery, teleoperation, and
virtual environments.

DANIEL M. MERFELD is Professor of Otolaryngology-Head & Neck Surgery at the

Ohio State University College of Medicine and the Senior Vestibular Scientist at the
Naval Medical Research Unit Dayton. Much of his research career has been spent
studying how the brain combines information from multiple sources (i.e., multisen-
sory integration), with a specific focus on how the brain processes ambiguous sensory
information from the vestibular system in the presence of noise. Translational work
includes research developing: (a) new methods to help identify people more likely
to fall so that we can intervene before the fall, (b) new methods to diagnose patients
experiencing vestibular symptoms, and (c) vestibular implants for patients who have
severe problems with their vestibular labyrinth.
Contents
1 Introduction 3
1.1 S
 ensation & Perception:
Welcome to Our World 3
1.2 T
 hresholds and the
Dawn of Psychophysics 6
Psychophysical Methods 8
Scaling Methods 10
Signal Detection Theory 13
1.3 S
 ensory Neuroscience and
the Biology of Perception 16
Nerves and Specific Nerve Energies 17
2 The First Steps in Vision: From Light to Neural Signals
2.1 A Little Light Physics 33
2.2 Eyes That Capture Light 35
Focusing Light onto the Retina 37
The Retina 39
What the Doctor Saw 39
Retinal Geography and Function 41
2.3 Dark and Light Adaptation 44
Pupil Size 44
Photopigment Regeneration 45
The Duplex Retina 45
Neural Circuitry 46
•Sensation & Perception in Everyday Life
When Good Retina Goes Bad 46
Neuronal Connections 20
Neuronal Firing: The Action Potential 22
Neuroimaging 24
1.4 M
 odeling as a Method:
Math and Computation 28
Computational Models: Probability,
Statistics, and Networks 28
Deep Learning 30
Summary 31
33
2.4 Retinal Information Processing 47
Light Transduction by Rod and Cone
Photoreceptors 47
Lateral Inhibition through Horizontal
and Amacrine Cells 49
Convergence and Divergence of
Information via Bipolar Cells 49
Communicating to the Brain via
Ganglion Cells 50
•Scientists at Work
Is One Photon
Enough to See? 54
Summary 54
 CONTENTS   ix

3 Spatial Vision: From Spots to Stripes 57

3.1 V
 isual Acuity: Oh Say,
Can You See? 58
A Visit to the Eye Doctor 61
More Types of Visual Acuity 62
Acuity for Low-Contrast Stripes 63
Why Sine Wave Gratings? 65
3.2 R
 etinal Ganglion Cells
and Stripes 67
3.3 The Lateral Geniculate Nucleus 68
3.4 The Striate Cortex 70
The Topography of the Human Cortex 71
Some Perceptual Consequences
of Cortical Magnification 72
3.6 Columns and Hypercolumns 78
3.7 S
 elective Adaptation:
The Psychologist’s Electrode 81
The Site of Selective Adaptation
Effects 84
Spatial Frequency–Tuned Pattern
Analyzers in Human Vision 85
3.8 The Development of Vision 88
•Sensation & Perception in Everyday Life
The Girl Who Almost Couldn’t See
Stripes 89
Development of the Contrast Sensitivity
Function 90

3.5 R
 eceptive Fields in •Scientists at Work Does the Duck’s Left
Eye Know What the Right Eye Saw? 91
Striate Cortex 75
Orientation Selectivity 75 Summary 91
Other Receptive-Field Properties 76
Simple and Complex Cells 77
Further Complications 78

4 Perceiving and Recognizing Objects 93

4.1 F
 rom Simple Lines and Edges to Parts and Wholes 115
Properties of Objects 93 Summarizing Mid-Level Vision 115
• Scientists at Work Rüdiger von der Heydt, From Metaphor to Formal Model 116
Border Ownership, and Transparency 96
4.2 What and Where Pathways 97
•Sensation & Perception in Everyday Life
Material Perception: The Everyday Problem
of Knowing What It Is Made Of 117
4.3 T
 he Problems of Perceiving and
Recognizing Objects 100 4.5 Object Recognition 118
Can We Build It? 120
4.4 Mid-Level Vision 103
Multiple Recognition Committees? 124
Finding Edges 103
Faces: An Illustrative Special Case 125
Texture Segmentation and Grouping 107
Figure and Ground 111 Summary 127
Dealing with Occlusion 114
x  CONTENTS
5 The Perception of Color
5.1 B
 asic Principles of
Color Perception 131
Three Steps to Color Perception 132
5.2 Step 1: Color Detection 132
5.3 Step 2: Color Discrimination 133
The Principle of Univariance 133
The Trichromatic Solution 134
Metamers 135
The History of Trichromatic Theory 136
A Brief Digression into Lights, Filters,
and Finger Paints 137
From Retina to Brain: Repackaging
the Information 138
Cone-Opponent Cells in the Retina
and LGN 139
5.4 Step 3: Color Appearance 140
Three Numbers, Many Colors 140
•Sensation & Perception in Everyday Life
Picking Colors 141
6 Space Perception and Binocular Vision
6.1 M
 onocular Cues to
Three-Dimensional Space 170
Pictorial Depth Cues 171
Occlusion 171
Size and Position Cues 172
Aerial Perspective 175
Linear Perspective 176
Seeing Depth in Pictures 177
6.2 T
 riangulation Cues to
Three-Dimensional Space 179
Motion Cues 179
Accommodation and Convergence 181
6.3 Binocular Vision and Stereopsis 181
Stereoscopes and Stereograms 186
•Sensation & Perception in Everyday Life
Recovering Stereo Vision 188
Random Dot Stereograms 189
131
The Limits of the Rainbow 142
Opponent Colors 143
Color in the Visual Cortex 146
5.5 Individual Differences in
Color Perception 146
Language and Color 146
Genetic Differences in Color Vision 149
Does Everyone See the Same Colors?
The Special Case of Synesthesia 151
5.6 F
 rom the Color of Lights to
a World of Color 152
Adaptation and Afterimages 153
Color Constancy 155
The Problem with the Illuminant 156
Physical Constraints Make Constancy
Possible 157
5.7 What Is Color Vision Good For? 158
•Summary
Scientists at Work Filtering Colors 163
164
167
Using Stereopsis 190
Stereoscopic Correspondence 191
The Physiological Basis of Stereopsis
and Depth Perception 193
•Scientists at Work
a Hunting Insect
Stereopsis in
197
6.4 Combining Depth Cues 197
The Bayesian Approach Revisited 198
Illusions and the Construction of
Space 199
Binocular Rivalry and Suppression 202
6.5 D
 evelopment of Binocular
Vision and Stereopsis 204
Abnormal Visual Experience Can Disrupt
Binocular Vision 207
Summary 209

7 Attention and Scene Perception 211
7.1 Selection in Space 213
The “Spotlight” of Attention 215
7.2 Visual Search 216
Feature Searches Are Efficient 217
Many Searches Are Inefficient 217
•Scientists at Work How Would You Study
Visual Search by a Fish? 219
Guided Searches in the Real World 220
The Binding Problem in Visual Search 222
Binding between the Senses 224
7.3 A
 ttending in Time: RSVP and
the Attentional Blink 224
7.4 T
 he Physiological Basis of
Attention 226
Attention Could Enhance
Neural Activity 227
Attention Could Enhance the Processing of
a Specific Type of Stimulus 227
Attention and Single Cells 228
Attention May Change the Way Neurons
Talk to Each Other 230
8 Visual Motion Perception 249
8.1 Motion Aftereffects 249
8.2 Computation of Visual Motion 251
Apparent Motion 253
The Correspondence Problem: Viewing
through an Aperture 254
Detection of Global Motion in
Area MT 256
•Sensation & Perception in Everyday Life
The Man Who Couldn’t See Motion 258
Second-Order Motion 258
8.3 Using Motion Information 260
Going with the Flow: Using Motion
Information to Navigate 260
Avoiding Imminent Collision:
The Tao of Tau 261
Something in the Way You Move:
CONTENTS   xi
7.5 Disorders of Visual Attention 231
Neglect 231
Extinction 232
•Sensation & Perception in Everyday Life
Selective Attention and Attention Deficit
Hyperactivity Disorder (ADHD) 233
7.6 P
 erceiving and Understanding
Scenes 234
Two Pathways to Scene Perception 235
The Nonselective Pathway Computes
Ensemble Statistics 236
The Nonselective Pathway Computes Scene
Gist and Layout—Very Quickly 236
Memory for Objects and Scenes Is
Amazingly Good 238
But … Memory for Objects and Scenes
Can Be Amazingly Bad: Change
Blindness 241
What Do We Actually See? 242
Summary 245
Using Motion Information to Identify
Objects 262
Motion-Induced Blindness (MIB) 263
8.4 Eye Movements 263
Physiology and Types of Eye
Movements 265
Eye Movements and Reading 267
Saccadic Suppression and the
Comparator 267
Updating the Neural Mechanisms for
Eye Movement Compensation 269
8.5 D
 evelopment of Motion
Perception 270
• Scientists at Work Guess Who’s Coming
to Dinner 270
Summary 271
xii  CONTENTS

9 Hearing: Physiology and Psychoacoustics 273

9.1 The Function of Hearing 273 •Scientists at Work Why Don’t Manatees
Get Out of the Way When a Boat Is
9.2 What Is Sound? 274
Coming? 295
Basic Qualities of Sound Waves: Frequency
Frequency and Pitch 296
and Amplitude 274
Sine Waves and Complex Sounds 276 9.5 Hearing Loss 297
Types of Hearing Loss 297
9.3 B
 asic Structure of the Mammalian
Auditory System 277 Causes of Hearing Loss 298
Outer Ear 278 Treating Hearing Loss 299
Middle Ear 278 Using versus Detecting Sound 301
Inner Ear 280
The Auditory Nerve 286
•Sensation & Perception in Everyday Life
Electronic Ears 302
Auditory Brain Structures 291 Summary 303
9.4 B
 asic Operating Characteristics of
the Auditory System 293
Intensity and Loudness 293

10 Hearing in the Environment 305

10.1 Sound Localization 306
Interaural Time Difference 307
Interaural Level Difference 310
Cones of Confusion 311
Pinnae and Head Cues 312
10.3 Auditory Scene Analysis 323
Spatial, Spectral, and Temporal
Segregation 324
Grouping by Timbre 326
Grouping by Onset 327

•Scientists at Work Vulcan Ears 315

Auditory Distance Perception 316
Spatial Hearing and Blindness 318
•Sensation & Perception in Everyday Life
Sounds from Wind Farms 319
10.2 Complex Sounds 320
Harmonics 320
Timbre 321
Attack and Decay 322
When Hearing Dominates Vision 328
When Sounds Become Familiar 328
10.4 C
 ontinuity and
Restoration Effects 329
Restoration of Complex Sounds 330
10.5 Auditory Attention 331
Summary 333
 CONTENTS   xiii

11 Music and Speech Perception 335

11.1 Music 335 Speech Perception 348

•Sensation & Perception in Everyday Life

Music and Emotion 336
•Scientists at Work
Cochlea 353
Tickling the

Musical Notes 336 Learning to Listen 354

Making Music 340 Speech in the Brain 358

•Sensation & Perception in Everyday Life

Sonic Seasoning 343
Summary 362

11.2 Speech 343

Speech Production 344

12 Vestibular Sensation 365

12.1 Vestibular Contributions 367
12.2 E
 volutionary Development and
Vestibular Sensation 368
12.3 M
 odalities and Qualities of Spatial
Orientation 369
Sensing Angular Motion (“Rotation”), Linear
Motion (“Translation”), and Tilt 369
•Sensation & Perception in Everyday Life
The Vestibular System, Virtual Reality,
and Motion Sickness 370
Basic Qualities of Spatial Orientation:
Amplitude and Direction 370
12.4 The Vestibular Organs 374
Hair Cells: Mechanical Transducers 374
Semicircular Canals 376
Otolith Organs 382
12.5 S
 patial Orientation
Perception 384
Rotation Perception 385
Translation Perception 386
Tilt Perception 387
12.6 Multisensory Integration 388
Visual-Vestibular Multisensory
Integration 388
12.7 B
 eyond Multisensory Integration:
Active Sensing 390
12.8 Reflexive Vestibular Responses 391
Vestibulo-Ocular Responses 392
Vestibulo-Autonomic Responses 394
Vestibulo-Spinal Responses 396
12.9 M
 ultisensory Spatial Orientation
Cortex 399
Vestibular Thalamocortical Pathways 400
Cortical Influences 400
12.10 W
 hen the Vestibular System
Goes Bad 401
Falls and Vestibular Function 402
Mal de Debarquement Syndrome 402
•Scientists at WorkVestibular Aging 403
Ménière’s Syndrome 403
•Sensation & Perception in Everyday Life
Amusement Park Rides—Vestibular Physics
Is Fun 404
Summary 405
xiv  CONTENTS

13 Touch 407
13.1 Physical Inputs to Touch 408 13.3 Haptic Perception 434
Touch Physiology 408 Perception for Action 434
Touch Receptors and Neural Fibers 408 Action for Perception 435
From Skin to Brain 416 The What System of Touch: Perceiving
Pain 422 Objects and Their Properties 437

•Scientists at Work Tickling Rats 424

Neural Plasticity of Somatosensation 427
13.2 Tactile Sensitivity and Acuity 429
How Sensitive Are We to Mechanical
Pressure? 429
How Finely Can We Resolve Spatial
Details? 430
How Finely Can We Resolve Temporal
Details? 432
Do People Differ in Tactile Sensitivity? 432
The Where System of Touch:
Locating Objects 441
Tactile Spatial Attention 443
Social Touch 444
Interactions between Touch and
Other Modalities 445
•Sensation & Perception in Everyday Life
Tangible Phones and Tablets 447
Summary 448

14 Olfaction 451
14.1 Olfactory Physiology 452 Psychophysical Methods for Detection,
Odors and Odorants 452 Discrimination, and Recognition 473
The Human Olfactory Apparatus 452 Identification: Olfaction and Language 475
How Well Do We Smell? 455 Individual Differences 476

•Sensation & Perception in Everyday Life

Anosmia and Conditions that Affect
Olfactory Function 456
14.2 Neurophysiology of Olfaction 459
The Genetic Basis of Olfactory
Receptors 461
The “Feel” of Scent 463
14.3 From Chemicals to Smells 464
Theories of Olfactory Perception 464
The Importance of Patterns 467
Is Odor Perception Synthetic or
Analytical? 468
Nasal Power 471
Odor Imagery 471
14.4 O
 lfactory Psychophysics,
Identification, and Adaptation 472
Detection 472
Discrimination and Recognition 473
Adaptation 478
•Scientists at Work A New Test to Diagnose
Parkinson’s Disease 479
Cognitive Habituation and Odor
Consciousness 482
14.5 Olfactory Hedonics 483
Familiarity and Intensity 483
Nature or Nurture? 483
An Evolutionary Argument 485
Caveats 486
14.6 A
 ssociative Learning and Emotion:
Neuroanatomical and Evolutionary
Considerations 486
The Vomeronasal Organ, Human
Pheromones, and Chemosignals 488
•Sensation & Perception in Everyday Life
Odor-Evoked Memory and the Truth
behind Aromatherapy 490
Summary 492
 CONTENTS   xv

15 Taste 495
15.1 Taste versus Flavor 495 15.5 Genetic Variation in Bitter 511
Localizing Flavor Sensations: Supertasters 512
The Role of Taste 497 Health Consequences of Variation in
•Sensation & Perception in Everyday Life
Volatile-Enhanced Taste: A New Way to 15.6 H
Taste Sensations 513
 ow Do Taste and Flavor Contribute to
Sweeten Foods 498 the Regulation of Nutrients? 514
15.2 A
 natomy and Physiology of
the Gustatory System 499 •Scientists at Work The Role of Food
Preferences in Food Choices 514
Taste Myth: The Tongue Map 501 Taste 515
Taste Buds and Taste Receptor Cells 502 Flavor 516
Non-Oral Locations for Taste Is All Olfactory Affect Learned? 517
Receptors 502
15.7 The Nature of Taste Qualities 518
Taste Processing in the Central Nervous
Taste Adaptation and
System 503
Cross-Adaptation 519
15.3 The Four Basic Tastes? 504 The Pleasure of the Burn of Chili
Salty 505 Peppers 519
Sour 505 Summary 521
Bitter 506
Sweet 507
15.4 A
 re There More Than Four Basic
Tastes? Does It Matter? 509
Protein: The Umami Question 510
Fat 510

