COGNITION AND EMOTION

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On the interdependence of cognition and emotion

Justin Storbeck & Gerald L. Clore

To cite this article: Justin Storbeck & Gerald L. Clore (2007) On the interdependence of cognition
and emotion, COGNITION AND EMOTION, 21:6, 1212-1237, DOI: 10.1080/02699930701438020

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Published online: 28 Aug 2007.

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COGNITION AND EMOTION
2007, 21 (6), 1212
1237

On the interdependence of cognition and emotion

Justin Storbeck and Gerald L. Clore
University of Virginia, Charlottesville, VA, USA

Affect and cognition have long been treated as independent entities, but in the
current review we suggest that affect and cognition are in fact highly interdepen-
dent. We open the article by discussing three classic views for the independence of
affect. These are (i) the affective independence hypothesis, that emotion is processed
independently from cognition, (ii) the affective primacy hypothesis, that evaluative
processing precedes semantic processing, and (iii) the affective automaticity
hypothesis, that affectively potent stimuli commandeer attention and evaluation is
automatic. We argue that affect is not independent from cognition, that affect is not
primary to cognition, nor is affect automatically elicited. The second half of the
paper discusses several instances of how affect influences cognition. We review
experiments showing affective involvement in perception, semantic activation, and
attitude activation. We conclude that one function of affect is to regulate cognitive
processing.

Different views of the relationship between cognition and emotion can be

seen in the comments of two prominent psychologists upon receipt of
distinguished scientist awards.
Robert Zajonc (1980, p. 151) in a paper titled ‘‘Preferences Need No
Inferences’’, proclaimed: ‘‘Affect and cognition . . . constitute independent
sources of effects in information processing’’. A year later, upon receipt of
the same award, Gordon Bower (1981, p. 147), in a paper on ‘‘Mood and
Memory’’, stated, ‘‘I am a cognitive psychologist, and . . . the emotional
effects we have found so far seem understandable to me in terms of ideas that
are standard fare in cognitive psychology’’. These two pivotal papers both
argued for the importance of studying emotion, but they proposed different
meta-theories. Zajonc argued that affect and cognition are processed
independently and that affect has temporal priority over even basic cognitive

Correspondence should be addressed to: Justin Storbeck, 102 Gilmer Hall, PO Box 400400,
Charlottesville, VA 22904
4400, USA. E-mail: storbeck@[Link]
Support for this research is acknowledged from National Institute of Mental Health Grant
MH 50074 to GLC.
# 2007 Psychology Press, an imprint of the Taylor & Francis Group, an Informa business
[Link]/cogemotion DOI: 10.1080/02699930701438020
 INTERDEPENDENCE OF COGNITION AND EMOTION 1213

processes. In contrast, Bower argued that cognitive processing could be used

to understand emotional phenomena. We agree with Bower’s conception,
and we extend it to suggest that cognitive processes are necessary for the
processing, elicitation, and experience of emotions.
The concepts of ‘‘cognition’’ and ‘‘emotion’’ are, after all, simply
abstractions for two aspects of one brain in the service of action. Zajonc
believed that emotion is independent from cognition. Our own view is that
the study of emotion and cognition should be integrated, because the
phenomena themselves are integrated (Dewey, 1894; Parrott & Sabini, 1989).
We argue against the notion that discrete emotions have separate and
distinct areas in the brain (Duncan & Barrett, 2007 this issue). Rather,
emotions emerge from a combination of affective and cognitive processes
(see Moors, this issue).1 Moreover, in agreement with Bower (1981), we
suggest that emotion can be studied using cognitive paradigms. Both
laboratory findings and everyday observation suggest a unity and inter-
relatedness of cognitive and affective processes, and that trying to dissect
them into separate faculties would neglect the richness of mental life
(Roediger, Gallo, & Geraci, 2002, p. 319). We suggest, like others, that the
interconnections found within the brain provide no obvious basis for
divorcing emotion from cognition (Erickson & Schulkin, 2003; Halgren,
1992; Lane & Nadel, 2000; Phelps, 2004).
This article has two parts. The first part is a critical review of what
recent neuroscience and social psychological research tells us about three
popular ideas about cognition and emotion.2 These include the affective
independence hypothesis, that emotion is processed independently of
cognition via a subcortical ‘‘low route’’, the affective primacy hypothesis,
that affective and evaluative processing takes precedence over semantic
processing, as evident in the mere exposure effect and affective priming,
and the affective automaticity hypothesis, that affectively potent stimuli
commandeer attention and that affective processes are especially likely
to be automatic. In the second part, we suggest that rather than
being processed independently, affect modifies and regulates cognitive
processing, as illustrated in a review of some recent research from our
own lab.

1
That is, we, along with others (e.g., Barrett, 2006), suggest that there is not a brain centre
dedicated to specific emotions such as fear, happiness, etc. But, there are specific areas critically
involved in emotion processing. For instance, the amygdala is critically involved in the emotion
of fear, but is not specifically dedicated to fear.
2
The conception of emotion we raise, affective independence and affective primacy, comes
mainly from Zajonc (1980, 2000). The affective automaticity derives from arguments made by
Bargh and colleagues (Bargh, 1997; Ferguson & Bargh, 2003).
1214 STORBECK AND CLORE

