British JournaI of PsychoIogy (1992), 83, 449-471 Printed in Great Britain 449

0 1992 The British Psychological Society

Motion sickness and perception : A reappraisal

of the sensory conflict approach

Lucy Yardley*
Medical Research ,Council Human Movement and Balance Unit, Section of Neuro-otology,
National Hospital, Queen Square, London W C l N 3BG, UK

This review examines the role of activity and perceptual learning in motion sickness
by means of a survey of the two kinds of recent research relevant to this topic. The
first is a body of literature concerned not with motion sickness as such, but with
perception of orientation and self-motion under the conditions of ‘sensory conflict ’
which are thought to provoke motion sickness. The second consists of
investigations into the prediction and prevention of motion sickness itself. A major
weakness is identified in the methodologies employed in both types of research:
namely, a neglect of the way in which responses to unusual and disorienting
environments, whether nauseogenic or not, may be affected by the activities, skills
and strategies of the perceiver. New directions are outlined for future research into
immediate reactions and longer-term adaptation to such environments.

‘Motion sickness ’ is the blanket term commonly given to the syndrome provoked by
many forms of travel, by fairground rides, and by various unusual forms of motion
which have been developed in the laboratory. Two partly independent classes of
symptom can be distinguished. The first group consists of the direct consequences
of disruption to perceptual and sensorimotor activities involving the vestibular
system, such as disorientation, disequilibrium, and inappropriate vestibulo-ocular
or vestibulo-spinal reflexes. The second group embraces a constellation of mainly
autonomic symptoms (commonly including pallor, drowsiness, salivation, sweating,
nausea and vomiting) which also appear to have a perceptual origin, since they are
triggered by the same unusual motions or disorienting perceptual conditions. Almost
all individuals eventually adapt to motions or situations which initially provoke
sickness ; continued exposure to a particular nauseogenic environment leads to a
gradual reduction in the disorientation and associated symptomatology, usually
culminating in recovery of both sensorimotor control and well-being. However, the
speed and pattern of adaptation can vary widely between individuals and in differing
environments.
Since susceptibility and adaptation to motion sickness are intimately linked to the
perception of orientation and self-motion, hypotheses concerning the onset and
cessation of motion sickness also serve as a useful testing-ground for the predictions
derived from more general theories of perception and perceptual learning. Some
fundamental theoretical issues are thus addressed in this review through a critique of
* Requests for reprints.

19 P S Y 83
450 Lmy Yardley
the ideas and research associated with the ‘sensory conflict’ theory of motion
sickness. In particular, this paper seeks to explain and give structure to the, hitherto
unstated, change in emphasis in research into orientation which has been occurring
gradually over the past decade. The motivation for this shift, from a preoccupation
with psychophysical quantification of responses to sensory stimuli towards a
growing interest in sensorimotor activities and capabilities, can be traced to the
investigative findings and frustrations described below.
Since the mid-l970s, perception of ‘sensory conflict ’ induced by exposure to
unusual combinations of visual and vestibular stimuli has widely been regarded as the
cause of motion sickness. The first part of this paper therefore outlines the sensory
conflict theory of motion sickness, and then critically examines the methodology and
outcome of investigations into perception under conditions of sensory conflict
(referred to below as the ‘perception’ studies). The second part of the paper
integrates the more atheoretical and pragmatic work on the prediction and
prevention of motion sickness (the ‘sickness ’ literature). The processes which may
contribute to susceptibility and adaptation to nauseogenic environments are
considered, drawing on the understanding of perception in disorienting conditions
gained from the review of the perception studies.

The sensory conflict theory bf motion sickness

Until the latter half of this century it was generally believed that motion sickness was
caused by overstimulation of the vestibular system. This was a natural supposition,
as many of the situations which provoke sickness involve turbulent motion or
unusual force environments, and motion sickness bears a remarkable resemblance to
the symptoms induced by vestibular malfunction (Benson, .1984; Brandt & Daroff,
1979). Moreover, although there is a great deal of variability in the susceptibility and
syniptomatology displayed by different individuals, the only group of people who are
completely resistant to the syndrome under all circumstances are those who have
no functioning vestibular system (Money, 1990). However, the ‘vestibular over-
stimulation’ hypothesis was discredited when it was noted that motion sickness
could be induced in a variety of situations where the sufferer experienced hardly any
physical movement or active stimulation at all; for example; in cineramas, or in fixed-
base aircraft simulators.’
In the mid-1970s a theory developed by James Reason (Reason, 1978; Reason &
Brand, 1975) identified ‘sensory conflict’ as the factor common to all the situations
provoking motion sickness. He proposed that the syndrome was triggered when the
pattern of dovariance between the incoming signals from the vestibular system and
the other sensory systems monitoring self-motion did not accord with the pattern of
signals expected on the basis of previous motion experience. Thus, sickness in
This statement is open to dispute, firstly, because the constellation’of symptoms provoked by visual field motion
(and also weightlessness) differs somewhat from that provoked by vestibular stimulation, and, secondly, because it
has been argued that discrepant visual input actually causes an intra-vestibular conflict via central vestibular pathways
(see Crampton, 1990). However, symptoms.provoked by visual conditions and weightlessness are generally regarded
as manifestations of motion sickness, and so these qualifications do not contradict the basic contention that motion
sickness is triggered by perceptual disorientation rather than simple sensory over-stimulation. The peculiarly
nauseogenic effect of ‘vestibular stimulation’ in the form of low frequency vertical oscillation (for example, at sea)
can also be explained in perceptual terms, since the otoliths register repetitive changes in the gravity vector, yet
neither the canals nor the somatosensory system indicate any head movement.
 Motion sickness and perception 451

simulators could be attributed to the unexpected absence of vestibular signals to