Glossary G-1
References R-1
Index I-1
Preface
This is an unusual time to be writing the Preface to the Sixth
Edition of our Sensation and Perception textbook. While
working on the Sixth Edition we find ourselves in the midst
of the COVID-19 shutdown. Because of the pandemic, we
are all locked out of our labs and classrooms, working from
home and interacting over the internet. Campuses have
been emptied, so virtually all of the students who are using
the Fifth Edition of our text are using it from home. Thus, it
feels like a bit of an act of faith to be writing to readers of the
sixth edition who, we may hope, are reading this in a world
that has returned to normal or to whatever the new normal
looks like going forward.
Why We Wrote this Book
So, why did we write this book and why did we revise it for a
Sixth Edition? The coronavirus actually provides an example
of the answer to the second question. It turns out that anos-
mia, an inability to smell, may be an important symptom of
the disease, sometimes the first symptom. Of course, much
of what we know about sensation and perception has been
known for many years. But we write new editions because the
science keeps changing and circumstances around science
keep changing.
We wrote the original version and continue to revise it
because we are fascinated by the human senses. We want to
know the answers to fundamental questions about the senses:
How does our brain create a three-dimensional perception
of the world from two-dimensional images, formed on the
back of each eye? Why do some substances taste “sweet?”
Why does music sound “musical?” In our own labs, we study
perceptual questions that arise from important problems in
the world. How do radiologists find cancer in X-rays? Why
is anosmia more disabling than you might think? Loss of a
sense of smell can have a major impact on diet and even on
social interactions.
We really love this material. We wrote this undergradu-
ate textbook in the hope that we might spread some of our
enthusiasm to you, our reader. In service of that goal, each of
the 15 chapters of this book aims to tell a coherent and inter-
esting story that will give the reader enough background and
exposure to current research to understand why these topics
are interesting and how they might be further investigated.
The author of each chapter is an expert in the topic who is
actively researching in the area. For every topic in the book,
we are acutely aware that there is vastly more information
than we can squeeze into a chapter. Moreover, we are not
naive or immodest enough to believe that you will devour a
chapter on “The Perception of Color” or “Perceiving and Rec-
ognizing Objects” in the way that you might devour a good
novel. However, we do hope that you will find each chapter
to be more than a compilation of facts. It is our hope that
this book teaches enough to inspire the reader to want to
know more. In service of these goals, each chapter includes
a Scientists at Work feature that gives a bit of detail about the
way that a specific topic has been studied as well as a section
on Sensation & Perception in Everyday Life that brings the
material out of the lab.
It is our intention to have produced a textbook that is
reasonably comprehensive while still being digestible. It is
possible that you, the student, may not think that a chapter
on motion perception, for example, is particularly digestible at
3:00 a.m. the day before the final exam, but that was the goal.
We want to present a coherent introduction to the important
topics in our field. As noted, we can’t cover everything. If you,
the instructor, or you, the interested student, think we missed
something that should be in here, please feel encouraged to
drop us an email.
New to this Edition
Each time we revise the book, we add some new topics and
we take some material out. Before we start writing, we trade
a few rounds of emails, asking ourselves if there are any
overarching topics that we should try to touch on in every
chapter. For this edition, one of the new points of emphasis
across chapters is multisensory integration.
Multisensory Integration
When you sit down to dinner, that is an experience with gus-
tatory (taste) and olfactory (smell) components, of course; but
it also involves vision, touch, and hearing (even if you are all
alone and hearing only the crunch of your carrot stick). All
those senses interact. If the carrot stick makes a squelchy
sound and puts up no resistance to your bite, your experience
of its ‘taste’ will be quite different. And if you are dizzy

(Chapter 12), well, the whole experience could be very differ-
ent. We need to write about hearing as hearing and olfaction
as olfaction, but, recognizing the multisensory nature of ex-
perience, we have tried to get out of our sensory silos and show
you some of the interactions between the senses. In the Sixth
Edition, these multisensory elements are called out in the text
with this symbol: Thus, if you look in Chapter 11, for ex-
ample, you will find a discussion of how music affects taste.
New material is not restricted to multisensory tidbits.
For instance, Chapter 1 has a new section on mathematical
and computational modeling in perception. Chapter 4 adds
recent findings on the “visual word form area” in the brain.
In Chapter 9, there is enhanced coverage of the functions of
the outer hair cells of the inner ear, along with a revised and
updated section on the treatment of hearing loss. In Chapter
10, there’s new material about how sound affects vision. Ves-
tibular sensation, the topic of Chapter 12, is a complicated
six-dimensional business. We have made renewed efforts
to provide that chapter with clear figures to illustrate how
we sense changes in the motion and position of the head
and body. Chapter 13 (Touch) adds new material on bionic
hands, pleasant touch, and the star-nosed mole (not all in
one paragraph). Chapters 14 (Olfaction) and 15 (Taste) have
always been more multisensory than most since when we
colloquially talk about the “taste” of a banana muffin, we are
talking about a sensory experience that involves the sensory
receptors in the nose and on the tongue. Smell is about more
than food and Chapter 14 includes new discussion of various
ways in which olfaction is involved in daily life. Chapter 15
takes on the multisensory perception of food and offers some
new ideas about what we mean when we talk about “taste”
and “flavor.” Maybe those are not synonyms.
Enhanced e-book
The enhanced e-book engages students in an interactive
environment with content such as integrated activities that
lead students through important processes, phenomena, and
structures, as well as self-assessment questions with imme-
diate feedback.
Accessible Content
Every opportunity has been taken to ensure that the content
herein is fully accessible to those who have difficulty perceiving
color. However, some of these Figures and activities will be
less accessible to some readers due to the intrinsic nature of
the colors and activities.
Acknowledgments
In trying to convey our enthusiasm for this material, we
wanted to create a beautiful book. If we have succeeded, it is
in no small part due to our publisher, Sinauer Associates, an
PREFACE   xvii
imprint of Oxford University Press. The people at Sinauer/
Oxford produce beautiful books and we have enjoyed working
with their talented editors to produce a book that strives to
be both aesthetically and intellectually appealing.
With the Sixth Edition, we welcome our new editor, Joan
Kalkut, who has successfully kept us on task and on time. We
also thank our previous editor, Sydney Carroll who oversaw
the last several editions and whose hand can still be seen in
the new edition. Lou Doucette and Carol Wigg have done ex-
cellent jobs as copy editors. They caught our errors, improved
our words, and made our text fit for public use. Mark Siddall
came up with an endless set of clever ideas for better images
and photos (check out Figure 7.33). Johannah Walkowicz co-
ordinated and oversaw the entire production process, as well
as developed figures, helped clarify prose, and corrected any
leftover inconsistencies or unclear writing. We also wish to
thank the entire production department at Sinauer, especially
Joan Gemme, who provided exceptional support and overall
project management, ensuring the highest quality and timely
production; Beth Roberge, who created an elegant book de-
sign and cover; both Beth and Michele Ruschhaupt created
beautiful page layouts; and Mike Demaray and Craig Durant
at Dragonfly Media Group created the stunning art program
of this text. Beyond the book, we would like to acknowledge
Suzanne Carter for managing and overseeing the digital proj-
ects. Many of these digital resources have been designed by
Evan Palmer of San Jose State University. He has created
and/or curated a host of great demonstrations of phenomena
discussed in the text. Moreover, as noted earlier, with all that
is new, we always need to prune out some old material in order
to keep the book from growing too long. That often makes us
sad because we are deleting good material. When we get too
sad, Evan tools the material into online essays.
Finally, it is always a question whether or not anyone ac-
tually reads the preface. If you did and you are reading this,
please send a note to jwolfe@bwh.harvard.edu, whether you
are a student or faculty. It is fun to hear from our “users.”
If you are reading this for a course, tell us who is teaching.
Odds are that one of us knows your instructor. More im-
portantly, please also feel encouraged to send us notes and
comments about the actual text. Hearing from readers is
an important way for us to make the book better. Thanks.
We are grateful to our colleagues who reviewed one or
more of the chapters of this book. It is extremely helpful to
have the wisdom of other experts in the field and of those
who use the text in classroom.
Nicole D. Anderson, MacEwan University
Benjamin Balas, North Dakota State University
Dirk Bernhardt-Walther, University of Toronto
Kent D. Bodily, Georgia Southern University
xviii  PREFACE
Amanda Carey, Simmons University
Shao-Ying I. Cheng, The University of Texas at Austin
Colin Ellard, University of Waterloo
Ahren Fitzroy, Mount Holyoke College
Alexis Green, Charleston Southern University
Alexis Grosofsky, Beloit College
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Thanasis Panorgias, New England College of Optometry
David Pittman, Wofford College
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T. C. Sim, Sam Houston State University
Jeffrey Stowell, Eastern Illinois University
D. Alexander Varakin, Eastern Kentucky University
Nicholas Watier, Brandon University
Mareike Wieth, Albion College
Meagan M. Wood, Valdosta State University
The following reviewers read and critiqued drafts and/or
previous versions of the text, and we are grateful for their
expert assistance:
Nicole D. Anderson, MacEwan University
Jeffrey Andre, James Madison University
Martin Arguin, University of Montreal
Simona Buetti, University of Illinois at Urbana-Champaign
Cheryl A. Camenzuli, Molloy College
Leslie Cameron, Carthage College
Linda C. Carson, University of Waterloo
Kathleen Cullen, McGill University
Thomas A. Daniel, Westfield State University
Nicolas Davidenko, University of California, Santa Cruz
Christopher DiMattina, Florida Gulf Coast University
Joshua Dobias, Rutgers, The State University of
New Jersey
Colin Ellard, University of Waterloo
Stephen Emrich, Brock University
Rhea Eskew, Northeastern University
Danielle Gagne, Alfred University
Carmela Gottesman, University of South Carolina,
Salkehatchie
Michael E. Hildebrand, Carleton University
Eric Jackson, University of New Mexico
Aaron Johnson, Concordia University
Ingrid S. Johnsrude, Western University
Jane Karwoski, University of Nevada, Las Vegas
Brock Kirwan, Brigham Young University
Roger Kreuz, University of Memphis
Leslie D. Kwakye, Oberlin College
Michael Landy, New York University
Michael Lantz, Concordia University at Loyola
Glenn Legault, Laurentian University
Max Levine, Siena College
Olga Lipatova, Christopher Newport University
Zili Liu, University of California, Los Angeles
Alejandro Lleras, University of Illinois at
Urbana-Champaign
Justin A. MacDonald, New Mexico State University
Kristen L. Macuga, Oregon State University
Janice C. McMurray, University of Nevada, Las Vegas
John Monahan, Central Michigan University
Richard Murray, York University
Gina O’Neal-Moffitt, Florida State University
Michael Owren, Emory University
Jennifer Peszka, Hendrix College
Robert Remez, Barnard College, Columbia University
Adrián Rodríguez-Contreras, The City College of
New York
Lisa Sanders, University of Massachusetts, Amherst
Eriko Self, California State University, Fullerton
Kevin Seybold, Grove City College

Steve Shevell, University of Illinois at Chicago
Rachel Shoup, California State University, East Bay
T. C. Sim, Sam Houston State University
Joel Snyder, University of Nevada, Las Vegas
Miriam Spering, University of British Columbia
Kenneth Steele, Appalachian State University
Julia Strand, Carleton College
William Stine, University of New Hampshire
Greg Stone, Arizona State University
Duje Tadin, University of Rochester
Jeroen van Boxtel, University of California, Los Angeles
Rachel Walker, Charleston Southern University
Dirk B. Walther, University of Toronto
Scott N. J. Watamaniuk, Wright State University
Laurie Wilcox, York University
PREFACE   xix
Many, many colleagues have sent us reprints and answered
questions about points both specific and general. We grate-
fully acknowledge their help even if we cannot list all of their
names (and even if we may still have failed to get things exactly
right). We are also indebted to the users of the text, students,
and faculty who pointed out errors, typos, and other short-
comings in the first five editions. We hope we caught them
all and we hope that the readers of this edition will continue
to offer us assistance. As noted earlier in the preface, if you
find a flaw or if you have any other comment—even a posi-
tive one—please feel encouraged to let us know. You can use
jwolfe@bwh.harvard.edu as a point of contact for all of us.
Digital Resources for
Sensation & Perception, Sixth Edition
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DIGITAL RESOURCES   xxi
Ideal for self-study, the Sensation & Perception, Sixth Edi-
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Activities and Essays
Throughout the textbook, you will see references to interactive resources in red text. These refer to specific activi-
ties and essays that are relevant to the topic being discussed. Below is the full list of activities and essays, by chapter.

CHAPTER ACTIVITY ACTIVITY ESSAY

1.1 Psychophysics 1.3 Neurons 1.1 Senses of Reality Through the Ages
1 1.2 Sensory Areas in the Brain
2.1 Visual System Overview 2.5 Retinal Structure 2.1 How Many Quanta Does It Take?
2.2 From Sun to Eye 2.6 Phototransduction 2.2 Clinical Case: The Man Who
2 2.3 Eye Structure 2.7 Acuity versus Sensitivity Couldn’t Read
2.4 Simulated Scotoma 2.8 Ganglion Receptive Fields 2.3 Seeing Illusory Stripes and Spots

3.1 Visual Angle 3.4 Fourier Analysis 3.1 Hyperacuity

3 3.2 Foveal Acuity
3.3 Gabor Patches
4.1 Object Substitution Masking
4.2 Gestalt Grouping Principles
4 4.3 Object Ambiguity
4.4 Infant Object Perception
5.1 The Principle of Univariance
5.2 Trichromacy
5 5.3 Color Mixing
6.1 Monocular Depth Cues
6.2 Binocular Disparity
6 Correspondence
7.1 Attentional Cueing
7 7.2 Visual Search
7.3 The RSVP Paradigm
8.1 Motion Aftereffects
8.2 Motion Detection Circuit
8 8.3 Types of Motion
3.5 Striate Receptive Fields
3.6 Hypercolumns
4.5 Pandemonium
4.6 Viewpoint Effects
4.7 The Face Inversion Effect
5.4 Afterimages
5.5 Color Constancy
5.6 Illusions of Lighting
6.3 Stereoscopes and
Stereograms
6.4 Stereoscopic
7.4 The Attentional Blink and
Repetition Blindness
7.5 Change Blindness
7.6 The Attentional Bottleneck
8.4 Motion Correspondence
8.5 Eye Movements
3.2 The Whole Brain Atlas
3.3 Seeing Images on the Cortex
4.1 The Role of Knowledge in
Figure-Ground Assignment
4.2 Dynamic Occlusion
4.3 Bayesian Analysis
4.4 Face Blindness
5.1 More About Opponent Processing
in Color Vision
5.2 The Philosophical Problem of
“Inverted Qualia”
5.3 Experiencing Color Blindness
5.4 Color Constancy in the Lab
6.1 Making the Implicit Explicit
6.2 Stereo Images on the Web
6.3 Stereo Movies, TV, and Video Games
6.4 Stereoscopic Mammograms
in Breast Cancer
6.5 Stereopsis in the Cuttlefish
6.6 The Moon Illusion
7.1 Balint Syndrome
7.2 Boundary Extension
7.3 Attentional Capture
8.1 Perceiving Motion in Static Images
8.2 Motion Illusions
8.3 Beyond Second-Order Motion
8.4 Biological Motion
 ACTIVITES AND ESSAYS   xxiii

CHAPTER ACTIVITY ACTIVITY ESSAY

9.1 What We Hear
9 9.2 Structure of the Auditory System
10.1 Auditory Localization Cues
10 10.2 The Missing-Fundamental
Effect
10.3 Timbre
11.1 Notes, Chords, and Octaves
11 11.2 Categorical Perception
12.1 A Guided Tour of the Vestibular
System
12 12.2 Sinusoidal Motion
13.1 The Need for Touch
13.2 Somatosensory Receptors
13 13.3 The Sensory Homunculus
14.1 Olfactory Anatomy
14 14.2 Odor Adaptation and
Habituation
15.1 Taste without Smell
15
9.3 Equal-Loudness Curves
10.4 Auditory Stream
Segregation
10.5 Continuity and Restoration
Effects
11.3 Word Breaks
11.4 The McGurk Effect
12.3 Observing Torsional Eye
Movement
13.4 The Rubber Hand Illusion
13.5 Two-Point Touch Thresholds
13.6 Haptic Object Recognition
14.3 Sensory Memory Cues
15.2 Gustatory Anatomy
9.1 Outer Hair Cells and Electromotility
10.1 Reverberations and
the Precedence Effect
11.1 Studying Brain Areas for Language
Processing
12.1 Gravity versus Linear Acceleration
12.2 Canal-Otolith Multisensory
Integration
12.3 Space Motion Sickness
13.1 Living without Kinesthesis
13.2 Body Image
13.3 Phantom Limbs
13.4 Lego Blocks Front and Back
14.1 Smell-O-Vision
14.2 Olfactory Lateralization
14.3 Verbal-Olfactory Interactions
15.1 Water Tastes
Chapter
1

© Oleg Shupliak art

Oleg Shupliak, Self-Portrait Under the Lime Trees, 2011

Introduction
Questions to Contemplate
Think about the following questions as you read this chapter.
By the chapter’s end, you should be able to answer and discuss them.
● How can scientists study something as personal as your internal
sensations and perceptions?
● Are there laws that relate the physics of the world to your
subjective experience?
● What is happening when you think that you might have heard
something or felt something but you’re not sure? If a stimulus
is detectable, why isn’t it detectable all the time?
● Are there rules that relate the activity of your brain to your
subjective experience?
● How do sensation and perception change over the life span?

