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Personality and Individual Differences 26 (1999) 583626
The personality theories of H. J. Eysenck and J. A. Gray:

a comparative review
Gerald Matthews a, *, Kirby Gilliland b
a
Department of Psychology, University of Dundee, Dundee DD1 4HN, Scotland, U.K.

b
Department of Psychology, University of Oklahoma, 455 West Lindsay, Norman,
OK 73019, U.S.A.
Received 6 November 1997
Abstract
Hans J. Eysenck and Jerey A. Gray have proposed inuential theories of the biological bases of
personality traits. Eysenck's theory concerns the extraversion, neuroticism and psychoticism traits,
whereas Gray proposes the use of new, rotated axes of impulsivity and anxiety. Eysenck uses
multiple arousal systems as the central explanatory constructs, whereas Gray describes more specic
systems related to behavioural inhibition and activation. This article reviews the evidence relating to
these theories provided by studies of c.n.s. and a.n.s. psychophysiology, subjective aect,
conditioning and attention and performance. It discusses key predictive successes and failures and
methodological problems which may impede theory-testing. It is concluded that there is a solid core
of predictive support for the Eysenck theory in some paradigms, such as the moderator eect of
stimulation level on individual dierences in phasic electrodermal response and eyelid conditioning.
In other settings, the theory fails to explain empirical data adequately, especially in studies of
subjective response and attention and performance. Gray's theory has advanced research through
stimulating interest in moderation of personality eects by motivational variables. It also provides a
better explanation than Eysenck's theory for certain data, such as instrumental conditioning to
reward stimuli and the positive aectivity of extraverts. Overall, however, Gray's theory explains a
narrower range of ndings than Eysenck's. There is little evidence that Gray's revised personality
axes are generally more predictive of psychophysiological and performance criteria than Eysenck's
original dimensions. Finally, it is suggested that the assumptions of the biological approach to
personality are in need of reassessment. It is possible that the biological theories may be improved
through developments in methodology or through discriminating multiple systems underpinning
traits. For example, extraversion may have distinct ``reticulocortical'' and ``dopaminergic'' aspects.
Alternatively, the biological approach may not in fact be adequate for explaining behavioural
* To whom all correspondence should be addressed. E-mail: g.matthews@dundee.ac.uk

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G. Matthews, K. Gilliland / Personality and Individual Dierences 26 (1999) 583626
correlates of traits. In this case, trait research should place more emphasis on cognitive or social
bases for personality. # 1999 Elsevier Science Ltd. All rights reserved.
1. Introduction
Hans Eysenck and Jerey Gray have been among the foremost exponents of the hypothesis
that personality traits provide a window on individual dierences in brain functioning. Both
theorists start from the assumption that we can characterise brain processes by means of a
simplied ``conceptual nervous system'' comprising the key circuits relevant to personality and
behaviour. The multitude of associations between personality and behaviour established
empirically may derive from individual dierences in quite simple parameters of brain function.
Gray's personality theory began as a modication to the Eysenck theory, but is now usually
seen as an alternative theory (cf. Gray, 1981). In this paper, we review data from experimental
studies testing predictions from the two theories and conclude with some thoughts on their
respective merits.
1.1. Conceptual nervous systems for personality
Eysenck (Eysenck, 1967; Eysenck and Eysenck, 1985) identies two principal brain systems
as the key components of his conceptual nervous system: reticulocortical and reticulolimbic
circuits. The reticulocortical circuit controls the cortical arousal generated by incoming
stimuli, whereas the reticulolimbic circuit controls response to emotional stimuli. Under
strong emotional stimulation, limbic system activity may spread to the cortex. Extraversion
introversion (E) relates to arousability of the reticulocortical circuit, so that introverts are
typically more aroused than extraverts. However, methodological analyses of extraversion
studies (notably Gale, 1973) have illuminated two basic problems for theory testing. First,
people actively seek a moderate level of arousal, so that relationships between personality and
arousal may also reect individual dierences in strategies for seeking or avoiding stimulation.
For example, in sensory deprivation studies, extraverts seem to raise their arousal by moving
around (Eysenck and Eysenck, 1985) and extraverts generally prefer more arousing activities
(Furnham, 1981).
Second, under high levels of stimulation, a protective ``transmarginal inhibition'' (TMI) may
lead to paradoxically reduced arousal. According to Eysenck (1994), increasing stimulation
provokes increasing c.n.s. reactivity until an optimal point is reached, beyond which inhibition
and decreasing reactivity set in. Eysenck (1994, p. 161) states that `Ìn the case of the most
arousing testing conditions we might suspect that the optimal point for introverts, but not for
extraverts, had already been passed, so that extraverts' arousal was still growing, introverts'
declining''. Hence, introverts may be higher, lower or equal to the arousal level of extraverts
according to complex interactions of personality type and environmental manipulation.
Careful, theory-driven control of experimental parameters is thus essential (H. J. Eysenck,
1981).
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Neuroticism-stability (N) is associated with arousability of the limbic circuit, such that
neurotics become more aroused than stable individuals as a consequence of emotion-inducing
stimulation. Hence, individual dierences in N may only be evident in emotional or stressful
contexts. The neuropsychology of Eysenck's third dimension of Psychoticism (P) has not been
worked out in detail. H. J. Eysenck (1992) suggests that P is inversely related to serotonergic
function, but, more recently (Eysenck, 1997), P is linked to dopamine.
Gray's theory modies both the psychometric alignment of the major personality dimensions
and, to some extent, their biological bases. Originally, Gray (1970) proposed two new
dimensions of anxiety (Anx) and impulsivity (Imp) oriented at 308 to Eysenck's E and N,
respectively. Gray's model is often assumed to posit a 458 rotation, but Gray (personal
communication, 18/11/97) points out that this is a misconception derived from the way the
model was presented graphically (e.g. Gray, 1981). Here, we shall assume a ``308 rotation
model'', which aligns Anx most closely with N and Imp most closely with E. Anx also
correlates to some degree with introversion and with low P, so that Anx may be contrasted
with risk-taking ``primary'' psychopathy (Gray, 1987a). A second, orthogonal Imp dimension
correlates most highly with E, but also correlates to some extent with N and P. Gray's (1987a)
conceptual nervous system is shown in Fig. 1. Anxiety is associated with sensitivity of one of
these components, the behavioural inhibition system (BIS), which is activated by fear and
novelty stimuli and signals of punishment and non-reward. The core of the system is a septohippocampal comparator which detects mismatch between the actual and predicted state of the
world. Mismatch detection results in inhibition of ongoing behaviour and increased arousal
and attention. The comparator works in conjunction with other brain structures, such as the
prefrontal cortex, permitting some cognitive control over anxiety in humans and various
ascending aerents from more primitive levels of the brain. Gray (1987a) discusses the neural
system supporting Imp, the behavioural activation system (BAS), more briey. It is sensitive to
signals of reward and non-punishment and inuences probability of approach behaviour. Gray
(1987a, 1991) sees the BAS as associated with mesolimbic dopaminergic pathways ascending
Fig. 1. A conceptual nervous system for personality (adapted from Gray, 1987a).
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from nucleus A10 of the ventral tegmentum of the brainstem (see also Gray et al., 1991). Gray
agrees with Eysenck that a third major dimension of personality is required, aligned at least
approximately with psychoticism, which Gray (1991) relates to the ght/ight system.
1.2. Levels of description within the theories
The two theories dier in the level of physiological construct preferred, as well as in the
specic circuits linked to personality. Eysenck's reticulocortical and limbic systems are
broadly dened and system components such as the ascending reticular activating system
(ARAS) may be comprised of a number of structures and pathways. Gray (1987a) identies
arousal as a system separate from BIS, BAS and ght/ight, although these systems are in
dynamic interaction. However, Gray (1987a) uses `àrousal'' in a more restricted sense than
Eysenck, to refer to the dorsal noradrenergic bundle (DNAB), bres which ascend from the
locus coeruleus to innervate many forebrain structures. Although some researchers (e.g.
Panksepp, 1982) relate noradrenergic activity to a general emotional arousal function, the
DNAB does not seem to constitute the generalised arousal system envisaged by the pioneers of
arousal theory. Robbins (1986) suggests that the DNAB may relate to a compensatory
mechanism which maintains eciency of selective attention and other functions during states
of stress. Hence, some functions traditionally related to arousal may not be attributable to
Gray's arousal system, including (1) motor responsiveness, which perhaps relates most closely
to dopaminergic pathways (cf. Gray et al., 1991) and (2) sensory and attentional functions
related to central cholinergic aerents to the cortex (Beatty, 1986), whose place in Gray's
conceptual nervous system is unclear.
One of the strengths of Eysenck's theory is that the use of arousal as a mediating variable
allows personality to be linked to many qualitatively dierent response indices. Gray's theory
trades o this generality of application for more ne-grained neuropsychological description
and greater specicity of prediction of a narrower range of behaviours. Often, however, it is
dicult to establish correspondences between human behavioural responses and the outputs of
Gray's systems. In such cases, Eysenck's arousal theory may provide a basis for prediction
where Gray's more neurologically detailed theory is silent. Hence, in contrasting the Eysenck
and Gray theories we are not altogether comparing like with like. Clearly conicting
predictions may sometimes be drawn from the two theories, but in other circumstances the two
theories may be referring to the same underlying physiology at dierent levels of description,
especially where the slippery construct of arousal is concerned.
1.3. Criteria for theory testing: scope of the review
Research on the two personality theories has adopted several strategies for theory testing,
including psychometric analyses, the comparison of normal and clinical populations and
experimental tests of predictions derived from neuropsychological theory. Eysenck (e.g.
Eysenck, 1994) has also emphasised personality correlates of clinical conditions, but the
theories do not seem to generate clearly contrasting predictions concerning the relationship
between personality and disorder, especially as Eysenck (1997) and Gray et al. (1991) agree on
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a dopaminergic mechanism for psychosis. In this article, we assess the theories against the
following criteria, each of which has its own advantages and disadvantages.
1.3.1. Comparison of dimensional models
Gray (1981, pp. 247252) highlights studies which investigated which sets of personality axes
correlate best with psychophysiological and behavioural criteria. He suggested that Imp is
more predictive than sociability (Soc) of various criteria. Hence, a rough and ready way to
distinguish between the theories is in terms of the predictive power of the trait variables they
specify. If we nd that Imp and Anx are more strongly and coherently related to dependent
measures, then we have an argument in favour of the Gray theory. In essence, we are seeking a
type of ``simple structure'': the set of traits which align most closely with sets of response
measures. Given the lack of comparative work on N and Anx, most of the evidence is
concerned with comparisons between E, Imp and Soc. In recent years, several researchers have
attempted to assess the Gray dimensions using new scales (e.g. Wilson et al., 1989;
MacAndrew and Steele, 1991) or by reinterpreting scales developed for other purposes.
However, there have been insucient psychometric and experimental studies for any of these
newer measures to be convincingly validated as measures of BIS and BAS function.
1.3.2. Comparison of moderating factors
The theories dier not just in their choice of dimensional axes, but also in the moderating
factors which they identify as critical. A further indication of theory validity is provided by
testing whether personality eects are primarily moderated by level of stimulation or arousal
(as Eysenck predicts) or by reinforcement signals (as Gray predicts). The diculty in applying
this criterion is that the arousing and motivating properties of experimental manipulations may
be mutually confounded.
1.3.3. Direct tests of theory
Comparison of dimensions and moderating factors is a somewhat indirect approach to
theory evaluation. A stronger criterion is the outcome of tests of conicting predictions derived
from the two theories within specic experimental paradigms. However, because of the
methodological and conceptual diculties previously described, it is dicult to nd suitable
indices of the underlying conceptual neural systems. Additional hypotheses are required to link
neuropsychology to overt psychophysiological and behavioural response, such as the Yerkes
Dodson Law in the case of studies of arousal and performance. Disconrmation of prediction
may reect failure of the additional hypothesis, rather than failure of the theory (see H. J.
Eysenck, 1981).
The present article is mainly concerned with experimental studies which aord direct tests of
theory. We will be concerned primarily with studies of E and N on the one hand and Anx and
Imp on the other, because the two theories have a similar view of P. This review is structured
around a number of dierent topic areas, within which the two theories make diering
predictions. For each area, we will assess the validity of the two theories by comparing how
successfully their predictions match empirical data. We will discuss whether the predictive
failures may be attributed to failures of additional hypotheses linking responses to neural
systems, rather than to the personality theory itself. We begin with psychophysiological
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correlates of personality, which we divide into central and peripheral nervous system studies.
Next, we consider studies of the specialised branch of psychophysiology which deals with
emotional and other subjective states. Two areas of behavioural research are reviewed:
conditioning and learning and attention and performance. The behaviours concerned become
progressively more distant from the neural substrates described by Eysenck and Gray and so
provide a progressively greater challenge for neuropsychological theory.
2. Psychophysiology of the central nervous system