accompany the visual field motion (a visual-vestibular conflict), whilst canal-otolith
conflict was held responsible for the sickness provoked, even in the blind or
blindfolded, by the unusual force environments encountered during air and space
travel.’
Reason derived his description of the perceptual processes mediating detection of
and adaptation to conditions of sensory conflict from von Hoist's (1954) ‘reafference
principle’, and modifications of this concept developed by Held (1961, 1965).
Following Held, he proposed that every active movement was accompanied by both
a command signal (efference), and a resultant pattern of input from the orientation
senses (reafference). Traces of these patterns of covarying efference and reafference
were retained in a neural store, and during active movement the efference-copy
enabled rapid retrieval of the reafferent trace combinations previously associated with
it. Reason hypothesized that under unusual sensory conditions a discrepancy between
the current and previous patterns of reafference would be detected, and a ‘mismatch
signal ’ generated which would trigger the mechanisms mediating. the motion
sickness syndrome. Subsequent adaptation to the nauseogenic conditions could be
explained by exactly the same principles. With continued exposure to these unusual
sensory conditions, new patterns of covarying efference and reafference would
gradually be established. Consequently, the discrepancy between the incoming and
previous patterns of reafference would be steadily reduced, and eventually motion
sickness would no longer be triggered.
While there is little theoretical or empirical cause to doubt the perceptual basis of
motion sickness, various difficulties are associated with Reason’s characterization
of the precise processes by which the individual detects and adapts to unusual
perceptual conditions. Some of these problems originate from weaknesses in the
theories from which his was derived. In particular, the idea that movements can be
described in terms of fixed patterns of efference and reafference has been called into
question (Lackner, 1981). Bernstein (1967) pointed out that the relationship between
efferent activity and the physical movement and reafference resulting from it must
vary constantly. This is because the actual effect of any particular motor command
is constrained by the context created by such factors as the position and state of the
limb to be moved. As early as 1932 Bartlett had also observed that no movement
is ever duplicated exactly, and further noted that this posed considerable storage
problems for theories which postulated a bank of neural traces corresponding to each
motion (see Bootsma, 1988). Some theorists have therefore proposed a more flexible
basis for sensorimotor coordination, based on skills adapted to the constraints of the
environment and human abilities, rather than simple neural traces (e.g. Lackner,
1981, 1985; Whiting, 1980).
The most recent reformulation of the sensory conflict theory, a ‘heuristic
mathematical model’ developed by Oman (1982), avoids the problem of a lack of
invariance between motor commands and their effect. This model characterizes the

* The sensory conflict theory seeks to explain how, but not, of course, why the precise symptoms of motion sickness
are triggered. Triesman (1977) has suggested that motion sickness is an accidental manifestation of an originally
adaptive response to ingesting poisons which disrupt coordination and perception through their effects on the
nervous system; vomiting would expel the poison, nausea would cause aversion to it, and the general malaise would
limit activity while dangerously uncoordinated.
19-2
452 Lucy Yardfey
central nervous system as an information processing system which can produce
continuously corrected predictions of sensory feedback using estimates of the current
state of the ‘control system’ based on efferent and afferent information as well as
inbuilt knowledge about the characteristics of that system. ‘Sensory conflict ’ is then
signalled by a discrepancy between predicted and actual feedback.
The predictive utility of such a model, in terms of its ability to identify the degree
of sensory conflict (and, therefore, motion sickness) that will be experienced in a
given situation, depends ultimately on its ability to represeot all the relevant internal
and external sources of information employed by the control system concerned - the
human individual. The way in which this information is integrated to determine
orientation and self-motion must also be accurately modelled. The experiments
reviewed in the following section attempted to define these parameters by
systematically documenting the ‘normal ’ sensory response to various combinations
of visual and vestibular stimuli. By employing a variety of unusual physical and
visual field motions the ‘perception ’ studies artificially created conditions of sensory
conflict, although the dependent variable upon which they focused was the
perceptual response to these conditions, rather than the actual provocation of
sicknes~.~ Since Oman (1982) explicitly stated that his model was intended to combine
Reason’s qualitative sensory conflict theory of motion sickness with the mathematical
approach to characterizing orientation adopted by the perception research, these
studies should have furnished the empirical data required in order to implement (and
validate) his model.
The purpose of the critique of these studies presented below is not to deny the
value of a detailed knowledge of the dynamics, constraints and biases of the sensory
systems which contribute to orientation, but to question whether it is appropriate,
or indeed possible, to characterize perception in unusual motion environments solely
in terms of normative sensory responses to visual and vestibular stimuli. A
reappraisal of the perception studies suggests that integral aspects of the human
control system, such as purposive behaviour and acquired knowledge and skills,
have tended to be ignored or excluded in the interests of experimental control and
the measurement of a ‘pure’ sensory response. Consequently, the description of
perception of orientation and self-motion derived from these investigations is, at
best, incomplete. Moreover, it is evident that the wealth and complexity of
information available to the perceiver, together with the flexible way in which this
information may be selected and utilized, militate against the development of a set
of universal ‘rules’ for the prediction of responses to the unusual perceptual
environments which provoke motion sickness.

The ‘perception’ research

Table 1 summarizes a representative, though by no means exhaustive, sample of
investigations conducted in an effort to determine the precise sensory conditions
under which the sensations of motion generated by the visual channel would
In many of these studies frank sickness would not actually have been provoked, owing to the limited extent and
duration of exposure to disorienting conditions; however, visual-vestibular conflict of this kind is also used to study
susceptibility and adaptation to motion sickness itself (e.g. Dobie & May, 1990; Matsnev, Kuzimin & Zakharova,
1986).
 Motion sickness and perception 453
dominate, sum with, or be overshadowed by the signals for motion arising from the
vestibular channel. During these experiments either the subjects, the visual field they
are viewing, or both, undergo a variety of different physical motions. The perception
resulting from each combination of visual field and self-motion is assessed by one of
several techniques which seem to be treated as essentially equivalent. These include
monitoring either the subject’s psychophysical estimation of motion or the
compensatory motion the subject judges necessary to ‘null ’ the motion experienced,
and measurement of compensatory eye-movements or postural adjustments.

Table 1. A representative sample of the ‘perception’ studies

Authors Motions employed

Huang & Young (1981) Subject rotated about an earth-vertical

Keller & Henn (1984) axis : stationary visual field.
Melcher & Henn (1981)
Probst, Straube & Bles (1985)
Wong & Frost (1981)
Young, Dichgans, Murphy & Brandt (1973)
Zacharias & Young (1981)
Brandt, Dichgans & Koenig (1973) Subject stationary, visual field rotated
Held, Dichgans & Bauer (1975) about various axes.
Reason, Mayes & Dewhurst (1982)
Young, Oman & Dichgans (1975)
Berthoz, Pavard & Young (1975) Subject linearly translated, with moving
Huang & Young (1987) or stationary field.
Lestienne, Soechting & Berthoz (1977)
Pavard & Berthoz (1977)
Huang & Young (1988) Other combinations of subject and/or
Lessard & Wong (1987) visual field motion.