I. ASSESSING THREE HYPOTHESES ABOUT AFFECT

AND COGNITION
A. The affective independence hypothesis: The low route
Zajonc (2000) lists ten ways in which affect and cognition differ and suggests
that they arise from separate systems. Papers in this tradition sometimes cite
LeDoux’s (1996) proposal that the amygdala can elicit emotion before
information reaches the cortex. In this section, we suggest (a) that the low
route does not play a role in processing the complex stimuli typically used in
social and emotional research (e.g., faces, ideographs, objects), and (b) that
the amygdala, and emotion in general, does not function independently of
perceptual and cognitive processes.
The ‘‘low route’’ (LeDoux, 1996; LeDoux, Romanski, & Xagoraris, 1989)
is a pathway that allows stimulus processing without cortical influence as
studied in rats. When light hits the retina, the signal is relayed to the
amygdala through the thalamus without going first to the visual cortex. The
pathway is adequate to support fear conditioning between illumination
changes and fear-invoking events (e.g., shock). Based on studies with the
auditory cortex, which have similar relay pathways as the visual cortex,
information can reach the amygdala within 20 ms (Quirk, Armony, &
LeDoux, 1997; Quirk, Repa, & LeDoux, 1995).
LeDoux and colleagues used rat models to examine the low route to
emotion, and the question remains whether a similar pathway exists in
humans. First, comparative anatomical studies (Linke, De Lima, Schwegler,
& Pape, 1999) and behavioural studies (Shi & Davis, 2001) using rat-based
models suggest that this pathway may be functionally relevant only
when cortical areas have been lesioned or damaged. In humans, converging
evidence suggests that the low route may not be functionally important for
emotion processing (Halgren, 1992; Kudo, Glendenning, Frost, &
Masterson, 1986; Rolls, 1999; Shi & Davis, 2001; Storbeck, Robinson, &
McCourt, 2006). However, the existence of the low route in humans is still
debated (see LeDoux, 2001).3

3
In particular, the strongest evidence for such a route comes from affective blindsight
individuals. Individuals have damage to area V1 of the visual cortex and as a result have no
conscious perception of the world. However, these individuals still demonstrate affective
reactions to fear-inducing visual stimuli. In the literature though, this is still a debated issue.
First, the pathways involved are unclear. That is, although information may not be visually
conscious to blindsight individuals, areas of the visual cortex still receive visual information
(area V4 and extrastriate) from subcortical structures such as the pulvinar and superior
collicolus. Therefore, although the area V1 is damaged, areas of the visual cortex still receive the
same visual information. Storbeck, Robinson, and McCourtt (2006) examine this issue more
extensively.
 INTERDEPENDENCE OF COGNITION AND EMOTION 1215

Can the low route discriminate emotional vs. non-emotional stimuli

without cortical involvement? That is, can the low route sufficiently dis-
criminate a snake from a bunny without cortical involvement? We say No.
The low route has limited capacity for stimulus discrimination. Fear
conditioning studies that find support for the low route typically require
only detection of the presence or absence of a stimulus (Duvel, Smith, Talk,
& Gabriel, 2001; LeDoux et al., 1989; Shi & Davis, 2001). When the task
requires discriminating one stimulus from another (e.g., CShigh freq.
tone, CSlow freq. tone), then cortical analysis appears to be necessary
(Butler, Diamond, & Neff, 1957; Duvel et al., 2001; Komura et al., 2001;
McCabe, McEchron, Green, & Schneiderman, 1993; Nicholson & Freeman,
2000; Thompson, 1962).
One way to test the affective independence hypothesis is to temporarily
inactivate the visual cortex to determine whether it is necessary for the
amygdala to determine affective significance of stimuli. Fukuda, Ono, and
Nakamura (1987) did just that in awake, behaving monkeys. They observed
that when the visual cortex (representing early cognitive processing) was
temporarily inactivated, monkeys failed to learn and failed to demonstrate
appropriate affective associations to visual cues of edible vs. inedible objects.
But, the amygdala was still intact, because the monkeys could determine the
affective significance of the same stimuli based on taste. This study suggests
that in order to determine whether a peanut or a baseball is edible based on
visual properties and prior experience, the visual cortex is necessary.
Another way to determine whether the amygdala is independent of initial
cognitive processing is to record single-cell activity within the visual cortex
and the amygdala. Again, if the amygdala can process visual stimuli
independent of cortical input, then amygdala neurons should still remain
active to visually presented objects that have affective significance when the
visual cortex is temporarily inactivated. Nishijo, Ono, Tamura, and
Nakamura (1993) recorded such vision-relevant neurons in the amygdala
of monkeys. They discovered that these neurons failed to respond to
affectively significant visual stimuli when the visual cortex was temporarily
inactivated. They suggested that amygdala activity is not related directly to
sensory inputs, but rather it relies on view-invariant representations of
objects from the visual cortex. That is, the goal of the visual cortex is to
create unique neural signatures for unique objects regardless of its
orientation, background lighting, etc. Therefore, the same affective sig-
nificance can be retrieved regardless of the visual state the object is in.
Halgren offers similar conclusions as Nishijo et al. by suggesting that when
‘‘the amygdala performs emotional evaluation, it does so within the
cognitive system. This could explain why it has been so difficult to dissociate
emotional from cognitive processing in humans’’ (Halgren, 1992, p. 212).
1216 STORBECK AND CLORE

Faces have always received special attention in the study of emotion due
to their possible evolutionary connection to survival (Davey, 1995; Öhman,
1997). Adolphs (2002) suggested that processing and recognising a fear face
requires a network of various structures and that the low route alone is
incapable of such processing. He proposed that the visual cortex first grossly
identifies the face, and in particular determines whether the face contains an
expression or not. Support for this model comes from the fact that there are
two areas in the monkey visual cortex dedicated to face processing, areas
STS and IT. Area STS (superior temporal sulcus) is involved in encoding
facial expressions, while the IT (inferotemporal cortex) is involved in
encoding facial identity (see Allison, Puce, & McCarthy, 2000; Kanwisher,
McDermott, & Chun, 1997; Narumoto, Okada, Sadato, Fukui, & Yone-
kura, 2001; Rotshtein, Malach, Hadar, Graif, & Hendler, 2001, for related
literature on humans). Both areas, STS and IT, have strong reciprocal
connections to the amygdala, suggesting the amygdala receives highly
processed facial information pertaining to both facial identity and facial
expression (Baylis, Rolls, & Leonard, 1987; Fukuda et al., 1987; Nishijo,
Ono, & Nishino, 1988b; Rolls, 1992). For example, Rotshtein et al. (2001)
found that lateral occipital cortex in humans, which processes facial
expressions, is concerned with the configuration for each expression, rather
than with its affective value. That is, the visual cortex does not code for
affective significance (Rolls, 1999; Rotshtein et al., 2001), but rather codes
for facial configurations and these configurations are sent to the amygdala
for affective processing. Thus, the visual cortex is needed for the amygdala to
correctly identify and respond to emotional stimuli.
For emotional stimuli used by psychologists (e.g., snakes; emotional
faces) the processing capacities of the low route would appear to be
inadequate (see Rolls, 1999; Smith, Cacioppo, Larsen, & Chartrand, 2003,
for similar concerns). But, the low route is still cited to help explain
particular affective phenomena (e.g., Bargh, 1997; Berkowitz & Harmon-
Jones, 2004; Zajonc, 2000). However, we suggest that the fact that the
amygdala relies on cortical input to make an evaluation requires reconsi-
deration of whether emotion at initial levels of processing can be dissociated
from cognitive processing. We should also note that the same processes
occur regardless of whether stimuli are presented subliminally or supralim-
inally (Rolls & Tovee, 1994; Rolls, Tovee, Purcell, Stewart, & Azzopardi,
2004; Storbeck et al., 2006).