2.1. Psychophysiology of personality: theoretical and methodological issues
Owing to the biological nature of both Eysenck and Gray's theories, psychophysiological
measures would appear to oer the most direct and ecient method for investigation.
However, there are various diculties in making and testing predictions. Gale's (e.g. 1973,
1983) series of methodological reviews of the EEG refers to problems with subject selection,
bias within the subject or experimenter, a lack of theoretical sophistication, over-attention to
outcome at the expense of process, poor psychometrics or psychophysiology, trivial
experimentation and procedural insensitivity (see also critical reviews by O'Gorman, 1984;
Eysenck, 1994). Four fundamental diculties are as follows:
1. It is hard to obtain an accurate translation from theory-relevant elements of the
neurological substrate to the level of discrete psychophysiological measures.
Psychophysiology is not neurophysiology (Gale, 1973; Zuckerman, 1991).
2. Gale (1973) identies a multitude of confounding variables that inuence arousal and the
integrity of specic psychophysiological measures, so that careful experimental control of
the experimental context is essential.
3. The validity of specic psychophysiological responses as arousal indices is threatened by
phenomena such as directional fractionation and response stereotypy (Lacey, 1967).
4. Understanding of both personality traits and the neuroanatomical substrates of
psychophysiological indices changes over time as constructs are claried, but researchers do
not always make methodologically and theoretically informed selections of personality scales
and psychophysiological indices.
These diculties may leave the impression that the enterprise of seeking relationships
between personality and psychophysiology is much like aiming at a moving target with
wobbling sights. Nevertheless, several commentators consider that robust, theoretically
signicant relationships between personality and psychophysiological responses may be
obtained using suitable methods (see Geen, 1983; Eysenck and Eysenck, 1985; Stelmack, 1990;
Eysenck, 1994 for reviews). More recent studies seem to be more rigorous than those criticized
in Gale's reviews, in general, although some continue to nd shortcomings (Zuckerman, 1991).
In this article, we focus on the more programmatic, theory-driven and methodologically
rigorous studies.
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We have organized the studies reviewed into two broad categories: those using central
nervous system (c.n.s.) measures and those using autonomic nervous system (a.n.s) measures.
These categories are both convenient and theoretically relevant. (We have omitted discussion of
biochemical and brain-scanning studies as being too far removed from the main theoretical
issues.) As discussed above, the Eysenck theory predicts that both E and N should relate to
c.n.s. and a.n.s. arousal. In the case of E, relationships should be moderated by level of
stimulation and in the case of N by levels of ``stress'' or emotion.
Prediction from the Gray theory is more dicult. If both BIS and BAS activity feed into
greater noradrenergic arousal, we expect impulsives (and extraverts) to show more arousal in
response to reward signals, whereas anxious individuals (and neurotics) should be more
responsive to punishment cues. However, it is somewhat unclear which arousal responses are
sensitive to DNAB activity (as opposed to other arousal systems). Furthermore, there may be
psychophysiological responses which are directly driven by the BIS or BAS, without
involvement of the separate arousal system, although, again, Gray does not specify these
responses in detail. Fowles (1980) has argued that electrodermal measures are probably a good
index of the BIS and that heart rate is a good measure of the BAS.
2.2. The electroencephalogram
There are numerous psychophysiological measures of c.n.s. activity. Perhaps the most well
known is the electroencephalogram (EEG), the measure of the raw electrical activity produced
by the brain, derived from passive electrodes placed on the scalp. The complex EEG waveform
is often analyzed by decomposing the waveform through spectrum analysis into various ranges
such as delta (<4 cps), theta (48 cps), alpha (813 cps) and beta (>13 cps). The alpha
frequency range has been classically associated with low states of arousal or relaxation, while
the beta range is most often associated with wakeful, active states.
Eysenck (1994) has referred to the EEG, especially the alpha range, as the ``standard
measure of cortical arousal, ever since the discovery of the ARAS'' (p. 167). Studies of the
relationship between raw EEG measures and E have been reviewed almost exhaustively (e.g.
Gale, 1983; O'Gorman, 1984; Zuckerman, 1991; Eysenck, 1994). While it is unnecessary to
recount this body of literature yet again, some attention to the reviews and their conclusions is
protable.
The earliest of the systematic reviews was a landmark analysis by Gale (1973). According to
Gale, several studies supported the hypothesis that introverts are higher in cortical arousal
than extraverts. However, a similar number found no dierences and three studies found
results that contradicted the theory. Gale (1973) observed that the arousal inducing properties
of the testing environment appeared to mediate the ndings of the studies reviewed.
Gale argued that moderate arousal-inducing environments, including opening and shutting the
eyes upon instruction, were the most amenable to testing predictions of Eysenck's (1967)
theory. Low arousal-inducing environments, such as when the subject was simply lying with
eyes closed, were believed to result in paradoxical arousal, especially in extraverts and high
arousal-inducing environments involving task performance demands were believed to result in
possible over-arousal, again especially for extraverts. After post hoc reordering of the studies
according to the arousal inducing properties of the testing environment, Gale revealed that in
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studies incorporating moderate arousal levels, introverts exhibited greater evidence of alpha
activity reecting lower levels of arousal, as compared to extraverts, as predicted. A later
review by Gale (1983) developed both the methodological critique and reviewed results from a
new decade of conicting EEG research on E, arriving at similar conclusions to the earlier
review.
O'Gorman (1984) contended that Gale may have made some errors of omission and
classication. His own, alternative analysis of the EEG studies found that use of Eysenck's
original E scales or similar measures, was a more important inuence on outcome than arousal
level (see Gale, 1984, for a rejoinder). Zuckerman (1991) has been less than enthusiastic about
the level of support that past EEG studies have provided for the cortical arousal hypothesis of
extraversion, although he points out that studies using female subjects or equal numbers of
both sexes seem to more often support Eysenck's theory.
There have been a few more recent studies that are noteworthy. In a sample of 180 subjects,
Matthews and Amelang (1993) measured power values in three bands in three experimental
conditions, whose levels of stimulation corresponded to Gale's three environment types.
Signicant correlations between personality and EEG measures averaged across environments
were low in magnitude, not exceeding 0.20, but broadly matched expectation. Extraverts and
impulsives showed more slow-wave activity (but not more alpha) and neurotics showed more
beta activity. Analyses of the individual environments showed little systematic dierence in
personality correlates across dierent levels of stimulation. In the ``moderate arousal''
condition considered by Gale most likely to support the Eysenck theory, both E and Imp were
unrelated to alpha, but positively correlated with beta. Matthews and Amelang suggest that
personality traits are only weakly predictive of the spontaneous EEG and most published
studies have far too few subjects for assessment of potential moderators of personalityEEG
associations.
Smith et al. (1995) reported that introverts were generally found to produce lower levels of
alpha activity reecting higher levels of arousal, but there were also complex hemisphere by
gender interactions. Two groups showed asymmetry of alpha: both female introverts and
extraverted males exhibited higher alpha activity on the right side. These ndings were
complicated by the fact that subjects were selected to be high in N because the study focused
on emotional reactivity. Aside from providing, once again, some conditional support for the
arousal hypothesis of E, these ndings suggested that concerns about sex of subjects may be
important (Zuckerman, 1991), as well as electrode placement (Gale, 1983).
Finally, Stenberg (1992) attempted to contrast potential dierences between the E and N
dimension of Eysenck and the Imp and Anx dimensions of Gray in a study which manipulated
positive and negative emotional imagery. Subjects were classied according to the EPI and the
Karolinska Scales of Personality. Factor analysis of the multiple EEG sites used revealed two
factors associated with arousal and three factors that were associated with responses to
emotional reactivity. The EPI E and N dimensions were not signicantly related to any of the
EEG factors. However, factor analysis of the joint personality scales revealed factors that
resembled Gray's Imp and Anx dimensions. More impulsive subjects showed signs of lower
EEG arousal than low impulsive subjects and more anxious subjects showed greater right-side
frontal theta activity across all conditions, suggesting higher emotionality. Furthermore, high
anxious subjects evidenced higher beta rhythm activation to the negative emotional condition,
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but the high impulsive subjects did not show a corresponding reaction to the positive
emotional condition. These results suggest use of Gray's dimensions is preferable, but it is
unclear to what degree the derived dimensions actually represented Gray's axes and the sample
size (40) was insucient for factor analysis. Stenberg concludes that there is much more
support for the view that Imp is associated with low arousal than with facilitation of positive
aect.
So what can be said of the EEG studies of E? Gale's (1973) original critique has led to a
well articulated debate on the many sources of experimental error in this area. Nevertheless,
despite the conicting results, methodological problems and thoughtful and well-reasoned
arguments to the contrary (O'Gorman, 1984; Zuckerman, 1991), there is support for an
arousal hypothesis for E. Reviewing eects of N and Anx, Zuckerman (1991) concludes that
the precise nature of the relationships between these traits and the EEG is unclear. Signicant
eects of N are not uncommon but their direction varies from study to study and N frequently
interacts with E. There is some evidence (O'Gorman and Lloyd, 1987) that Imp may be
especially predictive of the EEG, in line with Gray's theory. However, in the absence of
manipulation of motivational signals, the data have limited implications for the theory. A new
generation of investigations that dierentiate between the Gray and Eysenck theories is
required, in which the stimulating and reinforcing properties of experimental manipulations are
clearly distinguished.
2.3. Event-related potentials
Another index of c.n.s. activity is the event-related potential (ERP) or evoked response,
derived from averaging EEG samples across trials (see Coles et al., 1990). The majority of ERP
studies have investigated responses that occur in the rst 100500 ms following a stimulus,
although time periods as short as 10 ms are associated with the auditory brainstem response.
The major component waveforms are usually related to cognitive processing as follows. Early
components (N100, P200) relate to sensory properties of stimuli and to selective attention
process. Later components are more ``cognitive''. The P300 has been associated with processes
related to classifying or updating memory representations of stimuli. The amplitude of the
P300 increases as the signicance of the event and its relevance to the subject increases and as
the demand for cognitive resources increases. The latency of the P300 measure has been related
to the time needed to categorize and evaluate the stimulus, which appears to be independent of
the time needed for response-related processes. Most researchers using the P300 see it as an
index of cognitive rather than arousal processes (Coles et al., 1990), but, as Eysenck (1994) has
noted, ``The fact that the activities associated with the P300 (habituation, orienting responses,
stimulus classication) are closely related to the concept of cortical arousal suggests that tests
of the introversion-arousal hypothesis could use the P300 paradigm with advantage'' (p. 172).
Reviews of ERP measures and extraversion (Geen, 1983; Zuckerman, 1991; Eysenck, 1994;
Stelmack and Houlihan, 1995) reveal the same degree of variation in subjects, testing
conditions, stimulus characteristics and other variables as those seen in the EEG studies. Early
studies of ERP measures were also notable for their inconsistent results, probably owing to
numerous experimental control factors. More recent studies show a trend for amplitudes of
both the earlier somatosensory components and the later P300 to be higher in introverts than
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extraverts. The nding of higher P300 amplitudes in introverts has typically been interpreted as
suggesting that introverts are experiencing greater levels of attentional demand (Stelmack and
Houlihan, 1995), a result which may, in turn, be suggestive of higher arousal in introverts
(Eysenck, 1994). However, failures to replicate eects even across similar studies is a cause for
concern (Zuckerman, 1991). In addition, the increased P300 amplitude seen in introverts may
not be evident in initial trials or sessions with small numbers of trials. Rather, this dierence
may be the result of greater habituation in extraverts (Di Traglia and Polich, 1991) and thus
more likely to occur in sessions with a larger number of trials or in later trial blocks.
The modal nding is for larger P300 amplitude in introverts, but a few studies show the
reverse relationship. Stenberg's (1994) review of these ndings suggests that P300 is larger in
extraverts when the task is of brief duration or the task is complex and cognitively demanding.
Brocke et al. (1997) varied level of stimulation experimentally and showed that the negative
association between E and P300 amplitude became positive when subjects were exposed to
60 dB noise. This eect might be attributed to TMI, which is initiated at lower levels of
stimulation in introverts (H. J. Eysenck, 1981). There is little evidence for any consistent
dierence between extraverts and introverts in P300 latency, but faster P300 latency has been
reported for high N individuals (Stelmack and Houlihan, 1995).
Few ERP studies have attempted dierential tests of Eysenck and Gray's theories. However,
Bartussek and his colleagues reported three investigations that were specically designed to test
Gray's theory. In one study, ERPs were recorded from introverts and extraverts to neutral
tones and tones signifying gains or losses in a betting task (Bartussek et al., 1990a). In a
second study, ERPs were recorded to emotionally positive, negative or neutral adjectives
(Bartussek et al., 1990b), so as to dierentiate between Gray's BAS (reward seeking) and BIS
(punishment avoidance) components. In both studies, introverts had greater amplitude of the
ERP in the neutral condition and extraverts had greater amplitude of the ERP in both the
reward/positive and loss/negative conditions. These results failed to support Gray's theory and
could be viewed as supporting Eysenck's more general arousal model, provided emotional
stimuli induce TMI in introverts. In a third study, auditory ERPs were recorded to stimuli that
signalled winning or losing in a gambling task (Bartussek et al., 1993). A statistically signicant
personality by condition (win/lose) interaction was found for the ERP P200 component that
conformed to the predictions of Gray's theory, i.e., extraverts had higher amplitude in the win
condition and about equal amplitude in the loss condition, as compared to introverts. There
was a complex interactive eect of E, N and condition on P300 amplitude, such that data were
consistent with Gray's theory only in high N subjects. However, De Pascalis et al. (1996)
conducted a similar study and failed to nd any eect of E, although they found some
relationships between scales of the GrayWilson questionnaire (Wilson et al., 1989) and some
wave components. The study also showed that high N subjects showed larger N800 amplitudes
to punishment stimuli, a result which seems compatible with Gray's theory, although, as the
authors point out, the functional signicance of N800 is unclear.
Finally, Bartussek et al. (1996) tested for interactions between E and motivational stimuli in
two studies, one requiring processing of words of diering valence and one using a startle
response paradigm. In both studies complex interactions involving E, stimulus valence,
electrode site and other factors were found, for P300 in the rst study and P200 in the second.
Eects bore little resemblance to eects predicted from the Gray theory. Bartussek et al. (1996)
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suggest that extraverts tend to develop stronger arousal at frontal sites in response to
emotional stimuli, irrespective of valence. Such an eect might be seen as a further instance of
the TMI predicted by the Eysenck theory. However, the Bartussek studies provide little
evidence for enhanced response to emotional stimuli in high N subjects, although N
participated in various complex interactions. According to the Eysenck theory, N rather than E
should be the main factor moderating response to emotional stimuli. While any denitive
conclusion based on these studies remains elusive, they do represent experimental eorts of
precisely the type needed if we are to advance an understanding of either or both theories.
Another ERP method that has received increasing attention is the auditory brainstem evoked
response (ABR: see Jewett et al., 1970 and Bullock and Gilliland, 1993 for reviews). The ABR
produces a remarkably stable and replicable series of seven waves that occur in the rst 10 ms
following an auditory stimulus. Its various wave peaks relate to specic neural generators at
successively higher levels of the auditory pathway, ranging from the acoustic nerve (waves I
and II) to initial cortical projection activity (wave VII). Faster peak latencies or interpeak
latencies (indexing conduction times) reect higher levels of neural activity. For many years,
reports of the stability of ABRs across subjective arousal states and environmental
manipulations of arousal suggested that the ABR was better suited to exploring peripheral
rather than central nervous system processes (Eysenck, 1994; Stelmack, 1990). However, the
potential of the ABR for also exploring midbrain c.n.s. processes (Bullock and Gilliland, 1993)
is shown by its sensitivity to changes in cognitive demand (e.g. Lukas, 1980) and to certain
drug manipulations that act on c.n.s. arousal centers (e.g. Church and Shucard, 1987). Thus,
the ABR may be useful in reecting the inuence of dierential ARAS arousal or perhaps
individual dierences in more general ARAS arousal levels, even though it is less sensitive to
arousal states than other psychophysiological measures of c.n.s. activity. If reliable dierences
in ABR activity between introverts and extraverts can be demonstrated, these dierences may
provide additional evidence for the physiological basis of E.
Bullock and Gilliland (1993) briey reviewed early studies of ABR activity and E and
concluded that the most consistent nding was that introverts appeared to have faster wave V
latencies and faster wave IV conduction times than extraverts. Bullock and Gilliland (1993)
reported a study in which ABRs were recorded for introverts and extraverts during placebo
and two levels of caeine administration, as well as during resting and two levels of response
time task diculty. This study included an analysis of many control factors that may have
aected previous investigations. Introverts had faster wave V latencies than extraverts, as well
as faster wave IIII and IV conduction times. Stelmack et al. (1993a,b) found no dierences
between introverts and extraverts for any ABR wave component during sleep or wakefulness.
However, the one consistent nding, although nonsignicant, was that wave V was faster for
introverts, as compared to extraverts, for all conditions. Wave V relates to inferior colliculus
activity, which is one of the auditory pathway nuclei that adjoins the ARAS. In a study across
the full range of E scale scores, Swickert (1996) replicated the signicant positive correlation
between E and ABR wave V latency. Among the two subscales of E, Soc was found to
correlate positively with Wave V latency, while Imp was not. This nding appears inconsistent
with Gray's view that Imp is the primary causal component of E, although Gray has not
explicitly addressed ABR studies. Finally, one study (Stelmack and Wilson, 1982) reported
faster wave I latency, which led the authors to suggest peripheral neural mechanisms as the
594
possible site of extravertintrovert dierences in the ABR. To the extent that wave V latency
may be interpreted as a c.n.s. measure, the results of more recent ABR studies generally
support Eysenck's theory evidence that is particularly interesting because the ABR may be
anatomically consistent with locations and neural mechanisms which Eysenck relates to E.
There have also been a few studies of contingent negative variation (CNV), an EEG
response emitted in anticipation of a motor response. To measure CNV researchers typically
employ a response time procedure that includes both a warning stimulus followed by an
imperative stimulus to which the subject responds. During the brief interval between the
warning and imperative stimuli, there is a negative shift in the baseline of EEG activity that
includes an orienting component (O) and an expectancy or readiness for action (E) component
(Tecce, 1972). The CNV has been viewed as an indicator of attention or arousal functions
(Tecce, 1972), which oers the logical bridge for its use in explorations of E. In general, as the
Eysenck theory predicts, extraverts do have higher CNV amplitude as compared to introverts
(Werre, 1986). Varying environmental and intrinsic conditions can aect this relationship with
optimal results being provided by subjects who were young, well motivated, isolated from
extraneous stimulation and were performing a novel reaction task in the morning (Werre,
1986). Because of its links to attentional processes, the CNV has probably provided evidence
for the arousal hypothesis of E more like that from behavioral measures of attention and
arousal, in contrast to raw EEG measures that are generally viewed as having more direct links
to c.n.s. processes.
3. Psychophysiology of the peripheral nervous system