Some consistent patterns of results have emerged from these studies. Visual field
movement tends to be more effective in inducing illusory self-motion (vection) when
presented to peripheral vision, and the vection tends to become more pronounced as
the area and velocity of the visual field motion is increased (Brandt, Dichgans &
Koenig, 1973; Dichgans & Brandt, 1978; Held, Dichgans & Bauer, 1975). Visual
field motion also seems to have a more compelling influence on self-motion
perception at constant velocities and low accelerations, which the vestibular system
cannot reliably detect (Dichgans & Brandt, 1978; Melcher & Henn, 1981). However,
these broad principles simply provide indications of evolutionary, physiological and
developmental constraints or biases in the way that information from the visual and
vestibular systems is commonly utilized.
Many of the perception studies attempted to go beyond such generalities and
establish precise psychophysical laws or mathematical models of the form of sensory
integration and resultant perception induced by the various combinations of visual
454 Luy Yardley
and vestibular stimulation. Yet very different models have often been derived under
perceptual conditions which appear extremely similar in terms of their physical
characteristics. For example, Zacharias & Young (1981) exposed subjects to a visual
surround rotating at a constant velocity, and then superimposed physical yaw motion
(rotation about an earth-vertical axis). They found that the vestibular signal
generated by the physical motion dominated perception of self-motion at all
accelerations above 0.2-1 .0°/s2, and when the vestibular signal was not in agreement
with the sensation of movement induced by the rotating surround this sensation
was abruptly cancelled. However, when Probst, Straube & Bles (1985) conducted a
very similar experiment, they found that the visual input predominated over a
contradictory vestibular input of up to 250/s2 in determining the perceived direction
of self-rotation.
Probst attributes the discrepancy between his results and those of Young to
differences in the area of the moving visual field presented in the two experiments,
since both the total area and the extent of peripheral visual field motion employed
were less in Young's experiment. If the precise characteristics of the visual field
have such a pronounced influence on perception, it follows that the exact effect of
these factors, in isolation and in combination, must in turn be determined.
Unfortunately, attempts to tie the relative.weight attached to the visual channel to
specific physical parameters of the visual field have themselves encountered
difficulties. For example, the stronger influence of peripheral compared with central
field motion noted by Brandt and his colleagues has not always been replicated
(Paulus, Straube & Brandt, 1984). It can be removed or reversed simply by dividing
the visual field into several sectors (Held e t al., 1975). In addition, lamellar optic flow
has been shown to have much the same effect on postural control whether presented
to central or to peripheral vision (Stoffregen, 1985, 1986). It has also proved difficult
to determine whether perceived velocity is principally influenced by the total area
of the moving visual field (Held e t al., 1975), its spatial frequency (Diener, Wist,
Dichgans & Brandt, 1976), or the total boundary length of the elements within it
(Reason e t al., 1982). Even the three broad principles cited above were contravened
by an optical display created by Andersen & Braunstein (1985). They induced both
linear vection and motion sickness by presenting an expanding pattern of dots,
subtending less than 30" of visual angle, to central vision ; moreover, increasing the
area of the display did not increase the illusion of self-motion, while increasing the
speed of the display sometimes led to a reduction in vection.
Besides these changes in perception resulting from slightly differing combinations
of visual and vestibular stimuli, even when sensory conditions are held constant,
large unexplained inter- and intra-individual. differences are commonly observed.
Marked variability in response to linear motion of the visual field has been attributed
(post hoc) to learning effects (Berthoz, Pavard & Young, 1975), mental activities and
unknown individual differences (Lestienne, Soechting & Beahoz, 1977). Moreover,
the true extent and importance of individual differences in perception is often masked
by selective sampling, Fdr example, Huang & Young (1987) based their mathematical
model of linear self-motion perception on the responses of just five out of an original
sample of 11 normal subjects, since the remaining six were quite unable to judge their
self-motion under the unusual perceptual conditions employed in their experiment.
 Motion sickness and perception 455
Similarly, to derive their description of the combined effects of vestibular and visual
stimuli on the vestibulo-ocular reflex, Koenig, Allum & Dichgans (1978) preselected
just four subjects (from an initial group of 40) whose reflexes they considered suitably
stable.
The purpose of focusing on the anomalous results of these attempts to document
perceptual ‘rules’ €or the integration of visual and vestibular stimuli is not simply
to highlight the enormous difficulty and complexity of such a task. In fact, these
difficulties, and some of the explanations offered for them, suggest that the
investigators, in their quantification of the physical characteristics of the motions and
displays employed, have failed to describe adequately the parameters actually
effective in influencing perception in these studies. Individual differences, learning
effects and complex interactions between various features of the visual field and
physical motions do indeed have a fundamental impact upon perception, and must
consequently be given formal and explicit consideration. Some preliminary
indications of the role played by such factors are described below.

Relational perceptual structures

The notable, but not absolute, influence on perception of crude physical parameters
such as the area, velocity and retinal eccentricity of a moving visual display suggests
that these features of the visual field might be indirectly influencing perception,
through their mediating impact upon a related property of the visual environment.
A clue to the possible identity of one such higher-order variable is provided by
Brandt’s own observation that a moving visual surround is less able to induce an
illusion of self-motion if a stable structure can be seen beyond it (Brandt, Wist &
Dichgans, 1975). Thus, it is the motion of the perceived background which appears
to dominate the perception of self-motion. Movement of a small area of optic
structure in the centre of the visual field is likely to be associated with object
movement, but motion of large areas of the visual field, particularly in the retinal
periphery, is normally concomitant with self-motion (Gibson, 1966). Therefore, it
seems reasonable to suggest that such parameters as the area or retinal eccentricity of
visual field movement may influence the extent to which such motion is perceived as
pertaining to the foreground or background, and it is this which then determines
whether the motion is perceived as consistent with object or self-motion. Two
experiments by Ohmi & Howard (1988; Ohmi, Howard & Landolt, 1987) provide
compelling support for this interpretation. They presented subjects with various
combinations of stationary and moving displays and found that vection was always
controlled by whichever display was perceived as the background. Moreover, the
perception of a display as background did not appear to be tied to any specific ‘depth
cues ’, and background stationary displays were shown to suppress vection by about
the same amount whatever their size and eccentricity.
Ohmi & Howard’s demonstration of the intimate relationship between the
perception of background and self-motion carries profound implications for the way
in which motion perception must be characterized and studied. The perception of
‘background motion’ cannot be an automatic response to a particular sensory
stimulus. An integration of many kinds of information contributes to this perception,
456 L u y Yardley
and, as Ohmi’s studies have shown, no one source of information is necessarily either
indispensable or absolutely compelling. Consequently, the factors ‘controlling ’
perception, at least under the unusual perceptual conditions considered here, cannot
be described solely in terms of the physical characteristics of the environment, but
must also take into account the way in which the perceiver attends to, structures and
utilizes these characteristics.

