THE YEAR IN COGNITIVE NEUROSCIENCE 2009

Attention, Automaticity, and Awareness

in Synesthesia
Jason B. Mattingley
The University of Queensland, Australia

The phenomenon of synesthesia has occupied the thoughts of philosophers and artists
for decades. With the advent modern behavioral and brain imaging techniques, scientific
research on synesthesia has also moved into the mainstream of thought. Here I provide
a cognitive neuroscience perspective on the condition, with a particular emphasis on
grapheme-color synesthesia, the most common variant, in which individuals report vivid
and consistent experiences of color in association with numerals, letters, and words.
Behavioral studies have revealed several fundamental properties of induced synesthetic
colors. First, although they seem to arise automatically, without the need for voluntary
control, they are strongly modulated by selective attention. Second, they attain salience
relatively early in visual processing, and so can influence perceptual judgments and
guide focal attention in cluttered, achromatic displays. Third, brain activity during
synesthetic color experiences arises from within the ventral temporal lobe, including
color-selective area V4. It has been speculated that grapheme-color synesthesia arises
from disinhibited feedback or abnormal cross-wiring between brain regions involved
in extracting visual form and color.

Key words: attention; blood oxygen level-dependent (BOLD); color perception;

grapheme; event-related potential (ERP); functional magnetic resonance imaging
(fMRI); masking; Stroop effect; transcranial magnetic stimulation (TMS); visual im-
agery; visual search; V4

Introduction the phenomenon on television, radio, and in

The last two decades have seen a resurgence
of interest in the phenomenon of synesthesia
(Mattingley & Ward 2006; Rich & Mattingley
2002; Robertson & Sagiv 2005), a condition
in which individuals report unusual sensory,
cognitive, or affective experiences associated
with seen, heard, felt, smelled, or tasted stim-
uli. In the last two years alone there have been
more than 100 scientific papers on synesthesia
in the scientific literature (ISI Web of Science,
October 2008). Roughly the same number of
works again have appeared in the arts and hu-
manities (e.g., music, literature, creative arts),
and there have been countless stories about
Address for correspondence: Jason B. Mattingley, The University of
Queensland, Queensland Brain Institute and School of Psychology, St
Lucia, Queensland 4072, Australia. Voice: +61 7 3346 6331, fax. +61 7
3346 6301. j.mattingley@uq.edu.au
The Year in Cognitive Neuroscience 2009: Ann. N.Y. Acad. Sci. 1156: 141–167 (2009).
doi: 10.1111/j.1749-6632.2009.04422.x  C 2009 New York Academy of Sciences.
the print media worldwide. Clearly the condi-
tion has captured the imagination of scientists,
artists and the lay public alike, and perhaps not
surprisingly. There is something intriguing and
slightly unnerving about the idea that there is a
significant minority of people, individuals with
whom we live or work, whose sensory, cognitive
and emotional responses differ fundamentally
from the way most of us—that is to say, those
of us who are not synesthetes—experience the
world.
In this article I review evidence that bears
upon two central questions in research on
synesthesia. First, at what level does synesthe-
sia emerge in the perceptual processing hier-
archy, from the initial registration of inputs at
the sensory epithelia to the extraction of mean-
ing and the emergence of conscious experi-
ence? Second, what are the neurophysiolog-
ical correlates of synesthetic experience, and

