Handbook of Perception. Tasting and Smelling

314 Pages·1978·4.318 MB·English

by  Edward C. Carterette (Auth.)

HANDBOOK
OF PERCEPTION
VOLUME VIA
Tasting and Smelling
EDITED BY
Edward C. Carterette and Morton P. Friedman
Department of Psychology
University of California, Los Angeles
Los Angeles, California
ACADEMIC PRESS New York San Francisco London 1978
A Subsidiary of Harcourt Brace Jovanovich, Publishers
COPYRIGHT © 1978, BY ACADEMIC PRESS, INC.
ALL RIGHTS RESERVED.
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Library of Congress Cataloging in Publication Data
Main entry under title:
Tasting and smelling.
(Handbook of perception ; v. 6A)
Includes bibliographies and indexes.
1. Taste. 2. Smell. I. Carterette, Edward C.
II. Friedman, Morton P. [DNLM: 1. Smell. 2. Taste.
WL700H234 v. 6A]
QP456.T37 596\01'826 78-18286
ISBN 0-12-161906-0
PRINTED IN THE UNITED STATES OF AMERICA
LIST OF CONTRIBUTORS
Numbers in parentheses indicate the pages on which the authors' contributions begin.
LINDA M. BARTOSHUK (3), John B. Pierce Foundation Laboratory and
Yale University, New Haven, Connecticut 06519
M. G. J. BEETS (245), International Flavors & Fragrances (Europe),
Liebergerweg, Hilversum, The Netherlands
LLOYD M. BEIDLER (21), Department of Biological Science, Florida
State University, Tallahassee, Florida 32306
WILLIAM S. CAIN (197, 277), John B. Pierce Foundation Laboratory and
Yale School of Medicine, New Haven, Connecticut 06519
ROBERT C. GESTELAND (259), Department of Biological Sciences,
Northwestern University, Evanston, Illinois 60201
DONALD H. MCBURNEY (125), Department of Psychology, University of
Pittsburgh, Pittsburgh, Pennsylvania 15260
HOWARD R. MOSKOWITZ (157), MPi Sensory Testing Division of MPi
Marketing Research, Inc., New York, New York 10021
CARL PFAFFMANN (51), The Rockefeller University, New York, New
York 10021
SADAYUKI F. TAKAGI (233), Department of Physiology, School of
Medicine, Gunma University, Maebashi, Gunma-ken, Japan
IX
FOREWORD
The problem of perception is one of understanding the way in which the
organism transforms, organizes, and structures information arising from
the world in sense data or memory. With this definition of perception in
mind, the aims of this treatise are to bring together essential aspects of the
very large, diverse, and widely scattered literature on human perception
and to give a precis of the state of knowledge in every area of perception.
It is aimed at the psychologist in particular and at the natural scientist in
general. A given topic is covered in a comprehensive survey in which
fundamental facts and concepts are presented and important leads to
journals and monographs of the specialized literature are provided. Per­
ception is considered in its broadest sense. Therefore, the work will treat
a wide range of experimental and theoretical work.
This ten-volume treatise is divided into two sections. Section One deals
with the fundamentals of perceptual systems. It is comprised of six vol­
umes covering (1) historical and philosophical roots of perception, (2)
psychophysical judgment and measurement, (3) the biology of perceptual
systems, (4) hearing, (5) seeing, and (6) which is divided into two books
(A) tasting and smelling and (B) feeling and hurting.
Section Two, comprising four volumes, will cover the perceiving or­
ganism, taking up the wider view and generally ignoring specialty bound­
aries. The major areas include (7) language and speech, (8) perceptual
coding of space, time, and objects, including sensory memory systems
and the relations between verbal and perceptual codes, (9) perceptual
processing mechanisms, such as attention, search, selection, pattern rec­
ognition, and perceptual learning, and (10) perceptual ecology, which
considers the perceiving organism in cultural context, and so includes
aesthetics, art, music, architecture, cinema, gastronomy, perfumery,
and the special perceptual worlds of the blind and of the deaf.
The "Handbook of Perception" should serve as a basic source and
reference work for all in the arts or sciences, indeed for all who are
interested in human perception.