At a functional level, a considerable amount of peripheral nervous system activity is
regulated by c.n.s. processes. Within the peripheral nervous system, the a.n.s. is a highly
complex and interactive neural network that, at a very simple level, is organized as two
typically opposing subsystems, the sympathetic and parasympathetic nervous systems. Crudely,
the sympathetic nervous system responses assist the organism in meeting the demands of
arousal-inducing situations whereas the parasympathetic nervous system acts to conserve
bodily energy. Many personality researchers searching for methods of measuring bodily
function that reect arousal or arousability have focused on psychophysiological measures of
sympathetic or parasympathetic nervous system activity. The large majority of these studies
have included electrodermal or cardiovascular measures, while a far smaller number have
included an array of other measures.
3.1. Electrodermal activity
Electrodermal measures attempt to quantify the electrical activity of the skin. Typically, the
measurement of electrodermal activity (EDA) involves the application of a small amount of
electric current to the skin surface through one electrode, allowing measurement of skin
resistance level (SRL) between the electrodes (or its inverse, skin conductance level; SCL). SRL
or SCL are measures of baseline or tonic activity. Phasic EDA measures in response to a
known stimulus are referred to as skin resistance responses (SRRs) or skin conductance
595
responses (SCRs). There are many other EDA measures as well and each type of measure can
have several response characteristics, such as latency, amplitude, rise time, half recovery time
and habituation rate. It has been observed that electrodermal measures provide ``direct and
undiluted representation of sympathetic activity'' (Dawson et al., 1990, p. 310), which is
relatively free from other somatic inuence such as heart and respiration rate. EDA measures
have traditionally been viewed as a component of the orienting reex. Habituation of the
orienting reex has been a prime measure of inhibitory neural processes. Thus, measures of
EDA are theoretically relevant to personality research.
EDA has been investigated with respect to both E and N. Eysenck's theory makes the clearcut prediction that EDA should be higher in neurotics, especially under stress, but more
consistent ndings have been obtained with E. Electrodermal responses tend to be insensitive
to both N and trait anxiety (Zuckerman, 1991). Several reviews of EDA research on E have
been conducted in the past several years (e.g. O'Gorman, 1977; Stelmack, 1981; Geen, 1983;
Smith, 1983; Zuckerman, 1991). The general view is that EDA studies of tonic arousal
dierences between introverts and extraverts are quite inconsistent. However, in an interesting
naturalistic study, Wilson (1990) showed that introverts had higher SCLs than extraverts
throughout much of the day, but only after age dierences were statistically controlled. The
frequency of spontaneous EDA has also been viewed as a measure of tonic arousal. There is
an approximately equal number of studies that report no dierences in spontaneous EDA
activity between introverts and extraverts and studies that report higher levels of spontaneous
activity in introverts than in extraverts (Smith, 1983; Stelmack, 1990).
Studies of phasic EDA measures and E have shown more consistent trends. These studies
take many methodological forms, but most incorporate EDA measurement before, during and
after a test stimulus is presented (often auditory tones), sometimes in habituation or
dishabituation paradigms. Stimulus and environmental characteristics play an important role in
mediating the outcome of these studies (see Stelmack, 1981, 1990). For example, many
investigations show that introverts produce larger amplitude SCRs than extraverts when
moderate intensity auditory or visual stimuli are used. Personality has little eect in studies
using low level stimulation and at high levels of stimulation extraverts demonstrate higher SCR
amplitude, perhaps due to TMI (Stelmack, 1981, 1990). Similar moderating eects are found in
studies that have manipulated arousal level with caeine (see Smith, 1983), stimulus intensity
(Wigglesworth and Smith, 1976) and stress (Fowles et al., 1977). These studies uniformly
support higher levels of EDA in introverts as compared to extraverts at low levels of arousal,
but not when arousal level is high.
Habituation processes provide data that are less consistent. Early studies suggested faster
habituation in extraverts (O'Gorman, 1977), but more recent reviews concluded that the
evidence for dierences in trials-to-criterion during habituation between the personality groups
was simply not consistent enough to form a clear consensus (Geen, 1983; Smith, 1983). Post
habituation processes, especially dishabituation, seemed to yield fairly consistent ndings that
suggest introverts are more highly aroused than extraverts (Geen, 1983).
In summary, EDA measures of tonic arousal level have been either inconclusive or so
inconsistent as to add little to our understanding of arousal dierences. Studies using phasic
measures and tonic measures assessed during arousal manipulations have been far more
consistent and, in general, support Eysenck's arousal model of E. Studies in this area have not
596
been designed to discriminate the Eysenck and Gray theories, but Fowles (1980) has linked the
BIS to EDA. Unfortunately, there is general agreement that the EDA research in the area of
neuroticism has been `ùniversally negative'' (Zuckerman, 1991, p. 261; see also Fahrenberg,
1987; Naveteur and Freixa-i-Baque, 1987). Furthermore, trait anxiety and N do not appear to
predict either electrodermal orienting or habituation consistently, although state anxiety tends
to block habituation and relates to increased EDA (O'Gorman, 1977; Zuckerman, 1991).
Contrary to the GrayFowles prediction, E appears to be a better predictor of EDA than N is.
3.2. Cardiovascular and other autonomic measures
Measures of cardiovascular activity have also been used to explore arousal dierences
between introverts and extraverts, although, like many other autonomic measures, they are not
entirely under sympathetic control. For example, the cardiovascular system interacts highly
with the respiratory system. This requires greater skill in experimental design and the control
of confounding variables that might inuence these measures. Most studies used heart rate as a
dependent measure. In the majority of cases, no eect of E on baseline or tonic heart rate was
reported (e.g. Myrtek, 1984; Pearson and Freeman, 1991). However, heart rate changes in
response to stimuli or environmental changes have yielded more consistency. Greater increases
(or less deceleration) in heart rate or heart rate variability for introverts, as compared to
extraverts, have been reported during exposure to a series of tones (Orlebeke and Feij, 1979)
and during performance of a variety of tasks (e.g. Richards and Eves, 1991). There is little
evidence for elevated heart rate in subjects high in trait anxiety or N (e.g. Naveteur and Roy,
1990).
Fowles (1980) has suggested that `Ìf there is to be a relationship between HR [heart rate]
and one of the three arousal systems [of Gray] ...it must be with the BAS'' (p. 91). This, of
course, leads to the prediction that higher levels of Imp (and thus E) would exhibit higher
levels of cardiovascular activity, a prediction quite in opposition to Eysenck's predictions and
generally unsupported by the literature on cardiovascular activity that was reviewed above. As
with much of the EDA research, few studies using cardiovascular measures have focused on
Gray's theory. De Pascalis et al. (1996) proposed that heart rate deceleration to feedback
signals might be taken as an index of BIS activity. Neurotic subjects showed greater
deceleration irrespective of feedback, but not the predicted dierential sensitivity to feedback.
There was no interaction between E and N, but, in addition, extraverts were more sensitive to
reward signals, whereas introverts were more sensitive to punishment signals. These data
appear broadly consistent with the Gray theory, but it is curious that introverts but not
neurotics should be especially sensitive to punishment signals. This study indicates a need for
further investigation, but, overall, cardiovascular research leans somewhat in favor of
Eysenck's theory.
Finally, there should be brief mention of several other peripheral nervous system measures.
Support for the Eysenck arousal theory has been obtained in studies of regional cerebral blood
ow (e.g. Stenberg et al., 1990) and pupillary response (Stelmack, 1981). Pivik et al. (1988),
however, showed decreased spinal motoneural recovery in extraverts, which may be attributed
to increased dopaminergic activity. This association seems consistent with Gray's theory.
Mangan and Hookway (1988) showed that exposure to aversive lm clips induced higher heart
597
rate and skin conductance in extraverts than in introverts, but changes in muscle tension
showed the opposite pattern of eect. There were no eects of N or P. Finally, there is a
growing literature on the eyeblink component of the startle reex, measured
electromyographically (e.g. Corr et al., 1997; Kumari et al., 1996). Results seem to vary with
stimulus materials. Studies using unpleasant slides show enhanced response in anxious
individuals, a result compatible with both theories. Responses to pleasant slides (which might
dierentiate the theories) appear inconsistent across studies (Corr et al., 1997).
3.3. Psychophysiology: conclusions
The following inferences may tentatively be drawn from the plethora of studies. First,
associations between extraversion and tonic measures of c.n.s. and a.n.s. arousal are weaker
than might be expected from the Eysenck theory, although the overall trend is in the direction
predicted. Second, studies of phasic measures, including ERPs, SCRs and phasic cardiac
response, provide stronger support for greater arousability in introverts, consistent with the
theory. Third, some of the most consistent ndings concern the dependence of associations
between E and psychophysiological response on level of stimulation, as shown most clearly in
Smith's EDA work. Under high levels of stimulation, extraverts may actually appear more
aroused than introverts. These results are consistent with the Eysenck theory, but require the
additional assumption of TMI in introverts. Gale's hypothesis (Gale, 1973) that relationships
between E and electrocortical arousal were similarly mediated has not been clearly
substantiated (Matthews and Amelang, 1993). However, comparable eects have been obtained
in evoked potential data (Bartussek et al., 1996; Brocke et al., 1997). The problem with the
TMI hypothesis is a calibration problem across studies. The level of stimulation needed to
induce inhibition is never specied a priori and there is little agreement between studies. Brocke
et al.'s data on noise and ERPs suggest TMI operates at the moderate intensity of 60 dB. In
contrast, Fowles' study of EDA implies that the much higher intensity of 103 dB (with
additional stress) is necessary before TMI becomes evident in introverts. Fourth, there is little
evidence that N relates consistently to either arousal or arousability, even under stressful
conditions (Fahrenberg, 1987; Naveteur and Freixa-i-Baque, 1987), although there are sporadic
positive ndings (Eysenck, 1994) and complex, uninterpretable interactive eects. Fifth, there
have been few attempts to test Gray's theory directly and the evidence is mixed. Arguments
that Imp is particularly predictive of psychophysiological functioning (e.g. O'Gorman and
Lloyd, 1987) must be balanced against contrary ndings (e.g. Swickert, 1996). The attempt by
Fowles (1980) to link a.n.s. functioning to Gray's theory has been largely unsuccessful,
although the De Pascalis et al. (1996) heart rate data provide partial support. By and large,
Anx does not relate to increased EDA and Imp does not relate to increased heart rate.
Bartussek's ERP studies also fail to provide consistent support for the theory.
On balance, the evidence favours the Eysenck theory, at least for extraversionintroversion,
although reservations may be expressed about the level of inconsistency in the data. However,
as noted at the outset, reliability of outcome may be reduced by methodological problems and
diculties in establishing clear conceptual links between specic psychophysiological measures
and arousal theory (Eysenck, 1994). Diering reactions to these problems are possible. To the
extent that consistent results are obtained only in highly selected subjects under narrowly-
598
constrained testing conditions, the sceptic might reasonably wonder whether psychophysiological ndings tell us much about personality in the wider sense. The contrary view is
that individual dierences in brain function are more robust than individual dierences
in psychophysiological response. Eysenck's (1994) view was that: ``Given the enormous
complexity of the issues involved, the practical diculties outlined and the instrumental
problems of working at the boundary of the technically possible, I think we would have been
optimistic in the extreme had we expected results to have been more positive and in greater
agreement with theory'' (p. 198).
4. Mood and subjective states