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what light is shed on the phenomenon by the
application of techniques such as functional
brain imaging, electroencephalography, and
neurostimulation? Given the vast and grow-
ing literature in the area, I shall deliberately
focus on research that bears more or less di-
rectly on these two central questions. I shall also
focus almost exclusively on data pertaining to
so-called grapheme-color synesthesia, in which
individuals report vivid experiences of color
in association with graphemes (digits, letters,
and words in their printed form) (Baron-Cohen
et al. 1987; Galton 1880; Grossenbacher &
Lovelace 2001; Smilek et al. 2007; Ward et al.
2007). There is now good evidence that the
grapheme-color variant is by far the most com-
mon form of synesthesia, accounting for up to
90% of all self-reported cases (Barnett et al.
2008; Rich et al. 2005; Simner et al. 2006b).
I will not discuss recent work on the heritabil-
ity of synesthesia (Barnett et al. 2008; Baron-
Cohen et al. 1996; Rich et al. 2005; Simner
et al. 2006b; Smilek et al. 2005), or on the hy-
pothesized developmental determinants of the
condition (Marks & Odgaard 2005; Maurer &
Mondloch 2005, 2006), nor will I deal with
evidence concerning the causal link between
language and synesthetic experiences (Simner
2007; Simner et al. 2006a; Simner & Ward
2006; Ward et al. 2005). Readers interested in
learning more about these important topics are
referred to recent reviews by Hochel and Milán
(2008), Rich and Mattingley (2002), Robertson
(2003), and Simner (2007).
What Is Synesthesia?
It has proven remarkably difficult to arrive at
a definition of synesthesia that is both precise
and widely accepted. The word comes from
the Greek syn- meaning “together” and aisthe-
meaning “to feel, perceive” or “of the senses.”
The definition I offered at the outset (“a con-
dition in which individuals report unusual sen-
sory, cognitive, or affective experiences associ-
ated with seen, heard, felt, smelled or tasted
Annals of the New York Academy of Sciences
stimuli”) is perhaps not sufficiently precise,
but it is intended to capture the heterogene-
ity of synesthetic phenomena that have been
reported. The number of subtypes of the con-
dition seems virtually limitless (Grossenbacher
& Lovelace 2001; Rich et al. 2005).
Several studies have sought to catalogue
the various stimuli that trigger synesthesia
(so-called inducers) as well as the types of
sensory, cognitive, or emotional experiences
that are elicited by them (“concurrents”)
(Grossenbacher & Lovelace 2001). By far the
most common inducers, accounting for 80–
90% of all cases, are digits, letters, and words,
either heard, read, or evoked internally via im-
agery or thought (Barnett et al. 2008; Rich et al.
2005). On the other hand, synesthetic concur-
rents almost invariably involve color (around
90% of all cases, Barnett et al. 2008; Rich
et al. 2005; Simner et al. 2006b), either ex-
perienced “in the mind’s eye” or externalized
onto the body or into peripersonal or extrap-
ersonal space (Ward et al. 2007). Less com-
monly, individuals may report colors for music,
odors, tastes, or experiences of intense plea-
sure or pain (Rich et al. 2005). At a conser-
vative estimate there are probably more than
50 individual variants in the published litera-
ture. It is noteworthy, however, that a recent
study of 500 individuals recruited opportunisti-
cally from two universities in Scotland revealed
just nine different manifestations of synesthe-
sia, representing only 7% of the total possible
combinations of inducer and concurrent tested
for (Simner et al. 2006b). In that study the most
common manifestation of synesthesia was the
grapheme-color variant, which occurred in 1%
of a further opportunistic sample of more than
1,000 adults and children (Simner et al. 2006b).
Just as the types of inducers and concur-
rents vary considerably across individuals, so it
seems can the manner in which synesthetic con-
currents are experienced. Individuals with the
grapheme-color variant might perceive their
synesthetic colors as a “mist” or “haze” floating
out in space, some see their colors as hovering
above a printed character, or perhaps attached
Mattingley: Attention, Automaticity and Awareness in Synesthesia
to its surface, and many experience colors in
their mind’s eye as a kind of vivid mental image.
Considerable recent interest has focused on
whether these variations in subjective reports
might reflect fundamentally different types of
synesthesia, with distinct cognitive and neural
substrates. According to Dixon et al. (2004), in-
dividuals with grapheme-color synesthesia can
be divided into two distinct groups: “projec-
tors” and “associators.” Projectors, according
to their hypothesis, are those individuals who
experience synesthetic colors as “projected” out
beyond the body and into external space. By
contrast, associators are those who experience
synesthetic colors “internally,” or in the mind’s
eye, as a kind of mental image.
Note that the dichotomy suggested by
Dixon et al. (2004) concerns the manner
in which synesthetic concurrents are expe-
rienced. An analogous framework, suggested
by Ramachandran and Hubbard (2001a), has
been advanced for the way in which inducers
trigger synesthetic experiences. According to
this model the color experiences of “higher”
synesthetes are triggered at a conceptual or se-
mantic level of analysis. Thus, a person might
experience the same color green for the num-
ber “5,” the Roman numeral “V,” and a cluster
of five dots; or colors might be induced specif-
ically for ordinal sequences, such as days of
the week or months of the year (Rich et al.
2005). By contrast, the colors experienced by
“lower” synesthetes are assumed to arise from
an analysis of the physical form of the stimulus,
which might arise initially from visual process-
ing within the grapheme area of the ventral
occipito−temporal cortex (Ramachandran &
Hubbard 2001b).
Both the “projector−associator” dichotomy
and the “lower−higher” dichotomy have re-
ceived empirical support from behavioral and
brain imaging studies (Dixon et al. 2004;
Hubbard et al. 2005; Ramachandran & Hub-
bard 2001b), as discussed in the following sec-
tions. It should be noted, however, that not all
authors have agreed with these distinctions, at
least as they were originally framed by their
143
proponents (see Edquist et al. 2006; Ward et al.
2007). Moreover, most authors who have di-
vided their synesthete groups into projectors
and associators have done so on the basis of
subjective reports rather than empirical data,
and so these reports must be considered with a
degree of caution.
Despite the evident heterogeneity of synes-
thesia there are three underlying features that
seem relatively consistent across cases. First,
synesthetic experiences are typically reported
as arising involuntarily and without mental ef-
fort (Dixon et al. 2000; Mattingley et al. 2001;
Wollen & Ruggiero 1983). Second, the concur-
rents associated with particular inducers tend
to be consistent over time, as documented by
test−retest scores on questionnaire measures
taken over months or years (Asher et al. 2006;
Baron-Cohen et al. 1987). Third, synesthetic
experiences often begin in early childhood, and
might be associated with the development of
language abilities (Rich et al. 2005; Simner
2007; Simner et al. 2005). There have been at-
tempts to establish diagnostic criteria for synes-
thesia, but none is currently widely accepted,
and until more is known about the causes of
the condition it seems premature to accept
any particular taxonomy. The first two features
of synesthesia—its involuntary or “automatic”
nature and its consistency—have provided the
basis for the empirical work I discuss in this
article. If synesthesia arises automatically, with-
out the need for voluntary control, it should be
possible to measure the effect of its presence on
perceptual, cognitive, and affective tasks. Like-
wise, the fact that the link between inducers and
their concurrent experiences remains fairly sta-
ble over time means that synesthetes can be
tested repeatedly with the same stimulus sets,
thus allowing verification and replication.
How Common Is Synesthesia?
There have been numerous attempts to esti-
mate the prevalence of synesthesia, and these
have yielded highly variable outcomes. Two
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influential studies suggested a prevalence of 1
in every 2,000 people, with approximately six
females for every one male (Baron-Cohen et al.
1996; Rich et al. 2005). These studies counted
responses to newspaper articles about synes-
thesia and based their estimates of prevalence
on the average circulation of the newspaper on
the day the article appeared. The figure of 1
in 2,000 is likely to be a conservative estimate,
however, since this approach will not count in-
dividuals with the condition who do not re-
spond. More recently, a prospective study of
two large groups—one composed of university
students and the other of adults and children at-
tending a public science exhibition—estimated
a prevalence of around 4.0% overall and about
1% for the most common grapheme-color vari-
ant (Simner et al. 2006b). Interestingly, the
same study found no evidence for a difference
in the ratio of males to females, suggesting that
males might be less likely to respond to adver-
tisements or surveys than females.
There is, however, convincing evidence for
the long-held belief that synesthesia runs in
families (Rich et al. 2005; Ward & Simner
2005). In one recent investigation of 53 synes-
thetes, 42% of the probands reported at least
one first-degree relative with the condition
(Barnett et al. 2008). Interestingly, the same
study found that different manifestations of
synesthesia occurred within the same family,
and that the ratio of females to males was 4:1,
both in probands and their first-degree relatives
(cf. Simner et al. 2006b).
How Can We Measure
Synesthesia’s Effects on Behavior?
While early studies of synesthesia were con-
cerned principally with cataloguing its varia-
tions and the nature of the individuals’ subjec-
tive experiences, research over the last decade
has focused on providing objective, empiri-
cal evidence for its existence. The emphasis
initially was on verifying the consistency of
synesthesia by having individuals report their
Annals of the New York Academy of Sciences
experiences for a set of inducers, and then
retesting them weeks, months, or years later
(Asher et al. 2006; Baron-Cohen et al. 1987). As
mentioned previously, the data overwhelmingly
suggest that synesthetic experiences, at least by
self-report, are remarkably consistent over time.
Unfortunately, such self-report measures say
little about the mechanisms underlying synes-
thesia. Fortunately, however, the problem of
interrogating synesthetes’ unusual perceptual,
cognitive, or affective experiences has been
overcome by measuring their performance for
objective stimuli that have been manipulated
to have some consistent, formalized relation-
ship with their synesthetic experiences.
By far the most widely applied empirical
measure of synesthesia is based upon the clas-
sic Stroop task, in which observers are re-
quired to identify the color of letter strings
that compose familiar words, and in particular
color names (MacLeod 1991; Stroop 1935). In
the standard version of this task, color-naming
times are significantly slower for color names
printed in nonmatching (incongruent) colors
(e.g., the word RED displayed in green) than
for color names printed in matching (congru-
ent) colors (e.g., the word GREEN displayed
in green). The Stroop effect (Stroop 1935),
measured as the difference in response times
and errors for incongruent versus congruent
conditions, is thought to be due to interfer-
ence between an automatic and rapid word-
reading process, and a more controlled, effort-
ful, and therefore slower color identification
process (MacLeod 1991). Compatibility effects
of this sort are observed in numerous behavioral
domains, and researchers have determined that
they represent a general index of interference
between concurrent perceptual, cognitive, or
motor processes.
In a pioneering study Wollen and Ruggiero
(1983) had synesthetes perform rapid color-
naming of lists of letters selected to elicit synes-
thetic experiences of color. The investigators’
key manipulation was to color some of the
items congruently (i.e., the print color to be