Activity and perceptual exploration

Although in studies of visual-vestibular conflict a variety of responses have been
measured, from psychophysical judgements to postural adjustments, these often
appear to be regarded as interchangeable. Little attention has been paid to the
likelihood that quite different aspects or modes of perception may be involved (and
thus assessed) in each form of response. For example, a visual surround which rotates
about the line of sight can provoke subjective reports of what Brandt termed a
‘paradoxical perception ’ of continuous bodily rotation combined with a fairly steady
apparent tilt of the vertical. When subjects were asked to set a line to the vertical
under these conditions, the maximum average value of the tilt of the perceived
vertical was 15 degrees (Dichgans, Held, Young & Brandt, 1972). However, postural
compensation for this apparent tilt, using exactly the same apparatus and visual field
motion, seldom exceeded 3 degrees (ClCment, Jacquin & Berthoz, 1985). Clearly, the
activities of making judgements of the gravitational vertical and controlling posture
are quite Merent, and evoke different uses of the available perceptual information.
In the perception research very little attention has been paid to the role of
voluntary activity, which has generally been kept to a minimum. Activity emphasizes
information from the somatosensory system, the influence of which has been largely
neglected. Even during passive motion, Lackner & Graybiel (1978) found that the
perception of self-motion, and compensatory eye-movements, of blindfolded,
recumbent subjects rotated about the Z (long body) axis were more consistent with
the patterns of touch and pressure along the body surface than with the otolithic
signals (at rotational velocities above 12 rev/min). Active, voluntary movements
have a much more pronounced impact on perception. Indeed, the sensation of
rotation and postural adjustments which follow active turning are opposite in
direction to those induced by passive rotation, and serve to cancel out the vestibular-
induced illusory after-effects of constant velocity rotation (Bles, De Jong & De Wit,
1984a; Probst et al., 1985). Voluntary movement can be used to gain additional
information about orientation. For example, the vestibular information produced by
continuous head movements can delay and attenuate both the illusory self-motion
and the motion sickness induced by a rotating display (Lackner & Teixeira, 1977).
Activity also helps to direct attention and shape the selection of perceptual
information. The interdependent effects of attention, activity and somatosensory
information have been documented by extensive work on adaptation to distorting
prisms (see Welch, 1986 for a review of the relevant literature). For example, Welch,
Widawski, Harrington & Warren (1979) demonstrated that subjects who watched
their own arm movements while wearing displacing prisms were less susceptible to
the illusion that their arm position had changed if they made active, rather than
passive, movements. Movement affectedadaptation as well as immediate perception ;
 Motion sickness and perception 457
following active arm movements a shift in the perceived direction of gaze could be
measured, while passive movements produced only a shift in the felt position of the
arm. Various other effects on perceptual adaptation have been linked to factors such
as the instructions, activities and opportunities for detecting relevant information
incorporated into the experimental situation (Canon, 1970; Uhlarik, 1973;Warren &
Schmitt, 1978; Welch, 1978), and the variety of perceptual environments explored
(Redding, 1981).
Given the evident importance of activity and perceptual exploration for perception,
the validity of investigations into visual-vestibular conflict in which these are
virtually precluded must be questioned. Although many of the prism studies referred
to above were concerned more with adapted than immediate perception, in practice
it can prove extremely difficult to draw a dividing line between the two (which Welch
and his colleagues have, in any case, shown to be intimately related). For example,
Berthoz e t al. (1975) note that a marked change in perception occurred during the
course of their experiment on (initially) nalve perceptions of linear self-motion.
During the course of only 10 trials subjects apparently learned to attend to some
undefined feature of the experimental situation which helped them to detect their
true self-motion with ever increasing accuracy, despite the presence of visual field
movement which had initially caused severe disruption to such judgements.

Individual differences and perceptual learning

In the perception literature, marked individual differences in response are typically
ignored, and models of perception are based on averages derived from very widely
varying perceptual judgements. Yet, by glossing over this large unexplained inter-
subject variability in perception, valuable insights into the process of perception may
be lost. Drawing again on studies of adaptation to distorting prisms, it can be seen
that the amount of attention paid to a particular source of information can be
determined by an individual’s long-term perceptual skills or biases (McDonnell &
Duffetti, 1972; Melamed, Beckett & Halay, 1979). For example, Warren & Platt
(1975) were able to relate inter-subject differences in amount and type of adaptation
to prisms to pre-test measures of visual and manual abilities. Such variations in the
use of perceptual information may in turn result from an individual’s history of
sensorimotor experience. In a study of the effect of diving experience on perception
underwater, O’Reilly (1975) found that over a 24-minute period experienced divers
adapted to performing manual tasks underwater by a shift in the interpretation of the
visual information, whereas novice divers developed only a localized shift in the felt
position of the arm used to perform the task.