4.1. A framework for mood studies
Both Eysenck and Gray conceptualise aect or mood as a relatively direct output from brain
systems. There are considerable diculties, however, in establishing mappings between neural
systems and dimensions of mood due to the complexity of the physiological basis for aect
(Thayer, 1989, 1996). In reviewing data on personality and mood we shall use the framework
proposed by Matthews et al. (1990a), developed from Thayer's (1989) work on self-report
arousal. Thayer (1989) has shown that there are two quite distinct bipolar dimensions of
subjective arousal, which relate to autonomic arousal measures to a similar degree. Energetic
arousal contrasts feelings of vigour and tiredness and, according to Thayer (1989, pp. 130
131), relates to Eysenck's reticulocortical circuit. This hypothesis is supported by studies
showing facilitative eects of energy on demanding attentional tasks (Matthews, 1992a), in line
with theoretical accounts of cortical arousal and performance (Revelle, 1993). Tense arousal
(anxiety vs. calmness) may relate to limbic system arousal, which Eysenck sees as the basis for
emotion, especially negative emotion. Matthews et al. (1990a) add a further fundamental mood
dimension; hedonic tone or the pleasantness of mood, which contrasts feelings of happiness
and contentment with depression and sadness. Links between mood dimensions and brain
systems then support the following predictions. First, introverts should report greater energy:
Eysenck (1967, p. 83) stated that ``the concept of fatigue in relation to extraversion
introversion takes the place of the concept of emotion in relation to neuroticism-stability''.
Second, N should relate to tension, unpleasantness of mood and mood variability. There is no
basis for relating E to overall pleasantness of mood.
Gray (1987a) relates BIS activity (high Anx) to both subjective anxiety and neurotic
depression. Activity of dopaminergic reward systems, such as the BAS, may underpin positive
aect and subjective energy (e.g. Depue, 1995) and so Imp should tend to relate to positive
emotions. Presumably, noradrenergic arousal heightens the experience of aect. Fig. 2 shows
the dynamic interaction between BIS, BAS and the separate arousal system. Matthews et al.
(1990a) showed that energy and tension correlated with autonomic arousal but hedonic tone
was independent of autonomic indices. Plausibly, therefore, energy relates to the integrated
output of the BAS and the arousal system, tension to the BIS and arousal and hedonic tone
(HT) to the balance between the mutually inhibitory BIS and BAS systems. In terms of the
Eysenck dimensions, N should correlate with tension and unpleasant mood (negative HT),
599
Fig. 2. Possible inter-relationships between brain systems, mood states and personality traits, within Gray's (1987a)
theory.
especially when the BIS is activated, whereas E should relate to energy and pleasant mood,
especially when the reward system is activated.
Secondary predictions may be made from (1) Gray's rotation of axes and (2) interaction of
systems in the conceptual nervous system shown in Fig. 2. Because introversion is associated
with Anx, we expect introversion to relate, modestly, to negative emotions, i.e. increased
tension and lower HT. Similarly, the association between Imp and N implies high N subjects
should be somewhat more prone to positive hedonic tone. In addition, Fig. 2 shows that the
BIS and BAS are mutually inhibitory, but both systems tend to excite the arousal system.
Hence, N should relate to suppression of the BAS by the BIS and lower HT. The association
between N and pleasantness of mood may then be somewhat unstable, consistent with the high
mood variability of neurotics (Eysenck and Hepburn, 1989). Presumably, the dominant eect
of N varies with motivational context: N should be more positively related to pleasant moods
in situations which do not activate the BIS.
In summary, the two theories make similar predictions concerning aective correlates of N,
although the Gray theory also allows for variation of the association between N and positive
mood, depending on the dynamic interaction of the systems. However, the Eysenck and Gray
theories make conicting predictions concerning the direction of the relationship between E
and energy (Matthews et al., 1990a). The Gray theory also predicts that E should be associated
with higher HT and perhaps also with lower tension, via the two secondary eects described. A
diculty for the Eysenck theory is that extraverts' stimulation-seeking tendencies and moodregulation strategies might compensate for their lower alertness, especially in naturalistic
studies. Extraverts report desiring more activated and more pleasant mood states than do
introverts (Rusting and Larsen, 1995) and extraverts are more likely than introverts to use
exercise to increase feelings of vigour (Thayer et al., 1994).
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4.2. Correlational studies

There are two distinct branches of research on personality and mood, which relate to
psychometric and experimental traditions respectively. Illustrative results from some of the
larger-scale investigations are shown in Table 1. The rst four studies were concerned with
establishing relationships between personality and mood as psychometric constructs: the
Watson and Clark (1992) positive and negative aect dimensions are similar to energy and
tension, respectively. Typically such studies use a diary-based approach or require college
students to complete questionnaires for course credit. The studies show consistent correlations
between (1) E and energy/positive aect and (2) N and tension/negative aect, but correlation
magnitudes vary. Meyer and Shack (1989) and Watson and Clark (1992) have argued for a
particularly close correspondence between these pairs of dimensions. Other psychometricallyoriented studies suggest that both E and N relate to overall pleasantness of mood (HT).
Williams (1989) found that an elation scale correlated at 0.30 with E and 0.20 with N and
depression correlated at 0.31 with E and 0.45 with N (N = 172). The remaining studies listed
took mood measures in the context of experimental studies of performance and show
Table 1
Data from illustrative studies of personality and mood
Study
Costa and McCrae (1980)
575
Emmons and Diener (1986)
72
Meyer and Shack (1989)
231
Watson and Clark (1992)
532
Larsen and Ketelaar (1991)1
70
Adan and Guardia (1997)2
578
Adan and Guardia (1997)3
381
Matthews et al. (1990a)
158
Matthews et al. (in press)
762
Measures
EPI
PAS/NAS
EPI
ACL
EPQ
ACL
NEO PI
PANAS
EPQ
ACL
EPI
UMACL
EPI
UMACL
EPI
UMACL
EPQ-R
UMACL
Energy (PA)
Tension (NA)
11**
16**
35**
02
34**
32**
19*
50**
54**
25**
62**
03
Hedonic tone
(happiness)
E
01
11
52**
21**
10
29**
12
20**
18**
26**
08
27**
24**
06
11
18**
01
24**
23**
25**
13
23**
15
24**
12
13*
14*
38**
16**
26**
12**
24**
Neutral mood induction.2Additional data (personal communication, 30/6/97): subjects tested at 9 AM.3Additional
data (personal communication, 30/6/97): subjects tested at 9 PM.Correlation coecients multiplied 100, *p < 0.05,
**p < 0.01.PA = positive aect, NA = negative aect, EPI = Eysenck personality inventory, EPQ(-R) = Eysenck
personality questionnaire(-revised), PAS = positive aect scale, NAS = negative aect scale, PANAS = positive and
negative aect schedule, UMACL = UWIST mood adjective checklist, ACL = unpublished adjective checklist.
601
considerably weaker personalitymood associations. The data suggest that N is modestly

related to both higher tension and lower energy and to more depressed mood (lower hedonic
tone). E shows a weak but general tendency to be related to better mood.
Several factors may account for variation in results across studies (Dorn and Matthews,
1995). The rst is the time-frame of the mood rating, with longer time-frames giving stronger
correlations. Meyer and Shack's (1989) ``state'' measure asked for a mood rating for the past
day, for example, whereas the measure used by Matthews et al. (1990a); Matthews et al., in
press) emphasises feelings `àt the moment''. Second, the inuence of personality on mood
varies across situations (Brandstatter, 1994). Studies of college students in which the ``task'' is
no more than the completion of a set of questionnaires may simply pick up characteristic
reactions to university life. The happiness of extraverts may in part reect their greater
participation in social activities, which appears to be mood-enhancing (Watson et al., 1992).
Similarly, N relates to a greater frequency of negative life events (Bolger and Schilling, 1991),
although the relationship between N and emotional distress is robust in controlled settings
(Matthews and Deary, 1998). Thirdly, the personality measure used may inuence correlations.
The NEO-PI used by Watson and Clark includes two explicitly aect-laden ``facets'' (positive
emotions and activity). In one of their samples, Watson and Clark showed that omitting these
two facets in scoring E reduced the EPA correlation from 0.54 to 0.42, a substantial decrease
in the variance explained. Broadly, the correlational data suggest that neurotics are prone to
negative moods and less consistently, extraverts tend to experience positive moods.
4.3. Experimental studies
Experimental evidence on individual dierences in mood response to manipulated events is
limited. Blackburn et al. (1990) used the Velten technique to induce state depression and found
that N (but not E) predicted the increase in depression. Larsen and Ketelaar (1991) obtained a
similar result when subjects were required to imagine negative scenarios vividly. They also
showed mood enhancement due to imagining positive events was stronger in extraverts than in
introverts. Correlations from a neutral mood induction condition are given in Table 1 above.
With a negative mood induction, the NNA correlation increased slightly to 0.34, whereas a
signicant correlation of 0.32 between E and PA was obtained with the positive induction.
Rusting and Larsen (1997) report similar ndings.
There is a somewhat independent line of research based on interactionist models of anxiety.
There may be multiple anxiety traits related to dierent contexts, so that N and trait anxiety
may only moderate subjective responses to certain types of threat (Endler et al., 1991). Trait
anxiety may relate primarily to sensitivity to ego-threat (Eysenck, 1982). Studies conducted in
Endler's laboratory (e.g. Busch et al., 1994) suggest that both multiple anxiety traits and the
individual's cognitive appraisal of the nature of the threat must be taken into account in
predicting state anxiety change. Hence, a cognitive explanation for state anxiety change may be
more valid than a neuropsychological one. Neurotic/trait anxious individuals may have
particularly ready access to negative self-referent information in long-term memory, for
example (Wells and Matthews, 1994) and N relates to cognitive state variables such as
cognitive interference, lack of perceived control and low self-esteem in addition to negative
mood (Matthews et al., in press).
602
4.4. Mood: conclusions

N is reliably positively correlated with various aspects of negative mood, consistent with
both Eysenck and Gray theories. N is a considerably more reliable predictor of subjective
emotion than it is of autonomic arousal, even in the absence of an overt stressor. Hence, it is
unclear whether the trait-state relationship should be seen as dependent on neuropsychological
or cognitive processes (or some interaction between the two). With regard to E, the critical
issue is whether subjective energy is seen as an index of cortical arousal or of BAS activity. If
we interpret energy as reticulocortical arousal (Thayer, 1989), then its positive correlation
with E is strongly incompatible with the Eysenck theory and unpredicted by Gray. If we see
energy as a marker for a dopaminergic reward system, its association with E is unpredicted by
Eysenck, but highly consistent with the Gray theory. The Eysenckian counter-argument is that
extraverts tend to over-compensate for low arousability through stimulation-seeking. This
hypothesis receives some support from the attenuation of the correlation between E and energy
in laboratory settings which control for activity preference, but remains somewhat post hoc.
There is a marked discrepancy between the tendency of E to correlate negatively with
psychophysiological arousal measures, but positively with subjective energetic arousal.
The tendencies for E to relate weakly to reduced tension and N to reduced energy are
also compatible with the secondary predictions made from the Gray theory. The data on
measures of happiness and hedonic tone suggest that N and E may relate to the balance of the
BIS and BAS as Gray predicts. In experimental studies, we expect secondary eects of (1)
attenuation of negative aect response in extraverts, due to their lower Anx and (2)
enhancement of positive aect response in neurotics, due to their higher Imp. Larsen and
Ketelaar (1991) and Rusting and Larsen (1997) obtained the rst but not the second of these
eects. Contrary to Gray's theory, Emmons and Diener (1986) found that Soc was more
predictive than Imp of positive mood. Matthews et al. (1990a) suggested that, in mood studies,
it may be preferable to use the Eysenck dimensions, but interpret them in terms of Gray's
dimensions. Overall, mood data t the Gray theory quite well, but cognitive mechanisms
provide viable alternative explanations. Matthews (1997) reports that, across two studies,
correlations between E and mood covaried with the strength of correlations between E and
appraisal and coping scales.
5. Conditioning
Studies of conditioning provide one of the principal sources of evidence on both
theories. Recommended articles in this area include Levey and Martin's (1981) review of
methodological issues and earlier studies and Corr et al.'s (1995a) thoughtful discussion of the
use of conditioning paradigms to test the two theories. Prediction from Eysenck's theory
is based on the ``drug postulate'', that eects of extraversionintroversion on conditioning
should correspond to eects of depressant and stimulant drugs respectively. Pharmacological
studies indicate that arousal is correlated with rate of conditioning in both associative
and instrumental paradigms, implying that, in general, introverts should condition more
rapidly than extraverts (Eysenck, 1994). As ever, testing the prediction requires attention to
603
task parameters. In particular, in associative learning, TMI may disrupt conditioning if