named matched the synesthetic color elicited
Mattingley: Attention, Automaticity and Awareness in Synesthesia
Figure 1. Colored letters and the synesthetic
Stroop task. (A) Colors of letters experienced by
synesthete E.S. (B) Displays showing letters col-
ored congruently with E.S.’s synesthetic colors. (C)
Displays showing letters colored incongruently with
E.S.’s synesthetic colors.
by the letter) and to color others incongruently,
thus establishing a kind of “synesthetic Stroop”
effect (Fig. 1). The synesthetes were significantly
slower to name the print colors of incongruent
items than congruent items, whereas controls
showed no such difference. Their findings pro-
vided the first objective evidence that synes-
thesia has a measurable influence on behavior
and prompted a range of subsequent stud-
ies in which Stroop-like tasks have been used
to measure automaticity in synesthesia (Beeli
et al. 2005; Dixon et al. 2000; Mattingley et al.
2001). Almost without exception these studies
have reported interference effects analogous to
those observed by Wollen and Ruggiero’s study
(1983), thus providing one of the most repli-
cated behavioral effects in the area.
Some authors have objected that since
Stroop interference might in principle arise
anywhere in the processing hierarchy, from
the initial stages of sensory registration to
the final selection of an appropriate response,
such approaches cannot verify the “percep-
tual reality” of the condition (Palmeri et al.
2002; Ramachandran & Hubbard 2001a,
Ramachandran & Hubbard 2003). Indeed, it
is known that even nonsynesthetes can show
interference effects after training with arbi-
145
trary associations (Elias et al. 2003; MacLeod &
Dunbar 1988). While it is true that the conven-
tional application of the Stroop task cannot de-
termine whether synesthetic interference arises
early or late in processing, the paradigm lends
itself to modifications that can potentially reveal
the influence of fundamental processes such as
attention and cognitive set (MacLeod 1991), as
I describe below. Moreover, it is not clear that
any test yet devised can unambiguously deter-
mine whether synesthesia is experienced as a
sensory event, an overlearned association, or a
vivid mental image.
Dixon et al. (2004) used the Stroop paradigm
also to determine whether projector and as-
sociator synesthetes, classified on the basis of
subjective reports, can be distinguished objec-
tively based on cognitive performance. In one
task they had synesthetes name the print color
of individual graphemes presented on a com-
puter display, as per the original Wollen and
Ruggiero (1983) study. In a second task, they
asked synesthetes to name their synesthetic color
for each grapheme and to ignore its print color.
The projectors showed greater Stroop inter-
ference from synesthetic colors when naming
real colors than they did from real colors when
naming synesthetic colors, whereas the associ-
ators tended to show the reverse pattern (i.e.,
greater interference from real colors than from
synesthetic colors). Dixon et al. (2004) con-
cluded that for projectors the mechanisms giv-
ing rise to synesthetic concurrents are triggered
more automatically and thus earlier than those
supporting the naming of real colors, whereas
for associators real colors enjoy a processing
advantage over synesthetic colors.
Dixon et al. (2004) speculated that pro-
jected synesthetic colors might be more dif-
ficult to ignore than the internal colors ex-
perienced by associators because projected
colors are more perceptually intense, or be-
cause they occupy the same spatial position
as the colored graphemes that elicit them.
Indeed, Ward et al. (2007) have suggested
that the projector−associator dichotomy in
fact reflects the different spatial reference
146
frames within which synesthetes experience
their colors, rather than the internalized ver-
sus externalized nature of the experiences per
se. In their study Ward et al. (2007) observed
greater Stroop interference effects among in-
dividuals who projected their synesthetic col-
ors onto the viewing surface (e.g., a page
of text or a computer display) than among
those who projected their colors externally but
away from the surface upon which the in-
ducer was displayed. (Interestingly, the latter
group of projectors showed synesthetic Stroop
effects that were comparable to those of the
associators.)
Despite these demonstrations of objective
individual differences between projector and
associator synesthetes, there are some caveats
worth mentioning. First, for synesthetes who
project their colors onto the graphemes that
elicit them, one would expect significantly pro-
longed responses in the Stroop task when in-
dividuals have to identify the real color of an
incongruent grapheme, since in this condition
there should be more perceptual competition
than is encountered by associators or by in-
dividuals who do not see colors attached to
the graphemes that elicit them. In fact, Dixon
et al. (2004) found that response times in their
incongruent trials were the same for projec-
tors and associators (the reader can verify this
by comparing the left and right panels in their
Fig. 1). In fact, the group difference in the Dixon
et al. (2004) study occurred because the projec-
tors were significantly faster on congruent tri-
als than the associators, rather than slower on
incongruent trials, a result that is not readily
explained by perceptual interactions. A second
caveat is that many synesthetes spontaneously
change the way in which they describe their
color experiences, making classification based
on verbal reports particularly problematic. For
instance, Edquist et al. (2006) found that some
synesthetes could not say with any confidence
precisely where in space they experienced their
colors, while others changed from internalizing
to externalizing their colors, or vice versa, after
a 12-month interval.
Annals of the New York Academy of Sciences
Finally, a recent study found that the
magnitude of the interference effect elicited by
the Stroop task is larger for incongruent trials
when the hue of the grapheme and the hue
of the synesthetic color it elicits are opponent
colors (i.e., red−green, blue−yellow), compared
with incongruent trials in which the colors are
not from within the same opponent “channel”
(e.g., red−blue, green−yellow; Nikolić et al.
2007). This opponent-color effect, though
present in all six synesthetes tested, was
particularly pronounced in the two who were
classified as projectors based upon subjective
report. The authors suggest that the additional
interference generated in the opponent-color
condition was caused by competition within
early color-opponent areas of the visual system
(from V1 to V4) and that this in turn confirms
that the color experiences of projector synes-
thetes are truly perceptual in nature. Despite
the authors’ claims, however, several aspects
of their interpretation are rather speculative.
For instance, no explanation is offered for
how competition between opponent channels
within early color-selective visual neurons
could exert a knock-on effect for accessing the
appropriate color name from the lexicon, as
required by the Stroop task. Moreover, it is
entirely plausible that color names get stored
within, or are accessed from, the lexicon in a
manner that preserves the opponent organi-
zation of earlier perceptual stages of analysis.
Such an arrangement would only become
evident in synesthetes during Stroop-type tasks
in which there is competition for access to the
lexicon from both real and synesthetic colors.
In summary, the question of individual dif-
ferences in synesthesia remains controversial,
but research on the issue has yielded findings
that promise to drive the field forward.
Attention, Automaticity, and
Awareness in Synesthesia
Over the last 10 years a considerable
body of behavioral research has examined
Mattingley: Attention, Automaticity and Awareness in Synesthesia
issues concerning the potential influence of
synesthesia on perceptual processes, the extent
to which synesthetic concurrents can influence
attentional processes, and whether synesthesia
can be triggered without the synesthete’s
conscious awareness of an inducing stimulus.
A parallel line of work has used physio-
logical approaches such as scalp-recorded
event-related potentials (ERPs), functional
magnetic resonance imaging (fMRI), and
transcranial magnetic stimulation (TMS) to
uncover the neural correlates of synesthesia.
In the sections that follow I review recent
behavioral investigations relating to the issues
of attention, automaticity, and awareness
and draw upon relevant findings from brain
imaging and neurostimulation studies. In
doing so, I consider evidence from individuals
with grapheme-color synesthesia, since this is
the most commonly encountered form of the
condition and there are more published reports
on this synesthetic variant than any other.
Can Synesthetic Colors Act Like
Real Colors in Perceptual Tasks?
As mentioned earlier, several studies have fo-
cused on the question of whether synesthetic
experiences of color are akin to those involved
in the perception of real color in the world. In
an attempt to move beyond subjective reports
investigators have adapted well-known percep-
tual and cognitive tasks to answer questions
about the level at which synesthetic concurrents
are generated in the brain.
In an influential study, Ramachandran and
Hubbard (2001b) tested two synesthetes and
a group of nonsynesthetic controls on tests
of visual grouping and visual search. In their
grouping task, participants examined displays
containing a matrix of black digits arranged
in rows and columns on a white background.
The synesthetes tended to group the matrix
into rows or columns on the basis of the synes-
thetic colors elicited by the digits, whereas non-
synesthetes grouped the items based exclusively
147
on form and proximity. In the visual search
task, participants inspected an array of black
letters, a subset of which formed a single ge-
ometric shape of the same letter. Participants
had to identify the embedded shape, basing
their decision upon grouping of the local ele-
ments. The synesthetes tended to be faster than
controls in identifying the embedded shape, a
result that suggests the shape had a pop-out
effect for them by virtue of the unique synes-
thetic colors triggered by the letter stimuli. The
authors attributed their results to grouping pro-
cesses being driven by the sensory properties of
perceived synesthetic colors. It is equally plausi-
ble, however, that the synesthetes’ reports were
based upon semantic associations in the group-
ing study (much as one would recognize a fa-
miliar telephone number or PIN), and that pro-
longed viewing in the visual search task (one
second) permitted more efficient rejection of
distractor items based on their synesthetic color,
as opposed to a pop-out effect of the target
(Robertson 2003).
Despite these interpretative issues, subse-
quent studies have focused on whether synes-
thetic colors can act like “real” colors in per-
ceptual tasks. To date the findings have been
tantalizing, but not always easy to explain or en-
tirely convincing. For instance, Hubbard et al.
(2005) claimed that synesthetic colors can re-
duce visual crowding effects for graphemes
presented in the visual periphery, just as what
occurs for such tasks when the target and dis-
tractors are actually displayed in different col-
ors (Kooi et al. 1994). They had a group of
synesthetes try to identify a black target letter
positioned in peripheral vision. The target was
surrounded by black distractor letters of a dif-
ferent identity, thus inducing a crowding effect
that would normally render the target difficult
to identify. Three of the six synesthetes per-
formed better than nonsynesthetes on this task,
suggesting that somehow the unique color as-
sociated with the target helped unmask it from
the distractors. Superficially, the results imply
a direct effect of synesthetic colors on percep-
tion; but, for this conclusion to hold one must
148
assume that the color is triggered before the tar-
get grapheme is identified, and it remains un-
clear how any such precategorical effect might
arise (though see Smilek et al., 2001, for a clever
but untested account).
In a different approach to the question of
whether synesthetic colors arise early in sensory
processing, Kim et al. (2006) had two synes-
thetes judge the direction of motion in bistable
apparent motion displays. The approach in this