Y
ou’ve taken the plunge to read at least part of a textbook on “sensation
and perception.” You may be majoring in psychology or studying an allied
field, such as neuroscience or biology, or you may be simply curious. No
matter what interests you most, your understanding will be based on sensation
and perception.
“Why?” you ask. Most everything you know or think that you know about
the world around you depends on how you sense and how you perceive. These
foundational experiences began even before you were born. Your senses help you
to keep upright, stay warm or cool, avoid pain and poisonous things, and avoid
danger. Your experiences of the rich tapestry of life through movement, touch,
smell, taste, hearing, and vision inform most everything that you believe to be true.
It is small wonder that the questions posed in this textbook have been front and
center for big thinkers since the first written words, and probably earlier. Today, a
small army of researchers continue to pursue answers. This first chapter provides
an introduction to the sorts of questions that captivate the authors of this book and
the sorts of methods that researchers have developed to answer those questions.
These are only examples from the endless list of possibilities. The rest of the book
will introduce you to the vast array of questions that must occupy the attention of
anyone who really wants to know how we know what we think we know.

1.1 Sensation & Perception: Welcome to Our World

What does your phone feel as you run your finger down its touch screen (FIGURE
1.1)? What does it hear as you whisper into its receiver? We assume that these are
silly questions, though it would not be silly to ask about the lightest touch that the
phone’s screen could sense, or the faintest vibration in the air that its microphone
could sense. What does your cat feel as you run your finger down its back? That seems
a more reasonable question, though you have no access to the private experience of
 4  Chapter 1
Courtesy of m.h.siddall
FIGURE 1.1 Cell phone senses?
Would it make sense to ask what a cell
phone feels when you stroke its screen?
sensation The ability to detect a
stimulus and, perhaps, to turn that
detection into a private experience.
perception The act of giving
meaning to a detected sensation.
qualia In reference to philosophy,
private conscious experiences of
sensation or perception.
© Historical image collection by Bildagentur-online 2/Alamy Stock Photo
FIGURE 1.2 Étienne Bonnot de
Condillac French philosopher who
imagined the mental life of a statue
that had only a sense of smell.
the cat. You don’t even have access to the private sensations of a person whose back
you might stroke. Your own sensory experience is directly accessible only to you.
This book is titled Sensation & Perception. The ability to detect the pressure
of a finger and, perhaps, to turn that detection into a private experience is an
example of sensation. Perception can be thought of as the act of giving meaning
and/or purpose to those detected sensations. How do you understand the finger
that runs down your back? Is it a gesture of affection? Is it an officer at an airport
security checkpoint looking for contraband? This book will trace the path from
stimuli in the world, through your sense organs, to the understanding of the
world that you perceive.
Everything we feel, think, and do depends on sensations and perceptions. For
this reason, philosophers have thought, talked, and written about the topic in
profound and systematic ways for over two millennia. (See Essay 1.1: Senses of
Reality Through the Ages.) The idea that mental life depends on sensation and
perception has deep roots. The eighteenth-century French philosopher Étienne
Bonnot de Condillac (1715–1780) (FIGURE 1.2) famously asked his readers to
imagine the mental life of a statue with only a sense of smell. If we gave the statue
a whiff of the smell of a rose, Condillac said that the statue would be nothing but
that scent. With no other sense, it would have no idea of its own body, its place in
the world, or the existence of other objects. It would be the scent of rose. Period. If
more senses and more experience were provided to the statue, Condillac imagined,
a real mental life would develop.
If our mental life depends on information from our senses, then it follows that
the place for the study of the senses is within the science of human behavior and
human mental life—that is, within psychology. Of course, psychologists do not
have the topic entirely to themselves. Researchers studying topics in sensation and
perception can be found in biology, computer science, medicine, neuroscience,
and many other fields. Indeed, the authors of this book come from academic de-
partments of ophthalmology, radiology, optometry, speech, language and hearing
sciences, and community dentistry in addition to psychology. Critically, however,
we approach the study of sensation and perception as a scientific pursuit. As such,
it needs scientific methods. That’s why the next sections of this chapter are devoted
to an array of methods used in the study of the senses.
method 1: thresholds What is the faintest sound you can hear? How would
you know? What is the loudest sound you can hear? This last question is not as
stupid as it may sound, though it could be rephrased like this: “What is the loudest
sound you can hear safely or without pain?” If you listened to sounds above that
limit, perhaps by blasting your music too enthusiastically, you would change the
answer to the first question. You would have damaged your auditory system and
be unable to hear the faintest sound that you used to be able to hear. Your thresh-
old would have changed (for the worse). How would you measure that threshold?
As we’ll learn in this chapter, a variety of methods are available for measuring just
how sensitive your senses are.
method 2: scaling—measuring private experience When you say that
you “hear” or “taste” something, are those experiences—what the philosophers call
qualia (singular quale)—the same as the experiences of the person you’re talking
to? We can’t really answer the question of whether your qualitative experience of
“red” is like my qualitative experience of “green” or, for that matter, “middle C.” We
still have no direct way to experience someone else’s experiences. However, we
 Introduction  5

can demonstrate that different people do, in some cases, inhabit different sensory
worlds. Our discussion in this chapter will show how scaling methods can be used
to perform this act of mind reading.

FURTHER DISCUSSION of qualia can be found in Section 5.5.

method 3: signal detection theory—measuring difficult decisions

A radiologist looks at a mammogram, the X-ray test used to screen for breast
cancer. There’s something on the X-ray that might be a sign of cancer, but it is
not perfectly clear. What should the radiologist do? Suppose she decides to call it
benign, not cancerous, and suppose she is wrong. Her patient might die. Suppose
she decides to treat it as a sign of malignancy. Her patient will need more tests,
perhaps involving surgery. The patient and her family will be terribly worried. If
the radiologist is wrong and the spot on the mammogram is, in fact, benign, the
consequences may be less dire than those of missing a cancer, but there will be
consequences. This is a perceptual decision, made by an expert, that has real con-
sequences. Our discussion of signal detection theory will show how decisions of
this sort can be studied scientifically.

method 4: sensory neuroscience Grilled peppers appear on your table

as an appetizer. They have an appealing, smoky smell. When you bite into one,
it has a complex flavor that includes some of that smokiness. Fairly quickly you
also experience a burning sensation. There is no actual change in the tempera-
ture in your mouth, and your tongue is no warmer than it was, but the “burn”
is unmistakable. How does the pepper fool your nervous system into thinking
that your tongue is on fire? This chapter’s exploration of sensory neuroscience
will introduce the ways in which sensory receptors and nerves undergird your
perceptual experience.

method 5: neuroimaging—an image of the mind Suppose you ar-

range to view completely different images with different eyes. We might present a
picture of a house to one eye and of a face to the other (Tong et al., 1998). The result
would be an interesting effect known as binocular rivalry (see Section 6.4). The two
images would compete to dominate your perception: sometimes you would see a
house, and sometimes you would see a face. You would not see the two together.
One reason binocular rivalry is interesting is that it represents a dissociation of the
stimuli, presented to the eyes, and your private perceptual experience. Even if we
cannot share the experience, modern brain-imaging techniques enable us to see
traces of that experience as it takes place in the brain. Methods of neuroimaging
will be our final methodological topic in this chapter.

method 6: computational models If you’re your first language is En-

glish, Spanish ‘b’ and ‘p’ might both sound like English ‘b’ to your ears, and French
and German vowels are difficult to tell apart. Your ability to distinguish speech
sounds depends almost entirely on the kinds of speech sounds that you heard while
growing up. This is because, for all of your senses, perception is a combination
of things one is born with and things that are acquired only through experience.
Researchers might create a computational model to describe precisely how the
basic abilities that every infant possesses at birth become shaped by experience in
a particular language environment to predict the way you perceive speech sounds.
6  Chapter 1
FIGURE 1.3 Founder of experimental
psychology Thought by some to be the
true founder of experimental psychology,
Gustav Fechner is best known for his pio-
neering work relating changes in the phys-
ical world to changes in our psychological
experiences, thus inventing the field of
psychophysics.
dualism The idea that the mind has
an existence separate from the material
world of the body.
materialism The idea that the only
thing that exists is matter, and that all
things, including the mind and con-
sciousness, are the results of interaction
between bits of matter.
panpsychism The idea that the mind
exists as a property of all matter—that
is, that all matter has consciousness.
psychophysics The science of defin-
ing quantitative relationships between
physical and psychological (subjective)
events.
two-point touch threshold The mini-
mum distance at which two stimuli (e.g.,
two simultaneous touches) are just per-
ceptible as separate.
1.2 Thresholds and the Dawn of Psychophysics
Early on, study of the senses was a mix of experimental science and philosophy.
Fascinating work can be found in ancient Greek philosophy, in medieval Islamic
science, and in the writings of sages in China and India. We will start much later,
with the very interesting and versatile nineteenth-century German scientist-phi-
losopher Gustav Fechner (1801–1887) (FIGURE 1.3).
Before making his first contributions to psychology, Fechner had an eventful
personal history. Young Fechner earned his degree in medicine, but his interests
turned from biological science to physics and mathematics. By 1833, he was a full
professor of physics in Leipzig, Germany. Though this might seem an unlikely
way to get to psychology, events proved otherwise. He became absorbed with the
relationship between mind and matter. This pursuit placed him in the middle of a
classic philosophical debate between adherents of dualism and materialism. Dualists
hold that the mind has an existence separate from the material world of the body.
Materialists hold that the mind is not separate. A modern materialist position,
probably the majority view in scientific psychology, is that the mind is what the
brain does. Fechner proposed to effectively split the difference by imagining that
the mind, or consciousness, is present in all of nature. This panpsychism—the idea
that the mind exists as a property of all matter—extended not only to animals, but
to inanimate things as well. Fechner described his philosophy of panpsychism in
a provocative book (first published, in German, in 1848) entitled Nanna, or Con-
cerning the Mental Life of Plants. The title alone gives a pretty good idea of what
Fechner had in mind.
As a young scientist, Fechner overworked himself to exhaustion and severely
damaged his eyes by gazing at the sun while performing vision experiments (a
not-uncommon problem for curious vision researchers in the days before reliable,
bright, artificial light sources). The visually incapacitated Fechner had some form
of mental breakdown that left him sometimes unable to speak or eat. He resigned
his university position, withdrew from almost all his friends and colleagues, and
for 3 years spent almost all of his time alone with his thoughts.
Fechner apparently solved his eating problem with a diet of “fruit, strongly
spiced ham and wine” (Fancher, 1990, p. 133). His vision was also recovering. Then,
while lying in bed on October 22, 1850 (a date still celebrated as “Fechner Day” by
some), Fechner had a specific insight into the relationship between mental life and
the physical world. From his experience as a physicist, Fechner thought it should
be possible to describe the relation between mind and body using mathematics.
His goal was to formally describe the relationship between sensation (mind) and
the energy (matter) that gave rise to that sensation. He called both his methods and
his theory psychophysics (psycho for “mind,” and physics for “matter”).
In his effort, Fechner was inspired by the findings of one of his German col-
leagues, Ernst Weber (1795–1878) (FIGURE 1.4), an anatomist and physiologist
who was interested in touch. Weber tested the accuracy of our sense of touch by
using a device much like the compass one might use to draw circles. He used this
device to measure the smallest distance between two points that was required for a
person to feel touch on two points of the skin instead of one. Later, Fechner would
call this distance the two-point touch threshold. We will discuss two-point touch
thresholds, and touch in general, in Chapter 13.
For Fechner, Weber’s most important findings involved judgments of lifted
weights. Weber would ask people to lift one standard weight (that is, a weight that
stayed the same over a series of experimental trials) and one comparison weight that
differed from the standard. Weber increased the comparison weight in incremental
amounts over the series of trials. He found that the ability of a person to detect
 Introduction  7

the difference between the standard and comparison weights depended greatly on
the weight of the standard. When the standard was relatively light, people were
much better at detecting a small difference when they lifted a comparison weight.
When the standard was heavier, people needed a greater difference before they
could detect a change. He called the difference required for detecting a change in
weight the just noticeable difference, or JND. Another term for JND, the smallest
change in a stimulus that can be detected, is the difference threshold.
Weber noticed that JNDs change in a systematic way. The smallest change in
weight that could be detected was always close to 1/40 of the standard weight. Thus,
a 1-gram change could be detected when the standard weighed 40 grams, but a
10-gram change was required when the standard weighed 400 grams. Weber went
on to test JNDs for a few other kinds of stimuli, such as the lengths of two lines (for
which the detectable change ratio was 1:100). For virtually every measure—whether
brightness, pitch, or time—a constant ratio between the change and the standard
could describe the threshold of detectable change quite well. This ratio rule holds
true except for extreme stimuli—stimuli so small or large that they approach the
minimum or maximum of our senses. In recognition of Weber’s discovery, Fechner
called these ratios, or proportions, Weber fractions, and he called the mathematical FIGURE 1.4 Ernst Weber Weber
discovered that the smallest detectable
formula that described the general rule Weber’s law. Weber’s law states that the change in a stimulus, such as the weight
size of the just detectable difference (ΔI) is a constant proportion (K) of the level of an object, is a constant proportion of
of the stimulus (I). the stimulus level. This relationship later
In Weber’s observations, Fechner found what he was looking for: a way to became known as Weber’s law.

describe the relationship between mind and matter. Fechner assumed that the
smallest detectable change in a stimulus (ΔI) could be considered a unit of the mind
because this is the smallest bit of change that is perceived. He then mathematically
extended Weber’s law to create what became known as Fechner’s law (FIGURE 1.5): just noticeable difference (JND) or
difference threshold The smallest
S = k log R detectable difference between two
stimuli, or the minimum change in a
where S is the psychological sensation, which is equal to the logarithm of the phys- stimulus that enables it to be correctly
ical stimulus level (log R) multiplied by a constant, k. This equation describes the judged as different from a reference
fact that our psychological experience of the intensity of light, sound, smell, taste, stimulus.
or touch increases less quickly than the actual physical stimulus increases. With Weber fraction The constant of
this equation, Fechner provided us with at least one way to relate mind and matter. proportionality in Weber’s law.
Weber’s law The principle describing
the relationship between stimulus and
resulting sensation that says the just
noticeable difference (JND) is a constant
fraction of the comparison stimulus.
Fechner’s law A principle describing
the relationship between stimulus and
resulting sensation that says the magni-
tude of subjective sensation increases
Sensation intensity (S)

proportionally to the logarithm of the

stimulus intensity.