the unconditioned stimulus (UCS) is particularly strong, so theory-testing requires the use
of a weak UCS. Presumably, relatively weak stimuli are also desirable in instrumental
paradigms.
Corr et al. (1995a) contrast the ``HullEysenck'' perspective, that there is a general learning
mechanism, with the ``MowrerGray'' perspective, that there are two learning mechanisms
related to reward and punishment respectively. The simplest prediction from Gray's (1981)
theory is that in anxious individuals, sensitivity to conditioned aversive stimuli leads to
enhanced learning with such stimuli. Similarly, Imp relates to learning with conditioned reward
stimuli. However, some complications to these straightforward predictions should be
mentioned. First, Gray's theory does not deal with generalised sensitivity to reward and
punishment, but with sensitivity to specic cues (Zinbarg and Revelle, 1989; Pickering et al.,
1995). In conditioning tasks, it is easier to manipulate these cues than in other behavioural
paradigms. Hence, predictions concern conditioning to CSs rather than to USs: Gray (1987a,
p. 244) suggests that the Pavlovian association of neutral CSs with unconditioned reinforcers
may be a function of the ght/ight system. However, Gray (1987b, p. 501) cautions that the
degree of correspondence between individual dierences in conditioning to CSs and UCSs is an
open empirical question and Gray (1987a, p. 351) makes the explicit prediction that extraverts
will condition better with rewarding UCSs.
Second, Gray's theory does not specify the role of arousal in conditioning and it is unclear
how the empirical data on drugs and conditioning which provide the basis for Eysenck's drug
postulate are to be explained, unless it is supposed that stimulants act through their eects on
reward and punishment systems and their eects on arousal are incidental.
Third, the Gray theory not only proposes two learning factors (reward and punishment
systems), but also two learning processes for associative and instrumental phases of learning
(Corr et al., 1995a). Conditioned response to motivationally signicant stimuli involves an
initial stage of classical conditioning of a neutral CS to a UCS, followed by a stage of
instrumental conditioning of the CS to a response. Studies of instrumental learning have
tended to conate the two stages, but it may be important to separate them. In particular,
anxiolytic drugs fail to inuence aversive associative learning (Gray, 1982), so there is no
rationale for predicting anxiety eects on this phase of learning.
Fourth, there have been conicting statements over whether the theory predicts any form of
learning or not. Until recently, most statements of the theory explicitly related personality to
learning (e.g. Gray, 1981). Corr et al. (1995a, p. 52) state that ``...Gray's theory predicts that
two separate personality factors should aect associative learning under appetitive and aversive
stimuli...''. However, Gray (personal communication, 18/11/97) states that the animal studies
of anxiolytic drugs do not support predictions concerning anxiety eects on associative
conditioning. Instead, the theory is primarily concerned with the motivating properties of
associatively conditioned stimuli when the subject has learned a task to an asymptotic level
(Pickering et al., 1997). In fact, many human conditioning studies show personality eects on
learning following reinforcement cues, indicating a need for further theoretical restructuring, as
Pickering et al. (1997, pp. 5353) accept. Furthermore, animal studies of instrumental learning
have shown eects of anxiolytic drugs on acquisition but not asymptotic performance
(McNaughton, 1985), although these acquisition eects relate to response suppression rather
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than to conditioning per se (McNaughton, personal communication, 17/4/98). The problem is

really that conditioning is not a basic process, but the outcome of various factors including
sensitivity to motivational signals and pre-existing response tendencies. In this section, we will
focus on the version of the theory (Gray, 1981) actually addressed by most published
conditioning studies, which predicts individual dierences in learning. We will also highlight
the key aspects of Pickering et al.'s (1997) comments on these studies.
Overall, the strongest test of the Gray (1981) theory would be provided by studies of
instrumental conditioning to weak conditioned stimuli. Under such circumstances both the
Eysenck and Gray theories predict superior conditioning in passive avoidance and extinction
paradigms in introverts and, in stressful conditions, in individuals high in N. Within Gray's
current ``308 rotation'' model, we would expect N to be more strongly related than introversion
to BIS-mediated conditioning. However, with BAS-mediated conditioning, in approach and
active avoidance paradigms, Eysenck continues to predict superior conditioning in introverts
and stressed neurotics, but Gray (1981) predicts superior conditioning in extraverts and stable
individuals (especially extraverts). This clear predictive dierence has inspired much recent
work on conditioning.
5.1. Associative learning
The most popular associative conditioning paradigm has been the learning of the
conditioned eyeblink response, in which the UCS is an aversive air-pu. It seems wellestablished that with a weak UCS, introverts do indeed condition faster than extraverts, as
predicted by Eysenck (Levey and Martin, 1981). Extensive work on anxiety was conducted
within a drive theory framework in the 1950s and 1960s, although interest in this topic declined
sharply in subsequent decades. Anxious individuals condition faster than stable individuals in
stressful circumstances, such as those induced by noxious stimuli or ego-involving instructions
(Spence, 1964; Eysenck and Eysenck, 1985). These ndings are broadly consistent with the
Gray theory, to the extent that the stressors concerned activate the BIS. The reliability of the
extraversion eect is perhaps surprising, given that in Gray's current model E is only weakly
related to Anx.
However, more detailed scrutiny of individual studies often turns up ndings which are
dicult for either theory to accommodate, such as complex interactive eects of E, N and
experimental parameters (Levey and Martin, 1981). Eects of E on eyelid conditioning are
associated with narrow Imp and venturesomeness rather than with Soc or E per se (Eysenck
and Levey, 1972; Frcka and Martin, 1987). Gray (1981) infers that Imp rather than E may be
the key causal factor. However, the result does not seem to support Gray's theory either, in
that Imp should aect conditioning only with rewarding stimuli and eyelid conditioning
paradigm is usually seen as aversive. Frcka et al. (1983) showed a moderating eect of P in an
eyelid conditioning study. E was negatively related to CR frequency in low P individuals.
Among those high in P, extraverts emitted more CRs. Assuming that P is negatively related to
cortical arousal (Eysenck, 1994) the E P interaction is in the wrong direction. Introverts'
superior conditioning should be more reliable in the high P individuals, because of their lower
arousal. Another study providing somewhat ambiguous results is that of Barrett (1971), who
showed that both low Imp and high Anx were correlated with stronger conditioning. The data
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show eects of the Gray dimensions, but the problem for the Gray theory is that, with a
neutral CS and aversive UCS, it is unclear why BAS activity and Imp should inuence classical
conditioning.
Corr et al. (1995a) point out that the rather few studies of personality and associative
conditioning which have used appetitive stimuli have yielded mixed results. They cite two
studies which showed no eect of E on appetitive conditioning and two studies of sexual
conditioning which supported Gray's theory in showing that E was positively related to
conditioning with (presumably appetitive) sexual stimuli. Paisey and Mangan (1988) report a
study of electrodermal CRs employing both appetitive (sexual) and aversive stimuli, further
subdivided into weak and strong stimuli. They concluded that E relates negatively to
acquisition of the CR with weak appetitive stimuli and positively to acquisition with strong
appetitive stimuli, although results rest on a factor analysis of response indices conducted with
an inadequate sample size (31). The role of stimulus strength is similar to that found in
eyeblink conditioning (Levey and Martin, 1981) and, in this respect, the ndings are consistent
with the Eysenck theory. It is dicult for Gray's theory to explain why introverts should show
stronger conditioning than extraverts with weak sexual stimuli. However, given that extraverts
report more frequent and more diverse sexual encounters than introverts (Eysenck and
Eysenck, 1985), it is questionable whether sexual responses are ideal for theory-testing:
personality is confounded by prior learning. Evidence on the roles of Anx and N on appetitive
classical conditioning is even weaker. Paisey and Mangan (1988) cite three studies showing a
positive, a negative and no relationship between Anx or N and conditioning. Their own study
found N was negatively correlated with CR formation to appetitive stimuli, irrespective of
stimulus strength, a result which ts neither theory, especially if it is assumed that N is a
marker for high Imp.
Corr et al. (1995a; Experiment 2) used a task which required subjects to learn associations
between coloured lines (neutral CSs) and a subsequent visual display. The subject's response
was followed by an appetitive, neutral or aversive UCS, indicating loss or gain of money. E
was negatively correlated with learning in aversive, but not neutral or appetitive, conditions.
Corr et al. argue that this nding conicts with Eysenck's theory, in that personality eects
were moderated by type of reinforcement. From the perspective of Gray's theory, it is
surprising that it is E rather than N which moderated aversive learning, although there is an
argument that associative conditioning is outside the scope of Gray's theory (Pickering et al.,
1997).
5.2. Instrumental conditioning
Most studies of instrumental conditioning have used both positive and negative
reinforcement. The two most popular paradigms are the verbal operant paradigm and
discrimination learning of simple psychomotor responses. In verbal operant conditioning, the
subject is reinforced for producing attributes of sentences (e.g. `Ì'' or ``we''), within approach
or extinction paradigms. This task has been used mainly to investigate E, though results have
been somewhat mixed. Mangan's (1982) review of early work suggests that introverts tend to
condition more strongly than extraverts under rewarding conditions, consistent with Eysenck's
theory. However, several studies carried out by B. S. Gupta and his associates consistently
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showed that extraverts condition better with a rewarding stimulus, but introverts condition
better with a punishing reinforcer (e.g. Gupta and Shukla, 1989). These ndings t the Gray
theory better than the Eysenck theory. Gupta and Gupta (1984) also manipulated arousal
through administration of D-amphetamine. Results with a punishing reinforcer (electric shock)
showed that introverts conditioned more in a placebo condition, but amphetamine improved
conditioning in extraverts and impaired it in introverts. This nding is consistent with the
Eysenck theory, assuming TMI operates in introverts. It does not directly address the Gray
theory, given that shock is an aversive UCS rather than a CS. With a positive reinforcer
(experimenter says ``good''), extraverts showed greater conditioning than introverts in a
placebo condition. Amphetamine reduced conditioning in extraverts, but had no eect on
introverts. Placebo condition data are more consistent with Gray than with Eysenck, but the
drug eect is dicult for either theory to explain. In Gray's theory, both E and amphetamine
(as a dopaminergic agonist) should activate the BAS, with the combination of factors leading
to particularly good conditioning. If, as Eysenck claims, extraverts are under-aroused initially,
amphetamine should improve their conditioning. Comparison of Imp and Soc in some of these
studies has shown that both dimensions give similar results to analyses of E (Gupta, 1990).
Two studies using discrimination learning are of particular relevance to theory (Zinbarg and
Revelle, 1989; Corr et al., 1995a). Zinbarg and Revelle point out that moderation of learning
by cue type provides a more direct test of Gray's theory than moderation by type of
reinforcement. They required subjects to learn to respond to ``go cues'' or reward signals,
indicating reward or active avoidance of punishment and to ``no-go cues'' or punishment
signals, indicating omission of reward or passive avoidance of punishment. Reinforcement type
(reward vs. punishment) was also manipulated: subjects could win or lose points. Gray's theory
predicts interactive eects of Imp and Anx with cue type. The study was designed not to be
over-arousing, so Eysenck's prediction of generally better conditioning in introverts might be
tested. Across four studies, the most consistent learning eect was the interaction between cue
type, Imp and Anx. With go cues, high Anx facilitated learning among low impulsives, but
with no-go cues, Anx facilitated learning among high impulsives. This pattern of interaction is
inconsistent with Gray's theory, most simply because Anx should not inuence conditioning to
go cues and Imp should not aect performance with no-go cues. Zinbarg and Revelle modify
Gray's model to accommodate their ndings by incorporating constructs of expectancy and
processing resources, although they admit the modications are speculative. E and N showed
somewhat similar results to Imp and Anx respectively, but ndings with these dimensions were
less consistent, implying that the Gray dimensions may be more valid as predictors of
conditioning. Results provided little support for the Eysenck theory, with introverts showing
no general superiority in conditioning.
Corr et al. (1995a; Experiment 3) ran a somewhat similar discrimination learning study in
which subjects could win or lose small sums of money on each trial of a psychomotor task, by
modulating response speed following presentation of passive avoidance and approach cues.
Speed of response to approach cues was related to lower trait anxiety and frequency of
punishments in the passive avoidance condition was higher in impulsive subjects. These results
do not clearly t either theory. From Gray's perspective it is hard to see why Anx should
relate to approach behaviour, which is mediated by the BAS and Imp should relate to BISmediated passive avoidance. The authors suggest the nding should be attributed to
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inconsistency of response suppression during punishment. The interaction between Imp, Anx
and type of reinforcement obtained by Zinbarg and Revelle (1989) was also tested and found
to be non-signicant.
Pickering et al. (1997) describe the unpredicted eects of anxiety and impulsivity in the Corr
et al. (1995a) study as ``complementary-trait'' eects. Similar ndings suggesting that reward
signals increase behavioural activation in low anxious subjects have also been found in mazecrossing and card sorting tasks and in a startle reex study whose results failed to replicate
subsequently (see Kumari et al., 1996; Pickering et al., 1997). It is suggested that, in more
anxious individuals, individual dierences in BIS activity elicited by the experimental setting
tend to mask eects of positive reinforcement, because of the inhibitory eects of the BIS on
the BAS. This hypothesis does not explain the absence of eects of anxiety on passive
avoidance in the Corr et al. (1995a) study. Pickering (1997) suggests that neural network
modelling of the interaction between BIS and BAS systems may resolve anomalous ndings. A
simple mutual inhibition model failed to show the complementary-trait eect. A further model
included an arousal unit in the network and made the radical proposal that the BIS and BAS
aect response only via their facilitative eects on arousal. If it is assumed that (1) the BIS has
a stronger eect on arousal than the BAS and (2) arousal is related to response speed by the
YerkesDodson Law, then the complementary-trait eect for anxiety can be modelled. Given
the further modication of the revised Gray theory here, it is perhaps safest to view this result
as a demonstration of the potential of neural network modelling for explaining unexpected
eects, rather than as a strong explanation for the empirical ndings.
Various studies using other learning tasks show superior learning of extraverts under reward.
Pickering et al. (1995) list these studies and nd general support for greater sensitivity to
reward in extraverts and greater sensitivity to punishment in introverts. However, they point
out a variety of diculties in deriving clear predictions for learning tasks and detailed task
analysis may be necessary. There is evidence too for arousal eects on learning. Corr et al.
(1995b) report that caeine enhances procedural learning in extraverts, but impairs learning in
introverts, consistent with other performance research reviewed in the next section. This eect
was associated with Soc, but not Imp.
5.3. Conditioning: conclusions
First, ndings are only consistent within more ``traditional'' conditioning paradigms, notably
the eyeblink and verbal conditioning paradigms. It is disturbing that the well-designed and
thorough discrimination learning studies of Zinbarg and Revelle (1989) and of Corr et al.
(1995a) give entirely dierent results. In discrimination learning, subjects are explicitly
instructed to learn, which may make the task more dependent on high-level cognitive
mechanisms. Second, eyelid conditioning and verbal instrumental conditioning under
punishment are related to personality broadly as the Eysenck theory would predict, although
there are some anomalies in specic studies (Levey and Martin, 1981; Frcka et al., 1983).
Enhanced learning of introverts under these conditions may be explained by the Gray (1981)
theory to the extent that introversion is a proxy for high Anx. Third, results with verbal
operant conditioning under reward are more consistent with Gray than with Eysenck. Fourth,
some studies suggest that Anx and Imp are more reliable predictors of conditioning than E and
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N, but others fail to support this conclusion. Fifth, the most recent account of Gray's theory
(Pickering et al., 1997) muddies the waters by eschewing prediction of individual dierences in
learning and by emphasising interesting but unpredicted empirical ndings such as
``complementary-trait'' eects. It is dicult to evaluate post hoc interpretations for such eects
without further empirical work.
On balance, ndings appear to favour the Gray (1981) theory, in that the Gupta studies
conrm a primary prediction of the theory, enhanced instrumental conditioning to reward
signals. However, dierent reinforcers may vary in their eects on arousal and personality may
moderate the arousal response to motivationally signicant stimuli, so that manipulations
intended to test the Gray theory are confounded with arousal. The argument is sound, but
tends to be used in a rather inconsistent way. For example, Eysenck and Eysenck (1985, p.
245) suggest that appetitive conditioning may be more arousing than aversive conditioning (at
least with sexual stimuli). Corr et al. (1995a, p. 52), however, supposed that reward is less
arousing than punishment. They showed that the aversive CS induced more subjective tense
arousal on the UWIST Mood Adjective Checklist (Matthews et al., 1990a) than did the
appetitive CS. The nding that introversion related only to conditioning with an aversive CS is
thus problematic for the Eysenck theory, because introversion predicted conditioning only in
the most arousing of the three reinforcement conditions used. If (non-sexual) rewards only
induce moderate arousal, the superiority of extraverts on verbal operant conditioning under
reward becomes even more dicult to explain from the Eysenck theory. It is unfortunate that
so few studies have tested for possible confounds.
Some of the data may be explained by combining aspects of the Eysenck and Gray theories,
supposing that (1) arousability of extraverts and introverts is moderated by the incentive
value of the stimulus and (2) arousal is the principal inuence on conditioning. If, as the
subjective arousal data suggest, extraverts are more aroused by reward signals (or possibly
reward stimuli in general), then extraverts may show superior conditioning in rewarding
conditions simply because they are more aroused than introverts. Impairment of conditioning
in extraverts given amphetamine (Gupta and Gupta, 1984) might then reect TMI, although it
is curious that the apparent interaction between E and stimulus strength in conditioning with
sexual stimuli suggests greater TMI in introverts (Paisey and Mangan, 1988). Similarly,
introverts may be more aroused than extraverts under punishment conditions and during eyelid
conditioning.
6. Attention and performance