Summary and implications for research

This review of the perception research indicates that perception in unusual
environments does not consist of an automatic response to specific physical
parameters of that environment, but is influenced by the individual’s selective
utilization of the available perceptual information, which is in turn shaped by his or
her activities, and sensorimotor skills and biases. It therefore seems inappropriate to
employ a methodology which analyses the environment solely in terms of simple
458 Lug Yardiey
physical stimuli, prevents voluntary activity and perceptual exploration, and averages
important individual differences. Even though the situations which provoke motion
sickness normally involve passive modes of transportation, in these naturalistic
situations the sources of information from which one can select are actually quite
rich, and some exploratory movements can be made in an attempt to determine one’s
orientation or motion. Often the act of controlling some necessary activity, such as
maintaining balance or guiding a vehicle, can itself provide rapid feedback to the
perceiver about the accuracy and relevance of the various types of information. For
example, Kruk & Regan (1983) and Owen (1986) have identified subtle but highly
effective alterations in the selection of optic information by pilots for the differing
purposes of monitoring var’ious kinds of self-motion, or successfully tracking or
aiming at external targets. Martin, Riccio & Stoffregen (1988) have also shown that
the activity of maintaining balance itself constitutes a source of information about the
effective ‘vertical ’ which is quite independent of optic, vestibular and somatosensory
information. Future research into responses to novel perceptual environments
should therefore examine the selection of information from those environments in
relation to the various activities undertaken.
The way in which perceivers bring their knowledge and perceptual skills to bear
upon the available information is also relevant. Many skills and strategies are likely
to prove virtually universal. However, it is probable that, as in the studies of
adaptation to distorting prisms, some perceptual biases may prove specific to
identifiable groups of people, and may thus be able to account for some of the wide,
hitherto unexplained inter-subject variability observed in responses to various forms
of sensory conflict. A series of experiments initiated by Lackner has already furnished
some preliminary explorations of the impact of such perceptual processes (e.g. DiZio
& Lackner, 1986; Lackner, 1985). His findings suggest that when subjects are
confronted with apparently contradictory information they utilize their previous
experience of the regularities and constraints which exist in the environments
normally encountered to arrive at a plausible, albeit sometimes idiosyncratic,
interpretation of the ambiguous situation. The various solutions devised by his
subjects generally represent a compromise between the ‘evidence ’ of their senses,
and their knowledge of what is possible or probable.
In the following sections the insights into perception in novel environments
derived from this review of the perception literature are used to integrate and
interpret the results of investigations into susceptibility and adaptation to
nauseogenic conditions.

The ‘sickness’ research

Much of the research on motion sickness has consisted of diverse pragmatic, and
largely atheoretical, attempts to discover ways of predicting who is most likely to
become sick in the various nauseogenic situations encountered. Unfortunately, due
to this lack of theoretical integration several fundamental issues pertaining to the
reliability and validity of measures of susceptibility have not yet been resolved. The
issue of validity is closely related to questions concerning the factors which are
generally presumed relevant to the study and understanding of motion sickness. It
is also necessary to determine whether a person’s susceptibility is dominated by their
 Motion sickness and perception 459
immediate response to a novel motion environment, or by the ability to adapt to this
environment over time. In addition, the reliability with which susceptibility can be
predicted in different situations will depend upon the degree to which susceptibility
constitutes a global trait, rather than a local response to a particular set of perceptual
conditions.

Reliabilio and validio

Two main techniques are commonly employed for the practical purpose of
predicting susceptibility to motion sickness. The first is to administer a motion
sickness questionnaire (MSQ), which simply gathers inforniation about previous
responses to various types of travel, while the second consists of exposing the
individuals concerned to some laboratory-based nauseogenic motion. Future
susceptibility to motion sickness is then predicted on the basis of the degree of
malaise induced in the laboratory, or the score on the MSQ, which represents some
weighted summation of the individual’s reported history of reactions to motion.
Some authors report finding a significant relationship between observed
susceptibility to air or seasickness and scores on MSQs or provocative tests (Hixson,
Guedry & Lentz, 1984; Keinan, Friedland, Yitzhaky & Moran, 1981). However,
Lentz (1984) concluded a lengthy development and evaluation of five laboratory tests
of motion sickness susceptibility by commenting that ‘all of these tests have had
typically low correlations with field conditions ’ (pp. 29-7). Reschke, Homick, Ryan
& Moseley (1984) used a battery of provocative tests, MSQs and measures of
vestibular function to predict sickness during parabolic flight. No single test
correlated reliably with the response to parabolic flight, and the level of prediction
obtained using a linear equation derived from these data was only a little better than
chance. A subsequent logistic model developed from scores on a similar range of tests
could only correctly classify 65 per cent of a new sample of subjects into the
categories ‘sick’ or ‘nonsick’ (Lin & Reschke, 1987).
The limited predictive value of the commonly used tests of susceptibility does not
appear to be due to a lack of reliability in the techniques used to measure sickness.
Individual physiological measures such as gastric motility (Stern e t al., 1985) or
electrodermal activity (Isu, Koo & Takahasi, 1987; Warwick-Evans e t al., 1987)
show only moderate covariance with subjective symptoms, and therefore self- or
observer ratings of multiple symptoms are generally employed to assess tolerance of
a provocative motion. These rating systems usually show fair inter-rater agreement
(Lentz, 1984), and Calkins, Reschke, Kennedy & Dunlop (1987) have established a
relatively high test-retest reliability for symptom points scored on two tests
involving cross-coupled angular acceleration, and one consisting of exposure to
parabolic flight (between 0.70 and 0.88 for normalized data). The test-retest
reliability of MSQs is also good; Lentz & Collins (1977) report a coefficient of 34,
while Reason (1968) obtained a coefficient of .89, with an inter-test interval of six
months.
Since the reliability of existing measures of motion sickness susceptibility is
evidently fairly good, it seems probable that the inability of laboratory tests to predict
sickness under ‘field conditions ’ is due to more fundamental problems concerning
460 Luy Yardlg
their validity. The two aspects considered below are temporal and perceptual
characteristics.