study was to examine interactions between real
and synesthetic colors in successive visual dis-
plays, one of which contained pairs of achro-
matic graphemes that induced specific colors
and the other of which contained items that
appeared in real (but matching) colors. Non-
synesthetes perceived the direction of apparent
motion in these displays as completely ambigu-
ous (equally either clockwise or counterclock-
wise). By contrast, the synesthetes tended to
report a motion path in which synesthetic col-
ors induced by the achromatic display corre-
sponded with the real (matched) colors in the
chromatic display. While the findings appear to
suggest an early sensory influence of induced
colors on perception, the task used by Kim
et al. (2006) was not criterion-free, and thus
is open to the criticism that high-level strategies
could have influenced the synesthetes’ judg-
ments. For instance, a bias in apparent motion
judgments could readily arise from purely con-
ceptual grouping strategies across chromatic
and achromatic displays, per the grouping re-
sults of Ramachandran and Hubbard (2001b).
Another study by the same group (Blake et al.
2005) examined whether synesthetic colors as-
sociated with graphemes are able to induce the
McCollough effect, an orientation-contingent
color aftereffect that occurs when observers are
exposed to colored contours over several min-
utes (McCollough 1965). The authors showed
synesthetes achromatic adapting displays con-
taining vertical columns of letters that induced
a red color experience and horizontal rows of
letters that induced a green color experience.
After several minutes of adapting to these two,
achromatic, displays the synesthetes reported
Annals of the New York Academy of Sciences
faint opponent color aftereffects (a greenish
tinge or pinkish tinge) while looking at an
achromatic test display composed of nonalpha-
betic characters arranged in rows and columns.
These striking results for two individuals imply
that synesthetic colors might act like real colors
under some conditions. As highlighted earlier,
however, the findings must now be replicated
and quantified using a criterion-free measure of
performance. It will also be important to con-
sider the extent to which mental imagery might
contribute to such aftereffects, since some ac-
counts of synesthesia have suggested that the
condition is associated with abnormally vivid
visual imagery (Barnett & Newell 2008).
Another approach to examining whether
synesthetic colors behave like real colors in per-
ceptual tasks is to ask whether an achromatic
character experienced as colored can affect the
appearance of a nearby chromatic character.
It is known that perceived hue and brightness
are systematically influenced by light and color
from neighboring regions of the visual field,
and that these effects arise early in the visual
processing hierarchy (i.e., within primary vi-
sual cortex and earlier; Palmer 1999). Do synes-
thetic color experiences have analogous effects?
This question was addressed recently by Hong
and Blake (2008), who had four synesthetes
perform a variety of visual matching tasks de-
signed to assess brightness contrast and chro-
matic adaptation, two phenomena thought to
be resolved at the earliest stages of cortical
vision.
In one task, Hong and Blake (2008) found
that the perceived brightness of chromatic
test patches decreased as the brightness of a
surrounding annulus increased (a well-known
effect called brightness contrast), but that synes-
thetic colors induced by an achromatic charac-
ter showed no influence of their surround (i.e.,
the character was unaffected by brightness
contrast). In a second task the synesthetes were
adapted to an array of identical graphemes
over several minutes. Some of these arrays
contained chromatic characters, whereas
others contained achromatic characters that
Mattingley: Attention, Automaticity and Awareness in Synesthesia
induced a single synesthetic color. Following
adaptation, synesthetes adjusted the color of a
chromatic test patch until it was perceived as a
specified hue (known as “equilibrium yellow”)
(Shevell 1982). Whereas prolonged exposure
to an array of colored graphemes shifted the
participants’ perception of the test hue, expo-
sure to the achromatic, synesthesia-inducing
arrays had no such effect. There was also no
difference in equilibrium yellow judgments for
graphemes that induced synesthetic colors of
red and green compared with a neutral symbol
that induced no synesthetic color. Taken
together these results indicate that the neural
correlates of synesthetic color experiences
must arise downstream from the processes that
give rise to brightness contrast and chromatic
adaptation.
To summarize, evidence is somewhat mixed
on the question of whether synesthetic colors
are represented in the brain in the same way
as real colors. Clearly, synesthetic colors can
be used by synesthetes to influence decisions
about grouping and the direction of ambiguous
motion, but this does not necessarily imply an
early sensory locus for the effect. Based on sub-
jective reports, one may conclude that synes-
thetic colors can induce an orientation con-
tingent color aftereffect, but this result needs
to be quantified objectively and distinguished
from any potential influence of color imagery.
Finally, synesthetic colors seem to be immune
to brightness contrast and fail to induce chro-
matic aftereffects, suggesting that they are not
treated like real colors at the earliest stages of
visual processing.
Do Synesthetic Colors Pop Out
in Visual Search Displays?
Even if synesthetic colors do not arise from
neural processes akin to those associated with
the perception of real color, they evidently can
have a potent effect on the subjective sensory
lives of those who experience them. We have
had synesthetes describe to us their experiences
149
of being overwhelmed by colors “jumping out”
from signs and billboards in shopping malls.
If synesthetically induced colors are so salient,
they should exert a measurable influence on
behavior. Having synesthetes perform visual
search tasks, in which they are required to lo-
cate an achromatic target grapheme in an ar-
ray of achromatic distractors, provides a simple
way of addressing this question. If an achro-
matic target generates a synesthetic color pre-
attentively, and that color is distinct from any
colors elicited by the distractors, then the ob-
server should experience pop-out of the target.
Several investigators have asked whether synes-
thetic colors might be elicited preattentively in
otherwise cluttered displays, perhaps even in
the absence of any conscious experience of the
stimuli that induce them.
The visual grouping study of Ramachandran
and Hubbard (2001b), in which participants
had to identify a geometric shape made up of
letters of one identity (e.g., H) from among
distractor letters of a different identity (e.g.,
F and P), found an advantage for synes-
thetes over controls. But since the displays
were visible for a full second each, faster re-
sponses by the synesthetes might simply have
reflected their ability to extract the colors of
target and distractor items after a serial search,
rather than indicating true preattentive pop-
out of the target shape. In a subsequent study,
Palmeri et al. (2002) used a more traditional
visual search paradigm to test whether synes-
thetic colors confer some advantage for dis-
plays that would yield serial search in non-
synesthetes. They asked a synesthete, W.O., to
search for a 2 among 5s, which elicited or-
ange and green, respectively; or an 8 among
6s, which both elicited blue. The displays were
entirely achromatic, and the Arabic numer-
als were “digitalized” (i.e., composed of ver-
tical and horizontal line segments only) so that
they could not be distinguished on the basis of
individual features. For nonsynesthetes search
times increased over larger set sizes, as ex-
pected for this difficult serial search task and
for both target/distractor pairings. A similar
150
effect was found for W.O. when the target and
distractors elicited the same synesthetic color
(a blue 8 among blue 6s). But when the numer-
als elicited distinct synesthetic colors (an orange
2 in green 5s), W.O. showed a shallower search
slope. Interestingly, however, the search slope
was not completely flat, as is typically the case
in feature-search tasks. Thus, despite W.O.’s
spontaneous report that the target seemed to
pop out of the display, it is clear that his pro-
cessing of synesthetic colors did not occur in
parallel across the visual field.
At present the consensus is that synesthetic
colors do not arise preattentively during visual
search (Treisman 2005). Indeed, it seems likely
that in most synesthetes induced colors do not
normally confer any measurable advantage at
all in serial search tasks. Edquist et al. (2006)
tested 14 grapheme-color synesthetes using tai-
lored, achromatic displays of the kind intro-
duced by Palmeri et al. (2002) (see Fig. 2). They
found no evidence for shallower search slopes in
synesthetes compared with controls, either for
the group as a whole or for any of the individ-
uals, despite all participants showing reliable
pop-out effects for chromatic variants of the
same displays. One factor that might play an
important mediating role during visual search
is the location of the target item within the dis-
play. Synesthetes are seemingly more efficient
at searching within an entirely achromatic dis-
play if the target happens to fall at fixation or
within the current focus of attention (Laeng
et al. 2004; Sagiv et al. 2006).
Traditional visual search tasks, such as those
described above, aim to measure effects of in-
duced synesthetic colors on target detection
within entirely achromatic displays. By con-
trast, Smilek et al. (2001) examined whether
target identification could be affected by the re-
lationship between a synesthetic color and the
background upon which the inducer appears.
In one experiment the investigators asked their
synesthete, C., to identify a briefly flashed tar-
get digit that was immediately followed by a
pattern mask. The digit was achromatic, and
appeared on a background that was colored so
Annals of the New York Academy of Sciences
that it matched the synesthetic color induced by
the target (the congruent condition) or so that it
was not matched (the incongruent condition).
The authors reasoned that a target appearing
on a congruently colored background might
be more difficult to identify, by virtue of some
sort of color camouflage, than one presented
on an incongruent background. C. did in fact
make more errors for congruent versus incon-
gruent displays, in support of the authors’ pre-
diction. In a further experiment, C. performed
a more conventional visual search task in which
she had to indicate the presence of an achro-
matic target digit (a 2 or 4) among achromatic
distractors (8s). Once again, the target elicited
a synesthetic color that was either congruent
or incongruent with respect to the background
color. C. showed a somewhat shallower search
slope for the incongruent condition relative to
the congruent condition, though this difference
was not statistically reliable. A subsequent study
(Smilek et al. 2003), which employed a similar
paradigm with a different synesthete, yielded a
significant slope difference, with search being
more efficient for incongruent than congruent
trials.
The findings of these “color camouflage”
experiments are intriguing but difficult to in-
terpret unambiguously. Superficially, they sug-
gest that synesthetic colors are triggered prior
to character identification, or at least that the
two representations emerge in parallel. This
seems counterintuitive because it implies that a
synesthetic color that depends on an identified
grapheme for its existence can itself render the
inducing grapheme difficult to perceive against
a similarly colored background. The authors
offer a clever explanation for the effect, one
which posits an important role for feedforward
and feedback pathways within the extrastriate
visual system; but this account remains to be
tested. Although the existence of such pathways
is not disputed, their putative role in synesthesia
has yet to be established. Several aspects of the
data themselves are also puzzling. For instance,
C.’s performance in the congruent conditions
of the two experiments, in which the target item
Mattingley: Attention, Automaticity and Awareness in Synesthesia 151