EDWARD C. CARTERETTE
MORTON P. FRIEDMAN
xi
PREFACE
Nearly two and one half millenia ago Democritus explained taste qual­
ities by their atomic shapes—angular shapes gave rise to sourness, spher­
ical shapes to sweetness, and spherical shapes with hooks to bitterness.
This theory of molecular feature detection has a very modern ring! In­
deed, there is often a close relationship between the molecular structure
of sweet and bitter compounds (Beidler, p. 45, this volume). But De­
mocritus was far ahead of his time for, as Bartoshuk points out in her
History of Taste Research (Chapter 1) "little real progress by today's stan­
dards" was made between the fifth century B.C. and the nineteenth cen­
tury. Bartoshuk recounts the rise of the psychophysics of taste in Ger­
many in the nineteenth century. Later there was a general decline of
psychophysics generally but following World War II there was an expo­
nential increase in knowledge of the sensory coding of taste. Research
was stimulated by the "new psychophysics" of S. S. Stevens beginning
about 1960, a period which Bartoshuk calls the "Renaissance of Human
Taste Psychophysics."
From antiquity to now the intersection of all descriptions converges on
the homely four qualities: salty, sour, sweet, and bitter. Most of what we
know about taste mechanisms has come from studies of the electrical
events associated with the taste cells and their innervating axons when
various chemicals are placed on the tongue. Beidler's review of the
Biophysics and Chemistry of Taste (Chapter 2) cautions us that the electrical
record of a single fiber of the taste nerve bundle reports information from
many taste cells of several taste buds. Studies of taste stimulation and taste
cell transduction have not yet revealed the exact nature of the molecular
structure of the membrane, so that we can only surmise how chemicals and
receptor sites interact. Nonetheless, Beidler argues, the design of exper­
iments and prediction of events have been facilitated by a simple quanti­
tative adsorption theory of taste cell stimulation which he stated in 1954.
Beidler touches briefly on the (unsolved) problem of isolation of taste
receptor proteins and ends with a discussion of the properties of chemicals
associated with salty, sour, sweet, and bitter.
XI11
XIV PREFACE
The river is within us, the sea is all about us;
The sea is the land's edge also, the granite
Into which it reaches, the beaches where it tosses
Its hints of earlier and other creation:
The starfish, the hermit crab, the whale's backbone;
The pools where it offers to our curiosity
The more delicate algae and the sea anemone*
_T. S. ELIOT
PfafTmann in Chapter 3, The Vertebrate Phytogeny, Neural Code, and
Integrative Processes of Taste, treats us to an extensive review of one of
the chemosensory systems, taste, and its phylogenetic basis in vertebrates.
In vision and hearing, both receptor complexity and the amount of
cortical representation increase from fishes to birds to mammals, a con­
trast with ''the sense of taste which has undergone relatively little evo­
lutionary development as regards the structure of the receptor and the
morphology of the central neural connection subserving their reception
functions [p. 65]/' PfafTmann covers the central neural taste system in­
cluding its involvement in discrimination and conditioning. Recent studies
of neural taste coding show that there exist some four basic clusters of
sensitivity in animals, thus supporting the views of early sensory psychol­
ogists and physiologists. The final section discusses the significance of
taste stimulation in eliciting hedonic responses of animals and men.
Flavor is a complex of sensations and includes not only taste but those
of pressure, pain, touch,and smell. McBurney in Chapter 4, Psychological
Dimensions and Perceptual Analyses, restricts his survey to the nature of
taste qualities, the psychophysical methods of studying them, and the
influence on taste sensation of factors such as intensity, duration and area
of stimulation. The important phenomenon of adaptation is well covered,
with attention to the role of water, a substance now known to be a taste
stimulus, thereby complicating earlier views of adaptation and cross-
adaptation of taste.
From simple taste to complex flavor we turn with Moskowitz in Chap­
ter 5 to Taste and Food Technology: Acceptability, Aesthetics, and Prefer
ence. Hedonic tone means roughly degree of pleasantness or unpleasant­
ness. The hedonics of simple Aristotelian taste stimuli in solution—sugar,
acid, salt, quinine—do not tell much, alas, about the hedonics of food.