Consisteny of susceptibilig across situations and over time

Laboratory tests differ from most field conditions in that they generally consist of a
fairly brief exposure to a highly nauseogenic motion, and thus measure immediate
susceptibility. The target situations, in contrast, typically involve longer-term
exposure to milder motions, and so may require an ability to adapt quite quickly to
the new sensory conditions, rather than an immediate resistance to them. The
importance of distinguishing between initial susceptibility and ability to adapt has,
in fact, been stressed by Reason, who found rate of adaptation rather than immediate
response to be the principal correlate of both MSQ scores and persistence of
symptoms during prolonged exposure to laboratory tests (Reason & Graybiel, 1972).
Evaluating a programme designed to prevent in-flight sickness in susceptible airmen,
Graybiel & Kiepton (1978) noted that an individual’s rate, retention and transfer of
adaptation to provocative motions were much better predictors of a successful return
to flying than was his initial reaction. Similar conclusions were reached by Graybiel
& Lackner ( 1 9 8 0 ~ who) ~ exposed subjects to a ‘vestibulovisual’ test on four
occasions, separated by a few days. They drew the general conclusion that, although
tolerance of the test on the first trial was not a good predictor of adaptation to it,
tolerance remained fairly consistent from the second trial onwards.
Graybiel & Lackner performed no statistical analysis on the results of the study
reported above, but my own post hoc analysis of their published data not only confirms
their conclusions, it also suggests a second possible cause of invalidity in laboratory
tests of susceptibility. The vestibulovisual test consisted of a series of sudden stops
from constant velocity rotation, first with eyes closed and then with eyes open;
however, half the subjects failed to reach the ‘eyes open’ stage of the test on the first
trial, although all subjects were exposed to this condition during the later trials. For
the purposes of my analysis these subjects were allocated to two groups, according
to whether or not they had reached the ‘eyes open ’ stage of the test on the first trial,
and the Spearman’s rank order correlations between the scores obtained on each trial
were calculated for each group independently. The intercorrelations between scores
on the second, third and fourth trials ranged from .75 to .97 for both groups. For
those subjects who completed the whole test on the first trial the range of correlations
between the first and later trials was .49-.68, confirming that the initial response
to the test was a relatively poor indicator of later reactions. However, for those
subjects who did not reach the ‘eyes open’ on the first trial, the correlation between
the first and subsequent trials ranged from only .19 to .24. This suggests that the
initial susceptibility with eyes closed may have been a particularly inadequate
predictor of the level of adaptation finally achieved, with the aid of visual
information.
The results of this reanalysis can be readily understood in the light of the
conclusions derived from the review of the perception literature, which indicated
that wide variations in responses to novel conditions are to be expected if the
availability and utility of the perceptual information differs in the situations
compared. Further support for this contention is provided by the failure of attempts
 Motion sickness and perception 461
to predict space sickness on the Spacelab 1 mission (von Baumgarten, 1986; Oman,
Litchenber, Money & McCoy, 1986; Young, Shelhamer & Modestino, 1986).
Predictions were based on the responses of the four astronauts to six tests, including
four types of constant velocity rotation. During the constant velocity tests no
informative visual cues were available, signals from the semicircular canals were
misleading, and so otolithic and somatosensory signals provided the most accurate
guide to self-motion. However, the utilization of information required for orientation
in space is almost opposite to that needed for accurate perception of self-motion on
these rotation tests ; weightlessness alters the otolithic and somatosensory infor-
mation, necessitating a greater reliance on vision for orientation. The rank ordering
of susceptibility to the rotation tests was highly consistent, but the rank order of
severity of symptoms in space proved to be the exact reverse of that predicted from
these tests.
The evidence regarding the degree to which motion sickness susceptibility is
situation-specific is at present inconclusive. Lentz & Guedry (1978) and Stott &
Bagshaw (1984) have both observed subject-motion interactions in the susceptibility
of chronically motion sick individuals. However, whereas the inter-test correlations
obtained by Reschke e t al. (1984) were mainly non-significant, Lentz & Guedry
(1978) found that the inter-test correlations of normal subjects on three tests of
susceptibility ranged from .38 to .70, and were all significant. Moreover, the ability
to adapt to nauseogenic environments over time appears to be less situation-specific
than the initial susceptibility. Graybiel & Lackner (1983) have presented evidence of
a qualitative consistency in the rate and retention of adaptation in 14 subjects exposed
to a series of vestibulovisual tests and parabolic flights, despite differing initial
susceptibilities to the two conditions. A definite relationship between sea, car and air
sickness has also been reported by Pethybridge (1982) ; as this conclusion was drawn
from a questionnaire survey, scores were presumably based on the persistent
reactions of the individuals concerned over a period of time.

Summary and implicationsfor research

The preceding overview of measures of motion sickness susceptibility suggests that
many of the existing tests, while reasonably reliable, fail to predict sickness in specific
situations owing to problems of validity. Some of these problems may be due to a
lack of correspondence between the perceptual conditions used to test susceptibility
and those that exist in the situations actually provoking sickness; immediate
responses to potentially nauseogenic environments certainly appear to exhibit some
situational specificity. The difficulty in generalizing from one set of perceptual
conditions to another that emerges from this review of the sickness literature closely
parallels the high degree of situational specificity that was found to characterize the
models derived from the perception studies. Moreover, as in the perception studies,
the lack of correspondence between ‘field’ and test conditions may also have been
aggravated by the prevention of any natural exploratory physical and perceptual
activities during most tests of motion sickness susceptibility.
The review of the perception literature concluded that perception in, and
adaptation to, unusual perceptual environments was significantly affected by the
activities, perceptual biases and strategies adopted by the perceiver. Clearly, the
462 Lucy Yardley
effectiveness of such strategies must vary as a function of the conditions pertaining
to any particular situation. An individual’s response to a potentially nauseogenic
environment might therefore also be expected to depend to some extent on the
availability, accuracy and utility of various sources of information in that
environment, defined in relation to the activities undertaken by the individual. The
impact of such factors must consequently be taken into consideration when
attempting to predict susceptibility, and could usefully be explored further.
However, the ability to adapt to provocative motions, rather than immediate
reactions to them, appears to be the most important factor determining motion
sickness susceptibility in the long term (unless, of course, the individual experiences
no initial sickness, in which case adaptation is unnecessary). Moreover, subjects seem
to display more cross-situational consistency in their ability to adapt to motion than
in their initial susceptibility. It was noted in the section on perception research that
immediate perception and adaptation are intimately linked, and may both be affected
by sensorimotor activity, experience, skills and strategies. Since it is evident that the
processes subserving adaptation should be the main focus for investigations into
resistance to motion sickness, the next section is devoted to a discussion of these
processes.
Adaptability and adaptation
The question of why and how some people adapt more easily than others to
potentially nauseogenic conditions is, of course, intimately linked to questions
regarding the processes which mediate the response to unusual perceptual
environments, and the nature of the changes subserving adaptation. In this section
three possible explanations for individual differences in resistance and adaptation to
motion sickness are considered, based on differences in either low-level responses,
central processes, or perceptual learning and sensorimotor skills.