Figure 2. Target and distractor stimuli used in the visual search tasks of Edquist et al.
(2006). In each trial participants indicated which of two target items (T1 or T2) was present
among 7, 15, or 23 distractors (D). (A) Chromatic and achromatic stimuli for each of the 14
synesthetes (Syn). Note that the colors used in the chromatic displays were selected to match
the synesthetic colors for each individual. (B) Example of an achromatic display for the largest
set size (24 items). (C) Example of a chromatic display for the largest set size. Reproduced
from Edquist et al. 2006.

should have been effectively camouflaged by
the real background color, was in fact no worse
than that of several of the nonsynesthetic con-
trols. In addition, it is not clear what outcome
one would expect in these tasks if chromatic
characters were presented on matching or non-
matching backgrounds. Such a condition was
not included in either investigation.
A recent study revisited the color camou-
flage effect in a group of seven grapheme-color
152
synesthetes. Instead of using colored back-
grounds, however, Gheri et al. (2008) used a
4 × 4 matrix of Arabic numerals within which
a single, achromatic target numeral appeared.
The identity of the target was chosen so that
it elicited a unique synesthetic color for each
individual. The identities of the distractors, on
the other hand, were chosen such that they
evoked a single but distinct color relative to the
target (in the “unique” condition), or so that
at least one distractor evoked the same color
as that of the target (the “nonunique” con-
dition). If synesthetic colors pop out, then in
the unique condition the distinct color of the
target, relative to the distractor(s), should have
rendered search more efficient. By contrast, the
nonunique condition should have yielded less
efficient search, due to competition between
similarly colored target and distractor items
(i.e., a color camouflage effect. Gheri et al.
(2008) in fact found no evidence for a differ-
ence in search efficiency between synesthetes
and controls across the two conditions, imply-
ing that synesthetic colors do not arise in par-
allel across the visual field.
To summarize, several different visual search
tasks have been designed to test whether synes-
thetic colors are sufficiently salient to pop out
from among achromatic distractors. To date
there is no compelling evidence for the kind
of pop-out effect that is known to occur for
targets defined by real colors. Moreover, the
weight of evidence suggests that the perfor-
mance of most synesthetes faced with achro-
matic displays is not reliably different from that
of controls. Thus, despite synesthetes’ subjec-
tive reports of vivid color experiences, the po-
tency of their experiences fails to translate into
measurable effects in the lab, at least for the
visual search tasks that have been implemented
to date.
Can Voluntary Attention Influence
Synesthetic Color Experiences?
Beyond the realm of visual search, sev-
eral studies have investigated the question of
Annals of the New York Academy of Sciences
whether limiting or diverting selective atten-
tion from an inducing character can modulate
the synesthetic concurrent. Some synesthetes
we tested admit that their colors often fade or
are absent altogether when they are absorbed
in reading a book or listening to dialogue on
the radio or during a movie. One of our synes-
thetes, KP, said:
It’s kind of like looking at your own nose. If you try,
you can see it clearly, but you don’t walk around
the whole time ‘seeing’ your nose. But it’s always
there and you can see it, just that you don’t unless
you’re attending to it. (Rich et al. 2005 p. 67)
Anecdotal observations like this one suggest
that synesthetic colors, while apparently elicited
involuntarily under some conditions (such as
the synesthetic Stroop task), are strongly sup-
pressed from awareness when the individual is
attending elsewhere.
A striking illustration of this effect in the
laboratory emerges from having synesthetes
view Navon-type stimuli, in which the local
characters (e.g., As) elicit one synesthetic color
(red) and the global figure (e.g., B) elicits an-
other (green). When synesthetes are instructed
to attend selectively to the scale of the smaller
elements, they report experiencing the color
triggered by the local elements only (red in this
example); by contrast, when they attend to
the global shape they experience the color
for the global figure (green) (Palmeri et al.
2002). This attentional effect has been quanti-
fied experimentally by having grapheme-color
synesthetes name the print color of local or
global letters that are colored congruently or
incongruently with the synesthetic colors they
elicit (Rich & Mattingley 2003, see Fig. 3).
Responses were slowest for stimuli in which
items at both spatial scales were colored incon-
gruently and fastest when items at both scales
were colored congruently. Response speed was
intermediate between these two conditions
when the local elements made a different
global letter and the display color was congru-
ent with only one of the two letters (either local
or global). A further notable finding: When
synesthetes actively ignored an incongruently
Mattingley: Attention, Automaticity and Awareness in Synesthesia 153

