See it with feeling: affective predictions during
L. F. Barrett1,2,*and Moshe Bar2
1Boston College, Chestnut Hill, MA 02467, USA
2Massachusetts General Hospital/Harvard Medical School, Charlestown, MA 02129, USA
People see with feeling. We ‘gaze’, ‘behold’, ‘stare’, ‘gape’ and ‘glare’. In this paper, we develop the
hypothesis that the brain’s ability to see in the present incorporates a representation of the affective
impact of those visual sensations in the past. This representation makes up part of the brain’s
prediction of what the visual sensations stand for in the present, including how to act on them in the
near future. The affective prediction hypothesis implies that responses signalling an object’s salience,
relevance or value do not occur as a separate step after the object is identified. Instead, affective
responses support vision from the very moment that visual stimulation begins.
Keywords: affect; emotion; perception; prediction; amygdala; orbitofrontal cortex
Michael May lost the ability to see when he was 3 years
old, after an accident destroyed his left eye and
damaged his right cornea. Some 40 years later, Mr
May received a corneal transplant that restored his
brain’s ability to absorb normal visual input from the
world (Fine et al. 2003). With the hardware finally
working, Mr May saw only simple movements, colours
and shapes rather than, as most people do, a world of
faces and objects and scenes. It was as if he lived in two
different worlds: one where sound, touch, smell and
taste were all integrated, and a second world of vision
that stood apart. His visual sensations seemed foreign,
similar to a language he was just learning to speak. As
time passed, and Mr May gained experience with the
visual world in context, he slowly became fluent in
vision. Two years after his surgery, Mr May commen-
ted: ‘The difference between today and over 2 years ago
is that I can better guess at what I am seeing. What
is the same is that I am still guessing.’ (p. 916, italics in
What Mr May did not know is that sighted people
automatically make the guesses he was forced to make
with effort. From birth, the human brain captures
statistical regularities in sensory–motor patterns and
stores them as internal representations. The brain then
uses these stored representations, almost instan-
taneously, to predict continuously and unintentionally
what incoming visual sensations stand for the in-world
(Bar 2003, 2007, 2009; Kveraga et al. 2007a,b). When
the brain receives new sensory input from the world in
the present, it generates a hypothesis based on what it
knows from the past to guide recognition and action in
the immediate future. This is how people learn that the
sounds of a voice come from moving lips on a face, that
the red and green blotches on a round Macintosh apple
are associated with a certain tartness of taste, and that
the snowy white petals of an English rose have a velvety
texture and hold a certain fragrance.
External sensations do not occur in a vacuum, but in
a context of internal sensations from the body that
holds the brain. The sensory–motor patterns being
stored for future use include sensations from organs,
muscles, joints (or ‘interoceptive’sensations), as well as
representations of sights, sounds, smells, tastes and
touch (‘exteroceptive’ sensations). Although bodily
sensations are rarely experienced consciously or with
perfect fidelity (Barrett et al. 2004), they are continually
detected and internally represented (Craig 2002,
2009). Psychology refers to these internal bodily
changes as ‘affective’. In addition to directly experien-
cing changes in their breathing, muscle tension or
stomach motility, people routinely experience more
diffuse feelings, pleasure or discomfort, feeling worked
up or slowed down. Therefore, in addition to learning
that the sounds of a voice come from moving lips,
people also learn whether or not they liked the sound of
the voice (Bliss-Moreau submitted) and the person that
it belongs to (Bliss-Moreau et al. 2008); they learn that
they enjoy tartly flavoured red and green (Macintosh)
apples or the milder tasting yellow apples (Golden
Delicious); and they learn whether or not they prefer
the strong fragrance of white English roses over the
milder smell of a deep red American Beauty, or even
the smell of roses over lilies. When the brain detects
visual sensations from the eye in the present moment,
and tries to interpret them by generating a prediction
about what those visual sensations refer to or stand
for in the world, it uses not only previously encount-
ered patterns of sound, touch, smell and tastes, as well
as semantic knowledge. It also uses affective
representations—prior experiences of how those exter-
nal sensations have influenced internal sensations from
the body. Often these representations reach subjective
awareness and are experienced as affective feelings,
but they need not (e.g. Zajonc 1980; for a recent
Phil. Trans. R. Soc. B (2009) 364, 1325–1334
One contribution of 18 to a Theme Issue ‘Predictions in the brain:
using our past to prepare for the future’.
*Author and address for correspondence: Boston College, Chestnut
Hill, MA 02467, USA (firstname.lastname@example.org).
This journal is q 2009 The Royal Society
discussion, see Barrett & Bliss-Moreau in press).
Guessing at what an object is, in part, requires knowing
In this paper, we explore the hypothesis that the
brain routinely makes affective predictions during
visual recognition. We suggest that the brain’s predic-
tion about the meaning of visual sensations of the
present includes some representation of the affective
impact of those (or similar) sensations in the past. An
effective prediction, in effect allows the brain to
anticipate and prepare to act on those sensations in
the future. Furthermore, affective predictions are made
quickly and efficiently, only milliseconds after visual
sensations register on the retina. From this perspective,
sensations from the body are a dimension of
knowledge—they help us to identify what the object is
when we encounter it, based, in part, on past reactions.
If this hypothesis is correct, then affective responses
signalling an object’s salience, relevance or value do
not occur as a separate step after the object is
Figure 1. A descriptive map of core affect.
Figure 2. (a–d) The affective workspace. The neural workspace for core affect includes a broadly distributed set of
interconnected cortical and subcortical brain areas that are not functionally specific to affect, but that realize affective responses
as a network (for imaging evidence, see Kringelbach & Rolls 2004; Barrett et al. 2007a; Wager et al. 2008; for a discussion, see
Barrett & Bliss-Moreau in press). Some areas are traditionally considered to be ‘emotional’, such as the amygdala (rose) and
ventral striatum (green). Other areas were (until recently) considered ‘cognitive’ (cf. Duncan & Barrett 2007; for a similar view,
see Pessoa 2008), such as the orbitofrontal cortex (OFC; blue and purple). Typically, researchers use the term ‘orbital frontal
cortex’ to refer to anywhere within the orbital sector of the prefrontal cortex. This includes the lateral parts (blue) that are
bounded by the ventrolateral prefrontal cortex (vlPFC) and agranular insula (yellow), as well as the medial portions that wrap to
include the ventromedial prefrontal cortex (vmPFC; purple) extending back to the sub/pregenual portions of the anterior
cingulate cortex (ACC; brown and gold). Other components of this workspace include the hypothalamus (light green) and
autonomic control centres in the midbrain and brainstem (turquoise and maroon). This neural reference space for effect is
meant to be non-specific without sounding vague, in that different assemblies of neurons across these areas realize momentary
representations of value (Ghashghaei & Barbas 2002; Barbas et al. 2003). Photographs taken from DeArmond et al. (1989),
pp. 5, 7, 8 and 43.