Basic processes
Resistance to motion sickness might plausibly be thought to be mediated by a simple
reduction in the autonomic responsivity which subserves the secondary symp-
tomatology associated with the syndrome. Adaptation and susceptibility have been
linked to patterns of physiological responses, including heart rate, galvanic skin
response, respiration rate and blood volume pulse (Cowings, Naifeh & Toscano,
1990; Cowings & Toscano, 1982). However, the precise pattern of autonomic
response associated with reactions to nauseogenic motions remains a matter for
debate (Cowings, Suter, Toscano, Kamiya & Naifeh, 1986; Graybiel & Lackner,
1980b). It is also unclear whether the patterning of autonomic responses should be
regarded as a cause or a mere symptom of susceptibility; the close association
between symptoms of sickness and perceptual illusions observed by Reason &
Graybiel (1972) certainly suggests that the autonomic reaction is not independent of
the perceptual response to motion.
A great deal of research into individual differences in resistance to motion has been
devoted to an unsuccessful search for variations in sensory sensitivity, or low-level
sensory integration and reflex control. In view of the central role in the development
of motion sickness played by the vestibular system, an obvious candidate was
 Motion sickness and perception 463
vestibular sensitivity. However, investigations into possible links between sus-
ceptibility and vestibular sensitivity eventually concluded that no such relationship
existed (Clark & Stewart, 1973; Dobie, 1969). In view of the evidence that motion
sickness susceptibility seems to be dominated by a general ability to adapt to unusual
perceptual circumstances, rather than by immediate responses to particular sensory
conditions, it is hardly surprising that individual differences in resistance to motion
sickness are apparently not reducible to simple variations in basic sensory processing.
The visual-vestibular integration believed to subserve vestibulo-ocular reflex
(VOR) control has also received considerable attention. Here again, while some
researchers claim to have found associations between susceptibility and various slight
abnormalities of VOR control (Matsnev e t al., 1986; Shirabe, Soda, Kawano &
Shiraishi, 1986), others have failed to observe any systematic differences between the
reflexes of normal and sick or susceptible subjects (Lentz, 1976; Peterka & Black,
1986; Peterka, Black & Schoenhoff, 1987). Most recently, DiZio & Lackner (1991)
have reported a significant correlation between motion sickness susceptibility and the
effect of head movements on the post-rotatory VOR. However, the status of the
VOR as an indicator of the processes subserving motion sickness is again unclear.
Compensatory eye movements are not the product of a totally reflexive integration
of visual-vestibular stimuli. Instead, like most other responses to motion, they can be
influenced by auditory and somatosensory information, and by attention and mental
strategies (Guedry & Benson, 1983; Melville-Jones, Berthoz & Segal, 1984; Moller,
White & Odkvist, 1990). Thus, there is no reason to suppose that minor variations
in the VOR are either the fundamental cause of motion sickness, or even a privileged
indicator of visual-vestibular integration. Rather, VORs constitute just one of the
many facets of sensorimotor control in unusual environments which may be used to
assess the underlying perceptual and motor coordination or disorientation in that
environment.

Central processes and neural traces

Since it appears unlikely that resistance to motion sickness can be attributed to basic
autonomic or sensory processes, a reasonable alternative hypothesis is that this
resistance is mediated by more complex, central processing. This approach was
adopted by Reason, who proposed that adaptation proceeded through the gradual
laying down of neural traces corresponding to the precise patterns of covarying
efference and reafference that obtained under the novel sensory conditions provoking
sickness. Once the pattern of neural traces pertaining to the new sensory conditions
was sufficiently consolidated a mismatch between the expected and actual sensory
inputs would no longer exist, and so motion sickness would not be triggered.
The idea that perception is such a stereotyped process of sensory integration that
the perception of a particular motion in a particular environment can be represented
by one fixed set of neural traces seems somewhat incompatible with the indications
which emerged from the perception studies, that perception involves an active,
flexible and sometimes idiosyncratic selection from many potential sources of
information. The theoretical objections to the concept of a store of neural
representations of the efference and afference corresponding to every movement have
been discussed above. Moreover, Reason’s theory of adaptation should virtually
464 Lug Yardfey
preclude the possibility of transfer of protective adaptation acquired under a
particular set of conditions to another provocative environment. However, a review
of the empirical evidence on this point (presented below) suggests that some form
of transfer of adaptation may, in fact, occur.
A further consequence of Reason’s characterization of adaptation concerns the
processes hypothesized to mediate ‘adaptability ’. Since adaptation is presumed to
consist of a relatively automatic updating of neural traces, Reason could only
speculate that individual differences in adaptability must be due to internal variations
in the ability to acquire, store or retrieve these traces (Reason & Brand, 1975). This
view of adaptability unfortunately led to an apparent investigative dead-end. While
inter-subject differences in the rate and retention of adaptation could certainly be
documented, no other observable correlates of the hypothetical underlying neural
processes were suggested. Thus, although Reason himself concluded that ‘perceptual
adaptation is brought about by something very akin to learning’ (Reason & Brand,
1975, p. 165), his characterization of this learning was both inflexible and circular. If,
however, the learning mediating resistance to motion sickness could be shown to
involve the acquisition of sensorimotor skills and abilities, then a whole new range
of possibilities for the investigation of these processes presents itself.