Figure 3. Examples of the four stimulus types used to study the effect of attentional com-
petition between multiple synesthetic inducers, using the colors of synesthete E.S. The task
was to name the stimulus color as quickly as possible. Note that participants never had to
name the alphanumeric characters, which were irrelevant and nonpredictive with respect to
the display color. (A) Local and global letters consistent; display color matches the synesthetic
color elicited by the letter. (B) Local and global letters consistent; display color is different
from the synesthetic color elicited by the letter. (C) Local and global letters inconsistent; dis-
play color matches the synesthetic color elicited by the local letters but is incongruent with the
global letter. (D) Local and global letters inconsistent; display color matches the synesthetic
color elicited by the global letter but is incongruent with the local letters. Reproduced from
Rich and Mattingley (2003).

colored global stimulus and focused instead on
congruently colored local elements, the Stroop
effect was significantly attenuated relative to
when participants were not cued to the local
level in advance. The same reduction was
apparent when synesthetes ignored the incon-
gruently colored local elements and were cued
to attend to just the congruently colored global
form. These findings indicate that voluntary
attention modulates induced color experiences,
as measured by the synesthetic Stroop task.
It should be noted, however, that there was
still some residual cost of having incongruently
colored items at the ignored level, implying
that attentional control might not fully sup-
press synesthetic concurrents, a finding that
has emerged from other studies, as outlined
below.
Another approach to examining the effect of
attention on synesthetic colors is to have par-
ticipants engage in a secondary task while they
view inducing characters, and to measure the
magnitude of any color experience with and
without the secondary task. In one such study
(Mattingley et al. 2006), we had grapheme-
color synesthetes perform a variant of the synes-
thetic Stroop task. In each trial an achromatic
letter prime appeared briefly (for 400 ms) at fix-
ation, followed by a congruent or incongruent
color patch that participants had to name as
154 Annals of the New York Academy of Sciences