1326 L. F. Barrett & M. Bar
Phil. Trans. R. Soc. B (2009)
identified—affective response assists in seeing an object
as what it is from the very moment that visual
stimulation begins. Whether or not he was aware of
it, Mr May’s predictions about the visual world
probably became more effective, in part, as his affective
state became more responsive to his visual sensations.
Learning to see meant experiencing visual sensations as
value added. Whether experienced consciously or not,
a pleasant or unpleasant affective change in response to
visual stimulation might have helped Mr May (as it
helps all of us) to recognize moving lips as a particular
person speaking, a blotch of red and green as an apple
or white silky petals as a rose.
2. AFFECT DEFINED
In English, the word ‘affect’ means ‘to produce a
change’. To be affected by something is to be
influenced by it. In psychology, affect refers to a
specific kind of influence—something’s ability to
influence a person’s body state. Sometimes, the
resulting bodily sensations in the core of the body are
experienced as physical symptoms (such as being
wound up from drinking too much coffee, fatigued
from not enough sleep or energetic from exercise).
Much of the time, sensations from the body are
experienced as simple feelings of pleasure or displea-
sure with some degree of activation, either alone or as
Figure 3. (a) Medial and (b) lateral networks of the OFC. Adapted from O¨ngu ¨r et al. (2003) based on the evidence from Barbas
(1993, 2000), Carmichael & Price (1995), McDonald (1998), Cavada et al. (2000), Ghashghaei & Barbas (2002) and
Stefanacci & Amaral (2002). Orbital and medial views of the brain are shown. The medial network is shown in blue and has
robust reciprocal connections to all limbic areas (including many nuclei within the amygdala and the ventral striatum), as well as
to the hypothalamus, midbrain, brainstem and spinal cord areas that are involved in internal state regulation. The medial OFC
has few direct connections to sensory cortices. The lateral network is shown in purple and has robust connections with unimodal
sensory association areas as well as the cortical aspects of the amygdala (including the basolateral complex, which also receives
sensory input from unimodal association areas). The lateral OFC has few direct projections to autonomic and endocrine control
centres in the hypothalamus, midbrain and brainstem, but has some influence on those autonomic centres via projections to the
intercalated masseswithin the amygdalathat have the net effect disinhibiting (i.e. activating) these nuclei. The areas that connect
the two networks are shown in rose.
visceromotor network dorsal stream
sensory integration network
Figure 4. Connections between the OFC and visual networks. ‘S’ represents visual information that reaches the dorsal ‘where’
stream and the ventral ‘what’ stream both from early visual areas and from the thalamus. The medical OFC is part of a
‘visceromotor’ network that helps to control internal body state. The lateral OFC is part of a ‘sensory integration network’ that
joins together sensory representations from outside and inside the body.
L. F. Barrett & M. Bar1327
Phil. Trans. R. Soc. B (2009)
an emotion (figure 1; see Russell & Barrett 1999;
Barrett & Bliss-Moreau in press). At still other times,
bodily changes are too subtle to be consciously
experienced at all. Whether consciously felt or not, an
object is said to have affective ‘value’ if it has the
capacity to influence a person’s breathing, or heart rate,
hormonal secretions, etc. In fact, objects are said to be
‘positive’ or ‘negative’ by virtue of their capacity to
influence a person’s body state (just as objects are said
to be ‘red’ if they reflect light at 600 nm).
When affect is experienced so that it is reportable, it
can be in the background or foreground of conscious-
ness. When in the background, it is perceived as a
property of the world, rather than as the person’s
reaction to it. ‘Unconscious affect’ (as it is called) is
why a drink tastes delicious or is unappetizing
(e.g. Berridge & Winkielman 2003; Winkielman et al.
2005; Koenigs & Tranel 2008), why we experience
some people as nice and others as mean (Li et al. 2007)
and why some paintings are beautiful while others are
ugly (for a discussion of affect and preferences, see
Clore & Storbeck 2006). When in the foreground, it is
perceived as a personal reaction to the world: people
like or dislike a drink, a person or a painting. Affect can
be experienced as emotional (such as being anxious at
an uncertain outcome, depressed from a loss or happy
at a reward; cf. Barrett 2006a,b).
3. AFFECT AND PERCEPTION
For centuries, philosophers have believed that every
moment of waking life is to some degree pleasant or
unpleasant with some degree of arousal that affect is a
basic ingredient of mental life. This idea continues to
be incorporated into contemporary perspectives on
consciousness, including Damasio’s somatic marker
hypothesis (Damasio 1999), Edelman’s theory of
neural Darwinism (Edelman 1987; Edelman & Tononi
2000), Searle’s theory of consciousness (Searle 1992,
2004) and Humphrey’s (2006) theory of conscious
sensation. This idea can also be found in early
psychological writing of Spencer (1855), James
(1890), Sully (1892) and Wundt (1998). During the
behaviourist era in psychology, scientists no longer
regarded affect as central to perception and thought,
and this trend continued as psychology emerged from
the grips of behaviourism during the cognitive revolu-
tion. Affective responses were ignored in cognitive
science altogether and questions about affect were
relegated to the studyof emotion, where it was assumed
that affect occurred after object perception and in
reaction to it (e.g. Arnold 1960). First, a red, round,
hard object is perceived as an apple, and only then is the
object related to the past experiences of enjoying the
crunchy sweetness of the first bite, or to a breezy trip to
the farm for apple picking on a fine autumn afternoon.
As it turns out, philosophers and early psychologists
were right, at least when it comes to the place of affect
in mental life. We contend that prior affective reactions
to apples might actually help the brain to predict that
visual sensations refer to an apple in the first place. Not
only does the brain draft the prediction of an apple into
a conscious perception of an apple, but the resulting
affective response may also become conscious, so that
the apple is experienced as pleasant or perceivers
experience themselves as reacting to the apple with a
pleasant, mild, heightened state of arousal.