Cross-situational transfer of adaptation and sensorimotor strategies

The conditions under which adaptation to nauseogenic motions will transfer from
one situation to another has received little systematic exploration. This may be
because such transfer is not predicted by Reason’s model of adaptation, although
Reason himself recorded several instances of a long-term, but limited, generalization
of resistance to nauseogenic motion (Reason & Brand, 1975, ch. 6). Yet a degree of
generalization of adaptation has been documented by several other investigators (e.g.
Dobie & May, 1990; Graybiel & Knepton, 1978; Kennedy, Berbaum, Williams,
Brannan & Welch, 1987).
A generalized resistance to potentially nauseogenic vestibular stimulation is also
observed in various groups of people whose occupations involve repeated exposure
to unusual motion, such as seamen, pilots and athletes (Collins, 1974; Dowd &
Cramer, 1971 ;Hood, 1984). Dancers and skaters, who must learn to orient accurately
during and after pirouettes, seem able to suppress most of the compensatory eye-
movements, and virtually all the feelings of rotation, imbalance or nausea that are
normally induced by rotatory and caloric tests (Dix & Hood, 1969; McCabe, 1960;
Osterhammel, Terkildsen & Zilstorff, 1968). Yet such tests are very different, in
terms of physical activity and sensory input, from making self-induced spinning
movements about an earth-vertical axis. This resistance to the effects of motion is not
congenital, but acquired ;McCabe was actually able to chart the gradual development
of suppression of rotatory and caloric responses in a group of young skaters as they
were taught to spin over a five-month period.
Reason’s ‘neural trace ’ account of adaptation has difficulty in accommodating the
phenomenon of cross-situational transfer. However, the description of perception
and adaptation in novel environments emerging from the review of the perception
literature not only allows for such phenomena, but indeed provides a framework for
investigation into the processes which may be involved. If adaptation to potentially
 Motion sickness and perception 465
nauseogenic conditions involves the same processes as were shown to mediate
responses to ‘visual-vestibular conflict ’, then the response to a nauseogenic situation
will depend on the availability and utilization of various sources of information about
orientation and self-motion, defined in relation to the activities undertaken by the
individual.
Following from this approach, a plausible explanation of the phenomenon of
cross-situational transfer of adaptation is that some perceptual strategies or modes of
functioning developed in a particular nauseogenic environment will prove specific to
a restricted set of perceptual conditions, while others may prove useful in a wide
range of environments, or for several different activities. For example, post-flight
patterns of susceptibility in astronauts suggest both situation-specific and non-
specific components of adaptation (Oman e t a/., 1986). A short-term, relatively
environment-specific adaptation - perhaps the reinterpretation of otolithic infor-
mation (Parker, Reschke, Arrott, Homick & Lichtenberg, 1985) - provides
protection against certain nauseogenic motions (e.g. Coriolis head movements) but
temporarily increases susceptibility to others (e.g. seasickness). In addition, a longer-
lasting slight general reduction in motion sickness susceptibility is observed.
Variations in exploratory activity, and perceptual experience and skills, were also
shown to influence perception of, and adaptation to, the various unusual perceptual
conditions studied in the laboratory. There has been, to date, very little research into
the possible links between susceptibility to motion sickness and variations in
individuals’ activities or uses of perceptual information, but a few promising
examples can be found. One of these is a rare study of natural postural responses to
passive transportation by Fukuda (1975), who observed that bus drivers lean in the
opposite direction to passengers when the bus navigates a bend. It is well known that
individuals are much more resistant to motion sickness while driving a vehicle than
when they are passengers ; perhaps this resistance is associated with such postural
strategies. In addition, there is accumulating evidence to suggest that the ability to
select the appropriate source of information for orientation when some cues are
misleading may prove a useful index of the multisensory coordination required to
provide resistance to motion sickness. Several studies have demonstrated an
association between motion sickness susceptibility and the utilization of vision for
orientation even when vision is misleading (Bles, De Jong & Oosterveld, 1984b;
Frank & Casali, 1986; Yardley, 1990; Yardley, Lerwill, Hall & Gresty, 1992),
whereas dancers are apparently less influenced than most by vision-distorting prisms
(Kahane & Auerbach, 1973).

Sgmmay
Individual differences in resistance to motion have not yet been reliably linked to
any particular mode of low-level sensory or autonomic responding. Moreover, it is
debatable whether any variation in such basic processes as might be found could
justifiably be regarded as a cause, rather than a symptom, of motion sickness
susceptibility. Thus, Reason may have been correct in regarding the perceptual
learning which apparently subserves adaptation to motion sickness as the principal
factor influencing resistance to a nauseogenic motion. However, his characterization
of this learning consists of a somewhat circular definition in terms of hypothesized
466 Lug Yardle_y
central processes, and is incompatible with evidence of a degree of transfer of
adaptation from one set of voluntary motions and perceptual conditions to another.
The alternative conceptualization of adaptation derived from the review of the
perception studies suggests a potentially more fruitful analytic approach, involving
examination of the way in which the experience and sensorimotor skills of
individuals may influence both their susceptibility, and their motor and perceptual
activities and strategies in potentially nauseogenic environments.

Conclusions
The sensory conflict theory of motion sickness inspired a substantial body of research
into responses to various conditions of visual-vestibular conflict (the perception
literature), and also had a more remote and dilute influence on the diverse pragmatic
investigations into susceptibility and resistance to motion sickness (the sickness
studies). However, the review of both these literatures presented above reveals that
neither has yielded any decisive and comprehensive principles. The results of the
perception experiments have tended to be highly specific to the very artificial
situations in which they were obtained, while the sickness studies have so far failed
to develop any reliable methods of predicting or preventing the syndrome in most of
the target individuals and environments with which they have been concerned.
In this paper, the inconclusive results of both types of research into motion
sickness are traced to a common neglect of the role played by active perceptual
strategies, exploration and voluntary movement in reactions to unusual perceptual
environments. Despite the emphasis placed on activity and perceptual learning by
Held, from whose ideas the sensory conflict theory was derived, the perception and
sickness studies have tended to adopt a methodology which precluded any
examination of the impact of either of these factors. Yet it is clear from this review
that an individual's response to a novel environment can be strongly influenced by
the way in which the information available in that environment is selected and
utilized. This, in turn, depends on the activities, experience, biases and skills of the
individual in question. Thus, by utilizing unrepresentative perceptual conditions,
and controlling or ignoring the voluntary activities and perceptual strategies
employed by their subjects, the studies of motion sickness reviewed in this paper
have excluded the very factors most likely to influence responses to nauseogenic
conditions encountered in everyday life.
In conclusion, it is apparent that research into susceptibility and resistance to
environments provoking sickness must adopt a fresh approach. For several decades
it has been generally accepted that motion sickness is caused by an alteration in the
perceptual environment which disrupts the way in which information from the
senses is normally utilized to perceive, and control, orientation and self-motion.
Nevertheless, despite this widespread recognition of the perceptual basis for the
syndrome, principles derived from general models of perception and perceptual
learning have seldom been systematically applied to the problem of motion sickness.
In fact, not since Reason's attempt, over 15 years ago, to apply the ideas of von Holst
and Held to motion sickness has an explanation of motion sickness been grounded
on more general theories of perception and adaptation. In view of the considerable
theoretical advances that have since been made in relevant disciplines (e.g. Flach,
 Motion sickness and perception 467
1990; Runeson, 1988; Stoffregen & Riccio, 1988) a complete re-evaluation of the
problem of motion sickness seems overdue. This review has sought to provide a first
step in this direction, and some preliminary recommendations : future research
should be concerned, not with various parameters of the environment defined from
a physicalist perspective, and their effects upon a passive individual, but rather with
informative structures, which can only be defined in relation to the skills and
activities of the perceiver.

Acknowledgements
The first draft of this paper was completed at Southampton University; I would like to thank Dr Alan
Costal1 for his invaluable comments and advice, and Professor Anthony Gale, Dr Michael Griffin and
D r Michael Gresty for their support and encouragement.

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Received 27 March 1991; revised version received 6 January 1992