The overall judgment of pleasantness of a food depends on color, odor,
taste, texture, aroma, and temperature, and the relative weights of these
aspects will vary with a particular food. Thus, the acceptability of tuna
fish depended, in order of importance, on appearance, taste, and texture.
* [From T. S. Eliot, 'The Dry Salvages," in The Four Quartets, 1968. Reprinted by
permission of Harcourt Brace Jovanovich.]
PREFACE XV
Food attitude (preference) measurements seem to escape the specificities
of the laboratory. But preferences for individual foods are only one aspect
of the criteria for planning an entire menu. Moskowitz discusses the effect
of satiety and hunger on preference, the development of preferences, the
practical assessment of taste intensity, and the development of descriptive
systems for capturing the nuances of taste. And, at the end he suggests
ways of optimizing flavor and taste, an important notion theoretically and
practically.
From antiquity, smell has been an active field of research but only
recently have the workers persevered, says Cain in his History of Research
on Smell (Chapter 6). It was early understood that olfaction operated
either at a distance or in contact, and the correlations of odor with health,
disease and sexual attraction were catalogued. From Aristotle to the
twentieth century only Linnaeus's (1752) sevenfold classification of odors
survived to form the basis of Zwaardemaker's (1895) ninefold system.
Zwaardemaker's introspection was replaced by Henning's (1916) odor
prism based on (flimsy) experimental evidence. A contemporary failure is
Amoore's (1962, 1964) system elevated on assumed chemical properties
of odorants.
So, what is the stimulus for olfaction? Proposals for the physicochemi-
cal basis of olfaction make a long list, but do not answer this vital ques­
tion. Zwaardemaker's importance in odor research—on theory and
methods—justifies the extended coverage it receives. The final section
sketches the research leading to the conclusion "that information about
perceived odor quality can be revealed rather easily by neurophysiologi-
cal techniques." But poor success has ". . . obtained so far in pursuit of
the physicochemical correlates of neurophysiological responses."
The Biophysics of Smell (Chapter 7) is treated very succinctly by Takagi.
Apparently, only the olfactory cell among those of the olfactory
epithelium generate information about adsorbed molecules, although the
newly found blastema cell may alter this view. As with other receptor
systems, a variety of electrical potentials can be recorded from olfactory
cells. Some five EOG (electro-olfactogram) types are known and in­
clude the familiar on, off, and on-off types. Receptor potentials do not
arise unless odorous molecules contact receptors and recent work may
hold the key to clarifying how properties of the receptor site may suit the
molecular shape of odorants. The technique used is analogous to adapta­
tion methods used so widely in work on other senses. Takagi briefly sum­
marizes the mechanisms of the EOG and concludes that in the olfactory
cells and in the pathways of the olfactory nerve, discrimination is not very
selective.
According to Beets (Chapter 8, Odor and Stimulant Structure)
". . . the question whether a relationship between molecular structure
XVI PREFACE
and odor exists is meaningless. The only legitimate question is whether it
is simple enough to be detected.1' What the molecule knows is its struc­
ture and during the period of adsorption interacts with a site. Olfactory
discrimination is pattern recognition whose efficiency depends on the
match of molecule and site. The principles of molecular structure and
olfactory response are not the same for all molecules. To show the different
structural criteria Beets exhibits ketones, musks, and hydrocarbons; and
he mentions that entantiomers may elicit significant odor differences. In a
few cases1'. . . interesting relationships between olfactory response and
physical characteristics have been found."
Olfaction is the most illusive and mysterious of the senses says Geste-
land (Chapter 9, The Neural Code: Integrative Neural Mechanisms). The
central problem is how to represent olfactory space. The evidence is
against a single modality or continuum of quality. Yet it is not reasonable
to assign a modality to each distinguishable odor. So, Gesteland suggests
that the affinity of a receptor for two different odors may be very close.
This notion would lead to, say, a number of continua (modalities), thus
giving fewer modalities than odorants. "This does not put much of a limit
on the variety of theories which can be invented to account for olfac­
tion . . . [nor are there] any specific enough to be put to critical exper­
imental test [p. 260]."