Conclusions. Davidson (2003) claimed that one of seven deadly sins of

cognitive neuroscience is to assume that affect is independent from
cognition. We and several others agree that emotion should not be divorced
from cognition (Adolphs & Damasio, 2001; Barnard, Duke, Byrne, &
 INTERDEPENDENCE OF COGNITION AND EMOTION 1217

Davidson, 2007 this issue; Davidson, 2003; Duncan & Barrett, 2007 this
issue; Eder & Klauer, this issue; Erickson & Schulkin, 2003; Lane & Nadel,
2000; Lavender & Hommel, this issue; Lazarus, 1995; Parrott & Sabini,
1989; Phelps, 2004; Storbeck et al., 2006). For instance, based on anatomical
connections alone (Ghashghaei & Barbas, 2002) areas necessary for
cognition and emotion are highly interconnected, and these connections
are bidirectional, suggesting integrated processing of emotion and cognition.
Halgren (1992) suggests that emotion and cognition are so interconnected
that it is not practical to try to disentangle the temporal and casual relations
of emotion and cognition.

B. The affective primacy hypothesis

In addition to the idea that affect and cognition are independent sources of
influence, is the allied idea of affective primacy (Zajonc, 1980, 2000). The
mere exposure phenomenon (Zajonc, 1968) was an important source of
evidence for the hypothesis, because mere exposure involves an affective
reaction that is not dependent on conscious categorisation or identification
of the liked stimulus.
One problem with accepting the grand conclusions, that affective process-
ing occurs without cognitive processing, drawn from mere exposure research
is that they rest on an equation of consciousness with cognition (Lazarus,
1995). However, single-cell recording shows that the visual cortex can readily
identify stimuli presented below subjective thresholds. Therefore, lack of
conscious awareness has no bearing on whether the visual cortex categorises
and identifies a stimulus. In fact, studies have demonstrated that the only
difference between a stimulus presented for 30 ms as opposed to 1000 ms is
the strength at which a neural population fires (Rolls, 1999; Rolls & Tovee,
1994; Rolls et al., 1994). But the response pattern output remains the same,
which is thought to reflect a unique neural signature for a given stimulus
type. That is, the neural population identifies X whether presented for 30 ms
or 1000 ms, but the firing is stronger so that the system is more confident for
1000 ms presentations than for 30 ms presentations.
Another problem is that the mere exposure effect gets weaker the longer
stimuli are presented, as respondents have time to process the identity of the
stimulus and to realise that they have seen it before. The mere exposure effect
is, therefore, a mistake based on a misattribution of fluency as liking instead
of as familiarity (Winkielman, Schwarz, Fazendeiro, & Reber, 2003). Of
course, most real-world emotional reactions are not errors of this kind, so
that its appeal as a model of how affect is related to cognition is limited.
More importantly, Zajonc (2000) assumed that the mere exposure effect
does not rely on cognitive or cortical involvement, suggesting the effect may
rely on the low route to emotion. The low route to emotion would allow for
1218 STORBECK AND CLORE

affective processing without cortical or cognitive input. However, available

evidence suggests that the effect does not rely on processing by the
amygdala. For example, Greve and Bauer (1990) report the case of a
patient, GY, who had an accident that severed the connection between
the visual cortex and the amygdala. They found that GY showed the mere
exposure effect even though visual information was not getting to the
amygdala. Elliott and Dolan (1998) also report that the mere exposure effect
relies more on frontal cortical networks. They failed to observe any relevant
amygdala activation. These data suggest that the mere exposure effect is not
a phenomenon that requires the amygdala, let alone one that could be based
in the low route to the amygdala.

Affective priming as evidence for affective primacy. Another phenom-

enon that has been interpreted as demonstrating a special status for affect is
affective priming. Research suggests that people routinely evaluate objects in
their environment (Murphy & Zajonc, 1993; Niedenthal, 1990). For
example, in sequential priming studies, briefly presented prime words are
followed by target words that participants categorise (e.g., as positive or
negative) as quickly as possible (Bargh, Chaiken, Govender, & Pratto, 1992;
Bargh, Chaiken, Raymond, & Hymes, 1996; Fazio, Sanbonmatsu, Powell, &
Kardes, 1986). Priming is then seen when, for example, positive primes
facilitate responses to positive targets and interfere with responses to
negative targets. Such affective priming suggests that people may auto-
matically evaluate stimuli without an intention to do so.
In an effort to rule out the possibility that the response facilitation is
really due to some semantic dimension, investigators typically choose primes
and targets that have no association other than being similar or dissimilar in
evaluative meaning. However, Storbeck and Robinson (2004) point out that
this practice of limiting the relationship between primes and targets to
evaluation may force respondents into evaluative priming. If so, it would lose
its value as evidence that affect is independent of cognition or has primacy
over semantic meaning.
To test this hypothesis, Storbeck and Robinson (2004) used ‘‘a comparative
priming method’’ in which words were selected to vary not only evaluatively
but also categorically. For example, primes or targets might be positively or
negative valenced animals (e.g., puppies, snakes) or positive or negatively
valenced texture words (smooth, rough). Thus, prime-target pairs were related
evaluatively (good vs. bad), but also descriptively (animals vs. textures).
Two tasks were used, an evaluation task and a lexical decision task, and
both revealed semantic but not affective priming. In addition, the same
result was found when they used pictures instead of words. To verify that the
practice of artificially limiting the relationship between primes and targets to
 INTERDEPENDENCE OF COGNITION AND EMOTION 1219