FIGURE 1.5 Fechner’s law As the intensity of a physical

Physical stimulus intensity (R)
stimulus increases (x-axis), a larger change in that physical
stimulus is required to produce a just detectable difference
in sensation (y-axis). This is seen graphically in the increasing
amount of space between the dashed lines on the x-axis is
required to produce evenly spaced dashed lines on the y-axis.
The relationship between X and Y values is logarithmic.
8  Chapter 1
absolute threshold The minimum
amount of stimulation necessary for
a person to detect a stimulus 50% of
the time.
method of constant stimuli A psycho-
physical method in which many stimuli,
ranging from rarely to almost always
perceivable (or rarely to almost always
perceivably different from a reference
stimulus), are presented one at a time.
Participants respond to each presenta-
tion: “yes/no,” “same/different,”
and so on.
● TABLE 1.1 Absolute thresholds in the real world
Sense Threshold
Vision Stars at night, or a candle flame 30 miles away on a dark, clear night
Hearing A ticking watch 20 feet away, with no other noises
Vestibular A tilt of less than half a minute on a clock face
Taste A teaspoon of sugar in 2 gallons of water
Smell A drop of perfume in 3 rooms
Touch The wing of a fly falling on your cheek from a height of 3 inches
Source: From E. Galanter. 1962. In New directions in psychology. T. Newcomb et al. (Eds.), Holt,
Rinehart and Winston: New York.
Even if mind and matter are related, we take care to distinguish between units
of physical entities (such as light or sound) and measures of people’s perception
(“brightness,” “loudness”). For example, the physical intensity of a sound—the
sound pressure level—is a physical entity we can measure in decibels, whereas a
person’s perception of “loudness” is psychophysical and subjective (see Section 9.2).
Similarly, frequency is a measure of the rate of fluctuations of the physical sound
pressure, while the “pitch” of a musical note describes a psychophysical response to
that physical phenomenon. Frequency and pitch are not the same thing, although
they are closely correlated. Over a wide range, as frequency increases, so does pitch,
though it is unclear whether there is a perception of pitch for the highest audible
frequencies (Green, 2005).
Fechner was the first to objectively measure psychological events through new
ways to measure what people see, hear, and feel (Wixted, 2020). As such he can
be considered to be the true founder of experimental psychology (Boring, 1950),
even if that title is usually given to Wilhelm Wundt (1832–1920), who began his
work sometime later. All of Fechner’s methods are still in use today. In explaining
these methods here, we will use absolute threshold as an example because it is
perhaps the most straightforward; but we would use the same methods to determine
difference thresholds such as ΔI. An absolute threshold is the minimum intensity
of a stimulus that can be detected (TABLE 1.1). This returns us to the question we
raised earlier: What is the faintest sound you can hear? Of course, we can ask the
same question about the faintest light, the lightest touch, and so forth. (See Activity
1.1: Psychophysics.)
Psychophysical Methods
How can we measure an absolute threshold in a valid and reliable manner? One
method, known as the method of constant stimuli, requires creating many stimuli
with different intensities in order to find the tiniest intensity that can be detected
(FIGURE 1.6). If you’ve had a hearing test, you had to report when you could and
could not hear a tone that the audiologist played to you over headphones, usually
in a very quiet room. In this test, intensities of all of the tones were relatively low,
not too far above or below the intensity where your threshold was expected to be.
The tones, varying in intensity, were presented randomly, and tones were presented
multiple times at each intensity.
The “multiple times” piece is important. Subtle perceptual judgments such as
threshold judgments are variable. The stimulus varies for physical reasons. The
observer varies. Attention waivers and sensory systems fluctuate for all sorts of
reasons. As a consequence, one measure is almost never enough. You need to repeat
the measure over and over and then average the responses or otherwise describe the
 Introduction  9

(A) (B)
100 100

“I hear it.”

Percentage of times reported present

Percentage of times reported present

75 75

Threshold
50 50

25 25

“I don’t hear it.”

0 0
7 8 9 10 11 12 7 8 9 10 11 12
Stimulus level (arbitrary units) Stimulus level (arbitrary units)

FIGURE 1.6 The method of constant stimuli (A) We might expect the threshold to be
a sharp change in detection from never reported to always reported, as depicted here, but
this is not so. (B) In reality, experiments measuring absolute threshold produce shallower
functions relating stimulus to response. A somewhat arbitrary point on the curve, often
50% detection, is designated as the threshold (dashed line).

pattern of results. Some experiments require thousands of repetitions (thousands method of limits A psychophysical
of “trials”) to establish a sufficiently reliable data point. method in which the particular dimension
of a stimulus, or the difference between
Returning to our auditory example, as the listener, you would report whether two stimuli, is varied incrementally until
you heard a tone or not. You would always report hearing a tone that was relatively the participant responds differently.
far above threshold, and almost never report hearing a tone that was well below
threshold. In between, however, you would be more likely to hear some tone inten-
sities than not to hear them, and you would hear other, lower intensities on only a
few presentations. In general, the intensity at which a stimulus would be detected
50% of the time would be chosen as your threshold.
That 50% definition of absolute threshold is rather interesting. Weren’t we
looking for a way to measure the weakest detectable stimulus? Using the hearing Trial series

example, shouldn’t that be a value below which we just can’t hear anything (see 1 2 3 4 5 6 7 8
Figure 1.6A)? It turns out that no such hard boundary exists. Because of variability 20 Y Y
in the nervous system, stimuli near threshold will be detected sometimes and
19 Y Y Y Y
missed at other times. As a result, the function relating the probability of detection
with the stimulus level will be gradual (see Figure 1.6B), and we must settle for 18 Y Y Y Y
Intensity (arbitrary units)

a somewhat arbitrary definition of an absolute threshold. We will return to this 17 Y Y Y Y

issue when we talk about signal detection theory.
16 Y Y Y Y Y
The method of constant stimuli is simple to use, but it is an inefficient way to
conduct an experiment, because much of the listener’s time is spent with stimuli 15 Y Y Y Y Y Y Y Y
that are well above or below threshold. A more efficient approach is the method 14 Y N Y N Y N Y Y
of limits (FIGURE 1.7). With this method, the experimenter begins with the
13 N N Y N Y N N Y
same set of stimuli—in this case, tones that vary in intensity. Instead of random
presentations, tones are presented in order of increasing or decreasing intensity. 12 N N N N N N

11 N N N N
FIGURE 1.7 The method of limits Here the listener attends to multiple series of trials.
For each series, the intensity of the stimulus is gradually increased or decreased until the 10 N N N N
listener detects (Y) or fails to detect (N), respectively, the stimulus. For each series, an esti- 13.5 14.5 12.5 14.5 12.5 14.5 13.5 12.5
mate of the threshold (red dashed line) is taken to be the average of the stimulus level just
before and after the change in perception (i.e., the average “crossover value”). Crossover values (average = 13.5)

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Wolfe6e_01.06.ai 3.03.2020
10  Chapter 1
method of adjustment A method of
limits in which the participant controls
the change in the stimulus.
magnitude estimation A psychophys-
ical method in which the participant
assigns values according to perceived
magnitudes of the stimuli.
Stevens’s power law A principle
describing the relationship between
stimulus and resulting sensation that
says the magnitude of subjective sensa-
tion is proportional to the stimulus mag-
nitude raised to an exponent.
When tones are presented in ascending order, from faintest to loudest, listeners are
asked to report when they first hear the tone. With descending order, the task is to
report when the tone is no longer audible. Data from an experiment such as this
show that there is some “overshoot” in judgments. It usually takes more intensity
to report hearing the tone when intensity is increasing, and it takes more decreases
in intensity before a listener reports that the tone cannot be heard. We take the
average of these crossover points—when listeners shift from reporting hearing the
tone to not hearing the tone, and vice versa—to be the threshold.
The third and final of these classic measures of thresholds is the method of
adjustment. This method is just like the method of limits, except the person being
tested is the one who steadily increases or decreases the intensity of the stimulus.
The method of adjustment may be the easiest method to understand, because it is
much like day-to-day activities such as adjusting the volume dial on a stereo or the
dimmer switch for a light. Even though it’s the easiest to understand, the method of
adjustment is not usually used to measure thresholds. The method would be perfect
if threshold data were like those plotted in Figure 1.6A, but a graph of real data
looks more like Figure 1.6B. The same person will adjust a dial to different places
on different trials, and measurements get even messier when we try to combine
the data from multiple people.
Scaling Methods
Moving beyond absolute thresholds and difference thresholds, suppose we wanted
to know how strong your experiences are. For example, we might show you a light
and ask how much additional light you would need to make another light look
twice as bright. Though that might seem like an odd question, it turns out to be
answerable. We could give you a knob to adjust so that you could set the second
light to appear twice as bright as the first, and you could do it.
In fact, we don’t need to give the observers a light to adjust. A surprisingly
straightforward way to address the question of the strength or size of a sensation is
to simply ask observers to rate the experience. For example, we could give observers
a series of sugar solutions and ask them to assign numbers to each sample. We
would just tell our observers that sweeter solutions should get bigger numbers, and
if solution A seems twice as sweet as solution B, the number assigned to A should
be twice the number assigned to B. This method is called magnitude estimation,
and the approach actually works well, even when observers are free to choose their
own range of numbers. More typically, however, we might begin the experiment by
presenting one solution at an intermediate level and telling the taster to label this
level as a specific value—10, for instance. All of the responses should then be scaled
sensibly above or below this standard of 10. If you do this for sugar solutions, you
will get data that look like the blue “sweetness” line in FIGURE 1.8.
Inspired by his student Richard Held, a distinguished vision researcher whose
work you will learn about in Sections 6.4 and 13.1, Harvard psychologist S. S.
Stevens (1962, 1975) developed magnitude estimation. Stevens, his students, and
their successors measured functions like the one in Figure 1.8 for many different
sensations. Even though observers were asked to assign numbers to private expe-
rience, the results were orderly and lawful. However, they were not the same for
every type of sensation. That relationship between stimulus intensity and sensation
is described by what is now known as Stevens’s power law:
S = aI b
which states that the sensation (S) is related to the stimulus intensity (I) by an ex-
ponent (b). (The letter a is a constant that corrects for the units you are using. For
 Introduction  11

100 FIGURE 1.8 Magnitude estimation The lines on this

graph represent data from magnitude estimation experi-
90 ments using electric shocks of different currents, lines of
different lengths, solutions of different sweetness levels, and
lights of different brightness levels. The exponents for the
80 Electric
“power functions” that describe these lines are 3.5, 1.0, 0.8,
shock (3.5)
and 0.3, respectively. For exponents greater than 1, such as
70 for electric shock, Fechner’s law does not hold, and Stevens’s
Apparent power law must be used instead.
length (1.0)
Perceived intensity

60

50

40 Sweetness (0.8)

30

20
Brightness (0.3)
10

0
0 10 20 30 40 50 60 70 80 90 100
Stimulus energy

example, if you measured your stimulus in meters and then switched to measuring
it in centimeters, you would need to multiply by 0.01 [= divide by 100] to keep your
sensation numbers the same.) So, for example, experienced sensation might rise
with intensity squared (I × I). That would be an exponent of 2.0. If the exponent is
less than 1, it means that the sensation grows less rapidly than the stimulus—which
is what Fechner’s law and Weber’s law would predict.
Suppose you have some lit candles and you light 10 more. If you start with 1
candle, the change from 1 to 11 candles must be quite dramatic. If you start with
100 and add 10, the change will be modest. Adding 10 to 10,000 won’t even be
noticeable. In fact, the exponent for brightness is about 0.3. The exponent for
sweetness is about 0.8 (Bartoshuk, 1979). Properties like length have exponents near
1, so, reasonably enough, a 12-inch-long stick looks twice as long as a 6-inch-long
stick (S. S. Stevens and Galanter, 1957). Note that this length relationship is true
over only a moderate range of sizes. An inch added to the size of a spider changes
your sensory experience much more than an inch added to the height of a giraffe.
Some stimuli have exponents greater than 1. In the painful case of electric shock,
the pain grows with I3.5 (Stevens, Carton, and Shickman, 1958), so a 4-fold increase
in the electrical current is experienced as a 128-fold increase in pain!
Weber’s and Fechner’s laws have rather broad implications beyond questions
of apparent brightness or loudness. Some plants, for example, will respond to the
psychophysical experience of the bees that pollinate them (Nachev et al., 2017).
Suppose you are a plant pollinated by bees that are attracted to your flowers because
those flowers have sweet nectar. How much sugar do you need to put into that
nectar? After all, it’s going to cost you energy to produce sugar. If your nectar has 2
units of sugar and the neighbor flower only has 1, you probably have a competitive
advantage over the neighbor. However, if yours has 12 units and the neighbor’s has
11, that difference of 1 unit might fall below the bee’s Weber fraction for sweetness.
Thus, there will be greater evolutionary pressure to go from 1 to 2 than from 11 to
12. Similarly, a peacock with 51 feathers in his tail probably does not have much of

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Sensation & Perception 6E
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12  Chapter 1

cross-modality matching The ability a reproductive advantage over a 50-feather peacock. The peahen might not notice
to match the intensities of sensations the difference, putting an evolutionary brake on tail inflation (Farris, 2017).
that come from different sensory modal-
ities. This ability allows insight into sen-
At this point in our discussion of psychophysics, it is worth taking a moment
sory differences. For example, a listener to compare Weber’s, Fechner’s, and Stevens’s laws:
might adjust the brightness of a light 1. Weber’s law involves a clear objective measurement. We know how much
until it matches the loudness of a tone.
we varied the stimulus, and either the observers can tell that the stimulus
changed or they cannot.
2. Fechner’s law begins with the same sort of objective measurements as
Weber’s, but the law is actually a calculation based on some assumptions
about how sensation works. In particular, Fechner’s law assumes that all
JNDs are perceptually equivalent. In fact, this assumption turns out some-
times to be incorrect and leads to instances where the “law” is violated, such
as in the electric shock example just given.
3. Stevens’s power law describes rating data quite well, but notice that rating
data are qualitatively different from the data that support Weber’s law. We
can record the observer’s ratings and we can check whether those ratings
are reasonable and consistent, but there is no way to know whether they are
objectively right or wrong.
A useful variant of the scaling method shows us that different individuals can
live in different sensory worlds, even if they are exposed to the same stimuli. This
method is called cross-modality matching. In cross modality match-
ing, an observer adjusts a stimulus of one sort to match the perceived
Matching sensations
magnitude of a stimulus of a completely different sort (J. C. Stevens,
Strongest pain
1959). For example, we might ask a listener to adjust the brightness of
Loudest sound a light until it matches the loudness of a particular tone. Again, though
Brightest light the task might sound odd, people can do this, and for the most part,
Supertasters everyone with “normal” vision and hearing will produce a similar pattern
of matches of a sound to a light. We still can’t examine someone else’s
private experience, but at least the relationship of visual experience
Brightness of the sun
and auditory experience appears to be similar across individuals.
This similarity does not hold when it comes to the sense of taste.
Heat of scalding water
There is a molecule called propylthiouracil (PROP) that some people
experience as very bitter, while others experience it as almost tasteless.
Sound of a fire engine
Still others fall in between. This relationship between a chemical and
bitter taste can be examined formally with cross-modality matching
(Marks et al., 1988). When observers are asked to match the bitterness of
Pain of a severe headache
Sound of an airplane PROP to other sensations completely unrelated to taste, we do not find
Brightness of high-beam headlights the sort of agreement that is found when observers match sounds and
Smell of a skunk
Coldness of snow lights (FIGURE 1.9). Some people—we’ll call them nontasters—match
the taste of PROP to very weak sensations, like the sound of a watch or
a whisper. A group of “supertasters” assert that the bitterness of PROP
Medium is similar in intensity to the brightness of the sun or the most intense
tasters Brightness of low-beam headlights
Smell of bacon frying
Pain of a mild headache

Brightness of the moon/loudness FIGURE 1.9 Cross-modality matching The levels of bitterness of con-
of a conversation centrated PROP perceived by nontasters, medium tasters, and supertasters
of PROP are shown on the left. The perceived intensities of a variety of
everyday sensations are shown on the right. The arrow from each taster type
Loudness of a whisper indicates the level of sensation to which those tasters matched the taste of
Nontasters Sound of a watch PROP. (Data from K. Fast. 2004. Developing a Scale to Measure Just About
Anything: Comparisons across Groups and Individuals. Thesis Digital Library,
No sensation 3353. Yale University School of Medicine: New Haven, CT.)
 Introduction  13

pain ever experienced. Medium tasters match PROP to weaker stimuli, such as the signal detection theory A psycho-
smell of frying bacon or the pain of a mild headache (Bartoshuk, Fast, and Snyder, physical theory that quantifies the
response of an observer to the pre-
2005). As we will see in Section 15.5, there is a genetic basis for this variation, and sentation of a signal in the presence of
it has wide implications for our food preferences and, consequently, for health. noise. Measures obtained from a series
For the present discussion, this example shows that we can use scaling methods of presentations are sensitivity (dʹ) and
to quantify what appear to be real differences in individuals’ taste experiences. criterion of the observer.