It is easy to see evidence for this logic by referring
briefly to the neuronal workspace that realizes affective
circuitry isthe orbitofrontal cortex (OFC).Inthispaper,
we use the acronym ‘OFC’ to refer to the entire orbital
sector of the prefrontal cortex. The OFC is a
heteromodal association area that integrates sensory
input from the world and the body (i.e. from extra- and
intrapersonal space) to create a contextually sensitive,
multimodal representation of the world and its value to
the person at a particular moment in time (Barbas 1993,
2000; Carmichael & Price 1995; McDonald 1998;
Cavada et al. 2000; Mesulam 2000; Ghashghaei &
Barbas 2002). The OFC plays a role in representing
reward and threat (e.g. Kringelbach & Rolls 2004), as
well as hedonic experience (Kringelbach 2005;
Wager et al. 2008), but it also plays a role in processing
olfactory, auditory and visual information (see fig. 6
in Kringelbach 2005; also see Price 2007). The OFC’s
ongoing integration of sensory information from the
external world with that from the body indicates that
conscious percepts are indeed intrinsically infused with
affective value, so that the affective salience or signifi-
cance of an object is not computed after the fact. As it
turns out, the OFC plays a crucial role in forming the
the hypothesis that the predictions generated during
object perception carry affective value as a necessary and
normal part of visual experience.
4. EVIDENCE FOR AFFECTIVE PREDICTIONS
To formulate the affective prediction hypothesis, it is
important to consider a more general aspect about the
way the brain seems to predict. There is accumulating
evidence that during object perception, the brain
quickly makes an initial prediction about the ‘gist’ of
the scene or object to which visual sensations refer
(Schyns & Oliva 1994; Bar et al. 2001, 2006; Oliva &
Torralba 2001; Torralba & Oliva 2003). Like a Dutch
artist from the sixteenth or seventeenth century, the
brain uses low spatial frequency visual information
available from the object in context to produce a
rough sketch, and then begins to fill in the details using
information from memory (for a review, see Bar 2004,
2007, 2009). Effectively, the brain is performing a
basic-level categorization that serves as a gist-level
prediction about the class to which the object belongs.
With back and forth between visual cortex and the
areas of the prefrontal cortex (via the direct projections
that connect them), a finer level of categorization is
achieved until a precise representation of the object is
finally constructed. Like an artist who successively
creates a representation of objects by applying smaller
and smaller pieces of paint to represent the light of
different colours and intensities, the brain gradually
adds high spatial frequency information until a specific
object is consciously seen.
As the brain generates its initial prediction about an
object, it uses information from the OFC, supporting
the hypothesis that affect is an ingredient of prediction.
1328 L. F. Barrett & M. Bar
Phil. Trans. R. Soc. B (2009)
Studies that precisely measure the timing of neuronal
activity indicate that information about an object is
instantly propagated to the front of the brain. OFC
activation has been observed between 80 and 130 ms
after stimulus onset (when objects are presented in
isolation; for a review, see Lamme & Roelfsema 2000;
Bullier 2001; also see Thorpe et al. 1983; Bar et al.
2006; Rudrauf et al. 2008). Two studies indicate that
this early activity is driven by low spatial frequency and
magnocellular visual input characteristic of early-stage
prediction (Bar et al. 2006; Kveraga et al. 2007a,b).
This propagation of neuronal firing is sometimes called
a ‘feed-forward sweep’ (Lamme & Roelfsema 2000),
where sensory information from the world is projected
rapidly from the back to the front of the brain after an
image is presented to the visual system (Bar 2003,
2007; Bar et al. 2006). Many of these same studies
show another wave of OFC activity between 200 and
450 ms, which might represent the refinement of initial
Similar findings are reported in the face perception
literature. Early evoked response potentials (ERPs) in
the frontal regions begin at approximately 80 ms, but
are typically observed between 120 and 180 ms after
stimulus onset (depending on whether the face is
presented foveally or parafoveally). These early com-
ponents reflect the categorization of the face as a face
(versus a non-face) or as generally affective (neutral
versus valenced), as valenced (e.g. happy versus sad) or
as portraying some degree of arousal (for reviews, see
Eimer & Holmes 2007; Palermo & Rhodes 2007;
Vuilleumier & Pourtois 2007). Later ERP components
between 200 and 350 ms correspond to the conscious
perception of fear, disgust, sadness and anger (for a
review see Barrett et al. (2007b)).
5. THE CIRCUITRY FOR SEEING WITH FEELING
Neuroanatomical evidence provides the strongest
support for the notion that affect informs visual
perception and allows the affective prediction hypo-
thesis to be further specified. The two functionally
related circuits within the OFC (figure 3; for reviews,
see Barbas & Pandya 1989; Carmichael & Price 1996;
O¨ngu ¨r & Price 2000; O¨ngu ¨r et al. 2003) are differen-
tially connected to the dorsal ‘where is it’ visual stream
and the ventral ‘what is it’ stream (figure 4), suggesting
two different roles for affect during object perception.
In the next section, we describe how medial parts of
the OFC are connected to the dorsal ‘where’ visual
stream, and help to produce the brain’s gist-level
prediction by providing initial affective information
about what an object might mean for a person’s well-
being. With gist-level visual information about an
object, the medial OFC initiates the internal bodily
changes that are needed to guide subsequent actions on
that object in context. The ability to reach for a round
object and pick it up for a bite depends on the
prediction that it is an apple and that it will enhance
one’s well-being in the immediate future because it has
been done so in the past.
While the medial OFC is directing the body to
prepare a physical response (or, what might be called
‘crafting an initial affective response’), the lateral parts
of the OFC are integrating the sensory feedback from
this bodily state with cues from the five external senses.
Based on the anatomical connections that we review in
§7, we hypothesize that the resulting multimodal
representation helps to create a unified experience of
specific objects in context. The ability to consciously
see a particular apple not only requires integration of
information from the five senses, but also requires that
this information is integrated with the ‘sixth sense’ of
affect (e.g., that such apples are delicious).
6. BASIC-LEVEL AFFECTIVE PREDICTIONS IN
The medial portions of the OFC guide autonomic,
endocrine and behavioural responses to an object
(Barbas & De Olmos 1990; Carmichael & Price
1995, 1996; O¨ngu ¨r et al. 1998; Rempel-Clower &
Barbas 1998; Ghashghaei & Barbas 2002; Barbas et al.
2003; Price 2007). The medial OFC has strong
reciprocal connections to the lateral parietal areas
(MT and MST) within the dorsal ‘where is it’ visual
stream (Barbas 1988, 1995; Carmichael & Price 1995;
Cavada et al. 2000; Kondo et al. 2003; figure 4). The
dorsal stream carries achromatic visual information of
low spatial and fast temporal resolution through
posterior parietal cortex, processing spatial information
and visual motion, and providing the basis for spatial
localization (Ungerleider & Mishkin 1982) and visually
guided action (Goodale & Milner 1992).