evaluation had promoted affective priming in prior studies, Storbeck and

Robinson (2004) then repeated their experiment but removed any systematic
descriptive relationship between primes and targets, such that all words were
now animal exemplars. As expected, the usual affective priming results
reappear when participants are given only evaluation as a possible basis for
relating primes and targets.
Another comparative priming study was performed by Klauer and Musch
(2002). They used primes and targets that could be categorised based on
affect or another non-affective dimension, and manipulated only the task
demand (i.e., to evaluate or categorise). They concluded that affective
priming is not based on a special evaluation system. Rather, affective
priming relies on the same mechanisms responsible for semantic priming.
These experiments suggest that affective priming is not obligatory. The
evaluative meaning on which affective priming is based is represented within
a larger semantic network in which it is not the dominant mode of semantic
categorisation. Evaluation is doubtlessly a very basic level of analysis, but
evaluative meaning is not processed apart from other dimensions of
semantic meaning, nor does it invoke a special automatic evaluator.
The Storbeck and Robinson experiments used supraliminal exposures of
primes, but Erdley and D’Agostino (1988) used subliminal exposures. In a
very different paradigm, they too found that when both affective and
descriptive features were present, priming occurred along semantic rather
than evaluative lines, suggesting that categorisation may often have priority
over evaluation.
One might hypothesize that affect is elicited automatically at the onset of
a stimulus and degrades from that point. If so, the use of a relatively long
Stimulus Onset Asynchrony (SOA) (300 ms) by Storbeck and Robinson
(2004) might conceivably have prevented detection of affective priming. To
assess this possibility, studies might again use the comparative-priming
approach and shorten either stimulus durations or response times, either of
which might allow early components of the priming process to be visible. A
study by Klinger, Burton, and Pitts (2000) satisfied these two requirements,
and concluded that when primes are presented subliminally and response-
window procedures are used, finding semantic or affective priming depends
mainly on task requirements and response competition.
The Klinger et al. study is unique in the use of the response-window
procedure. Since spreading activation builds up over hundreds of milli-
seconds (Perea & Rosa, 2002), such procedures tend to reduce any effects
due to spreading activation within the semantic network, making it likely
that any priming effects obtained are due to response compatibility, rather
than spreading activation. It is interesting to note, though, that both
semantic and affective priming were sensitive to similar task constraints,
suggesting that both result from similar mechanisms. Other studies using
1220 STORBECK AND CLORE

similar comparative methods and response-window procedures also found

both affective and categorical priming (Klauer & Musch, 2002; Klinger et
al., 2000). However, crucially, these studies failed to equate semantic and
affective features, and in each, affect was the most salient feature. Together
these studies suggest that with response-window procedures, regardless of
prime duration, priming is driven by response compatibility.
Since the use of a response window shortens the time available for effects
due to spreading activation, what happens when spreading activation is
allowed to build up over time, by presenting primes subliminally without a
response window. Kemp-Wheeler and Hill (1992) performed such a study
with a lexical decision task, and found both affective and semantic priming.
But they also found that affective priming occurred mostly when people
could detect the prime. Such detection did not facilitate semantic priming.
They argued that affective priming is a subform of semantic priming and
occurs when more time is given to revealing the affective significance of
primes and targets.
Moving away from the priming procedure, Storbeck, Robinson, Ram,
Meier, and Clore (2004) examined evaluations and categorisations of single
target words using a response-window procedure. The window of response
varied from 100 ms to 2000 ms and the dependent measure was accuracy.
The experiment included nine participants over five days with over 500 trials
per day. This allowed us to produce predictive models for the rise of semantic
and affective accuracy. The results revealed that in shorter response
windows, participants were more accurate in detecting semantic information
than affective information. Other studies have found that semantic distinc-
tions can occur as early as 80 ms, while evaluative distinctions start around
100 ms (Van Rullen & Thorpe, 2001).
EEG measures can also be used to discriminate semantic and affective
aspects of processing without involving motor output processes. Cacioppo,
Crites, and Gardner (1996) and Ito and Cacioppo (2001) found that ERP
potentials always tracked semantic relations, even when semantic analysis
was not the focus of the task. ERPs also tracked affective features, but only
when the task had an explicitly evaluative focus, unless the evaluative
components were quite potent. More critically, evidence suggests that
the same discriminative processing based on semantic features performed
by the visual cortex occurs whether stimuli are presented subliminally or
supraliminally, regardless of conscious experience (Dehaene et al., 2001;
Rolls & Tovee, 1994; Rolls et al., 1994; Stenberg, Lindgren, & Johansson,
2000). ERP and single-cell recordings both demonstrate that semantic
information appears to be represented regardless of the task at hand and
whether or not there is conscious perception of the stimuli. That is, semantic
information always gets activated, regardless of the explicit task, whereas
 INTERDEPENDENCE OF COGNITION AND EMOTION 1221

affective information is processed mainly when evaluation is an explicit part

of the task or a highly salient aspect of the stimulus.
To be clear, in this view, the system needs an identification stage before an
evaluation stage, and identification occurs in later stages of processing in the
visual cortex. Even in classical conditioning, some kind of identification is
required by the cortex (e.g., visual cortex) to discriminate a conditioned
stimulus from all other stimuli. By ‘‘identification stage’’ we mean simply
that a view invariant, neural signature of an object is activated in the visual
cortex. Only then can the object activate affective and other associations.