Signal Detection Theory

Let’s return to thresholds—particularly to the fact that they are not absolute. An
important way to think about this fact and to deal with it experimentally is known
as signal detection theory (D. M. Green and Swets, 1966). Like so much of mod-
ern psychophysics, even signal detection theory was anticipated by Fechner over a
hundred years earlier (Wixted, 2020). Signal detection theory begins with the fact
that the stimulus you’re trying to detect (the “signal”) is always being detected in
the presence of “noise.” If you sit in the quietest place you can find and put on your
best noise-canceling headphones, you will find that you can still hear something.
Similarly, if you close your eyes in a dark room, you still see something—a mottled
pattern of gray with occasional brighter flashes. This is internal noise, the static
in your nervous system. Many neurons in the brain are firing all the time, even
when nothing is happening. For example, many neurons in the auditory system
fire up to 50 times per second when there is no sound at all, and you will learn in
Chapter 12 that neurons in the vestibular system fire 100 times per second even
when you’re perfectly motionless. When you’re trying to detect a faint sound or
flash of light, you must be able to detect it in the presence of such internal noise.
Near your threshold, it will be hard to tell a real stimulus from a random surge of
internal noise.
There is external noise, too. Consider again that radiologist reading a mammogram
looking for signs of breast cancer. As you can see in FIGURE 1.10, the mammogram
contains lots of similar regions; the marked white region is the danger sign. We can Cancer
think of the cancer as the signal. By the time it is presented to the radiologist in an
X-ray, there is a signal plus noise. Elsewhere in the image, and in other images, are
stimuli that are nothing other than noise. The radiologist is a visual expert, trained
to find these particular signals, but sometimes the signal will be lost in the noise
and missed, and sometimes some noise will look enough like cancer to generate a
false alarm (Nodine et al., 2002). Thus, the radiologist will be faced with uncertainty,
introduced both as external and internal noise. Not cancer

© Xray Computer/Shutterstock.com
Of course, sometimes neither internal nor external noise is much of a problem.
When you see this dot, , you are seeing it in the presence of internal noise, but the
●

magnitude of that noise is so much smaller than the signal generated by the dot that
the noise has no real impact. Similarly, the dot may not be exactly the same as other
dots, but that variation—the external noise—is also too small to have an impact. If
asked about the presence of a dot here, , and its absence here, , you will be correct
●

in your answer essentially every time. Signal detection theory exists to help us un-
derstand what’s going on when we make decisions under conditions of uncertainty. FIGURE 1.10 Differentiating signal
Because we are not expert mammographers, let’s introduce a different example from noise Mammograms—X-rays of
the breast—are used to screen women for
to illustrate the workings of signal detection theory. You’re in the shower. The water
breast cancer. The solid white region is
is making a noise that we will imaginatively call “noise.” Sometimes the noise sounds the signal of a cancerous growth; however,
louder to you; sometimes it seems softer. We can plot the distribution of your per- the mammogram contains many similar
ception of noise as shown in FIGURE 1.11A . On the x-axis, we have the magnitude regions (“noise”). Reading such images is a
difficult perceptual task. Even for a radiol-
of your sensation from “less” to “more.” Imagine that we asked you, over and over
ogist trained to discriminate and identify
again, about your sensation. Or imagine we took many repeated measures of the particular signals, there is always uncer-
response in your nervous system to the sound. For some instances, the response tainty due to internal and external noise.
 14  Chapter 1

(C)

+
(A) (B)
NO Criterion YES

Number of instances
Shower “noise” alone

Ringtone + noise

Less More Less More Less More

Your perception Sounds like phone Sounds like phone

(D) Correct rejection (E) Hit (F) False alarm (G) Miss
NO Criterion YES NO Criterion YES NO Criterion YES NO Criterion YES
Number of instances

Less More Less More Less More Less More

Sounds like phone Sounds like phone Sounds like phone Sounds like phone

FIGURE 1.11 Detecting a stimulus using signal detection theory (SDT) ( A) SDT
assumes that all perceptual decisions are made against a background of noise (the red curve)
generated both in the world and in the nervous system. (B) Your job is to distinguish ner-
vous system responses due to noise alone (dotted, red curve) or to signal plus noise (solid,
blue curve). (C) The best you can do is establish a criterion (solid black line) and declare that
you detect something if the response is above that criterion. (D–G) Signal detection theory
includes four classes of responses. (D) “Correct rejection” (you say “no” and there is, indeed,
no signal). (E) “Hits” (you say “yes” and there is a signal). (F ) “False-alarm errors” (you say
“yes” to no signal). (G) “Miss errors” (you say “no” to a real signal).

would be “less.” For some, it would be “more.” On average, it would lie somewhere
in between. If we tabulated all of the responses, we would get a bell-shaped (or
“normal”) distribution of answers, with the peak of that distribution showing the
average answer that you gave.
Now a ringtone plays. That will be our “signal.” Your perceptual task is to
detect the signal in the presence of the noise. What you hear is a combination of
the ringtone and the shower. That is, the signal is added to the noise, so we can
imagine that now we have two distributions of responses in your nervous system:
a noise-alone distribution and a signal-plus-noise distribution (FIGURE 1.11B).
For the sake of simplicity, let’s suppose that “more” response means that it
sounds more like the phone is ringing. So now your job is to decide whether it’s
time to jump out of the shower and answer what might be the phone. The problem
is that you have no way of knowing at any given moment whether you’re hearing
criterion In reference to signal
noise alone or signal plus noise. The best you can do is to decide on a criterion level
detection theory, an internal thresh- of response (FIGURE 1.11C). If the response in your nervous system exceeds that
old that is set by the observer. If the criterion, you will jump out of the shower and run naked and dripping to find the
internal response is above criterion, phone. If the level is below the criterion, you will decide that it is not a ringtone
the observer gives one response
(e.g., “yes, I hear that”). Below cri-
and stay in the shower. Note that this “decision” is made automatically; it’s not
terion, the observer gives another that you sit down to make a conscious (soggy) choice. Thus, in signal detection
Wolfe
response (e.g., “no, I hear nothing”). theory, a criterion is a value that is somehow determined by the observer. Within
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 Introduction  15

(A) No sensitivity (B) Moderate sensitivity (C) High sensitivity

Number of instances
Number of instances
Shower “noise” alone d′ = ~0 d′ = ~1 d′ = ~4

Ringtone + noise

Less More Less More Less More

Sounds like phone Sounds like phone Sounds like phone

FIGURE 1.12 Sensitivity (dʹ) in SDT Your sensitivity to a stimulus is illustrated by the
separation between the distributions of your response to noise alone (dotted, red curve)
and to signal plus noise (solid, blue curve). This separation is captured by the measure
dʹ (d-prime). (A) If the distributions completely overlap, dʹ is almost 0 and you have no
ability to detect the signal. (B) If dʹ is intermediate, you have some sensitivity but your
performance will be imperfect. (C) If dʹ is big, then distinguishing signal from noise is easy.

the observer, a response above the criterion will be taken as evidence that a signal sensitivity In reference to signal detec-
is present. A response below that level will be treated as noise. tion theory, a measure that defines the
ease with which an observer can tell the
There are four possible outcomes in this situation: You might say “no” when
difference between the presence and
there is no ringtone; that’s a correct rejection or true negative (FIGURE 1.11D). absence of a stimulus or the difference
You might say “yes” when there is a ringtone; that’s known as a hit or true positive between Stimulus 1 and Stimulus 2.
response (FIGURE 1.11E). Then there are the errors. If you jump out of the shower
when there’s no ringtone, that’s a false alarm or false positive (FIGURE 1.11F). If
you miss the call, that’s a miss or false negative (FIGURE 1.11G).
How sensitive are you to the ringtone? In FIGURE 1.12, the sensitivity is graphed
as the separation between the noise-alone and signal-plus-noise distributions. If the
distributions are on top of each other (Figure 1.12A), you can’t tell noise alone from
signal plus noise. A false alarm is just as likely as a hit. By knowing the relationship of
hits to false alarms, you can calculate a sensitivity measure known as dʹ (d-prime),
which would be about zero in Figure 1.12A. In Figure 1.12C we see the case of a
large dʹ. Here you could detect essentially all the ringtones and never make a false
alarm error. The situation we’ve been discussing is in between (Figure 1.12B).
Now suppose you’re waiting for an important call. Even though you really don’t
want to miss the call, you can’t magically make yourself more sensitive. All you can
do is move the criterion level of response, as shown in FIGURE 1.13. If you shift
your criterion to the left, you won’t miss many calls, but you will have lots of false

(A) “Gotta get that call!” (B) “Is that the phone?” (C) “That’s not the phone.”
Number of instances

Shower “noise” alone

Ringtone + noise

Less More Less More Less More

Sounds like phone Sounds like phone Sounds like phone

FIGURE 1.13 Criterion in SDT For a fixed dʹ, all you can do is change the pattern of
your errors by shifting the response criterion. If you don’t want to miss any signals, you
move your criterion to the left (A), but then you have more false alarms. If you don’t like
false alarms, you move the response criterion to the right (C), but then you make more
Wolfe
miss errors.
Sensation In all these
& Perception 6Ecases (A–C), your sensitivity, dʹ, remains the same.
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 16  Chapter 1

Correct rejection Pr(N|n) FIGURE 1.14 Receiver operating characteristic (ROC) curves
1.0 0
1.0 0 Theoretical ROC curves for different values of dʹ. Note that dʹ = 0 when
performance is at the chance level. Higher values of dʹ indicate that
the probability of hits and correct rejections increases, and the proba-
bility of misses and false alarms decreases. Pr(N|n) = probability of the
2
=

response “no signal present” when no signal is present (correct rejec-

d′

5
1.

tion); Pr(N|s) = probability of the response “no signal present” when

=
d′

signal is present (miss); Pr(S|n) = probability of the response “signal

75
0.

present” when no signal is present (false alarm); Pr(S|s) = probability of

=

ce
d′

the response “signal present” when signal is present (hit).

an

Miss Pr(N|s)
rm
Hit Pr(S|s)

fo
r
pe
e
nc

alarms (Figure 1.13A). That’s annoying. You’re running around

ha
C

0 1.0
0 1.0
False alarm Pr(S|n)
receiver operating characteristic
(ROC) curve In reference to stud-
ies of signal detection, the graphical
plot of the hit rate as a function of the
false-alarm rate. If these are the same,
points fall on the diagonal, indicating
that the observer cannot tell the dif-
ference between the presence and
absence of the signal. As the observer’s
sensitivity increases, the curve bows
upward toward the upper left corner.
That point represents a perfect ability to
distinguish signal from noise (100% hits,
0% false alarms).
doctrine of specific nerve energies
A doctrine, formulated by Johannes
Müller, stating that the nature of a
sensation depends on which sensory
fibers are stimulated, rather than how
they are stimulated.
naked, dripping on the floor, and traumatizing the cat for no good
reason. If you shift your criterion to the right, you won’t have those
annoying false alarms, but you will miss most of the calls (Figure
1.13C). For a fixed value of dʹ, changing the criterion changes the
hits and false alarms in predictable ways. If you plot false alarms
on the x-axis of a graph against hits on the y-axis for different
criterion values, you get a curve known as a receiver operating
characteristic (ROC) curve (FIGURE 1.14).
Suppose you were guessing (the Figure 1.12A situation). You
might guess “yes” on 40% of the occasions when the phone rang,
but you would also guess “yes” on 40% of the occasions when the phone did not ring.
If you moved your criterion and guessed “yes” on 80% of phone-present occasions,
you would also guess “yes” on 80% of phone-absent occasions. Your data would fall
on that “chance performance” diagonal in Figure 1.14. If you were perfect (the Figure
1.12C situation), you would have 100% hits and 0% false alarms and your data point
would lie at the upper left corner in Figure 1.14. Situations in between (Figure 1.12B)
produce curves between guessing and perfection (the green, purple, and blue curves
in Figure 1.14). If your data lie below the chance line, you did the experiment wrong!
Let’s return to our radiologist. She will have an ROC curve whose closeness
to perfection reflects her expertise. On that ROC, her criterion can slide up and
to the right, in which case she will make more hits but also more false alarms,
or down and to the left, in which case she will have fewer false alarms but more
misses. Where she places her criterion (consciously or unconsciously) will depend
on many factors. Does the patient have factors that make her more or less likely to
have cancer? What is the perceived cost of a missed cancer? What is the perceived
cost of a false alarm? You can see that what started out as a query about the lack of
absolute thresholds can become, quite literally, a matter of life and death.
Signal detection theory can become a rather complicated topic in detail (e.g.,
what happens if those noise and signal + noise curves are not exactly the same
shape?). To learn about how to calculate dʹ and about ROC curves, you can take
advantage of many useful websites and several texts (e.g., Macmillan and Creelman,
2005; see Burgess, 2010 if you’re interested in the application to radiology).
1.3 S
 ensory Neuroscience and
the Biology of Perception
Many of you reading this book will have had some introduction to neuroscience.
Here, we review very briefly some of the neuroscience that is relevant to the study
of sensation and perception. If this is your first encounter with neuroscience, you
may want to consult a neuroscience text to give yourself a more detailed background
than we will provide.
 Introduction  17

During the nineteenth century, when Weber and Fechner were initiating the cranial nerves Twelve pairs of nerves
experimental study of perception, physiologists were hard at work learning how (one for each side of the body) that orig-
inate in the brain stem and reach sense
the senses and the brain operate. Much of this work involved research on animals. organs and muscles through openings
It’s worth spending a moment on a key assumption here: studies of animal senses in the skull.
tell us something about human senses. That may seem obvious, but the assumption olfactory (I) nerves The first pair of
requires the belief that there is some continuity between the way animals work and cranial nerves. The axons of the olfactory
the way humans work. sensory neurons bundle together after
The most powerful argument for a continuity between humans and animals passing through the cribriform plate to
form the olfactory nerve, which conducts
came from Darwin’s theory of evolution. During the 1800s, Charles Darwin impulses from the olfactory epithelia in
(1809–1882) proposed his revolutionary theory in On the Origin of Species (1859). the nose to the olfactory bulb.
Although many of the ideas found in that book had been brewing for some time, optic (II) nerves The second pair of
controversy expanded with vigor following Darwin’s provocative statements in cranial nerves, which arise from the ret-
The Descent of Man (1871), where he argued that humans and apes evolved from ina and carry visual information to the
thalamus and other parts of the brain.
a common ancestor. If there was continuity in the structure of the bones, heart,
and kidneys of cows, dogs, monkeys, and humans, then why wouldn’t there be vestibulocochlear (VIII) nerves
The eighth pair of cranial nerves, which
continuity in the structure and function of their sensory and nervous systems? connect the inner ear with the brain,
An inescapable implication of the theory of evolution is that we can learn much transmitting impulses concerned with
about human sensation and perception by studying the structure and function of hearing and spatial orientation. The
our nonhuman relatives. vestibulocochlear nerve is composed
of the cochlear nerve branch and the
vestibular nerve branch.
Nerves and Specific Nerve Energies
oculomotor (III) nerves The third pair
At the same time that Darwin was at work in England, the German physiologist of cranial nerves, which innervate all the
Johannes Müller (1801–1858) (FIGURE 1.15) was writing his very influential Hand- extrinsic muscles of the eye except the
book of Physiology during the early 1830s. In this book, in addition to covering lateral rectus and the superior oblique
muscles, and which innervate the ele-
most of what was then known about physiology, Müller formulated the doctrine
vator muscle of the upper eyelid, the
of specific nerve energies. The central idea of this doctrine is that we cannot be ciliary muscle, and the sphincter muscle
directly aware of the world itself, and we are only aware of the activity in our nerves. of the pupil.
Further, what is most important is which nerves are stimulated, and not how they trochlear (IV) nerves The fourth
are stimulated. For example, we experience vision because the optic nerve leading pair of cranial nerves, which innervate
from the eye to the brain is stimulated, but it does not matter whether light, or the superior oblique muscles of the
eyeballs.
something else, stimulates the nerve. To prove to yourself that this is true, close your
eyes and press very gently on the outside corner of one eye through the lid. (This abducens (VI) nerves The sixth pair
of cranial nerves, which innervate the
works better in a darkened room.) You will see a spot of light toward the inside of lateral rectus muscle of the eyeballs.
your visual field by your nose. Despite the lack of stimulation by light, your brain
interprets the input from your optic nerve as informing you about something visual.
The cranial nerves leading into and out of the skull illustrate the doctrine of
specific nerve energies (FIGURE 1.16). The pair of optic nerves is one of 12 pairs of
cranial nerves that pass through small openings in the bone at the base of the skull;
these nerves are dedicated mainly to sensory and motor systems. Cranial nerves
are labeled both by names (in boldface type) and by Roman numerals that roughly
correspond to the order of their locations, beginning from the front of the skull.
Three of them—olfactory (I), optic (II), and vestibulocochlear (VIII)—are exclusively
dedicated to sensory information. The vestibulocochlear nerve serves two sensory
modalities: the vestibular sensations that support our sense of equilibrium (see
Chapter 12) and hearing (discussed in Section 9.3). Three more cranial nerves—
oculomotor (III), trochlear (IV), and abducens (VI)—are dedicated to muscles that
move the eyes. In addition to instructions from the visual system, these nerves