Studies of human performance have long been of interest to researchers testing Eysenck's
arousal theory. In recent years, there has been an upsurge of interest in the implications of
Gray's theory for performance. Eysenck links personality to performance on the basis of three
quite separate hypotheses. First, introverts and stressed neurotics tend to be chronically high in
cortical arousal during performance. Second, cortical arousal is related to performance by the
inverted-U curves specied by the YerkesDodson Law. Third, subjects are motivated to seek
an intermediate level of arousal, which may aect their strategy for response. Hence, eects of
E on performance should be moderated by arousal. In particular, the normally under-aroused
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extravert should perform more eciently under stimulating conditions, whereas de-arousing
environments favour the introvert. Gray's theory makes fewer predictions outside the
conditioning paradigms already discussed. Much of the work to be reviewed is concerned with
the implications of ``sensitivity'' to reward and punishment cues for performance on tasks
which do not correspond closely to those used in animal research. It is often unclear whether
sensitivity to motivational signals should be expressed as performance enhancement or as
distractibility.
The literature on personality and performance is far too extensive to review in full (see
Eysenck and Eysenck, 1985; Matthews, 1992b, 1997; Matthews and Dorn, 1995). In this
section, we consider key studies in four areas of particular theoretical relevance: the interaction
of extraversion and arousal, attentional systems, memory and responsiveness. These studies
have focused mainly on E. In the nal sub-section, we briey review the theoretical
implications of studies of Anx and N conducted outside the Eysenck and Gray frameworks.
6.1. Extraversion: moderating eects of arousal and task factors
The simplest prediction from Eysenck's theory, that extraverts should tend to out-perform
introverts in arousing conditions, is quite well-supported (Eysenck and Eysenck, 1985). Studies
of perceptual thresholds provide striking evidence. Introverts show higher auditory and visual
thresholds than extraverts in baseline conditions, but additional stimulation in a dierent
modality leads to threshold increasing in introverts, but decreasing in extraverts (Shigehisa and
Symons, 1973; Shigehisa et al., 1973). Extraverts' superiority in performance on attentionally
demanding tasks (other than vigilance) noted by M. W. Eysenck (1981) is also consistent with
theory. According to the YerkesDodson Law, more dicult tasks have a lower optimal level
of arousal for performance and so extraverts should be advantaged. At one level, this line of
research supports the Eysenck theory, in that Gray's theory provides no straightforward
explanation for these performance eects.
However, the assumptions made in linking data on E arousal interactions to arousal have
been increasingly challenged. Typical factorial designs are notoriously weak as a means for
testing arousal theory (Hockey, 1984) and, as discussed previously, the evidence for higher
tonic arousal in introverts in performance assessment settings is unconvincing. At an empirical
level, three areas of diculty for arousal theory have emerged: the role of time of day, the role
of task factors and the lack of evidence for arousal-mediation. Interaction between E and
arousal appears to vary with time of day (Revelle et al., 1980). In the evening, extraverts
perform better under low arousal, implying they are over-aroused. Revelle et al. (1980) suggest
that eects of E are driven by Imp, which relates to a phase dierence in circadian arousal
rhythm, such that high impulsives are indeed more aroused than low impulsives in the evening.
There are diculties for this theory too, such as the weakness of the association between E and
morningness-eveningness (Matthews, 1988; Tankova et al., 1994). The reliability of the
superiority of Imp over Soc measures as predictors of performance has also been questioned
(Amelang and Ullwer, 1991). Nevertheless, the Eysenck theory oers no explanation for the
moderating eect of time of day.
Another type of moderating factor is the nature of the task. The YerkesDodson Law
concerns generalised cortical eciency, such that task diculty but not the precise information-
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processing demands of the task inuence the arousal-performance relationship. In fact, task
demands appear to be of critical importance. The characteristic E arousal interaction is
largely restricted to simple tasks requiring encoding of easily-perceived stimuli or to more
complex tasks with a routine encoding component (Matthews and Dorn, 1995; Matthews,
1997). Attentionally demanding tasks do not show the eect, although the YerkesDodson
Law predicts that the over-arousal of introverts should be particularly deleterious on dicult
tasks. Matthews and Harley (1993) relate the interaction specically to spreading activation
processes related to low-level encoding.
A nal diculty is evidence that eects of E are not in fact mediated by individual
dierences in cortical arousal. Matthews (1992b) reviews studies in which measures of all
relevant constructs were taken arousal, E and performance and concludes that they
provide little support for the arousal-mediation hypothesis. In studies of self-report arousal,
interactions between E and arousal show eects of E with arousal statistically controlled
(Matthews, 1985). In other words, arousal moderates rather than mediates the E eect. Similar
ndings have been obtained when arousal is measured through EEG alpha power (Matthews
and Amelang, 1993).
6.2. Studies of attention
Attentional paradigms are frequently used in personality research, but many studies neglect
the distinctions between dierent systems developed by cognitive neuropsychologists (e.g.
Posner and Rothbart, 1991). The Eysenck theory predicts a general enhancement of attention
in more aroused individuals (i.e. introverts), but more specic predictions may be derived for
the Gray theory.
6.2.1. Spatial attention
Orienting attention in space is a functionally and anatomically distinct aspect of attention.
Typical experimental studies use a cue to draw the subject's attention to a peripheral location,
followed by a target which may or may not be at the cued location. Posner and Rothbart
(1991) describe a ``posterior'' orienting system which is innervated by the DNAB, such that
release of noradrenaline facilitates speed of orienting and slower disengagement from the
attended stimuli (Posner and Raichle, 1994). Gray's theory implies that impulsives given
reward signals and anxious individuals exposed to punishment cues should show greater
DNAB arousal and enhanced spatial orienting. Consistent with prediction, Avila (1995)
and Derryberry and Reed (1997) showed that N and Anx are associated with slower
disengagement of attention following orienting to a cued location in space. In Derryberry and
Reed's (1997) study the eect was contingent upon most trials being negative. However, Anx
fails to predict speed of moving to a cued location, even when cues are negative (Derryberry
and Reed, 1997).
Derryberry and Reed (1994) used two target locations, one associated with reward and one
with punishment (gaining or losing points). They argued that Gray's theory predicts that
personality should interact with cue type (i.e. cue at positive location vs. negative location).
They also manipulated post-trial feedback. Results were complex, but the most consistent eect
across the various studies was an interaction between E, cue type and feedback on the previous
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trial. Personality eects were strongest following negative feedback. Under these conditions,
extraverts were relatively slow following a positive cue, but introverts tended to show greater
costs of the negative cue. These eects tended to be stronger in high N individuals. Derryberry
and Reed (1994) suggest that disengagement is slowed in neurotic introverts (high Anx) given
negative feedback and in neurotic extraverts (high Imp) given positive feedback.
Results from spatial attention studies provide some support for Gray, but there are problems
also. First, Gray's theory does not provide a rationale for the moderating role of feedback,
which is an actual outcome rather than a signal of reinforcement (Derryberry and Reed, 1994).
Second, Derryberry and Reed (1994) report briey that analysis in terms of E and N provided
a more coherent pattern of results than analysis using Imp and Anx factors. Third, the eect
seems specic to disengagement; Gray's theory provides no rationale for distinctions between
the dierent components of spatial orienting. It is dicult to accommodate ndings on spatial
attention within the Eysenck theory in any simple way.
6.2.2. Target detection
It is often supposed that the analysis of a stimulus required for object identication or
discrimination (``what is it?'') is distinct from analysis of its spatial location (``where is it?'').
Posner and Raichle (1994) describe an anterior system for detection of events located in areas
of midprefrontal cortex, inuenced by dopaminergic aerents. It is active during visual target
detection and conscious eortful control of attention.
Tasks in which frequent targets are delivered through multiple channels are expected to
activate the anterior system. In fact, extraverts seem to enjoy a general performance advantage
on multiple-channel tasks, especially with verbal or symbolic stimuli (M. W. Eysenck, 1981;
Matthews, 1997). This eect does not appear to be arousal-mediated: Eysenck and Eysenck
(1979) demonstrated superior dual-task performance in extraverts irrespective of manipulated
arousal (noise). Tentatively, this eect might be attributed to dopaminergic inuences on
Posner's anterior system, consistent with Gray's hypothesis that Imp (and hence E) correlates
with dopaminergic activity. There appears to be anatomic overlap or near-overlap between the
paralimbic structures, such as parts of the cingulate cortex, which contribute both to the event
detection system and to the BIS function of inhibition of motor plans. Furthermore, eects of
subjective energy on demanding attentional tasks are often considerably stronger than those of
E and energy may reect activity of dopaminergic circuits (Matthews and Davies, 1998).
Anx and N frequently impair demanding, non-spatial attentional tasks (e.g. M. W. Eysenck,
1992). However, psychobiological explanations may be misplaced. Performance decit is
associated with worry rather than with emotion and bodily symptoms and anxiety interacts
with a variety of information-processing parameters in ways unpredictable from arousal
theory, leading most researchers to adopt cognitive theories of anxiety eects (M. W. Eysenck,
1992). Derryberry and Reed (1997) have shown enhanced attentional focusing to targets in
negative incentive conditions, although, in contrast to spatial attention, there were no
moderating eects of the incentive value of the individual trial or of feedback. The eect was
found only for right visual eld targets, supporting an explanation in terms of ``tonic
activation'' of dopaminergic projections from the ventral tegmental area to left hemisphere
object processing pathways. This account of the anxiety eect is at odds with Gray's
conception of anxiety.
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6.2.3. Vigilance
The traditional vigilance task requires detection of infrequent targets over prolonged periods
of work. Arousal tends to fall during performance and so, according to the Eysenck theory,
extraverts should be particularly prone to under-arousal and performance decrement. Posner
and Rothbart (1991) relate the DNAB to a vigilance function, the clearing of consciousness to
maintain readiness for detecting incoming stimuli. Impulsives and high anxious individuals are
more prone to arousal of the DNAB (see Fig. 1) and so these two groups should tend to show
superior vigilance. Vigilance tasks tend to be somewhat unpleasant, so we might expect Anx
eects to predominate. In this case, the evidence favours Eysenck over Gray, at least
supercially. Introverts show an advantage in detection rate and perceptual sensitivity on
visual, but not auditory, vigilance tasks (Koelega, 1992) and extraverts' performance decit is
sometimes eliminated by arousal manipulations (Davies and Parasuraman, 1982). Giambra et
al. (1989) found that such eects relate to Soc rather than Imp. Unfortunately, results from
studies which have taken an independent measure of arousal suggest that eects of E are not
directly mediated by individual dierences in cortical arousal (Matthews, 1992b). High event
rate tasks used in recent research are sensitive to arousal, but rarely sensitive to E (Matthews
and Davies, 1998). The prediction from Gray's theory that Anx should enhance vigilance is not
supported. Davies and Parasuraman's (1982) review concludes that eects of N are ``minimal''
and state anxiety may actually impair vigilance (Geen, 1985). An alternative means for
investigating Posner's vigilance system is through assessing the phasic alertness generated by a
warning signal in RT studies. Derryberry (1987) predicted that, according to Gray's theory,
introverts should show enhanced alerting to a negative warning signal, but failed to nd any
eect of E on alerting processes across two studies.
6.3. Studies of memory
Eects of E on memory frequently depend on retention interval. Broadly, extraverts show
superior recall up to intervals of about 5 min, after which introverts show an increasing recall
advantage as retention level increases (e.g. Howarth and Eysenck, 1968). Eysenck and Eysenck
(1985) explain extraversionintroversion eects on memory on the basis of action-decrement
theory, which claims that high arousal inhibits retrieval in the short-term so as to enhance
consolidation and long-term retention. As in the case of E arousal interactions, we have a
phenomenon for which the Eysenck but not the Gray theory has a ready explanation.
However, as M. W. Eysenck (1981; 1982) discusses, action decrement theory is unsatisfactory,
because there are a variety of instances in which arousal eects on memory are not as
predicted. As with E arousal interactions, there are also concerns over the role of time of day
(Revelle and Loftus, 1992), task factors (M. W. Eysenck, 1981) and the mediating role of
arousal (Matthews, 1992b). The exact neural basis for personality eects on memory is also
unclear.
Other personality factors and the emotional content of stimuli may also inuence memory.
Recent evidence suggests that eects of E on incidental free recall may be moderated by
stimulus valence (Bartussek, 1996), but there is a general lack of both theory and data in this
area. N and Anx tend to be associated with impairment of memory at both short and long
retention intervals, a result which is inconsistent with action-decrement theory (M. W. Eysenck,
613
1981). Contemporary explanations for such eects focus on cognitive mechanisms such as
reduced elaboration of encoding and allocation of working memory to processing worries
(M. W. Eysenck, 1992; Wells and Matthews, 1994).
6.4. Responsiveness
Both theories provide a rationale for personality inuencing response rate. According to
Eysenck (1967), extraverts should be prone to frequent response as a strategy to raise their
arousal towards the hedonic optimal, especially in low arousal environments, whereas
introverts should be reluctant to respond. Stressed neurotics should also avoid response in
order to lower arousal. According to Gray, vigour of response is one of the principal outputs
of the arousal system, whereas probability of response relates to the BAS and BIS systems.
Because both the BAS and BIS activate the arousal system, both Imp and Anx might be
expected to raise responsiveness, especially under the appropriate reinforcement conditions,
with the proviso that Anx eects may be countered by behavioural inhibition.
Extraverts tend to show faster response on short tasks, riskier speed-accuracy tradeo and
lower response criterion (Eysenck, 1967). A series of RT studies conducted by Brebner and
Cooper (1985) suggested that extraverts appear to be ``geared to respond'' and introverts seem
``geared to inspect''. However, eects of E on responsiveness do not seem to be very reliable
across dierent tasks and are not apparent in many well-designed studies (Amelang and
Ullwer, 1991; Matthews, 1992b). Koelega's (1992) meta-analysis of vigilance showed no
signicant eects of E on false positives and RT, but, in extreme-group studies, response
criterion (indexed by beta) was lower in extraverts. When eects are observed, there is
surprisingly little evidence on their relationship to individual dierences in arousal. Matthews
et al. (1990b) found that, in the morning, energetic arousal was positively related to
responsiveness (i.e. low beta) in extraverts but not introverts, with the eect tending to reverse
or disappear in the evening. As with the performance data previously discussed, the
E arousal interaction suggests that arousal moderates rather than mediates extraversion
eects. Arousal theory also ignores evidence that speed-accuracy tradeo may be inuenced by
a variety of independent processing stages. Dickman and Meyer (1988) used a stage analysis of
a visual comparison task to show that Imp did not have a general eect on speed-accuracy
tradeo, but inuenced a feature comparison stage taking place relatively early in processing.
RT studies sometimes distinguish ``decision time'', which aggregates stages up to and including
response selection and ``movement time'', which refers to response execution. Stelmack et al.
(1993a,b) found that E was reliably negatively related to movement time on various speeded
response tasks, but not to decision time. Doucet and Stelmack (1997) replicated this nding.
Newman et al. (e.g. Wallace et al., 1991) tested a modied version of the Gray theory in
studies of responsiveness. They identied the arousal system with N and the balance of the BIS
and BAS with E, so that N tends to amplify the reactivity of extraverts to reward stimuli and
the reactivity of introverts to threat stimuli. The dominant response set also moderates
personality eects. In a study of pattern matching, Nichols and Newman (1986) found an
overall tendency for extraverts to respond relatively rapidly in reward-only conditions, but for
introverts to respond more quickly in punishment-only conditions. This eect might be seen as
mediated by individual dierences in arousal dependent on the relative sensitivities of BAS and
614
BIS. Hernaiz-Sanders (1991) failed to replicate the eect. She identied two diculties with the
Nichols and Newman paradigm. First, it is dicult to distinguish eects of unconditioned
reinforcement from reinforcement signals. Second, the consequences for responsiveness of BIS
activation are hard to predict, because its outputs include behavioural inhibition as well as
arousal. Nichols and Newman also found that extraverts tended to speed up following
punishment, but introverts tended to be faster after reward, when both reward and
punishments are delivered across trials. As discussed above, Derryberry and Reed (1994)
obtained similar eects in a spatial orienting paradigm and argued that incentive processes may
aect ability to disengage from stimuli.
N is not related to low responsiveness, contrary to the Eysenck theory. Stelmack et al.
(1993a,b) found that N related to faster decision time, but was unrelated to movement time.
State anxiety may lead to a more impulsive strategy under stressful conditions (Leon and
Revelle, 1985). Wallace and Newman (1990) demonstrated `ànxious impulsivity'' using a circle
tracing task requiring continuous response. Response was faster in neurotic introverts (high
Anx subjects) than in stable extraverts (low Anx subjects) when goals are uncertain, a
condition which may activate the arousal rather than the inhibition response of the BIS. With
a denite behavioural goal, neurotic extraverts (high Imp subjects) traced faster than stable
introverts (low Imp subjects). Behavioural goals may activate the BAS. Given Newman's
assumptions about the task, these data seem quite consistent with Gray's theory, although little
attempt was made to consider alternative explanations. The main diculty is that response
criterion and response latency are not direct indices of some basic reactivity, but depend on
strategic decisions based on a statistical model of the task performed (Matthews, 1996). Within
signal detection theory, response vigour (RT) and response probability (beta) are closely
related (Davies and Parasuraman, 1982). It is hard to interpret responsiveness data without an
information-processing analysis of the task.
6.5. Attention and performance: conclusions
In our view, both theories are having diculties in accommodating the diverse empirical
ndings emerging from dierent paradigms. The Eysenck theory succeeds fairly well in
explaining some of the general trends in the extraversion data, such as arousal-related
impairments in introverts and the responsiveness of extraverts. On some tasks, such as sensory
threshold tasks and paired associate learning, the correspondence between data and prediction
is impressive (Eysenck, 1997). However, studies of attention and memory tasks exposes the
limitations of arousal theory, such as the failure of any study of performance to demonstrate
directly that eects of E are in fact arousal-mediated (Matthews, 1992b). There are also large
areas of performance research in which the theory either makes no predictions or even basic
predictions from the theory are not supported. Arousal theory, in Eysenck's formulation, does
not explain personality eects on performance in the afternoon and evening (Revelle et al.,
1980), on spatial orienting (Derryberry and Reed, 1997), on demanding target detection tasks
(Matthews et al., 1990b) and on a variety of memory tasks (M. W. Eysenck, 1992). Eects of
N in moderating eects of E on performance are often broadly compatible with arousal theory
(see Eysenck and Eysenck, 1985), but, for the most part, contemporary work on Anx and N
makes little reference to arousal and focuses on cognitive and attentional mechanisms.
615
Tests of Gray's theory have been somewhat stymied by diculties in deriving testable
predictions on typical performance tasks. The theory receives some support from motivational
moderation of eects of E. However, the most reliable eect of this kind, associated with
individual dierences in reactions to feedback on a previous trial, relates to motivational
outcomes rather than cues (Derryberry and Reed, 1994) and so relevance of the theory is
uncertain. Some other ndings are broadly compatible with the theory, such as `ànxious
impulsivity'' (Wallace and Newman, 1990). Speculatively, extraverts' superiority in multichannel performance and greater responsiveness might be attributed to the BAS and
dopaminergic activity, although apparently impulsive response style may actually relate to
individual dierences in stimulus analysis (Dickman and Meyer, 1988). Gray (e.g. Gray, 1982)
deserves credit for predicting the bias in selective attention characteristic of anxiety (Wells and
Matthews, 1994) prior to its demonstration in experimental studies. However, there are
considerable diculties in mapping BIS functions onto human selective attention functions
(Wells and Matthews, 1994, pp. 327332) and, embarrassingly for Gray's theory, cognitive
therapy abolishes the bias in selective attention in anxiety patients, but anxiolytic drugs do not
(Golombok et al., 1991). In addition, attentional bias is found in panic disorder (McNally et
al., 1990), a condition Gray (1987a) attributes to the ght/ight system rather than the BIS.
At best, neither theory seems likely to account for more than a subset of observed
personality eects on performance. The areas of research to which the theories fail to
contribute are suciently extensive that predictive failures cannot simply be attributed to
inappropriate choice of task parameters. Furthermore, there are acceptable cognitive
mechanisms for explaining many of the ndings problematic for the psychobiological theories
(e.g. M. W. Eysenck, 1992; Matthews, 1997). If either theory is to have a future role in
performance research, it must either specify improved methods for identifying behavioural
indices controlled by the brain systems of interest or tackle the dicult problem of how
outputs from those systems may interact with information-processing mechanisms.
7. Conclusions
The evidence reviewed raises several interlocking issues. First, the theories disagree on the
key traits and moderating factors. Eysenck emphasises E, N and level of stimulation, whereas
Gray focuses on Imp, Anx and motivational signals. We will consider whether the data suggest
one set of variables is more predictive of response than the other set. Second, evidence relating
to direct tests of theory will be assessed. Finally, the wider implications for the
psychobiological approach to personality theory will be discussed.
7.1. Indirect tests of theory: choice of constructs and moderating factors
Consistent with Gray's (1981) position, there are some areas of research where Imp does
seem to be somewhat more predictive than either Soc or E, including the spontaneous EEG
(Stenberg, 1992) and interactive eects of personality and caeine on verbal ability (Revelle et
al., 1980). However, these Imp-driven eects seem very specic: for example, the EEG result
fails to generalise to brainstem EPs (Swickert, 1996). Often, dierences between Imp and Soc
616
are either absent or inconsistent across studies (Gupta, 1990; Amelang and Ullwer, 1991; Corr
et al., 1995a). Hence, although Imp may be more predictive in certain paradigms, overall there
is little compelling reason to favour the Gray axes over the Eysenck ones. More general
psychometric evidence favours the Eysenck and Eysenck (1985) structural model, of course.
The other choice between constructs concerns the factors which moderate relationships
between traits and response. Probably the single most secure conclusion from biologicallybased trait research is that level of stimulation or arousal moderates associations between E
and response (Eysenck and Eysenck, 1985). The question raised by Gray's theory is whether
there are moderating eects of motivational signals over and above these arousal eects.
Clearly, motivational manipulations moderate personality eects, but in some cases these
manipulations may operate through their stimulating eects (Eysenck and Eysenck, 1985;
Bartussek et al., 1996). Other paradigms, notably mood induction and instrumental
conditioning, show dierential eects of reward and punishment manipulations broadly
compatible with Gray's theory. However, it is unclear that reward and punishment cues (CSs)
have a stronger moderating eect than actual rewards and punishments (USs), as the theory
implies (cf. Derryberry and Reed, 1994) and it may be dicult to discriminate US from CS in
human subjects. Motivational eects may be confounded with arousal, but, as we suggested in
the section on conditioning, this argument seems to be used in a rather post hoc way.
Personality may moderate the arousing eects of motivational stimuli, a possibility which has
received insucient attention. The data suggest that motivational and arousal-related factors
may both be important as moderator variables, but no rm conclusions may be drawn.
7.2. Direct tests of theory