Through largely magnocellular pathways, the medial
OFC receives the same low spatial frequency visual
information (Kveraga et al. 2007a,b), devoid of specific
visual detail, that is used to create a basic-level category
representation about the object’s identity (Bar 2003,
2007). Through strong projections to hypothalamic,
midbrain, brainstem and spinal column control
centres, the medial OFC uses this low spatial frequency
visual information to modify the perceiver’s bodily state
to re-create the affective context in which the object
was experienced in the past (to allow subsequent
behaviour in the immediate future). Based on its
neuroanatomical connections to the lateral parietal
cortex, we hypothesize that the OFC’s representation
of these autonomic and endocrine changes is relayed
back to the dorsal ‘where is it’ stream as an initial
estimate of the affective value and motivational
relevance. With bidirectional processing between the
medial OFC and the lateral parietal cortex, a person’s
physiological and behavioural response is coordinated
with the information about the spatial location of the
respond to an object (based on this gist-level prediction)
even before the object is consciously perceived.
There is some additional neuroimaging evidence to
support our hypothesis that affective changes are part
of a basic gist prediction during object perception.
Areas in the medial OFC, including the ventromedial
prefrontal cortex (vmPFC) and the portion of the
rostral anterior cingulate cortex (ACC) beneath the
corpus callosum, typically show increased activity
during processing of contextual associations, where
an object triggers cortical representations of other
objects that have predicted relevance in a particular
L. F. Barrett & M. Bar 1329
Phil. Trans. R. Soc. B (2009)
situation (Bar & Aminoff 2003; Bar 2004). For
example, a picture of a traffic light activates visual
representations of other objects that typically share a
‘street’ context, such as cars, pedestrians and so on.
These findings suggest that an object has the capacity
to reinstate the context with which it has been
associated in prior experience. Given that the vmPFC
and rostral ACC project directly to autonomic and
endocrine output centres in the hypothalamus, mid-
brain, brainstem and spinal cord, it is likely that this
reinstatement includes reconstituting the internal
affective context that is associated with past exposures
to the object. The internal world of the body may
be one element in the ‘context frame’ that facilitates
object recognition (for a discussion of context frames,
see Bar 2004).
7. AFFECTIVE PREDICTIONS IN VISUAL
The lateral portions of the OFC receive information
about the nuanced body changes that occur as the
result of gist-level affective predictions (from vmPFC
and anterior insula). Lateral OFC then integrates this
bodily information with sensory information from the
world to establish an experience-dependent represen-
tation of an object in context. Many neurons within
the lateral OFC are multimodal and respond to a
variety of different sensory inputs (Kringelbach &
Rolls 2004). The lateral OFC has robust reciprocal
connections to the inferior temporal areas (TEO, TE
and temporal pole) of the ventral ‘what’ visual stream
(figure 4; Barbas 1988, 1995; Carmichael & Price
1995; Cavada et al. 2000; Rolls & Deco 2002; Kondo
et al. 2003). The ventral stream carries higher
resolution visual details (including colour) through
interior temporal cortex that give rise to the experi-
ence of seeing.
Through largely parvocellular pathways, the lateral
OFC receives this high spatial frequency visual
information full of rich detail about the visual features
of objects used to create a specific representation of an
object. The internal sensory information received from
the anterior insula is itself important for the conscious
experience of affect (Craig 2002, 2009). The resulting
multimodal representation then influences processing
in the ventral ‘what’ stream, and with additional back
and forth a specific, polysensory, contextually relevant
representation of the object is generated. This con-
scious percept includes the affective value of the object.
Sometimes, this value is represented as a property of
the object, and other times it is represented as a
person’s reaction to that object.
8. COORDINATING AFFECTIVE PREDICTIONS IN
On the basis of neuroanatomical evidence, we have,
thus far, proposed that affect plays two related roles in
object perception. The medial OFC estimates the value
of gist-level representations. A small, round, object
might be an apple if it is resting in a bowl on a kitchen
counter, associated with an unpleasant affective
response if the perceiver does not enjoy the taste of
apples, a pleasant affective response if he or she is an
apple lover and is hungry, and even no real affective
change in an apple lover who is already satiated. The
medial OFC not only realizes the affective significance
of the apple but also prepares the perceiver to act—to
turn away from the apple, to pick it up and bite, or to
ignore it, respectively. The lateral OFC integrates
sensory information from this bodily context with
information from other sensory modalities, as well as
more detailed visual information, producing the visual
experience of a specific apple, ball or balloon. These
two aspects of affective prediction do not occur in
stages per se, but there might be slight differences in the
time at which the two are computed.
Autonomic, hormonal or muscular changes in the
body that are generated as part of a gist-level
prediction via the medial OFC might be initiated
before and incorporated into the multimodal rep-
resentation of the world that is represented in the
lateral OFC. Visual information arrives more quickly
to the medial OFC owing to a ‘magnocellular
advantage’ in visual processing (term by Laylock
et al. 2007). Magnocellular neurons projecting from
the lateral geniculate nucleus (in the thalamus) rapidly
conduct low spatial frequency visual information to V1
and the dorsal ‘where’ stream areas (compared with
the parvocellular neurons that carry highly specific and
detailed visual information to V1 and to the ventral
‘what’ stream). In humans, magnocellular neurons in
V1 fire from 25 ms (Klistorner et al. 1997) to 40 ms
(Paulus et al. 1999) earlier than parvocellular neurons in
neurons within the dorsal stream that receive input
directly fromthe lateral geniculatenucleus (e.g.V5/MT;
Sincich et al. 2004) become active even before V1
(Ffytche et al. 1995; Buchner et al. 1997). As a result,
low spatial frequency visual information about an object
arrives to the medial OFC before high spatial frequency
visual information arrives to the lateral OFC. Consistent
in the prefrontal cortex became active at approximately
10 ms after neurons in the dorsal ‘where’ stream,
but coincident with the activation in the ventral
A magnocelluar advantage extending to the medial
OFC would help to resolve the debate over how the
brain processes affective value of objects and faces
when they are unattended or presented outside of
visual awareness (either because of experimental
procedures or brain damage). Some researchers argue
for a subcortical route by which low spatial frequency
visual information about objects and faces can bypass
V1 via the amygdala to represent affective value in the
body and behaviour (e.g. LeDoux 1996, 2000;
Weiskrantz 1996; Morris et al. 1998; Catani et al.