FIGURE 1.15 Johannes Müller The German physiologist Johannes Müller formulated
the doctrine of specific nerve energies, which says that we are aware only of the activity in
our nerves, and we cannot be aware of the world itself. For this reason, what is most import-
ant is which nerves are stimulated, not how they are stimulated.
18  Chapter 1

I Olfactory
Smell III Oculomotor
All eye muscles except
II Optic superior oblique and
Vision lateral rectus

IV Trochlear
Superior oblique muscle

Sensory VI Abducens
Motor Lateral rectus muscle

V Trigeminal
Face, sinuses,
teeth

Jaw
muscles

XII Hypoglossal
Tongue muscles

XI Spinal accessory
Sternomastoid
and trapezius
muscles

VII Facial
Auditory Tongue,
nerve soft palate
X Vagus Vestibular
Heart, lungs, nerve Facial muscles,
gastrointestinal salivary glands,
tract, bronchi, Same tear glands
trachea, larynx

IX Glossopharyngeal VIII Vestibulocochlear

Posterior tongue, Spatial orientation,
tonsils, pharynx, Same
balance, hearing
pharyngeal muscles

FIGURE 1.16 Cranial nerves Twelve

pairs of cranial nerves pass through small
openings in the bone at the base of the
skull. These nerves conduct information for
sensation, motor behavior, or both. (After
S. M. Breedlove and N. V. Watson. 2013.
Biological Psychology: An Introduction to
Behavioral, Cognitive, and Clinical Neuro-
science, 7th ed., Oxford University Press/
Sinauer Associates: Sunderland, MA.)
have direct connections from the vestibular system so your eyes know where your
nose is pointed. The other six cranial nerves either are exclusively motor (spinal
accessory [XI] and hypoglossal [XII]), or convey both sensory and motor signals
(trigeminal [V], facial [VII], glossopharyngeal [IX], and vagus [X]).
The doctrine of specific nerve energies extends beyond the cranial nerves, as
illustrated especially well by our senses of hot and cold on the skin. Two specialized
types of nerve cells are warmth fibers and cold fibers, which respond to increases

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 Introduction  19

Central Motor Somatosensory cortex polysensory Referring to blending

fissure cortex (skin senses) multiple sensory systems.
sensory integration or multisensory
Parietal integration The process of combining
lobe different sensory signals. The senses
Frontal typically work together to learn about
lobe the world and to guide behavior. This
is not the same as the mathematical
Occipital process of integration learned in
Anterior lobe Posterior calculus (e.g., the integral of accelera-
tion is velocity).

Olfactory Visual cortex

bulb (vision)

Olfactory cortex
(smell)
Sylvian Temporal Auditory cortex
fissure lobe (hearing)

FIGURE 1.17 Cortex of the human brain The cortex is the outer layer of the cerebral
hemispheres. This lateral view shows the left hemisphere. The anterior (front of head) is on
the left and the posterior (back of the head) is on the right. The darkened areas show where
information from four of our sensory modalities first reaches the cortex. Also shown is the
Wolfe cortex, which is engaged in balance, touch, and some auditory processing.
motor
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and decreases in temperature on the skin (see Section 13.1). Capsaicin, a chemical
that occurs naturally in chili peppers, causes warmth fibers to fire, creating a sense
of increasing heat even though the temperature has not changed. On the other
hand, menthol, which imparts a minty flavor in cough drops, stimulates cold fibers
(Bautista et al., 2007), so skin feels cooler without getting physically colder. In
sufficiently high amounts, both capsaicin and menthol stimulate pain receptors in
the skin. Paradoxically, this is why ointments often contain capsaicin or menthol,
where their effect is to mask real physical pain (Green, 2005).
Just as different nerves are dedicated to individual sensory and motor tasks,
areas of the brainstem and cerebral cortex are similarly dedicated to particular
tasks. Areas of the cortex dedicated to perception actually are much larger than
the darkened areas in FIGURE 1.17. The areas depicted here are primary sensory
areas; more complex processing is accomplished across cortical regions that spread
well beyond these primary areas. For example, visual perception uses cortex that
extends both anteriorly (forward) into parietal cortex and ventrally (lower) into
regions of the temporal lobe (see Figures 4.2 and 4.4). In addition, as processing
extends beyond primary areas, cortex often becomes polysensory, meaning that
information from more than one sense is being combined in some manner. (See
Activity 1.2: Sensory Areas in the Brain.) While this textbook is organized into
sense-specific chapters, we always want to remember that we live in a multisensory
world. Researchers specifically study what is called sensory integration or multi-
sensory integration (Spence, 2018). Throughout this book, you will find multisensory
research, flagged with .
Hermann Ludwig Ferdinand von Helmholtz (1821–1894) (FIGURE 1.18), one of
the greatest scientists of the nineteenth century, was greatly influenced by Müller
at the Free University of Berlin. In an early effort, Helmholtz estimated that the FIGURE 1.18 Hermann von Helmholtz
Contributor to many fields of science, von
speed of signal transmission in the nerves in frog legs was about 90 feet per second, Helmholtz was one of the greatest scien-
comfortably within the range of ordinary physical events. Later he concluded that tists of all time. He made many important
human sensory nerves transmitted signals at speeds of between 165 and 330 feet discoveries in physiology and perception.
 20  Chapter 1

(A) (B)

FIGURE 1.19 Santiago Ramón y Cajal (A) Santiago Ramón y Cajal (looking tired; per-
haps this was not his happiest day in the lab) created meticulous drawings of brain neurons
(B) while peering into a microscope for many hours. Because of his painstaking care and
accuracy, his early drawings are still cited and reproduced today.

synapse The junction between neu-
rons that permits information transfer.
Neuroscience History Archives, Brain Research Institute, University of California, Los Angeles
per second. (See Activity 1.3: Neurons.) This transmission speed was slower than
many people believed at the time, probably because it feels like you are sensing what
is happening right now. In fact, you are experiencing recent (very recent) history.
To use Helmholtz’s example, a “whale probably feels a wound near its tail in about
one second and requires another second to send back orders to the tail to defend
itself ” (Koenigsberger, 1906/1965, p. 72). Some neurons are faster than others but,
regardless, it is interesting to realize that when you stub your toe, a measurable
amount of time elapses before you feel the consequences.

Neuronal Connections
The neuron that carries a signal from your toe to your brain needs to pass its in-
formation on to other neurons. How do neurons connect with each other? The
Spanish neuroanatomist Santiago Ramón y Cajal (1852–1934) (FIGURE 1.19A)
looked closely at thin slices of tissue and created some of the most painstaking and
breathtaking drawings of the organization of neurons in the brain. FIGURE 1.19B
shows an example. His drawings suggested that neurons do not actually touch
one another. Instead, he depicted neurons as separate cells with tiny gaps between
them. Sir Charles Scott Sherrington (1857–1952) (FIGURE 1.20) named the tiny
gap between the axon of one neuron and the dendrite of the next a synapse, from
the Greek word meaning “to fasten together” (FIGURES 1.21 and 1.22).

FIGURE 1.20 Sir Charles Scott Sherrington An English neurophysiologist, Sherrington

Wolfe earned a Nobel Prize for his pioneering discoveries concerning neural activities.
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 Introduction  21

Presynaptic neuron Postsynaptic neuron FIGURE 1.21 A typical neuron

Features of a typical neuron and their
functions are described. Axon length and
dendrite branching patterns vary greatly
across the different types of neurons.

Dendrites The cell body Information collected by The axon Axon terminals
receive contains the dendrites is integrated conducts action synapse with a
information nucleus and in the axon hillock, potentials away target cell.
from other most cell which generates from the cell body.
neurons. organelles. action potentials.

Presynaptic neuron Postsynaptic neuron

Axon

Axon terminal
(presynaptic)

Synaptic
vesicles

Dendrite
(postsynaptic)

na
Sy

ps
e
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Neurotransmitter
molecules Receptors
Wolfe6e_01.21.ai 3.27.2020
FIGURE 1.22 A synapse An axon terminal in the presynaptic cell communicates with a dendrite
of the postsynaptic cell. Neurotransmitter molecules are released by synaptic vesicles in the axon
and fit into receptors on the dendrite on the other side of the synapse, thus communicating from
the axon of the first (presynaptic) neuron to the dendrite of the second (postsynaptic) neuron.
 22  Chapter 1

Initially, people thought that some sort of electrical wave traveled across the
synapse from one neuron to the next. However, Otto Loewi (1873–1961) (FIGURE
1.23) was convinced that this could not be true. One reason is that some neurons
increase the response of the next neuron (they are excitatory), whereas other neurons
decrease the response of the next neuron (they are inhibitory). Loewi proposed
that something chemical, instead of electrical, might be at work at the synapse.
The chemicals turned out to be molecules, called neurotransmitters, that travel
from the axon across the synapse to bind to receptor molecules on the dendrite of
the next neuron. There are many different kinds of neurotransmitters in the brain,
and individual neurons are selective with respect to which neurotransmitters excite
or inhibit them from firing. Drugs that are psychoactive, such as amphetamines,
Wellcome Collection/CC BY 4.0

work by increasing or decreasing the effectiveness of different neurotransmitters.

Today, scientists use chemicals that influence the effects of neurotransmitters in
efforts to understand pathways in the brain, including those used in perception.

Neuronal Firing: The Action Potential

After Loewi’s discovery of neurotransmitters, scientists learned what it really
FIGURE 1.23 Otto Loewi The German means to have a neuron “fire.” Important early research took advantage of the fact
pharmacologist, Otto Loewi, received a that giant neurons of some squids have axons as thick as 1 millimeter. At their
Nobel Prize for demonstrating that neurons laboratory in the seaport of Plymouth, England, Sir Alan Hodgkin (1914–1998)
communicate with one another by releasing and Sir Andrew Huxley (1917–2012) (FIGURE 1.24) would study squid fresh off
chemicals called neurotransmitters.
the fishing boat. They would isolate a single giant neuron from the squid and
test how the nerve impulse traveled along the axon. With such large axons, they
could pierce the axon with an electrode to measure voltage, and could even inject
different chemicals inside. They learned that neuronal firing is actually electro-
chemical (FIGURE 1.25). Voltage increases along the axon are caused by changes
in the membrane of the neuron that permit positively charged sodium ions (Na+)
neurotransmitter A chemical
substance used in neuronal to rush very quickly into the axon from outside the cell. Then, very quickly, the
communication at synapses. membrane changes again in a way that pushes positively charged potassium ions

(A) Hodgkin (B) Huxley (C) Hodgkin and Huxley’s squid

Brain

Neuron:
Cell body
© Keystone Press/Alamy Stock Photo

© PA Images/Alamy Stock Photo

Neuron:
Giant axon

FIGURE 1.24 First recordings from inside a neuron British physiologists, Sir Alan
Hodgkin (A) and Sir Andrew Huxley (B), earned a Nobel Prize for discovering how the
action potential works to conduct signals along the axons of neurons. They moved from
Cambridge to the coastal city of Plymouth, England, to record voltages from inside the
giant axons of freshly caught Atlantic squid (C).
Another random document with
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and approaching near, saw it was a fine cobra, about six feet in
length. The reptile had its head raised nearly a foot, and was making
slight darts at the fowls which ventured near, but unable to do much,
for as soon as it attempted to move, several pecks were made at its
tail. A friend who was standing near, knocked the snake on the head,
and immediately all the inhabitants of the poultry-yard set upon it and
tore it to pieces.
After nine hours’ severe poling and hauling, we came to a stop for
the night at 4.25 p.m. I was desirous to reach the Madihit, but the
men were too exhausted to proceed farther. We have not had a
bearing of Molu for some time; we are anxious for it, in order to
determine our position. I got out all the salt fish that was left, and
divided it among the men, to whom it was most acceptable, as they
had had nothing but rice for many days.
After we had finished our supper, I called the whole party together
to discuss our future proceedings. I explained to them the
information I had been enabled to collect from the Muruts; that we
might reach the Adang landing-place by water, but that as we
advanced the country became more difficult; and that, in the present
state of the river, it was impracticable to face the rapids; that if we
attempted the water way and failed, we must return home, as it was
quite impossible to walk along its bank; the hills were too lofty and
far too steep; in fact, no one would ever attempt it who knew
anything of the country. The second plan was to ascend the Madihit,
which could not be very far off—a mile or two—then leave our boats,
and walk across, the Murut guides assuring us that we could do the
distance in seven days. I put it to the whole party to consider. Musa,
after a short consultation, said they would prefer walking to facing
even seven days of such work as we had had; and as it was two or
three days from the Adang landing-place to the houses, it was better
to start at once from the Madihit, leaving the principal portion of our
heavy baggage with the boats. My heart was slightly despondent
yesterday, but to-day’s progress has completely cheered me up.
Just at sunset, we noticed a few flying foxes far up in the sky; they
seemed very numerous, but were almost beyond view. Sometimes,
however, they fly low, pursuing their onward course with steady
flapping wings. For two hours I watched them at the entrance of the
Sibuyau river, passing us by tens of thousands, and all flying in one
direction, doubtless towards some place where they rest at night.
While preparing their tents, my men disturbed a huge frog that
appeared about nine inches long; it was of a dark gray colour. I was
assured, however, that a full-grown frog was double this size, and if
one could judge of their dimensions by their noise, when they are
heard in a marsh, one might readily believe the Muruts’ account. I
remember hearing the late Mr. Hayes of Siam say that he had seen
them there with bodies as large as a full-sized dinner plate.
8th.—We found the Madihit to be two miles farther up on the left
bank. The main stream continues broad, and is furnished with
islands and immense pebbly flats, reminding me of the portion near
the Madalam at very low water. The hills are now at some distance
apart, allowing broader and flatter strips of cultivatable land along the
banks. Our last night’s resting-place was quite a level point; we
brought up a little above it, under a jutting portion of the bank, which
was higher, and near it was a beautiful natural bower which would
have afforded shelter for all our boats, had the stream risen high;
some of the men encamped in it. The Madihit, a short distance from
the mouth, is but a shallow, pebbly torrent; and a little more than half
a mile up we were obliged to leave our large garei, and take to the
sampirs, most of the men following along the banks; and now, less
than two miles farther, we have to leave our boats, and take to
walking.
We noticed a very handsome tree, whose trunk shoots out almost
horizontally from the banks: its branches rise perpendicularly into the
air, but let fall on either side rows of long drooping twigs, covered
with elegant foliage. It was loaded with fruit, whose long wings are of
a beautiful pink colour, and amid this forest it had a gorgeous effect. I
believe it is of the order Dipterocarpæ. There is another very
remarkable one which grows on rocky soil; its bark strips naturally
from the trunk, leaving a brightish brick-red stem.
We have passed yesterday and to-day much young jungle;
indeed, except where the hills are steep, there are few old trees. The
fish are very plentiful in the river, but the rapidity of the stream
prevents the net from acting properly. Just before entering the
Madihit, I noticed a range of high hills, bearing south-east, said to be
called the Paya Paya in Malay, the very difficult hills, and round their
base the Limbang ran.
No rain last night or to-day. I find it impossible to continue writing,
from the cloud of sand-flies that are tormenting me. Having made a
smoky fire, I am at last rid of them. I have divided the remainder of
the rice, and find that the careful men have enough for seven or
eight days, while the improvident have not enough for five; so that
they must carry sago; and, to my regret, Ahtan reports that all my
tinmeats are left at home, but I have enough biscuit for seven days;
in fact, I shall leave some behind me for the journey back. Our
sportsmen again missed both a deer and a pig; so that, after having
had every day many chances, nothing has been secured, though we
have with us two of the most famous hunters in the Limbang. I never
had a shot myself, as my heavy boat was generally behind the
others. The rock that forms the base of many of these karañgans or
pebbly rapids is a dark gray sandstone. By the barometer, we are
637 feet above the level of the sea, and as we have been toiling up
rapids since we left the Damit stream, it accounts for this elevation.
The men have been working away, forming a cache for the things
that we must leave behind: it is raised on four poles, so as to be
quite secure from pigs.
To show how extensively the Chinese formerly spread over the
country, I may notice that they had pepper plantations even up the
Madihit as late as the remembrance of some of the oldest Muruts.
 CHAPTER IV.
MY LIMBANG JOURNAL—continued.