Figure 4. Sequence of displays in a typical trial of the attentional load task of Mattingley
et al. (2006). Participants attended to diagonally opposite sides of the diamond throughout
a block of trials. In this example, focusing attention on the top-left and bottom-right gaps
constituted the low-load condition (a relatively easy discrimination); focusing attention on the
top-right and bottom-left gaps constituted the high-load condition (a difficult discrimination).
Participants first named the color of the target display as quickly as possible (“green” in
this case), and then indicated which gap in the attended pair was larger. Reproduced from
Mattingley et al. (2006).

quickly as possible (Fig. 4). The letter prime
was encompassed within a diamond-shaped
surround, into which small gaps were inserted
midway along all four sides. These gaps, which
were used for the secondary task, appeared and
disappeared with the prime letter. At the begin-
ning of each block, participants were instructed
to attend to opposite limbs of the diamond (e.g.,
upper left and lower right) in order to judge
which of the attended gaps was larger. They
made an unspeeded response at the end of the
trial, after their speeded response to the color of
the target. “Low-attentional load” blocks were
those for which the sizes of the gaps were ad-
justed to be relatively easy to distinguish; “high-
attentional load” blocks were those where the
gaps were much harder to discriminate.
We measured the magnitude of the synes-
thetic Stroop effect under conditions of low and
high load in the secondary task, and compared
this with the effect when participants were in-
structed to ignore the gaps altogether (a “no
load” condition). We found a robust synesthetic
Stroop effect—that is, slower reaction times
(RTs) for incongruent versus congruent trials—
in the absence of any attentional load, consis-
tent with previous reports. Crucially, with the
introduction of a secondary task, the magnitude
of the synesthetic Stroop effect was significantly
smaller under high-attentional load than un-
der low-attentional load. Note that the displays
and task demands remained identical across
the two load conditions, ruling out any expla-
nation for the difference in terms of working
memory or response competition with the gap
discrimination task. The findings suggest that
diverting visual attention from a synesthetic in-
ducer attenuates synesthetic colors (or at least a
measurable effect of their occurrence), consis-
tent with many synesthetes’ subjective reports
(Rich et al. 2005).
Rather than having synesthetes divide their
perceptual resources across multiple elements
of a common display, investigators have also
measured the influence of the temporal allo-
cation of attention by embedding an inducer
within a rapid serial visual presentation (RSVP)
of distractor items. Using this design and having
Mattingley: Attention, Automaticity and Awareness in Synesthesia
participants identify an initial target item (T1)
in the stream enables the investigators to ask
if synesthetic colors can arise for inducers that
appear within the period of the “attentional
blink” (AB) (Raymond et al. 1992). Johnson
et al. (2007) had grapheme-color synesthetes
monitor an RSVP stream of non-alphanumeric
distractors, such as #, %, and ?, selected so as
not to elicit any synesthetic colors. Each stream
also contained a single, achromatic letter (T1),
chosen for each synesthete so that it would not
elicit a synesthetic color, plus a single achro-
matic digit (T2), at different lags, chosen from
a set of digits, each of which elicited a distinct
synesthetic color. Participants were to iden-
tify T1 and T2, and then to indicate whether
they had been aware of any color during the
trial. Note that the Johnson et al. (2007) de-
sign ensured that any synesthetic color should
only be attributable to the digit, which was
always T2.
Synesthetes and nonsynesthetic controls
showed comparable AB effects, both in terms
of duration and depth (roughly 88% correct
T2 identification). The crucial data, however,
concerned the synesthetes’ color responses.
When they correctly identified T2 they virtu-
ally always reported the appropriate synesthetic
color. When T2 was missed, the synesthetes
typically reported no color. But on a small pro-
portion of “miss” trials, a color was reported,
and this was significantly more likely to be the
color normally elicited by the T2 digit than
that elicited by any of the other possible digits.
Superficially, these findings would seem to sup-
port the authors’ claim that synesthetic colors
can arise unconsciously, during a time period
in which the relevant inducer was present but
rendered invisible due to the AB. On closer in-
spection, however, the results seem less convinc-
ing. First, the AB effect was very small overall,
and so the color-report data were based on just
138 of 3,200 trials over the group of 10 synes-
thetes. Moreover, only 4 of the 10 individuals
exhibited reliable color reports for unseen T2
digits. Finally, and perhaps most importantly,
the synesthetes might have been less willing to
155
guess the identity of a partially processed T2
digit, but were nevertheless prepared to guess
its color. Although the authors instructed their
participants to report stimuli only if they were
certain they had seen them, it is impossible to
rule out such a criterion shift. A second exper-
iment, in which the T2 digit was now colored
congruently or incongruently with the synes-
thetic color it elicited, revealed that T2 was
more likely to be identified if it was colored
congruently than if it was colored incongru-
ently. This finding provides some evidence for
an interaction between real and synesthetic col-
ors during RSVP streams in which attention
must be rapidly deployed over multiple stim-
uli. However, the effect was reliable for only 3
of the 10 synesthetes, and it does not speak at
all to the issue of whether synesthetic colors can
be elicited by unseen inducers presented within
the period of the AB.
We recently examined this question (Rich &
Mattingley 2005) using an achromatic RSVP
stream of nonalphanumeric distractors within
which an inducing letter was positioned on 50%
of trials at a variable lag following presenta-
tion of T1 (an achromatic grating at one of
four orientations; see Fig. 5). Our aim was to
establish whether a T2 letter presented dur-
ing the AB would influence the time required
to identify a congruently or incongruently col-
ored target patch at the end of the stream. In
separate blocks of trials participants performed
one of three tasks. First, we verified an AB ef-
fect by having participants detect T2 after first
identifying the orientation of the T1 grating. In
this task both controls and synesthetes exhib-
ited a significant dual-task decrement at lags 2
and 3 (stimulus onset asynchronies [SOAs] of
233 and 350 ms). Second, we established that a
synesthetic Stroop effect was present for T2 let-
ters by having participants detect T2 and then
identify the color of the patch at the end of the
RSVP stream. For this task, synesthetes were
significantly slower to name incongruent ver-
sus congruent color targets across all SOAs (up
to 816 ms). In the critical task we determined
the influence of attention on the congruency
156 Annals of the New York Academy of Sciences