2003; Rudrauf et al. 2008), whereas others argue that
such a circuit is not functional in primates (Rolls 2000;
Pessoa & Ungerleider 2004). Either way, affective
predictions do not necessarily require a purely sub-
cortical route. Nor is a subcortical route necessary to
explain how objects presented outside of visual
awareness influence the body and behaviour, how
blindsighted patients can respond to the affective tone
of a stimulus despite damage to V1, or why patients
with amygdala damage have deficits in face perception.
1330 L. F. Barrett & M. Bar
Phil. Trans. R. Soc. B (2009)
And because OFC lesions have been linked to memory
deficits (for a discussion, see Frey & Petrides 2000), the
absence of a magnocellular advantage may also help to
explain why people suffering from agnosia can experi-
ence deficits in affective responses to visual stimulation
but not to other sensory stimuli (Bauer 1982; Damasio
et al. 1982; Habib 1986; Sierra et al. 2002). The
amygdala’s importance in object and face processing
(for a review, see Vuilleumier & Pourtois 2007) may
refer as much (if not more) to its participation in
affective predictions than to its ability to work around
early cortical processing in perception. These findings
also suggest that the enhanced activity observed in
fusiform gyrus in response to unattended emotional
faces (e.g. Vuilleumeir et al. 2001) may be influenced,
in part, by the OFC (which is strongly connected to the
amygdala) rather than to the amygdala per se.
That being said, there is other evidence to suggest
that both components of affective prediction happen
more or less simultaneously. There is something like a
parvocellular advantage in visual processing, in that
visual information reaches the ventral ‘what’ stream
very quickly, and like the input to the dorsal ‘where’
stream, arrives without the benefit of cortical proces-
sing in early visual cortex. The lateral geniculate
nucleus not only projects directly to the dorsal stream
but also appears to project directly to the anterior
temporal lobe areas that are connected to the ventral
stream (including the parahippocampal gyrus and
amygdala; Catani et al. 2003). As a consequence,
upon the presentation of a stimulus, some of the
neurons in the anterior temporal cortex fire almost
coincidently with those in the occipital cortex (e.g. 47
versus 45 ms, respectively; Wilson et al. 1983). Without
further study, however, it is difficult to say whether
these are magno- or parvocellular connections.
Taken together, these findings indicate that it maybe
more appropriate to describe the affective predictions
generated by the medial and lateral OFC as phases in a
single affective prediction evolving over time, rather
than as two separate ‘types’ of affective predictions
(with one informing the other). This interpretation is
supported by the observation that the medial and
lateral OFC are strongly connected by intermediate
areas (figure 3); in addition, the lateral OFC receives
some low spatial frequency visual information and the
medial OFC some high spatial frequency information;
and, magnocellular and parvocellular projections are
not as strongly anatomically segregated as was first
believed (for a review, see Laylock et al. (2007)).
Furthermore, there are strong connections throughout
the dorsal ‘where’ and ventral ‘what’ streams at all
levels of processing (Merigan & Maunsell 1993; Chen
et al. 2007). Finally, the OFC has widespread
connections to a variety of thalamic nuclei that receive
highly processed visual input and therefore can not be
treated as solely bottom-up structures in visual
processing.1As a result of these interconnections, the
timing differences in the medial and lateral OFC
affective predictions will be small and perhaps difficult
to measure with the current technology, even if they
prove to be functionally significant in the time course of
A tremendous amount of research has now established
that object recognition is a complex process that relies
on many different sources of information from the
world (e.g. contrast, colour, texture, low spatial
frequency cues). In this paper, we suggest that object
recognition uses another source of information:
sensory cues from the body that represent the object’s
value in a particular context. We have laid the
foundation for the hypothesis that people do not wait
to evaluate an object for its personal significance until
after they know what the object is. Rather, an affective
reaction is one component of the prediction that helps a
person see the object in the first place. Specifically, we
hypothesize that very shortly after being exposed to
objects, the brain predicts their value for the person’s
well-being based on prior experiences with those
objects, and these affective representations shape the
person’s visual experience and guide action in the
immediate future. When the brain effortlessly guesses
an object’s identity, that guess is partially based on how
the person feels.
Our ideas about affective prediction suggest that
people do not come to know the world exclusively
through their senses; rather, their affective states
influence the processing of sensory stimulation from
the very moment an object is encountered. These ideas
also suggest the intriguing possibility that exposure to
visual sensations alone is not sufficient for visual
experience. And even more exciting, plasticity in visual
cortex areas might require, at least to some extent, the
formation of connections between visual processing
areas in the back of the brain and affective circuitry in
the front. This suggests that affective predictions may
not be produced by feedback from the OFC alone. As
unlikely as it may seem, affective predictions might also
influence plasticity in the visual system so that visual
processing is changed from the bottom-up.
This, of course, brings us back to Mr May. Even
several years after his surgery, Mr May’s brain did not
have a sufficiently complex and nuanced cache of
multimodal representations involving visual sensations
to allow him to easily predict the meaning of novel
input from the visualworldaround him. Said adifferent
way, his paucity of knowledge about the visual world
forced him to think through every guess. Our
hypothesis is that the guessing became easier, and
more automatic, as Mr May’s visual sensations took on
affective value. He went from having what the
philosopher Humphrey (2006) called ‘affectless vision’
(p. 67) to seeing with feeling.
We have proposed that a person’s affective state has a
top-down influence in normal object perception.
Specifically, we have proposed that the medial OFC
participates in an initial phase of affective prediction
(‘what is the relevance of this class of objects for me’),
whereas the lateral OFC provides more subordinate-
level and contextually relevant affective prediction
(‘what is the relevance of this particular object in this
particular context for me at this particular moment in
time’). If this view is correct, then personal relevance
L. F. Barrett & M. Bar1331
Phil. Trans. R. Soc. B (2009)
and salience are not computed after an object is already
identified, but may be part of object perception itself.
Deep thanks to Michael May for sharing his thoughts and
experiences. We also thank Eliza Bliss-Moreau, Krysal Yu
and Jasmine Boshyan who helped with the preparation of
figures. Thanks to Daniel Gilbert and the members of the
Barrett and Bar laboratories who made helpful comments on
the previous drafts of this manuscript. Preparation of this
paper was supported by NIH grant R01NS050615 to M.B.,
and a National Institutes of Health Director’s Pioneer Award
(DP1OD003312), grants from the National Institute of
Aging (AG030311) and the National Science Foundation
(BCS 0721260; BCS 0527440) and a contract with the
Army Research Institute (W91WAW), as well as by a Cattell
Award and a fellowship from the American Philosophical
Society to L.F.B.
1For example, the midbrain’s superior colliculus and thalamic nuclei
such as the pulvinar and mediodorsal receive cortically processed
visual input from V1, the ventral visual stream (area IT) and the
sensory integration network in the OFC(Abramson & Chalupa1985;
Casanova 1993; Webster et al. 1993, 1995).