Start on the Land Journey—Our Course—Ascend the Rawan Torrent

—Attacked by Hornets—Native Remedy—Severe Effect of Sting
—Disturbed by Ants—Japer left behind—Fresh Traces of
Strangers in the Forest—Appearance of the Country—Water
Snake killed—Our Adventure with One—The Swimming Cobra—
Romantic but timely Meeting—Story of Pangeran Mumein and the
Murut Concubine—Malay Revenge—Punishment of an Offender
—Cause of the Borneans being converted to Mahomedanism—
Capturing the Daughter of Johore—Independent Position of the
Pablat Borneans—Reach the Wax-seekers’ Hut—Flesh of the
Wild Boar—The Adang Muruts—Their Sumpitans—The Poison
on the Arrows—Melted in hot Water—Weapons purchased, not
made by them—Dress of the Muruts—Japer rejoins us—Continue
our Journey with new Guide—Method of catching Fish—Effect of
Loss of Blood by Leeches—Extraordinary Insect—The
Freshwater Turtle—Its Description—Curious Fly—Poisoning the
River—Getting short of Provisions—Galton’s Method of dividing
Food—Adopt it—Improvidence of the Malays—Cry of the
Wahwah—Rejoin the Limbang—Omen Bird—Prepare for
Enemies—Quarrel among the Guides—Divide the Party—Hard
Walking—The Otter—A Fight with my Dogs—Still beyond the
Mountain—Find good Huts—The Stragglers—The last of the
Food—Ascend the Mountains—Exhaustion of the Guides—The
Remains of the Ham—Its Effects—Reach the Summit—Descend
to the Farms—Meet the Adang Muruts—Hearty Welcome—
Names—Recent History of these Villagers—Kayan Attacks—
Driven from the Limbang—The Geography of the Country—The
Houses—Cold, and Fire-places—Arrival of my Followers—Sir
James Brooke—The Friend of the Aborigines—His Fame had
preceded me—How Reports spread—The Tigers’ Cave—Curious
Story.
10th.—I was so stung by hornets yesterday that I was unable to
write in my journal, and even now it is a work of difficulty. After some
trouble, I got the sampirs hauled up on the bank, to be beyond the
reach of freshes; and all the things we intended to leave behind were
carefully stowed away. The sago we left in the boats, none of the
men choosing to burden themselves with any of it, though I warned
them not to take less than seven days’ provisions, but all warnings
were useless.
Having divided the packages, we got away at 9.15, and started in
an easterly direction for about an hour and a quarter, then southerly
for about two hours, when we joined the small stream of Rawan. Our
course led us from one bank to the other of the Madihit, crossing and
recrossing it nearly every reach, a most fatiguing operation, and
trying to one’s feet, over slippery sandstone rocks and pebbles. The
forest, though principally consisting of small trees, is tolerably clear,
and presented no particular difficulties, nor was it so infested by
leeches as in the neighbourhood of Molu. At 12.30 we began to
ascend a steep valley (easterly), generally making the bed of the
Rawan our path. A mountain torrent never affords very easy walking,
and three of our men were so fatigued that at 3.15, after six hours’
walking, I consented to encamp.
It was in following the bed of the Rawan that I was stung. Notice
was given by the guide to leave the direct path, and we all did; but I
suppose some one disturbed the hornets, as they attacked me with a
ferocity that appears incredible: many flew at me, but two fixed on
my arms and stung me through my double clothing. They poised
themselves a moment in the air, and then came on with a rush that it
was impossible to avoid. The pain was acute, but I saved my face. I
tumbled down the steep bank in a moment, and, throwing aside rifle
and ammunition, plunged up to my eyes in a pool, until the buzzing
ceased, and the hornets had returned to their nests.
Some of my men were also stung; they squeezed a little tobacco
juice on the wounds, and they say they felt no further inconvenience.
I tried it about an hour afterwards, but it did me no good. I had no
idea that the sting of this insect was so severe: my right arm swelled
up to double its natural size, and was acutely painful; now, on the
second day, it is much less so, but as the swelling continues, it is
impossible to use it much. In the night we were again unfortunate,
being attacked by the selimbada, a most poisonous ant, which
quickly drove us from the tents, and compelled us to take refuge on
a small pebbly flat near the stream. Fortunately for us it was a fine
star-light sky, as we had to spend the remainder of the night there. I
have called the sting poisonous; it is scarcely that, as, though the
pain is acute, it has but little permanent effect. The little sleep I had
was due to the last bottle of porter that remained. My men were so
heavily laden, that had I brought a stock of drinkables, I could not
have had them carried. I have with me but a couple of bottles of
whisky and one of brandy.
Having put my right arm, the most painful, into a sling, I was
enabled to make a start with the rest of the party, except Japer, who
had an attack of elephantiasis. So leaving two volunteers with him,
we continued our course; they promising to follow as soon as
possible after the fit was over; they will easily catch our loaded men.
I would gladly have stayed behind myself, as my arms were in a very
unfit state to be used, and I was compelled to carry my rifle in my left
hand; but the fear of running short of provisions made me push on.
Having left the tents at 7.30, we soon commenced ascending hills.
Our course was at first S.W., but on reaching the summit of a ridge,
we followed it to the S.S.E. The guides had some difficulty in
discovering the path, which was completely overgrown. We found
traces, however, of recent visitors.
Leaving the ridge, we descended a stream called Patra, where
the guide said he wished us to remain while he went ahead to look
for the path; so that, very unwillingly, I brought up at twelve, and our
guide started to explore. One of the traces we found in the course of
our walk was quite fresh, a human footstep in the mud, not a couple
of hours old, and many broken branches. This caused as great
excitement as ever Robinson Crusoe could have felt when he
discovered the one on the sand; and the whole party collected to
examine it. One man ventured to observe, “Perhaps there are
enemies in the neighbourhood.” At this I laughed, and suggested it
might be a wax-hunter, who, having caught sight of us, has started
off to give notice of the approach of our formidable party: the twenty
muskets must make it look formidable to him.
Our path to-day was rather difficult, as we had to ascend many
steep hills, and sometimes to move along their faces, which is
tiresome walking: it is only on the ridges that our advance is
pleasant. We had a view of the Molu range through the trees, and
also saw many other mountain ranges, but no sign of those
extensive plains that we were promised; perhaps, however, we have
not penetrated far enough into the country. No portion of Molu
appears to drain into the Limbang, except through the Madalam, the
waters of its southern and western faces being carried off by the
Tutu and the Millanau into the Baram. I have been trying to find
some flowers for Mr. Low, but we have as yet seen none, except of
the most trifling description. The waters of the Patra are very muddy,
there evidently having been a landslip in its upper course, in fact I
observed one to-day that left the surface of a whole hill quite bare.
Our Muruts have just killed a large rock snake; they are now
cleaning it preparatory to its forming a portion of their dinner.
They surprised their snake basking in the sun and cut off its head:
but some time before we were ascending a waterfall, and while
looking at some flowers, a friend accidentally touched a gray rock-
snake that lay lazily extended across our path; I saw him spring
aside, and had but a moment to get out of the way, as the reptile
dashed past. On our return, while choosing a good pool to bathe, we
observed the snake, which was about twelve feet long, vainly
endeavouring to escape up a rock, and finding it could not, it made a
dash at us. We thought at the time that it meant to attack us, but
probably it was only an endeavour to effect an escape. We drove it
back with boughs to the deep pool before us, and then hurled large
stones at it till it seemed to be dead; but on descending to the water
to get out the body to examine, it had disappeared, diving into some
hole or crevice in the rock. The Malays have a great dread of most
kind of snakes, but this they especially dislike.
The cobra is, however, the most disagreeable one, particularly
when met with in the water; one tried to swim across the river just
below our boat; as soon as he saw us, he directed his course our
way, not, I believe, to attack us, but simply as something to rest on;
but my men in great alarm gave way, and escaped the beast. I have
heard men say, that the cobra will not be frightened back by beating
the water with the paddles, but must be killed, or it will enter the
boat; if it succeed in doing so every Malay would instantly spring into
the water and leave their canoe to drift away.
4 p.m.—Our guide has just returned, and brought with him the
man whose traces have kept us on the qui vive. Our guide says that
after walking about half an hour, he was hailed, and after a distant
parley, the man approached, and to their mutual astonishment they
found that they were countrymen, and that it was, in fact, his brother-
in-law, in order to visit whose sister he had joined our party, the gay
gentleman having left his young wife to go and seek his fortune and
another wife in the neighbourhood of Brunei. Such meetings
sometimes take place in romance, but seldom in real life. The
stranger is one of a large party who are now pig-hunting in the
neighbourhood, so that my chief fear—that we should not find our
way—has left me. We may get a deer, as these hunters have just
secured a pig. The new comer says he thought we were a party of
head-hunters, therefore, did not make himself known to us. He
followed us for some time, and felt a great inclination to send a
poisoned arrow amongst us; but that we were too strong. Very
fortunate for him he did not do so, or it might have fared ill with him.
11th.—Our guide left us again last night to join the wax-seekers,
and we are now waiting his return. I will enter the following story
while fresh in my memory. The Pañgeran Mumein, the Shabandar’s
eldest son, took a Murut girl as a concubine; she, however, was to
stay with her father up country. He paid, as a bri-an or marriage
portion, a pikul of bedil (133 pounds of brass guns). When she had
had her first child, he, as usual, got tired of her, and told her father
he did not want her any more, unless she liked to follow him to
Brunei. This was objected to. A few days after, he said he should fine
the father for not allowing him to take the girl to Brunei, where he
would have sold her as a slave. He made the father pay him back
the pikul of guns, as well as two more pikuls, and then said, “Your
daughter may marry whom she pleases.”
After some time, she married one of her own countrymen.
Pangeran Mumein hearing of this, in a most unaccountable fit of
jealousy, determined to kill them all, and gave orders to Orang Kaya
Upit to seize them. The Orang Kaya hid himself, so the Pangeran
was obliged to employ the Bisayas, who caught the husband and
brought him to the noble. He had him tied up to the Orang Kaya’s
landing-place, and there cut him to pieces with his own hand, taking
his head and giving it as a present to the Gadong Muruts. The father
and girl escaped. The Pangeran remains unpunished, probably
unblamed. Of course, there are two sides to the story: he declares
he had not separated from the girl.
The following is an instance of the punishment of an offender:—
Salam, of Bukit Manis, together with his brother and his son, was
accused of stealing buffaloes, and doubtless had done so, but acting
generally as an agent of Mumein’s in his rascalities. The Sultan,
wearied of the constant complaints against Salam, gave orders that
he should be put to death: his house was immediately surrounded
and fired into, and his brother and son killed. He then came out into
his garden, begging that he might be taken to Brunei to be executed;
but after they had induced him to throw away his sword, and fasten
his own hands with his handkerchief, so that he could not
immediately seize his weapon, they rushed in and cut him down.
Difference of rank, difference of treatment.
There is an old Pablat man with us named Bujang: he says that
the people of his kampong, or parish, as well as those of Burong
Piñg-e, descend from Muruts, and that they turned Mahomedans
shortly after they captured the daughter of the Sultan of Johore. This
is the tradition or history: they were, as usual, cruising down the Gulf
of Siam, looking out for prey, when they observed a prahu, gaily
dressed out with banners, pulling along the coast. They gave chase,
and soon came up with her, and found the daughter of the Sultan of
Johore, surrounded by a bevy of pretty attendants; they seized them
and carried them off to Brunei, and presented the lady to their chief,
who married her.
When the father heard of it, he sent a great deputation of nobles
to entreat the Murut to turn Mahomedan, and marry his daughter
according to the custom of that religion. He made no difficulty, but,
on the contrary, invited the nobles to remain and take wives in the
country. Many did, and it soon became a great rendezvous for the
Malays; in fact, the other twenty kampongs are descended from
odds and ends of strangers, together with their wives, taken among
the aborigines. The rajahs all say they are of Johore descent.
Bujang affirms that the Pablats muster nearly a thousand fighting
men; and that, with the Kadayans, or Mahomedans of the Hills, they
have an offensive and defensive alliance, which enables them to
hold their own, and not be treated as the other kampongs are. He
was very proud that they would not suffer the rajahs to take their
women, except with the parents’ consent. It is a fact that an
unbetrothed girl, of decent appearance, can never be kept by her
parents. A rajah sees her, and orders her to be sent to his harïm,
that he may honour her by taking her as a concubine. They,
therefore, betroth their children in childhood, and then they are
usually safe. He says, and the Bisayas have a similar tradition, that
Brunei was formerly a lake, that burst through into the sea near the
island of Iñgaran.
3 p.m.—We have only walked two hours and a half to-day, as,
when we arrived at the hut of the Adang hunters, our Muruts were
unwilling to go farther, hoping to come in for a very good share of the
flesh of the wild pig that was hung up around: so we stopped, though
anxious to get to a river. It is a great luxury, after a walk, to get a
bathe in a fresh, running stream. Now we are encamped on an
elevation of 3,000 feet, with only a very tiny spring of water near.
With the party of Adang Muruts there is an old chief who has agreed
to return with us to his village, leaving his young men to gather
honey and wax. The few Adangs that we have met have rather a
heavy, forbidding appearance, except the old chief and another.
Nearly all of this party of hunters were armed with sumpitans,
which were as usual of dark hard red wood, and had a spear-head,
lashed on very neatly with rattans on one side of the muzzle, and an
iron sight on the other. The arrows were carried in very neatly-carved
bamboo cases, and were themselves but slips of wood, tipped with
spear-shaped heads cut out of bamboo. The poison looks like a
translucent gum, of a rich brown colour; and when dipped into water
of a temperature of one hundred and fifty degrees, it began to melt
immediately; but on being withdrawn and placed over the flame of a
lighted candle, it instantly became hard again. The butt of the arrow
is fixed in a round piece of the pith of a palm, which fits the bore of
the blow-pipe. The natives say also, that the juice from one kind of
creeper is even more virulent than that of the upas. On inquiry, I find
that none of the people of these countries can manufacture the
sumpitan themselves, but purchase them from traders, who procure
them at Bintulu and Rejang from the wild Punans and Pakatans, and
are therefore very dear, and highly prized, and no price offered will
induce a man to part with a favourite sumpitan.
These Muruts were furnished with war jackets and helmets. The
former were well padded, and thickly covered over with cowrie
shells; the latter was of the same material, with flaps hanging, so as
to protect the wearer’s neck from poisoned arrows. I heard that the
supplying of cowrie shells formed a sort of trade in the Trusan
districts. These Muruts, like those seen near the coast, often wear
their hair tied in a knot behind, and keep it in its place by a great pin,
fashioned something like a spear-head both in size as well as in
appearance, which is made, according to the means of the wearer,
either of brass or of bamboo.
Our walk to-day presented no feature of interest: a stiff climb to a
narrow ridge, and then along it rising gradually to the hunter’s
temporary hut. We but occasionally got glimpses of the country
through the trees, and it presents one general view of forest,
covering hill and valley. If we are to take such short walks as these,
the journey will be an easy one indeed. I did not regret our little
progress to-day, as it enabled old Japer to come up with us, his
acute attack having left him. I never was in such a country for bees:
they everywhere swarm in the most disagreeable manner, and ants
and other insects are equally numerous.
12th.—There was much thunder and lightning last night, with rain;
but, notwithstanding the continued drizzle, I got away before seven,
the chief sending his son. Our walk was more direct to-day, as the
path was well known to the guide, he having just used it. The course
was generally in a S. by E. direction; on the whole, over a very
similar country to yesterday; got no view whatever. We crossed
numerous streams, as the Sañgin, Ropan, and Gritang, and have
now stopped at a very pleasant one, the Lemilang, encamping on a
high bank about 100 feet above it (900 feet above the level of the
sea). We have done more work to-day, walking more than six hours.
Noticed some tracks of the tambadau. The Kalio hills, perhaps 5,500