Predictions from the Eysenck theory of extraversion are supported in several research areas.
Overall, the psychophysiological data are suggestive of lower arousability of extraverts in lowstimulation environments and greater arousability of extraverts in high-stimulation
environments, especially in ERP and EDA paradigms. E is a weak predictor of tonic arousal
measures. These eects are consistent with the theory if we allow that TMI develops in
introverts under high stimulation. In behavioural paradigms, similar eects have been
demonstrated across multiple studies in eyeblink conditioning (Levey and Martin, 1981) and in
sensory threshold studies (Shigehisa and Symons, 1973). Together, these results provide a solid
core of empirical support for the Eysenck theory of extraversion.
In other research areas, the Eysenck theory has been less successful. Sometimes E simply
fails to aect response measures at all, though such results might reect insucient statistical
power or other methodological problems. More seriously, there are many studies showing
interactions between E and other factors which cannot readily be explained by the theory,
although tentative, post hoc explanations are advanced in some cases. Such ndings suggest
incompleteness of the theory. Such problems are especially apparent in the performance
domain, within which eects of E do not seem directly contingent upon individual dierences
in arousal (Matthews, 1992b). Performance correlates of E are best conceptualised as a
``cognitive patterning'', associated with a variety of independent information-processing
mechanisms (Matthews, 1997).
617
The Eysenck theory has been less successful in providing an account of N. It predicts the
characteristic negative aect of neurotics and their greater sensitivity to negative mood
inductions. It also predicts superior associative conditioning in anxious individuals. However,
there is a striking mismatch between the robust subjective data on N and negative mood and
the inconsistency of associations between N and autonomic arousal, even when stress is
explicitly manipulated. As in the case of E, performance studies provide supercial support for
arousal theory, in that neurotics (or high Anx subjects) are especially disadvantaged on more
dicult tasks). Again, the arousal theory explanation breaks down when scrutinised in detail,
with most contemporary theories of anxiety making worry rather than arousal the central
explanatory construct (M. W. Eysenck, 1992; Wells and Matthews, 1994).
Overall, the Eysenck theory successfully explains an important subset of the evidence, but
not all of it. The operation of TMI is critical for tting outcomes to prediction, but we have
noted the problems in calibrating the levels of stimulation necessary for TMI across studies,
which prevent ante hoc prediction of TMI. The same problem emerges in comparing results
across dierent areas of research. For example, psychophysiologists assume that performance
of demanding tasks (Fowles et al., 1977) or moderate dosages of caeine induces TMI in
introverts (Smith, 1983). But in performance research, similar dosages are seen as arousing in
both extraverts and introverts (Revelle et al., 1980). Basic assumptions about arousal levels
vary radically across studies and so it is dicult to decide on the paradigms where arousal
theory might or might not be useful.
There are two questions for the Gray theory, given the partial success of the Eysenck theory.
First, can it provide an alternative explanation for the arousal-dependent eects of E which the
Eysenck theory accommodates? Second, are ndings inconsistent with the Eysenck theory
predicted by Gray's theory? Gray's theory might explain the arousal ndings as follows.
Introversion is correlated with sensitivity of the BIS, one of whose outputs is arousal. Hence,
given that most laboratory environments are likely to provide at least a low level of
punishment or novelty cues, we expect that BIS activation will feed into higher arousal in
introverts. This hypothesis has two diculties. Gray's arousal system is supported by the
DNAB, but there is little evidence that this system shows any equivalent of TMI, i.e. inhibition
under high levels of stimulation. Thus, the hypothesis fails to explain the role of stimulation
level in moderating correlations between E and arousal. Second, the 308 rotation model states
that the BIS is better indexed by N than by E. Hence, if introverts are typically somewhat
more aroused, due to activation of the BIS, then neurotic individuals should be markedly more
aroused than stable individuals, which is simply not the case. The classical extraversion/arousal
data t Eysenck's theory better than Gray's, although it might be argued that these eects are
outside the scope of the Gray theory.
The second issue for Gray's theory is its capacity to explain additional ndings. Gray
himself emphasises studies of conditioning to motivational stimuli as a source of evidence, but
the evidence is mixed. The single strongest piece of evidence in favour of Gray's theory derives
from the Gupta studies showing that extraverts condition better than introverts to verbal
reward. The post hoc explanation oered by arousal theory, that verbal reward induces TMI in
introverts seems a little contrived in this instance. However, the eect is very paradigm-specic:
personality eects on discrimination learning of positive cues do not provide strong support for
Gray (e.g. Zinbarg and Revelle, 1989). Other ndings which may support Gray's theory are the
618
associations between E and subjective energy and ``behavioural activation'' performance