Abramson, B. P. & Chalupa, L. M. 1985 The laminar
distribution of cortical connections with the tecto- and
cortico-recipient zones in the cat’s lateral posterior
nucleus. Neuroscience 15, 81–95. (doi:10.1016/0306-
Arnold, M. B. 1960 Emotion and personality. New York, NY:
Columbia University Press.
Bar, M. 2003 A cortical mechanism for triggering top-down
facilitation in visual object recognition. J. Cogn. Neurosci.
15, 600–609. (doi:10.1162/089892903321662976)
Bar, M. 2004 Visual objects in context. Nat. Rev. Neurosci 5,
Bar, M. 2007 The proactive brain: using analogies and
associations to generate predictions. Trends Cogn. Sci. 11,
Bar, M. & Aminoff, E. 2003 Cortical analysis of visual
context. Neuron 38, 347–358. (doi:1016/S0896-6273(03)
& Mendola, J. D. 2001 Cortical mechanisms specific
to explicit visual object recognition. Neuron 29, 529–535.
Bar, M. et al. 2006 Top-down facilitation of visual
recognition. Proc. Natl Acad. Sci. USA 103, 449–454.
Bar, M. 2009 The proactive brain: memory for predictions.
Phil. Trans. R. Soc. B 364, 1235–1243. (doi:10.1098/rstb.
Barbas, H. 1988 Anatomic organization of basoventral and
mediodorsal visual recipient prefrontal regions in the
rhesus monkey. J. Comp. Neurol. 276, 313–342. (doi:10.
Barbas, H. 1993 Organization of cortical afferent input to
orbitofrontal areas in the rhesus monkey. Neuroscience 56,
Barbas, H. 1995 Anatomic basis of cognitive-emotional
interactions in the primate prefrontal cortex. Neurosci.
Biobehav. Rev. 19, 499–510. (doi:10.1016/0149-7634(94)
Barbas, H. 2000 Connections underlying the synthesis of
cognition, memory, and emotion in primate prefrontal
cortices. Brain Res. Bull. 52, 319–330. (doi:10.1016/
Barbas, H. & De Olmos, J. 1990 Projections from the
amygdala to basoventral and mediodorsal prefrontal
regions in the rhesus monkey. J. Comp. Neurol. 300,
Barbas, H. & Pandya, D. N. 1989 Architecture and intrinsic
connections of the prefrontal cortex in the rhesus
monkey. J. Comp. Neurol. 286, 353–375. (doi:10.1002/
Barbas, H., Saha, S., Rempel-Clower, N. & Ghashghaei, T.
2003 Serial pathways from primate prefrontal cortex to
autonomic areas may influence emotional expression.
BMC Neurosci. 4, 25–37. (doi:10.1186/1471-2202-4-25)
Barrett, L. F. 2006a Solving the emotion paradox: categor-
ization and the experience of emotion. Pers. Soc. Psychol.
Rev. 10, 20–46. (doi:10.1207/s15327957pspr1001_2)
Barrett, L. F. 2006b Valence as a basic building block of
emotional life. J. Res. Pers. 40, 35–55. (doi:10.1016/j.jrp.
Barrett, L. F. & Bliss-Moreu, E. In press. Affect as a
psychological primitive. Adv. Exp. Social Psychol.
Barrett, L. F., Quigley, K., Bliss-Moreau, E. & Aronson,
K. R. 2004 Arousal focus and interoceptive sensitivity.
J. Person. Social Psychol. 87, 684–697.
The experience of emotion. Annu. Rev. Psychol. 58,
Barrett, L. F., Lindquist, K. & Gendron, M. 2007b Language
as a context for emotion perception. Trends Cogn. Sci 11,
Bauer, R. M. 1982 Visual hypoemotionality as a symptom of
visual-limbic disconnection in man. Arch. Neurol. 39,
Berridge, K. C. & Winkielman, P. 2003 What is an
unconscious emotion? The case for unconscious ‘liking’.
Bliss-Moreau, E., Barrett, L. F. & Wright, C. I. 2008
Individual differences in learning the affective value of
others under minimal conditions. Emotion 8, 479–493.
Bliss-Moreau, E., Barrett, L. F. & Owren, M. Submitted.
I like the sound of your voice: affective learning about
the human voice.
Buchner, H., Gobbele, R., Wagner, M., Fuchs, M.,
Waberski, T. D. & Beckmann, R. 1997 Fast visual evoked
potential input into human area V5. Neuroreport 8,
Bullier, J. 2001 Integrated model of visual processing.
Brain Res. Rev. 36, 96–107. (doi:10.1016/S0165-0173
Carmichael, S. T. & Price, J. L. 1995 Limbic connections of
the orbital and medial prefrontal cortex in macaque
monkeys. J. Comp. Neurol. 363, 615–641. (doi:10.1002/
Carmichael, S. T. & Price, J. L. 1996 Connectional networks
within the orbital and medial prefrontal cortex of macaque
monkeys. J. Comp. Neurol. 371, 179–207. (doi:10.1002/
Casanova, C. 1993 Response properties of neurons in area 17
projecting to the striate-recipient zone of the cat’s lateralis
posterior-pulvinar complex: comparison with cortico-
tectal cells. Exp. Brain Res. 96, 247–259. (doi:10.1007/
1332 L. F. Barrett & M. Bar
Phil. Trans. R. Soc. B (2009)
Catani, M., Jones, D. K., Donato, R. & Ffytche, D. H. 2003
Occipito-temporal connections in the human brain. Brain
126, 2093–2107. (doi:10.1093/brain/awg203)
Cavada, C., Company, T., Tejedor, J., Cruz-Rizzolo, R. J. &
Reinsos-Suarez, F. 2000 The anatomical connections of
the macaque monkey orbitofrontal cortex: a review. Cereb.
Cortex 10, 220–242. (doi:10.1093/cercor/10.3.220)
Chen, C. M., Lakatos, P., Shah, A. S., Mehta, A. D., Givre,
S. J., Javitt, D. C. & Schroeder, C. E. 2007 Functional
anatomy and interaction of fast and slow visual pathways
in macaque monkeys. Cereb. Cortex 17, 1561–1569.
Clore, G. L. & Storbeck, J. 2006 Affect as information about
liking, efficacy, and importance. In Hearts and minds:
affective influences on social cognition and behaviour (ed.
J. Forgas), pp. 123–142. New York, NY: Psychology Press.
Craig, A. D. 2002 Opinion: how do you feel? Interoception:
the sense of the physiological condition of the body. Nat.