feet high, have been on our left all day. The most active man I have
ever seen is a young Murut, who walked part of the way with us to-
day: he had a perfect figure, and is the only pleasant-looking man
besides the chief that belonged to the party of hunters.
13th.—I have little to enter, except that we walked five hours
south-east, then four east, then another north, passing over the
same kind of country as usual. The only noticeable event was
catching some fine fish in the Madihit, just below the junction of the
Rapaw and the Obar. The operation is simple: stones are hurled into
a pool in the river; the fish fly for concealment under the stones and
to the holes in the rocks; the men jump in and soon bring them out of
their hiding-places. They caught five large and above a dozen small
fish; one was eighteen inches long, and very fat. Ahtan was
successful in getting a fine one, which he is at present engaged in
cooking, so that we shall each dine off half of it to-night. The scales
are very large; it is the same kind as I have previously noticed in the
Madalam and Tampasuk.
After this good fortune, we crossed the river, and ascended the
steep hill of Pakong Lubfing, till we reached a little rivulet running
into the Obar. I really believe that the great loss of blood from
leeches is the cause of the faintness I feel to-night; I have pushed off
hundreds, and the wounds continue to bleed for some time. Perhaps
this, and the very heavy day’s work, may account for the strong
disinclination I feel to write my journal. The Muruts are beginning to
be full of dismal stories of enemies, saying they have suffered much
lately from the attacks of neighbouring tribes, who have shot at them
with their poisoned arrows, many dying, including some women and
children. However, there is not much reliance to be placed in such
stories when told in the woods. During the evening a very
disagreeable-looking insect kept attacking my candle. It looked like a
dead but branched twig, and an ordinary observer would scarcely
notice the difference till he saw it move; its legs are represented by
four dead branches. There is another more common, that has wings
like a couple of bright green leaves. (Our camp 1,500 feet.)
The Murut guides are but scantily provided with food, and search
eagerly for snakes, tortoises, and fresh-water turtle. The last would
be a grand find, as it is often three feet long, without including the
head and neck: its upper shell and chest are covered with a soft
skin, large openings being left for the limbs, each of which has five
fingers, three of them armed with thin but strong claws, those on the
thumbs being the largest; the fourth and fifth fingers have no claws,
and their joints are such as to admit of much movement in a lateral
direction, particularly the fifth, which, when the fin is distended, is at
right angles to the first three. The back is of an uniform dark gray;
the upper part of the head dark olive green, mottled with yellow
spots; the nose is prolonged into a non-retractile, pointed soft snout,
about an inch in length, and the nostrils are rather large; the tail is
about three inches long and very fleshy. Its stomach, when opened,
often contains many kinds of fruits, particularly wild figs and some
large kernels, which, though not harder than filberts, pass unbroken
through his horny jaws, which appear, however, well adapted for
cutting up food of this consistence. I may add, that the lower part of
its neck is covered with a fold of loose cartilaginous skin, into which it
withdraws its head, with the exception of the pig-like snout.
14th.—Walked to-day seven hours in a general east-north-east
direction; in fact, from the range that separates the Madihit waters
from the Limbang, the Adang mountains were pointed out, bearing
east-north-east. We had a distant view of some high peaks, bearing
due south, said to be those in the interior of Baram. We had also a
tolerable view of the Molu range, which enabled me to fix our
position. I found to-day, just as we were crossing the ridge, one of
the most curious insects I have ever seen; it appeared like a gigantic
moth, above four inches in length, and was of a brown colour, with a
band of bright green just across its neck; although it had the look of a
moth, on closer examination it looked like a great horse-fly. I have
little to enter to-day, as the walking has been over the same style of
ground, and there has been no incident to vary the ordinary routine.
We pass a good many abandoned huts, with an occasional deer-skin
hanging up to dry, the mark of the hunter; and, to our great
disappointment, we find the same men have been poisoning the river
with the beaten out roots of the tuba plant, capturing all the good fish
near the encampment. Some of the hungry ones strayed farther
down, and saw several of a very large kind, but they escaped into
such deep holes that it was impossible to get them out.
I am afraid our provisions are drawing to a close. I see some
hungry looks and other symptoms of uncomfortableness among
about half the men—all their own fault. The careful have still two
days’ rice; three, if they did not assist their companions: the greedy
ones have been trusting to my biscuits. However, as I could not sit
down to my dinner of plain boiled rice with so many envying me, I
told my servant to take sufficient for ourselves, and then divide the
remainder of the biscuits into twenty-three heaps. I remembered
Galton’s plan, and making one of the men turn his back to the little
piles, I pointed to a heap, and he cried out a name; so that they were
fairly and without favour distributed.
The non-careful men were very disappointed with this plan; they
thought I should have only given those that had no rice a share, but
that I explained to them would be encouraging improvidence. A
similar thing occurred when I lost my boat at Molu. The men, rather
than carry rice, preferred trusting to jungle produce, vainly hoping we
could do three days’ work in one. We are encamped on the Urud, a
tributary of the Limbang. The highest point crossed to-day was 2,500
feet: our present elevation is about 1,400 feet, after many ups and
downs. One of the most melancholy sounds in the forest is the cry of
the wahwah, and after sunset it sounded near us, to be answered by
a disconsolate companion not very far off.
15th.—An hour’s walk brought us to the Limbang, whose bed is
here, perhaps, seventy yards wide, very shallow, not reaching to the
hip. It flowed from the Siliñgid mountains, and is said, after skirting
their western face, to turn to the south-east to its sources in Lawi. Si
Nuri, one of our guides, pretended to hear a bad bird, and wanted to
return, though we were all struggling against the stream in the
middle of the river. As this was the second time he had done it to-
day, I would not stand it, so ordered him over; however, to satisfy
timid minds, I had the guns discharged, cleaned, and reloaded. He
said his añgei (omen bird) told him there was fighting at his house.
We then continued to Suñgei Kapaw, where we stopped to eat our
breakfast. The old chief’s son was very much astonished by the
rapidity and accuracy of fire of one of the revolving carbines. He had
never seen any fire-arms, unless they were common flint muskets.
Shortly after again starting, I observed a commotion among the
guides, and was told that the new man and Si Nuri, his brother-in-
law, wanted us to take a roundabout way, to avoid a country
swarming, they said, with enemies, who would shoot at us from the
jungle. This would entail the loss of another day, and the climbing of
an extra high mountain. We preferred the enemies to the extra work,
as our food will be all finished to-morrow; so the two guides left us,
and one of the remaining Adang men undertook to find a road. It
struck me afterwards that it was merely a dispute about whose
house we should go to first. The Orang Kaya Upit and four followers
wanted naturally to go to their relatives, while Si Nuri was anxious to
get as quickly as possible to see his wife and relations; but as he
could speak little Malay, I did not understand his explanation.
Our new guide, turning north for a short time, soon brought us to
the Ropo, a branch of the Limbang, which is, perhaps, a third less
than the main stream. From thence we continued our course north,
climbing up a steep mountain, about 2,500 feet in height; then
turning to the north-east, we continued that course till 4.10 p.m.,
when we reached the Bapangal stream. Including all stoppages and
petty rests, we walked about six hours and a half, and made about
eight miles. From near the summit of the high hill we had a view of
some fresh clearings which appeared about seven miles off in a
north-east direction. The villages are said to be on the other side of
the clearings, beyond a low hill. As we must have approached these
farms three miles, we cannot have more than eight miles of walking
before us; at all events, I shall do my utmost to reach the houses to-
morrow night, whether the men follow or not, as after the biscuit
division last night I have only rice for to-morrow, coffee, and the
remains of a bad China ham. Just as we were crossing the Ropo, an
animal slipt quietly into the water, which I think was an otter. A few
days before leaving my house, I was witness of a desperate fight
between my dogs and a very fierce female otter. They had
surrounded a pool, and kept her in it, but as it was very shallow a
dog would dash in and make a snap at her; at last she seized one by
the nose, and would not let go; the dog, a very small English terrier,
did not utter a cry, but struggled towards the bank, when the whole
pack fell upon the enemy, and tore it to pieces. Yet in death, it did not
let go its hold, and to free the dog’s nose its jaws had to be forced
open with a spear-head. (Resting-place 1,300 feet.)
16th.—Walking on till 4.30, after eight hours of regular hard work,
we have not reached the Adang villages, but have a prospect of
doing so to-morrow. We have only the range of mountains to cross:
yesterday’s clearings and low hills we have passed. When at 4.30 I
sat down on a rock, with the rain pouring down in torrents, and the
men standing shivering around me, I could not but feel a little
despondent when I asked the guide where were the houses, and he
replied over that high mountain: but observing the men were
watching me, I burst out laughing, a very unnatural sound it seemed,
and said I thought the Adang dwellings had vanished in the clouds.
Presently the Orang Kaya Upit came stealing back and said he
had found two fresh huts that belonged to a Murut tribe with whom
they were at war, that we must instantly retrace our steps half a mile
at least, and pitch our tents there. I was in a very bad mood to
receive such a proposition and told him that if the devil himself lived
in those huts, I would make him give me a share of them to-night,
and told him to lead the way. Tired as he was, I could scarcely
persuade him that I was in earnest; but, calling on my Malays to
follow, we soon made an advance towards the dreaded spot, and
then, after twice attempting to lead us astray, the Orang Kaya
brought us to the huts, which had evidently not been vacated many
hours; perhaps after they had got a distinct view of our party. The
ashes of the fires were still warm, and we had no difficulty in
rekindling them.
As the rain continued to descend in torrents, we were pleased
indeed to find warm dry quarters, and having extended the tent from
one of the huts, there was room for all. As old Japer and four of the
men had not reached us by half-past five, we discharged our fire-
arms one after the other to give them notice of our whereabouts, in
case they were wandering within hearing. It likewise served the
purpose of intimating to the Muruts concealed about that we were
not head-hunters, these seeking concealment and not publicity. My
best men having assisted the improvident have but a poor meal to-
night. Our walk to-day was over very uneven ground, steep hills and
numerous streams, among others the true Adang. The last candle is
going out, so I finish.
17th. and 18th.—My journal did not reach me till this evening, so I
must enter the two days together. Japer did not make his
appearance during the night. At dawn, finding I had a small biscuit a-
man left, I shared them out, and then started off at 6.30, hoping to
reach the houses, and from thence send assistance to the
stragglers. I gave notice that I expected no man to wait for his
neighbour, but to push on and do the best he could for himself. Our
route at first lay over the low ground that skirts the foot of the
mountain, then up a tolerably easy ascent, one of the spurs of the
Adang range, that appears to run north and south. After half an
hour’s walking, I observed the guides dart to an old tree; I followed,
and we were soon occupied in devouring mushrooms; after this
slight repast, we continued our course.
About 10 a.m. four of the Muruts dropped their heavy burdens,
intending to return for them next day, and started off at a running
pace. I was following when I heard Ahtan’s soft voice say, “May I
come too, sir?” I said, “Yes, if you can keep up.” This was very well
for a few hundred yards; when, to my surprise, I found a Murut drop
behind, then the Orang Kaya; at last the remaining guide stretched
himself on his back, and said he was done up. I tried all I could to
rouse them, but with no effect, until the arrival of the fifth Murut, who
was one of the baggage train, with half a ham, the last of the
provisions, and not good food unsoaked, without a drop of water for
miles, but I told the Muruts that if they would try and reach the top of
the mountain I would then cook and divide it.
This roused two, and after much trouble we arrived at the first
peak, there we stopped; a fire was lit, and with a chopper we divided
the ham into great bits. While cooking we shouted to the other three
Muruts to make haste, and presently we heard the Orang Kaya
begging us not to begin till he arrived. As none of the Malays would
touch it, we divided the meat into seven portions, and they were
soon devoured skin and all, and the bones crushed for the marrow,
Ahtan and myself eagerly joining in the meal and securing our share.
For a fortnight we had lived on very unsubstantial food, and the great
exercise we were taking gave us appetites that scorned being
satisfied with simple boiled rice.
A few minutes after the meal was over, the Muruts gave a grunt of
satisfaction and started off, and continued ascending till we had
reached the height of 5,000 feet; where we struck off to the north-
east, down to a dashing torrent, one of the feeders of the Trusan
river; here we bathed, then off again up a very steep hill. This was
too much for Ahtan. He turned an imploring look on me, and seemed
ready to burst into tears; but I spoke to him very angrily, asking him if
at the last moment he intended to disgrace himself, and relieving him
of my sword, he pushed on, and in a few minutes we were at the
summit. Here we sat down for half an hour; then on, generally
descending, sometimes excessively steep, and it was with the
greatest pleasure that at 5.30 p.m. I found myself at the edge of a
great clearing, still burning, with two village houses at its eastern
verge.
Our guides had a short consultation as to how we should
announce ourselves to the people, whom we could perceive working
on the farms. At length Orang Kaya Upit shouted out. There was
immediately a great commotion among the Muruts, but they soon
recognized the voices, and, as we forced our way through the
tangled trees that were felled in every direction, they came forward
to welcome my companions. They soon explained who I was, and I
was received with great civility and with symptoms of much curiosity.
No wonder; they had often heard of the white man, but I was the first
specimen that had ever reached their country. Half way across the
clearing we met crowds of women and children collected to hear the
news, all of us being supplied with sugar-cane to amuse us while a
meal was prepared.
 At 6 p.m., just at sunset, we reached the houses, and pleased
was I to be able to stretch myself alongside of one of their fires. This
was the hardest day’s walking we had had, ten hours of actual work,
crossing a double range of 5,000 feet in height; and I was both
hungry and tired, and exceedingly enjoyed the meal they prepared
for us. The only drawback was, that there were but Ahtan and one
Malay to enjoy it with me; two, however, came in afterwards, but
fourteen succumbed to the fatigue and did not show themselves.
However, I was relieved of much of my anxiety by the Chief
promising to send out a party at early dawn with provisions.
The Adangs were rejoiced to see my companions, plenty of arrack
was produced, and the news freely given and received. They kept it
up to a very late hour. One family heard of the death of a near
relative, and their wailing in the next room was very painful.
Gradually, however, the spirits they drank began to have an effect
upon my companions, and they all dropped off to sleep, in which I
was but too glad to join them.
The only one of my Malay followers who kept up with me, and
who, though one of the most improvident, had yet refused to touch
the ham, although no other Mahomedan was present, was called by
his companions Si Kurap, on account of a skin disease which
covered his body. The Malays very often give nicknames, referring to
some personal quality or defect, which, as in the above case, takes
the place of the real name. One man I knew, who lived at the town of
Sakarang, was called Sauh Besi, the iron anchor, on account of his
great strength, and immense muscular development. Others are
called from their low or high stature: Si Buntak, Mr. Short, or Si
Panjang, Mr. Long, or Si Juling, Mr. Squint, from having that defect in
his eyes. A very stout Chinese trader went always by the name of
Baba Lampoh, or Mr. Fat. The women are called after the same
fashion. Si Buntar, or the round, was the name given to a baby, on
account of its plump appearance; and when she grew to be a woman
and became thin, she never had another name. Some are called
Sulong and Bongsu, the eldest and the youngest born, to the day of
their death, never receiving any other appellation. They have also