Conclusions. These studies suggest that both semantic4 and affective

features are represented in a single semantic network, and that semantic
information (which is not to say lexical information, see footnote 3) has a
necessary priority. That is, we feel that affective priming is a special case of
semantic priming and can be obtained when affect is part of the task
demand, the salient feature of the stimuli, or the focus of attention (Storbeck
& Robinson, 2004). Under the right set of circumstances, affective relations
can be made more accessible than semantic relations (e.g., Bargh et al., 1992,
1996; Klinger et al., 2000; Storbeck & Robinson, 2004). For example,
Storbeck and Robinson (2004) found that when they crossed descriptive and
evaluative features of stimuli in an evaluative priming task, semantic but not
affective priming was observed. But when the relations between primes and
targets stimuli were limited to their evaluative features, then affective
priming was observed. Thus, under the right set of conditions, affective
priming can readily be observed, but such evaluative priming is in no way
obligatory. Thus, the fact that evaluative priming can be found when
evaluative meaning is made salient, provides little support for ideas about
affective primacy or about the separate nature of affective and cognitive
processing.

C. The affective automaticity hypothesis

Although the automatic
controlled distinction arose in cognitive psychol-
ogy (Shiffrin & Schneider, 1977), a special association is often assumed
between affect and automaticity. Perhaps the idea was that thoughts can be
more easily controlled than feelings has made affect seem to have a life of its

4
We will use the term ‘‘semantic’’ to describe the meaning analysis that we propose precedes
affective analysis. What we have in mind specifically are at least three achievements: (1) the
integration of multiple features of the object into a single ‘‘object’’ code; (2) the identification of
this object; and (3) the categorisation of the object (e.g., as animate or not). The term semantic,
then, refers somewhat more directly to the achievements of area IT (especially invariance,
identification, and categorisation) that seem to occur in order for a person to retrieve affective
associations.
1222 STORBECK AND CLORE

own. One can decide to think about one particular topic rather than another,
but one cannot decide to feel one way or another, except by guiding
thoughts. Is automaticity a key distinction that makes affect and emotion
separate from cognition?
Cognitive psychologists have recently become critical of the term
‘‘automaticity’’. Recent reviews have concluded that the initial demonstra-
tions of what was purported to be automaticity may actually have required
attention after all (see, Lavie & De Fockert, 2003; Logan, 2002; Pashler,
Johnston, & Ruthruff, 2001; Stolz & Besner, 1999). For example, Pashler
et al. (2001, p. 648) stated that, ‘‘A variety of proposals for ‘wired-in’
attention capture by particular stimulus attributes have been effectively
challenged; attention, it turns out, is subject to a far greater degree of
top-down control than was suspected 10 years ago’’. Generally, the relevant
data have come from studies of cognition rather than affect. In this section,
we suggest that the same conclusion applies in the case of affective stimuli.
Harris, Pashler, and Coburn (2004) examined whether affective words
could be processed automatically. Their data indicated that affective words
can slow responses down on a primary task, suggesting that affect may
capture attention. However, when the primary task was made difficult, thus
reducing attentional resources, affective words failed to slow responses,
suggesting that affect did not capture attention. These results suggest that
under high-load conditions, when attention is occupied, affective words
should not be expected to ‘‘grab’’ attention in a bottom-up manner. Instead,
affect appears to be processed by top-down networks. Similar results have
been found when emotional faces were used in a modified Posner cueing
paradigm (Fox, Russo, Bowles, & Dutton, 2001) and when threat-related
words and faces were used in a variation of the Stroop task (White, 1996).
Moreover, examining the affective pronunciation priming task, De Houwer
and Randell (2002) observed affective priming only when attention was
focused on the primes. When attention was not focused on the primes,
affective priming was not observed in the pronunciation paradigm.
These studies all presented evidence to suggest that affective stimuli
require attention and that they do not grab attention in a bottom-up
manner. However, Lundqvist and Öhman (2005) have argued that evolutio-
narily relevant threat stimuli (e.g., snakes, spiders, faces) should be especially
likely to be processed pre-attentively (see Davey, 1995, for a relevant
criticism to evolutionary preparedness account).
Relevant data are limited, but, the data available would suggest that even
faces require attention in order to be processed. As discussed above, Fox
et al. (2001) found that angry, happy, and neutral faces failed to capture
attention when the effects of attention capture versus disengagement were
disentangled. Narumoto et al. (2001) found that when faces were presented,
area STS, which processes facial expressions, was significantly activated only
 INTERDEPENDENCE OF COGNITION AND EMOTION 1223

when the task required facial expression discrimination, but not when identity
or gender discriminations were required for the emotional faces (see also
Critchley et al., 2000). Pessoa, Kastner, and Ungerleider (2002) performed a
study similar to the Harris et al. study, but they used pictures with facial
expressions and collected neuroimaging data. They observed that under low-
load conditions, amygdala activation was observed to task-irrelevant fear
faces. But, under high-load conditions, when processing resources were
limited, the amygdala failed to show significant activation to task-irrelevant
fear faces, suggesting that attention was driven by top-down influences. These
findings suggest that even the amygdala needs attentional resources in order to
process fear faces and that fear faces can fail to capture attention.

Amygdala evaluation requires attention. It has been suggested that

emotional stimuli are processed automatically, namely, without attention
(LeDoux, 1996; Öhman, Esteves, & Soares, 1995; Vuilleumier, Armony,
Driver, & Dolan, 2003), and that the amygdala plays a key role in automatic
stimulus evaluation (Morris, Öhman, Dolan, 1998; Whalen et al., 1998).
This process is often cited as the basis of affective primacy (e.g., Bargh &
Chartrand, 1999; Zajonc, 2000). However, cortical input appears to be more
important in amygdala processing than has sometimes been emphasised (as
discussed earlier), and the data reviewed below suggest that the amygdala
requires attention to process threatening and novel stimuli.
Several studies have tested the hypothesis that exposure to affective words
should elicit amygdala activation, reflecting the automatic evaluation
process (Beauregard et al., 1997; Canli, Desmond, Zhao, Glover, & Gabrieli,
1998). No evidence was found of the hypothesised amygdala activation
unless attention was explicitly drawn to the affective content of words by
asking participants to evaluate them. Such results suggest that the amygdala
does not continuously evaluate all incoming stimuli.
These studies involved lexical stimuli, but the same turns out to be true
for the evaluation of pictures.5 When presented with affective pictures,