Rev. Neurosci. 3, 655–666.
Craig, A. D. 2009 How do you feel—now? The anterior
insula and human awareness. Nat. Rev. Neurosci. 10,
Damasio, A. 1999 The feeling of what happens: body and
emotion in the making of consciousness. Fort Worth, TX:
Damasio, A. R., Damasio, H. & Van Hoesen, G. W. 1982
Prosopagnosia: antomic basis and behavioral mechanisms.
Am. Acad. Neurol. 32, 331–341.
DeArmond, S. J., Fusco, M. M. & Dewey, M. M. 1989
Structure of the human brain: a photographic atlas, 3rd edn.
New York, NY: New York University Press.
Duncan, S. & Barrett, L. F. 2007 Affect as a form of
cognition: a neurobiological analysis. Cogn. Emotion 21,
Edelman, G.M. 1987 Neural Darwinism: the theoryof neuronal
group selection. New York, NY: Basic Books Inc.
Edelman, G. M. & Tononi, G. 2000 A universe of
consciousness: how matter becomes imagination. New York,
NY: Basic Books.
Eimer, M. & Holmes, A. 2007 Event-related brain potential
Ffytche, D. H., Guy, C. N. & Zeki, S. 1995 The parallel visual
motion inputs into areas V1 and V5 of human cerebral
cortex. Brain 118, 1375–1394. (doi:10.1093/brain/118.
Fine, I., Wade, A. R., Brewer, A. A., May, M. G., Goodman,
D. F., Boynton, G. M., Wndell, B. A. & MacLeod, D. I. A.
2003 Long-term deprivation affects visual perception and
cortex. Nat. Neurosci. 6, 915–916. (doi:10.1038/nn1102)
Foxe, J. J. & Simpson, G. V. 2002 Flow of activation from V1
to frontal cortex in humans. A framework for defining
“early” visual processing. Exp. Brain Res. 142, 139–150.
Frey, S. & Petrides, M. 2000 Orbitofrontal cortex: a key
prefrontal region for encoding information. Proc. Natl
Acad. Sci. USA 97, 8723–8727. (doi:10.1073/pnas.
Ghashghaei, H. T. & Barbas, H. 2002 Pathways for emotion:
interactions of prefrontal and anterior temporal pathways
in the amygdala of the rhesus monkey. Neuroscience 115,
Goodale, M. A. & Milner, A. D. 1992 Separate visual
pathways for perception and action. Trends Neurosci. 15,
Habib, M. 1986 Visual hypo-emotionality and prosopagnosia
associated with right temporal lobe isolation. Neuropsycho-
logia 24, 577–582. (doi:10.1016/0028-3932(86)90101-6)
Humphrey, N. 2006 Seeing red: a study in consciousness.
Cambridge, MA: Harvard University Press.
James, W. 1890 The principals of psychology. New York, NY:
Klistorner, A., Crewther, D. P. & Crewther, S. G. 1997
Separate magnocellular and parvocellular contributions
from temporal analysis of the multifocal VEP. Vision Res.
37, 2161–2169. (doi:10.1016/S0042-6989(97)00003-5)
Koenigs, M. & Tranel, D. 2008 Prefrontal cortex damage
abolishes brand-cued changes in cola preference. Soc.
Cogn. Affect. Neurosci. 3, 1–6. (doi:10.1093/scan/nsm032)
Kondo, H., Saleem, K. S. & Price, J. L. 2003 Differential
connections of the temporal pole with the orbital and
medial prefrontal networks in macaque monkeys. J. Comp.
Neurol. 465, 499–523. (doi:10.1002/cne.10842)
Kringelbach, M. L. 2005 Linking reward to hedonic
experience. Nat. Rev. Neurosci. 6, 691–702. (doi:10.
Kringelbach, M. L. & Rolls, E. T. 2004 The functional
neuroanatomy of the human orbitofrontal cortex: evidence
from neuroimaging and neuropsychology. Prog. Neurobiol.
72, 341–372. (doi:10.1016/j.pneurobio.2004.03.006)
Kveraga, K., Boshyan, J. & Bar, M. 2007a Magnocellular
projections as the trigger of top-down facilitation in
recognition. J. Neurosci. 27, 13 232–13 240. (doi:10.
Kveraga, K., Ghuman, A. S. & Bar, M. 2007b Top-down
predictions in the cognitive brain. Brain Cogn. 65,
Lamme, V. A. & Roelfsema, P. R. 2000 The distinct modes of
vision offered by feedforward and recurrent processing.
Laylock, R., Crewther, S. G. & Crewther, D. P. 2007 A role
for the “magnocellular advantage” in visual impairments
Neurosci. Biobehav. Rev. 31, 363–376. (doi:10.1016/
LeDoux, J. E. 1996. The emotional brain. New York, NY:
Simon & Schuster.
LeDoux, J. E. 2000 Emotion circuits in the brain. Annu.
Rev. Neurosci. 23, 155–184. (doi:10.1146/annurev.neuro.
Li, W., Moallem, I., Paller, K. A. & Gottfried, J. A. 2007
Subliminal smells can guide social preferences. Psychol.
18, 1044–1049. (doi:10.1111/j.1467-9280.2007.
McDonald, A. J. 1998 Cortical pathways to the mammalian
amygdala. Prog. Neurobiol. 55, 257–332. (doi:10.1016/
Merigan, W. H. & Maunsell, J. H. R. 1993 How parallel are
the primate visual pathways? Annu. Rev. Neurosci. 16,
Mesulam, M. 2000 Behavioral neuroanatomy: large-scale
networks, association cortex, frontal syndromes, the
limbic system,and hemispheric
Principles of behavioral and cognitive neurology (ed. M.
Mesulam), pp. 1–120. 2nd edn. New York, NY: Oxford
Morris, J. S., Ohman, A. & Dolan, R. J. 1998 A subcortical
pathway to the right amygdala mediating “unseen” fear.
Proc. Natl Acad. Sci. USA 96, 1680–1685. (doi:10.1073/
Oliva, A. & Torralba, A. 2001 Modeling the shape of a scene:
a holistic representation of the spatial envelope. Int.
J. Comput. Vision 42, 145–175. (doi:10.1023/A:1011139
O¨ngu ¨r, D. & Price, J. L. 2000 The organization of networks
within the orbital and medial prefrontal cortex of rats,
monkeys and humans. Cereb. Cortex 10, 206–219. (doi:10.
L. F. Barrett & M. Bar 1333
Phil. Trans. R. Soc. B (2009)
O¨ngu ¨r, D., An, X. & Price, J. L. 1998 Prefrontal cortical Download full-text
projections to the hypothalamus in macaque monkeys.