measures such as faster movement time, although in these cases the link between the criterion
and the BAS is somewhat speculative.
In other studies of motivational moderators, support for the Gray theory is patchy, at best.
Psychophysiology provides occasional support (e.g. De Pascalis et al., 1996), but it is telling
that the most comprehensive series of studies testing the theory directly nds little consistent
support for it (Bartussek et al., 1996). In performance paradigms, Gray's theory, like
Eysenck's, has diculties in accommodating variation of personality eects with informationprocessing demands of the task. Motivational variables sometimes have moderating eects
broadly consistent with the theory, although there are concerns about replicability (HernaizSanders, 1991; Pickering et al., 1997) and discrimination of motivational USs and CSs (cf.
Derryberry and Reed, 1994). Gray's theory of Anx has similar problems to Eysenck's theory of
N, including lack of support from psychophysiology (Fahrenberg, 1987), lack of clear-cut
eects of Anx in instrumental learning studies and diculties in accounting for the various
information-processing factors which moderate Anx eects in performance studies (M. W.
Eysenck, 1992). The problems of anxiety studies are disappointing in view of the impressive
evidence for the BIS as the basis for anxiolytic drug eects in animals (Gray, 1982;
McNaughton, 1997). It is tempting to conclude that human anxiety is simply much more
susceptible to cognitive control than to the BIS: self-regulative processes are central to human
anxiety (Wells and Matthews, 1994), but may lack close analogues in the rat.
The Gray theory receives its strongest support from the role of motivational signals as an
important moderator of personality eects and from the broad association between E/Imp and
``behavioural activation''. Overall, however, our impression is that the theory has had less
predictive success than the Eysenck theory, even within those domains to which it most clearly
applies. Pickering et al. (1997, p. 63) admit that dierent post hoc explanations are needed to t
the theory to results from dierent studies. It is premature to dismiss the theory at this stage
and some specic paradigms provide qualied support for it. Recent commentary on the
theory (Pickering, 1997; Pickering et al., 1997) demonstrates conceptual diculties in linking
the BIS and BAS to behavioural measures, in predicting their interaction and in characterizing
the role of arousal. There seem to be dierences in opinion between Gray's coworkers
on development of the model: Pickering (1997) makes arousal the proximal inuence on
behaviour, whereas McNaughton's (1997) account of anxiety is based on hippocampal
computation of threat. The domain of application of Gray's theory is also uncertain: there
seems to have been a retreat from the wide-ranging account given by Gray (1981) to a search
for a small number of specic paradigms which might support the theory (Pickering et al.,
1997). Resolution of these diculties would facilitate theory testing.
Tentatively, the evidence reviewed suggests E may have two clusters of psychophysiological
and behavioural correlates. Table 2 distinguishes (1) a ``corticoreticular'' E, related to classical
arousal indices, conditioning and sensory thresholds and (2) a ``dopaminergic E'' associated
with motor responsivity, subjective energy, multiple-task performance (controlled by Posner's
event detection system) and reward sensitivity. The dopaminergic basis for modulation of
motor activity, positive emotion and reward-incentive behaviour in particular seems quite wellestablished (Depue, 1995). In addition, molecular genetic data show relationships between
extraversion and a polymorphism for a dopamine receptor gene (Ebstein et al., 1996). In a tidy
619
Table 2
Two types of correlate of extraversion
``Corticoreticular'' extraversion
``Dopaminergic'' extraversion
Low cortical arousability

Low autonomic arousability
Insensitivity to TMI
Poor eyelid conditioning
High sensory threshold
decreased motoneuronal excitability

conditioning to reward
faster movement time
multiple channel detection
subjective energy
world, these two sets of indices would correlate with distinct primary factors, but the data
reviewed provide little basis for doing so1. Contrary to prediction from Gray's theory, Imp
relates at least as strongly to factors associated with corticoreticular E as it does to
dopaminergic E. Specically, there are studies showing associations between Imp and lower
EEG arousal (O'Gorman and Lloyd, 1987; Stenberg, 1992), reduced phasic EDA in placebo
conditions (Smith et al., 1981) and eyelid conditioning (Eysenck and Levey, 1972). Smith et
al.'s EDA study (Smith et al., 1981) reported impulsivity caeine interactions suggestive of a
higher threshold for TMI in high impulsives.
7.3. Personality theory and the biological approach

Not so long ago, trait psychologists and social learning theorists such as Mischel (1968)
belonged to two opposing, sometimes antagonistic camps, as exemplied by the debates of the
1960s and 1970s. Trait psychology came with a sharply-dened package of assumptions,
including the heritability of traits and a causal model assuming feed-forward of individual
dierences in brain function into behaviour and learning. The resurgence of trait psychology
following on from Eysenck and Eysenck's (1980) decisive refutation of situationism has
provided considerable evidence in support of these assumptions (Matthews and Deary, 1998).
The behaviour and molecular genetic evidence conrm that biology plays some role in
personality traits. However, the ``Berlin Wall'' between trait approaches and social learning
approaches is crumbling. Many social learning theorists recognise that traits have at least some
validity and are exploring social cognitive explanations for their stability and inuence on
behaviour (e.g. Snyder, 1992). Conversely, trait psychologists are increasingly developing
information-processing models within which cognitive factors are more proximal causal
inuences on individual dierences than are biological factors (e.g. Revelle, 1993).
1
A reviewer of this paper suggested that, on the basis of the supposed dopaminergic basis for psychoticism
(Eysenck, 1997), ``dopaminergic E'' could equally well be regarded as ``dopaminergic P''. However, P does not seem
to be related to the criteria we have used to dene dopaminergic E, such as positive emotion (Matthews et al., in
press), motoneural excitability (Pivik et al., 1988) and movement time (Doucet and Stelmack, 1997). Corr et al.
(1995a) failed to show any signicant correlations between P and conditioning to reward stimuli. Conversely, P
relates to reticulocortical indices such as EEG measures (Matthews and Amelang, 1993) and eyelid conditioning
(Frcka and Martin, 1987).
620
In this new era of research, the central issues concern not the basic validity of traits, but the
nature of the explanatory constructs to be used in linking traits to individual dierences in
behaviour. Both the Eysenck and the Gray theories make a clear statement that neural
processes are the preferred explanatory construct and the Eysenck theory in particular has had
notable successes in explaining individual dierences in responses closely tied to brain function.
However, the diculties which both theories have experienced in explaining the human
performance data challenge the centrality of neural explanations. Furthermore, diculty in
specifying moderating factors leads to considerable predictive ambiguity. We can predict that
either extraverts or introverts are more aroused, depending on assumptions made about TMI.
Similarly, Gray's theory can predict that Anx is associated with either more or less motor
responsivity, depending on whether behavioural inhibition or arousal eects predominate. The
scope for post hoc rationalisation of results is obvious.
Thus, trait psychology has two central challenges to meet. First, should we retain individual
dierences in the functioning of broad neural systems as the principal explanatory construct?
Cognitive and social-cognitive models may provide viable alternatives, with more precise
specication of moderating factors. Second, both Eysenck and Gray assume that each trait
corresponds to a single, key underlying system, but there may not be any simple one-to-one
mappings between brain functions and traits (Zuckerman, 1991). The twin challenges posed by
cognitive theories and by multiple-systems models of traits may be met in various ways. The
best hope for research in the EysenckGray tradition is methodological improvements in
assessment of brain function. Perhaps current psychophysiological techniques simply do not
pick up the individual dierences in neural processes which drive behaviour. A second
approach, retaining the centrality of neural explanations, is to develop more complex
physiological models (e.g. Zuckerman, 1991), at the possible cost of loss of theoretical
coherence and testability. Within such an approach, both the Eysenck and Gray theories may
have partial validity. Perhaps E is independently related to corticoreticular arousability and to
a reward system. A third alternative (Revelle, 1993), is to extend the causal chain linking traits
to behaviour by introducing cognitive variables as additional mediating constructs, so that
neural processes become more distal. The nal possibility is to argue that the associations
between traits and broad neural systems are simply not strong enough to explain the
behavioural correlates of traits. Matthews (Matthews and Dorn, 1995; Matthews, 1997) has
argued that traits are associated with packages of somewhat independent neural and cognitive
functions which together support adaptations to specialised social and physical environments.
Cognitive constructs may be more appropriate than biological ones for explaining the majority
of behaviours, so that explanations of the kind oered by the Eysenck and Gray theories are
relevant to a restricted range of phenomena only. It remains to be seen whether either theory
provides a comprehensive explanation for the correlates of traits, but, in any case, personality
psychology owes an enormous debt to both scientists.
Acknowledgements
This article was written at the invitation of Hans Eysenck. We were greatly saddened by his
death while the article was in preparation. We thank Jerey Gray, Alan Pickering and Robert
621
Stelmack for helpful comments on a previous draft of this paper. We are also grateful to Neil
McNaughton for insights into animal studies of anxiety and to Ana Adan for making available
data on personality and mood.
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PERGAMON

Personality and Individual Differences 26 (1999) 583626

The personality theories of H. J. Eysenck and J. A. Gray:

Department of Psychology, University of Dundee, Dundee DD1 4HN, Scotland, U.K.

* To whom all correspondence should be addressed. E-mail: g.matthews@dundee.ac.uk

G. Matthews, K. Gilliland / Personality and Individual Dierences 26 (1999) 583626

G. Matthews, K. Gilliland / Personality and Individual Dierences 26 (1999) 583626

G. Matthews, K. Gilliland / Personality and Individual Dierences 26 (1999) 583626

G. Matthews, K. Gilliland / Personality and Individual Dierences 26 (1999) 583626

G. Matthews, K. Gilliland / Personality and Individual Dierences 26 (1999) 583626

2. Psychophysiology of the central nervous system

G. Matthews, K. Gilliland / Personality and Individual Dierences 26 (1999) 583626

G. Matthews, K. Gilliland / Personality and Individual Dierences 26 (1999) 583626

G. Matthews, K. Gilliland / Personality and Individual Dierences 26 (1999) 583626

G. Matthews, K. Gilliland / Personality and Individual Dierences 26 (1999) 583626

G. Matthews, K. Gilliland / Personality and Individual Dierences 26 (1999) 583626

G. Matthews, K. Gilliland / Personality and Individual Dierences 26 (1999) 583626

3. Psychophysiology of the peripheral nervous system

G. Matthews, K. Gilliland / Personality and Individual Dierences 26 (1999) 583626

G. Matthews, K. Gilliland / Personality and Individual Dierences 26 (1999) 583626

G. Matthews, K. Gilliland / Personality and Individual Dierences 26 (1999) 583626

G. Matthews, K. Gilliland / Personality and Individual Dierences 26 (1999) 583626

4. Mood and subjective states

G. Matthews, K. Gilliland / Personality and Individual Dierences 26 (1999) 583626

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4.2. Correlational studies

Costa and McCrae (1980)

Emmons and Diener (1986)

Meyer and Shack (1989)

Watson and Clark (1992)

Larsen and Ketelaar (1991)1

Adan and Guardia (1997)2

Adan and Guardia (1997)3

Matthews et al. (1990a)

Matthews et al. (in press)

G. Matthews, K. Gilliland / Personality and Individual Dierences 26 (1999) 583626

considerably weaker personalitymood associations. The data suggest that N is modestly

G. Matthews, K. Gilliland / Personality and Individual Dierences 26 (1999) 583626

4.4. Mood: conclusions

G. Matthews, K. Gilliland / Personality and Individual Dierences 26 (1999) 583626

task parameters. In particular, in associative learning, TMI may disrupt conditioning if

G. Matthews, K. Gilliland / Personality and Individual Dierences 26 (1999) 583626

than to conditioning per se (McNaughton, personal communication, 17/4/98). The problem is

G. Matthews, K. Gilliland / Personality and Individual Dierences 26 (1999) 583626

G. Matthews, K. Gilliland / Personality and Individual Dierences 26 (1999) 583626

G. Matthews, K. Gilliland / Personality and Individual Dierences 26 (1999) 583626

G. Matthews, K. Gilliland / Personality and Individual Dierences 26 (1999) 583626

6. Attention and performance

G. Matthews, K. Gilliland / Personality and Individual Dierences 26 (1999) 583626

G. Matthews, K. Gilliland / Personality and Individual Dierences 26 (1999) 583626

G. Matthews, K. Gilliland / Personality and Individual Dierences 26 (1999) 583626

G. Matthews, K. Gilliland / Personality and Individual Dierences 26 (1999) 583626

G. Matthews, K. Gilliland / Personality and Individual Dierences 26 (1999) 583626

G. Matthews, K. Gilliland / Personality and Individual Dierences 26 (1999) 583626

G. Matthews, K. Gilliland / Personality and Individual Dierences 26 (1999) 583626

G. Matthews, K. Gilliland / Personality and Individual Dierences 26 (1999) 583626

7.2. Direct tests of theory

G. Matthews, K. Gilliland / Personality and Individual Dierences 26 (1999) 583626