5
A host of fMRI studies have demonstrated the activation of the amygdala to masked fear
faces and other emotional stimuli. Such studies are interesting because individuals do not have a
conscious perception of the image. However, the amygdala only shows enhanced activation to
arousing images (e.g., fear faces), but not to non-arousing faces (e.g., houses). Although such
evidence suggests that amygdala activation can occur without perceptual awareness, we still
suggest that the visual system still codes that image and sends its input forward to the amygdala
in the same manner as if the stimulus was presented supraliminally. Moreover, imaging studies
have a weakness of comparative activity. Therefore, it is difficult to gage how much processing
is done between masked and non-masked fear faces. In addition, there is plenty of evidence
to suggest that the visual cortex processes masked and non-masked images in a similar manner.
(continued overleaf)
1224 STORBECK AND CLORE

Keightley et al. (2003) found no amygdala activation. When participants

were explicitly asked to evaluate affective stimuli, amygdala activation was
found only for negative information (Keightley et al., 2003; Lane, Chua, &
Dolan, 1999). For fearful faces, however, even passive viewing showed
amygdala activation (Critchley et al., 2000; Morris et al., 1998; Vuilleumier
et al., 2003; Whalen et al., 1998). However, with other face stimuli there was
no amygdala activation even when participants explicitly evaluated them
(Critchley et al., 2000; Keightley et al., 2003). Happy and angry faces also
showed no amygdala activation for either passive viewing or active
evaluation (Blair, Morris, Frith, Perrett, & Dolan, 1999; Morris et al.,
1998; Surguladze et al., 2003).

(Continued )
Moreover, evidence from single-cell recording suggests that the visual system can still determine
whether a face or a house was presented regardless of whether each image was presented with a
mask and subliminally. Therefore, studies demonstrating that the amygdala activates for a
subliminal, but not a supraliminal picture does not mean that the visual cortex did not send the
same information. There is no reason to believe that the categorisation processes performed by
area IT are conscious. Indeed, on the basis of ERP data, we might conclude that unconscious
categorisation routinely precedes conscious categorisation. Furthermore, unconscious
categorisation by the visual system may occur extremely quickly after stimulus exposure, in as
little as 48 ms for ‘‘global templates’’ (Sugase, Yamane, Ueno, & Kawano, 1999) and 70
80 ms
for classes of stimuli (Van Rullen & Thorpe, 2001). Interestingly, Van Rullen and Thorpe (2001)
also found that the initial (70
80) categorisation-related ERP component was not highly
correlated with a participant’s response to the task at hand, whereas an ERP component that
occurred at 190 ms post-stimulus onset was. Thus, categorisation appears to occur quite rapidly
and seems to occur independently of later, possibly more conscious, categorisation processes.
Relatedly, people can classify objects on the basis of category membership even with no
awareness of the distinct categories guiding their response (e.g., Reed, Squire, Patalano, Smith,
& Jonides, 1999). In summary, we conclude that categorisation occurs within later stages of the
visual cortex, specifically area IT. Moreover, other data suggest that these same visual areas are
not sensitive to the affective significance of objects (Iwai et al., 1990; Nishijo, Ono, & Nishino,
1988a; Rolls, 1999; Rolls, Judge, & Sanghera, 1977). Thus, within area IT and other later stages
of the visual cortex we appear to have considerable evidence for categorisation prior to affect
retrieval. Recall that studies have found distinct category-related ERPs within 70
80 ms post-
stimulus onset (e.g., Van Rullen & Thorpe, 2001). Object identification also appears to occur
rapidly, perhaps within 100 ms of stimulus onset (Lehky, 2000; Rolls & Tovee, 1994). These
findings suggest that categorisation tends to occur prior to identification. Nevertheless, studies
that present masked stimuli have demonstrated that even stimuli presented as briefly as 20
60
ms with pre- and postmasks are still sufficiently processed by area IT to support object
identification (Dehaene et al., 2001; Rolls, 1999; Vogels & Orban, 1996). In the latter connection,
Rolls, Tovee, Purcell, Stewart, and Azzopardi (1994) argued that such subliminal presentations
reduce the amplitude of neural responses to stimuli, but do not change fundamental neural
identification processes (see also Kovacs, Vogels, & Orban, 1995, for similar results). Thus, the
primary difference between subliminal and optimal viewing conditions pertains to the amplitude
of the neuronal responses within area IT, but sufficient processing still occurs to produce an
invariant neural code (i.e., identification). From this perspective, demonstrations of
‘‘unconscious’’ cognition or affect are not particularly special from a neurological point of view.
 INTERDEPENDENCE OF COGNITION AND EMOTION 1225

Conclusions. These results suggest that valence is not automatically

processed by the amygdala, but the amygdala may be sensitive to arousing
stimuli such as fearful faces. Other research groups have also suggested that
the amygdala is important for encoding arousal, but not the valence,
dimension of stimuli (Adolphs & Damasio, 2001; Adolphs, Russell, &
Tranel, 1999; Cahill et al., 1996; Lane et al., 1999; McGaugh, 2004; Morris
et al., 1998; Surguladze et al., 2003). Moreover, the evidence suggests that
when affect is salient and processing demands are relatively low, emotional
information may engage attention. But when processing demands are high
and affective stimuli are not of attention, affect will not ‘‘capture’’ attention.
Such findings limit the conditions for automaticity, and, as cognitive
psychology has already discovered, processing relies on attention, even for
affective stimuli.