J. Comp. Neurol. 401, 480–505. (doi:10.1002/(SICI)1096-
O¨ngu ¨r, D., Ferry, A. T. & Price, J. L. 2003 Architectonic
subdivision of the human orbital and medial prefrontal
cortex. J. Comp. Neurol. 460, 425–449. (doi:10.1002/cne.
Palermo, R. & Rhodes, G. 2007 Are you always on my mind?
A review of how face perception and attention interact.
Neuropsychologia 45, 75–92. (doi:10.1016/j.neuropsycho-
Paulus, W., Korinth, S., Wischer, S. & Tergau, F. 1999
Differential inhibition of chromatic and achromatic
perception by transcranial magnetic stimulation of the
human visual cortex. Neuroreport 10, 1245–1248. (doi:10.
Pessoa, L. 2008 On the relationship between emotion and
cognition. Nat. Rev. Neurosci. 2, 148–158. (doi:10.1038/
Pessoa, L. & Ungerleider, L. G. 2004 Neuroimaging studies
of attention and the processing of emotion-laden stimuli.
Prog. Brain Res. 144, 171–182. (doi:10.1016/S0079-
Price, J. L. 2007 Connections of orbital cortex. In
The orbitofrontal cortex (eds D. H. Zald & S. L. Rauch),
pp. 38–56. New York, NY: Oxford University Press.
Rempel-Clower, N. L. & Barbas, H. 1998 Topographic
organization of connections between the hypothalamus
and prefrontal cortex in the rhesus monkey. J. Comp.
Neurol. 398, 393–419. (doi:10.1002/(SICI)1096-9861
Rolls, E. T. 2000 Pre ´cis of “The Brain and Emotion”. Behav.
Brain Sci. 23, 177–191. (doi:10.1017/S0140525X00
Rolls, E. T. & Deco, G. 2002. Computational neuroscience of
vision. Oxford, UK: Oxford University Press.
Rudrauf, D., David, O., Lachaux, J.-P., Kovach, C. K.,
Martinerie, J., Renault, B. & Damasio, A. 2008 Rapid
interactions between the ventral visual stream and
emotion-related structures rely on a two-pathway archi-
Russell, J. A. & Barrett, L. F. 1999 Core affect, prototypical
emotional episodes, and other things called emotion:
dissecting the elephant. J. Pers. Soc. Psychol. 76, 805–819.
Schyns, P. G. & Oliva, A. 1994 From blobs to boundary
edges: evidence for time- and spatial-dependent scene
recognition. Psychol. Sci. 5, 195–200. (doi:10.1111/
Searle, J. 1992 The rediscovery of the mind. Cambridge, MA:
Searle, J. 2004 Mind: a brief introduction. New York, NY:
Cambridge University Press.
Sierra, M., Lopera, F., Lambert, M. V., Phillips, M. L. &
David, A. S. 2002 Separating depersonalisation and
derealisation: the relevance of the ‘lesion method’.
J. Neurol. Neurosurg. Psychiatry 72, 530–532.
Sincich, L. C., Park, K. F., Wohlgemuth, M. J. & Horton,
J. C. 2004 Bypassing V1: a direct geniculate input to area
MT. Nat. Neurosci. 7, 1123–1128. (doi:10.1038/nn1318)
Spencer, H. 1855 Principles of psychology. London, UK:
Stefanacci, L. & Amaral, D. G. 2002 Some observations on
cortical inputs to the macaque monkey amygdala: an
anterograde tracing study. J. Comp. Neurol. 451, 301–323.
Sully, J. 1892. Outlines of psychology, vol. 2. New York, NY:
Appleton and Company.
Torralba, A. & Oliva, A. 2003 Statistics of natural image
categories. Network 14, 391–412.
Thorpe, S. J., Rolls, E. T. & Maddison, S. 1983 The
orbitofrontal cortex: neuronal activity in the behaving
monkey. Exp. Brain Res. 49, 93–115. (doi:10.1007/
Ungerleider, L. G. & Mishkin, M. 1982 Two cortical visual
systems. In Analysis of visual behavior (eds D. A. Ingle,
M. A. Goodale & R. J. W. Mansfield), pp. 549–586.
Cambridge, MA: MIT Press.
Vuilleumier, P. & Pourtois, G. 2007 Distributed and
interactive brain mechanisms during emotion face percep-
tion: evidence from functional neuroimaging. Neuropsy-
chologia 45, 174–194. (doi:10.1016/j.neuropsychologia.
Vuilleumier, P., Armony, J. L., Driver, J. & Dolan, R. J. 2001
Effects of attention and emotion on face processing in the
human brain: an event-related fMRI study. Neuron 30,
Wager, T. D., Barrett, L. F., Bliss-Moreau, E., Lindquist, K.,
Duncan, S., Kober, H., Joseph, J., Davidson, M. &
Mize, J. 2008 The neuroimaging of emotion. In The
handbook of emotion (eds. M. Lewis, J. M. Haviland-Jones,
& L. F. Barrett), pp. 249–271, 3rd edn. New York, NY:
Webster, M. J., Bachevalier, J. & Ungerleider, L. G. 1993
Subcortical connections of inferior temporal areas TE and
TEO in macaque monkeys. J. Comp. Neurol. 335, 73–91.
Webster, M. J., Bachevalier, J. & Ungerleider, L. G. 1995
Transient subcortical connections of inferior temporal
areas TE and TEO in infant macaque monkeys. J. Comp.
Neurol. 352, 213–226. (doi:10.1002/cne.903520205)
6, 215–220. (doi:10.1016/S0959-4388(96)80075-4)
Wilson, C. L., Babb, T. L., Halgren, E. & Crandall, P. H.
1983 Visual receptive fields and response properties of
neurons in human temporal lobe and visual pathways.
Brain 106, 473–502. (doi:10.1093/brain/106.2.473)
Winkielman, P., Berridge, K. C. & Wilburger, J. L. 2005
Unconscious affective reactions to masked happy versus
angry faces influence consumption behavior and judge-
ments of value. Pers. Social Psychol. Bull. 1, 121–135.
Wundt, W. 1998 Outlines of psychology. Bristol, UK:
Thoemmes Press. (Trans L. C. H. Judd) (Original work
Zajonc, R. B. 1980 Feeling and thinking: preferences need no
inferences. Am. Psychol. 35, 151–175. (doi:10.1037/0003-
1334 L. F. Barrett & M. Bar
Phil. Trans. R. Soc. B (2009)