Gesteland details the several electrophysiological methods for studying
the receptors where the relations of neural activity and odor properties
"should be transparent." Several response measures yield similar results
for all cells if measured over a range of stimulus intensities, as recent
work shows. One outcome is that different cells respond optimally to
different intensities, and some cells are very sharply tuned. Also, all cells
respond with varying degrees of selectivity to many odorants. This leads
to the notion that a given odorant at a fixed intensity maximally activates a
small proportion of receptors, inhibits another small proportion, tran­
siently activates a larger proportion, and elicits little or no action from the
majority of receptors. It is the task of the nervous system to integrate the
meaning of these weak patterns.
At the olfactory bulb, neurons may be spatially selective for stimuli.
Certainly anatomy shows many possibilities for synaptic interaction, in
sharp contrast to the independence of olfactory receptors. Curiously the
highly connected mitral cells show spike activity patterns like those of
primary nerve fibers. "Why all of this structure for so little apparent
information processing?" The final section states the little that is known
about the central coding of olfactory information, "Above the level of the
bulb, [where] olfactory physiology rests squarely on the work of the
behaviorists."
PREFACE XV11
Cain's Chapter 10 on The Odiferous Environment and the Application of
Olfactory Research is the homolog of Moskowitz' chapter on taste and
food technology in Part II. At the heart of controlling odor pollution by
industry are the questions raised for the air-quality engineer. He must
know whether an effluent is an odorant, how odor depends on concentra­
tion of the odorant, and how odors adapt, irritate, mix, can be masked or
modified. These are questions for the psychophysics of odor, which leads
Cain to provide the psychophysical foundation for applications of the
answers to the questions. Cain deals with chemical signals which warn,
aid in communication and regulate physiology. Remarking on the perva­
sive role of odorants on the behavior of animals at every level of
phylogeny, Cain says, "If man does not communicate via chemical se­
cretions then he is an exception to a very general rule." In the context of
language and civilization man does not seem to be an osmatic animal.
Financial support has come in part from The National Institute of Men­
tal Health (Grant MH-07809), The Ford Motor Company, and The Re­
gents of The University of California.
Editors of Academic Press both in New York and in San Francisco have
been extremely helpful in smoothing our way.
Chapter 1
HISTORY OF TASTE RESEARCH*
LINDA M. BARTOSHUK
I. Introduction 3
II. Taste Research from the Time of the Ancient Greeks to the Late
Nineteenth Century 4
III. The Late Nineteenth Century: Kiesow and Öhrwall 7
IV. The Early Twentieth Century: Final Contributions from Classical Taste
Psychophysics 10
V. Electrophysiological Techniques: Sensory Coding of Taste Quality 12
VI. Renaissance of Human Taste Psychophysics 14
References 16
I. INTRODUCTION
This chapter traces specific periods in the development of our under­
standing of the taste sense, with particular emphasis on views of taste
quality. The first and longest period—from the fifth century B.C. to the
nineteenth century—displayed little real progress by today's standards.
Most of the concern during this time that seems meaningful to us now
focused on the number of elementary taste sensations and on the identifi­
cation of taste receptors and their neural connections. During the scien­
tific revolution of the nineteenth century, the establishment in Germany of
experimental psychology as an independent discipline led to great pro­
gress in the psychophysics of human taste. The two world wars marked
the end of German dominance of psychophysics. At the same time, new
electrophysiological techniques, which permitted recordings from single
fibers in taste nerves, stimulated interest in the study of taste across
species. This change in emphasis hastened still further the decline in
human psychophysics. The years following World War II were enor­
mously productive ones for understanding the sensory coding of taste. In
* This work was supported in part by National Institutes of Health Grant ES 00880.
Thanks to Ferenc A. Gyorgyey and Joseph E. Hawkins for help with historical material.
HANDBOOK OF PERCEPTION, VOL. VIA Copyright © 1978 by Academic Press, Inc.
All rights of reproduction in any form reserved.
ISBN 0-12-161906-0