II. THE AFFECT
COGNITION CONNECTION

Throughout history, people’s optimism or pessimism about the human
condition has often turned on their beliefs about the possibility of rational
thought unsullied by emotion. Gradually, however, cognition and emotion
are coming to be viewed as complementary rather than antagonistic
processes. Our current research is informed by an affect-as-information
approach (Schwarz & Clore, 1983; Clore & Storbeck, in press), which
assumes that affective reactions provide useful feedback both explicitly and
implicitly from emotional appraisal processes. Evidence in support of such a
view comes from observations that the inability to use affective feedback as a
result of brain damage has profoundly negative consequences for judgement
and decision making (Damasio, 1994). Conversely, expertise at using
affective information seems to be associated with effective personal and
social functioning (Mayer, Salovey, & Caruso, 2004).

A. Emotion modulates cognition6

In Part I, we argued against the idea that cognition and emotion involved
distinct brain areas or that they operate independently. The strongest claim
for independence relied on the ‘‘low route’’ to emotion (LeDoux, 1996), a
direct pathway from the sensory thalamus to the amygdala. However, by all
available evidence, the low route does not appear to be a candidate for
explaining any instance of human emotion. If it operates at all in humans, it
appears incapable of even basic affective discriminations without cognitive

6
The section title implies that cognition does not modulate emotion. We would suggest, like
others have, that in fact cognition does modulate emotion (e.g., Ochsner & Gross, 2005), but
such a discussion is beyond the scope of this article.
1226 STORBECK AND CLORE

input. Rather, the evidence from neuroscience suggests that evaluations of

the amygdala are dependent upon input from the visual cortex. We suggested
that affect probably does not proceed independently of cognition, nor
precede cognition in time.
How, then, do we see the relationship between emotion and cognition? At
the most general level, emotion modulates and mediates basic cognitive
processes. The brain, of course, accomplishes numerous tasks all at once,
including automatic processes (Barnard et al., 2007 this issue; Robinson,
1998). As the sensory cortex identifies stimuli in the environment, the visual
cortex processes it in a view-invariant manner, allowing it to determine
attributes of the object, including its affective significance, regardless of the
position the object happens to be in. Once the visual cortex creates a view-
invariant code for the object, it projects that information to other areas in
the brain.
One of the primary pathways of the visual cortex is to the amygdala, and
the role of the amygdala is in part to determine the urgency of the stimulus,
which eventuates in the marking of apparently important experiences
hormonally and in terms of experienced arousal. The amygdala retrieves
the affective value of the stimulus or determines that it is novel and guides
subsequent cognitive processing. The amygdala has extensive back projec-
tions to all areas of the visual cortex, which we believe modulate visual
perception, attention, and memory for affectively significant stimuli. Note
that the amygdala is probably not the only area involved in emotional
processing that can modulate cognition. The visual cortex also has extensive
projections to areas such as the orbitofrontal cortex, prefrontal cortex, and
cingulate cortex, all of which can guide cognitive processing based on
affective value.
In this section, we illustrate how we believe affect regulates cognition by
briefly reviewing several recent studies from our lab. The studies discussed
focus on two problems *the role of affect in perception and the affective
regulation of styles of information processing. We note that in performance
situations, emotional cues regulate cognitive processing, serving to adjust the
mix of cognition and perception. Of special interest are several recent
experiments that ask about affective consequences for implicit processes of
learning, memory, priming, and attitude.

B. The affective regulation of perception

The ‘‘New Look’’ in perception, a movement in the 1950s (Bruner, 1957),
maintained that rather than being a passive registration of reality, perception
reflected internal expectations and motivations as part of an adaptive
process. That movement quickly ran its course without having much impact,
but, today, research again suggests that perception of the physical world is
 INTERDEPENDENCE OF COGNITION AND EMOTION 1227

influenced by emotion and other internal factors. For example, Proffitt and
colleagues (e.g., Bhalla & Proffitt, 1999; Proffitt, Stefanucci, Banton &
Epstein, 2003; Witt, Proffitt, & Epstein, 2004) have found that hills appear
steeper and distances farther to people with reduced physical resources,
either from wearing a heavy backpack, being physically tired, or being
elderly. Recent research shows that emotion can have similar effects. In one
study (Riener, Stefanucci, Proffitt, & Clore, 2003) participants listened to
happy or sad music as they stood at the bottom of a hill. The results showed
that sadness can make mountains out of molehills. Sad mood led to
overestimation of the incline on verbal and visual measures, but not on a
haptic measure. That is, the sad individuals were more likely to say that the
hill was steeper compared to happy individuals, but both groups provided
similar haptic responses.
Affective feelings thus appear to inform explicit, but not implicit
measures of perception. That is, when asked to estimate the incline verbally
in degrees (i.e., verbal measure) and when indicating the incline analogically
with a sort of protractor (i.e., visual measure), individuals feeling sad
estimated the hill to be significantly steeper than individuals who were
feeling happy or who had not heard any music. Such perceptual measures
are thought to reflect conscious visual perception that relies on processing in
the ventral visual stream, or ‘‘what’’ system, concerned with visual
identification (Milner & Goodale, 1995). A reasonable argument can be
made for why this system might be sensitive to resources for coping with
inclines and distances (Proffitt, 2006). The third, haptic measure involved
tilting a palm board (without looking at it) to match the incline of the hill.
This haptic measure of incline is generally found to be quite accurate and to
be immune from the influence of resource depletion such as physical
exhaustion. It was also unaffected by sad mood. The measure is thought to
reflect unconscious visual perception and relies on processing in the dorsal
visual stream, or ‘‘how’’ system, engaged in the visual control of motor
behaviour. Whereas it might be adaptive for one’s perception of a hill to
reflect one’s resources, as decisions on whether to take action or not might
hinge on such information, but for regulation of one’s actual foot placement,
such overestimations might be disastrous.
In extensions of this work, Stefanucci, Proffitt, and Clore (2005) also
examined the effect of fear on hill estimates. They had individuals on top of
the hill and to manipulate fear, some individuals stood on a skateboard,
whereas others stood on a stable platform. They found that individuals on
the skateboard provided steeper verbal hill estimates again on both the
verbal and visual measures when compared to individuals standing on
the stable platform. As expected, the haptic measure was again unaffected by
the manipulation of emotion.
1228 STORBECK AND CLORE

C. Affective regulation of processing

At the beginning of the cognitive revolution, Jerome Bruner (1957) famously
concluded that people are active processors who typically ‘‘go beyond the
information given’’. A number of experiments have been conducted in our
lab over the past five years in which emotions and moods were added to
classic experiments in cognitive psychology. One way to summarise our
results is to say that happy affect appears to promote this ‘‘going beyond’’
through its influence on ‘‘relational processing’’. In contrast, negative affect
leads to more item-specific processing. Such results lead us to conclude that
Bruner’s dictum, and all that it implies, may not be applicable when
emotional cues of sadness are present. The experiments from our lab suggest
(perhaps ironically) that the cognitive revolution had a hidden emotional
trigger.
Many of the classic phenomena on which cognitive psychology was
founded turn out to depend on affect. For instance, we observed that
individuals in happy moods, but not those in sad moods, demonstrate
schema effects on constructive memory of the kind introduced by Bartlett in
1932 (Gasper & Clore, 2002). Other classic phenomena also turn out to be
more pronounced in happy moods than in sad moods. These include
semantic priming (Storbeck & Clore, 2006), script processing (Bless et al.,
1996), schema-guided memory (Gasper & Clore, 2002), stereotype use
(Isbell, 2004), heuristic reasoning (Gasper, 2000), the global superiority
effect (Gasper & Clore, 2002), and false memory generation (Storbeck &
Clore, 2005). These results do not arise from general performance deficits
caused by sad mood. On the contrary, general reaction times, overall
memory accuracy, and basic performance levels often show no mood-based
differences. Moreover, since the classic paradigms often rely on particular
errors to show the mediating role of knowledge structures, individuals in sad
moods may perform better in certain ways than those in happy moods.
These observations are compatible with findings demonstrating that
positive moods are associated with processing that is generative (e.g., Erez &
Isen, 2002), constructive (e.g., Fiedler, 2001), and broad (e.g., Fredrickson &
Branigan, 2005). Our own account of these effects emphasises the informa-
tional properties of affect. For example, during task performance, positive
affect may be experienced as efficacy and negative affect as difficulty. Feeling
that one is effective confers value on one’s own generative thoughts and
goals resulting in the reliance on them to process incoming information
(relational processing). On the other hand, experiences of difficulty and lack
of efficacy reduce the apparent value of one’s own cognitions and goals,
leading to a focus on more-specific, literal aspects of stimuli.
 INTERDEPENDENCE OF COGNITION AND EMOTION 1229

D. Affective regulation of implicit processes

Priming. In other research, Storbeck and Clore (2006) tested whether
this relational processing of associations can carry over to semantic
knowledge. They observed that happy individuals were more likely to relate
primes and targets together, demonstrating both category and evaluative
priming, depending on the nature of the task. However, sad individuals
failed to demonstrate priming on the same tasks, suggesting they were
impaired in relating the descriptive meaning from primes to targets. Again,
the results suggest that negative affective cues act as though they undermine
confidence in using accessible cognitions. In the implicit learning situation, it
prevented expression of what had been learned, and in the priming situation,
it allowed sad participants to respond to target stimuli independently from
the descriptive meaning of the primes.

False memory effects. To investigate further the hypothesis that negative

affect impairs the formation and use of implicit associations, Storbeck and
Clore (2005) induced positive and negative moods before a false-memory
task. The task produces false memories by presenting word lists in which the
lists are composed of words that are highly associated to a non-presented
word, referred to as the critical lure. False memories are engendered because
as individuals are relating the words from the list together, the critical lures
should come to mind and are then likely to be falsely recalled. We observed
that, in fact, negative moods led to a decrease in activation and subsequent
recall of critical lures compared to the positive mood group and the control
group. In addition, no differences were observed for the recall of presented
items between the three groups. Ironically, the observed effect demonstrated
that negative affect can improve memory performance by inhibiting the use
of lexical associations during learning. Such findings suggest that affect from
mood can influence the expression of implicit associations (Storbeck &
Clore, 2005, 2006).

Affective involvement in implicit attitudes. The previous experiments

show that affective states modulate the use of implicit associations in
cognitive performance situations. Extensive prior research has already
shown that affective states can influence evaluative judgements or attitudes
expressed in self-report measures. But the intense interest in implicit attitude
measurement raises the question of whether or not affect influences attitudes
when assessed on implicit measures such as the Implicit Association Test or
IAT (Greenwald, McGhee, & Schwartz, 1998).
Several experiments (Huntsinger, Sinclair, Dunn, & Clore, 2006) tested the
hypothesis that positive affect would serve as a ‘‘go’’ sign and negative affect as
a ‘‘stop’’ sign for acting on goals that were either chronically or temporarily
1230 STORBECK AND CLORE

activated. The goal in one experiment concerned taking an egalitarian stance

regarding sexist attitudes, and in the other experiment the goal was either to
adopt or not to adopt the racial attitudes held by an experimenter. Implicit
measures of attitude were employed (a lexical decision task and an IAT). An
elaborate set of effects neatly confirmed the predictions, showing that in each
case, happy moods prompted participants to act on their chronic or
temporarily activated goals, whereas sad moods interfered with goal expres-
sion. Importantly, the goals had been activated implicitly as a subtle part of
the social situation, and the attitudes were measured implicitly using two
different measures *see also DeSteno, Dasgupta, Bartlett, and Cajdric (2004)
for a demonstration of the effects of anger on implicitly measured prejudice
toward an outgroup. Thus, affective states appear to regulate not only the
expression of implicit learning and implicit lexical associations, but also the
expression of implicitly measured attitudes.
In summary, our main goal of this section was to demonstrate that affect
and cognition should be thought of as fundamentally interactive. In this
view, affect is potential moderator of all kinds of cognitive operations from
perception and attention to implicit learning and implicit associations (see
also, Duncan & Barrett, this issue). We have argued against conceptualising
emotion as a separate force in opposition to cognition in favour of viewing
cognition and emotion as inherently integrated. We included examples of
recent research in our own lab showing affective moderation of basic
cognitive processes.

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