Figure 5. Appearance and timing of displays in the attentional blink task of Rich and
Mattingley (2005). (A) Schematic representation of the trial sequence. Each display was
presented for 100 ms and was followed by a blank screen for 17 ms (not shown). The color
of the probe was either congruent or incongruent with the synesthetic color elicited by the
T2 letter-prime. (B) Timing of displays within a trial, and the number of target and distractor
items. SOA = stimulus onset asynchrony; ISI = interstimulus interval.

effect by measuring color-naming times when Can Synesthetic Color Experiences

participants first identified the orientation of
the T1 grating. We found a reliable but atten-
uated congruency effect for T2 items falling
outside of the AB, but not for T2 items that
appeared at the deepest point in the AB (i.e.,
at a T1-T2 SOA of 350 ms). Taken together,
the findings suggest that synesthetic colors, and
the effect they exert on color identification, can
be effectively eliminated when visual attention
is completely occupied in processing a visual
distractor.
Arise from Unconsciously
Processed Inducers?
The attentional blink studies reviewed above
yielded conflicting results on whether unat-
tended inducing characters can trigger synes-
thetic experiences of color, as outlined above.
Johnson et al. (2007) found that synesthetic col-
ors were reported occasionally when the induc-
ing character could not be overtly identified,
whereas Rich and Mattingley (2005) found that
Mattingley: Attention, Automaticity and Awareness in Synesthesia
synesthetic interference was eliminated for in-
ducers that appeared during the AB. In both
these studies participants were unable to con-
sciously identify the critical inducers because of
an attentional bottleneck in information pro-
cessing induced by the requirement to report
a preceding target within the RSVP stream
(Raymond et al. 1992). Despite this discrep-
ancy, it is widely agreed that many perceptual
and cognitive processes can proceed in the ab-
sence of subjective awareness. There is a wealth
of evidence, for example, that visual stimuli that
are presented briefly and masked from aware-
ness can nevertheless exert a measurable influ-
ence on behavior (Snow & Mattingley 2003).
Mattingley et al. (2001) addressed this issue
by having grapheme-color synesthetes perform
a variant of the synesthetic Stroop task, in which
achromatic inducing characters were flashed
briefly and masked to limit conscious percep-
tion. In the main experiment a forward mask
was followed immediately by a letter prime,
and then a colored backward mask that acted
as a target. Participants ignored the prime and
concentrated exclusively on naming the color
of the target mask, which could be congru-
ent or incongruent with the letter prime, as
quickly as possible. When the prime appeared
for 500 ms, and thus was clearly visible, there
was a robust synesthetic Stroop effect such that
color-naming times were slower in the incon-
gruent trials than in the congruent trials. For
primes of 56 and 28 ms duration, however, the
Stroop effect disappeared. To check whether
the primes could be identified explicitly at these
shorter presentation durations a further experi-
ment was conducted in which participants were
instructed instead to identify the prime and ig-
nore the colored target. In this experiment, par-
ticipants could readily identify the prime when
it appeared for 500 ms but were unable to do
so reliably at 56 and 28 ms. To verify that the
primes were being processed at these shorter
durations, a control experiment was performed
in which the colored target was replaced by
an uppercase letter that either matched or did
not match the identity of the lowercase prime.
157
Both synesthetes and matched nonsynesthetic
controls were significantly slower to name the
target letters on incongruent than congru-
ent trials, indicating that the unseen primes
were processed at least to the stage of letter
recognition.
The findings of Mattingley et al. (2001)
suggest that explicit recognition of inducing
characters is required for the induction of
synesthetic colors (or at least for any effects of
synesthetic colors to be measured objectively).
This conclusion has been challenged on the
grounds that the synesthetic priming effect
might be weak or variable in comparison with
the letter-identity priming effect measured in
their control task (Blake et al. 2005). If so, then
many more trials of the synesthetic priming
task might be needed to uncover a subtle effect
of an unconscious prime on color-naming
performance. A further issue is that there is
no reason to expect that all synesthetes will
exhibit an unconscious priming effect, and so
analysis of group data might miss any reliable
individual differences. We have examined the
performance of each individual who partici-
pated in the Mattingley et al. (2001) study and
found no evidence for reliable color priming
effects in any of the synesthetes. Nevertheless,
it would be useful to repeat the study in a
small number of synesthetes, using individually
titrated prime durations and many more trials,
to determine whether synesthetic colors can
be elicited by inducers that are not consciously
perceived.
How Might Synesthetic Experiences
Arise within the Brain?
In the context of the behavioral data re-
viewed above, it seems reasonable to ask how
synesthesia might arise from coordinated brain
activity. It is now more or less taken for granted
that grapheme-color synesthetes do not have
any kind of recognized psychiatric or neurolog-
ical illness (Rich et al. 2005). Moreover, de-
spite the evidence for significant Stroop-like
158
interference effects in laboratory studies, most
synesthetes clearly lead completely normal—if
slightly more colorful—lives. Early suggestions
of a higher incidence of left-handedness have
proven unfounded (Rich et al. 2005), and al-
though some synesthetes report difficulties with
tasks such as mathematical reasoning (Rich
et al. 2005), others have been shown objec-
tively to have superior abilities in areas such as
new learning and recent memory (Smilek et al.
2002).
Any neural explanation for synesthesia must
be able to account both for the unusual per-
ceptual experiences and for the absence of con-
comitant abnormalities in the perceptual, cog-
nitive, and affective domains. To date there
have been two dominant accounts. One sug-
gests that synesthesia arises as a consequence
of disinhibited feedback from higher associa-
tive areas of the cortex to lower perceptual
and sensory areas (Grossenbacher & Lovelace
2001); the other suggests that there is anoma-
lous or additional cross-wiring between one
or more brain areas responsible for processing
graphemes and color (Harrison & BaronCo-
hen 1996; Ramachandran & Hubbard 2001a).
The behavioral data reviewed here do not re-
ally offer direct evidence for or against either
model. Nor is it clear what kind of behav-
ioral experiment would actually allow a rig-
orous test of the models, since neither has
yielded predictions specific enough to permit
falsification. Since both the disinhibited feed-
back and anomalous cross-wiring hypotheses
are couched in terms of neural architectures,
a more fruitful approach might be to seek an-
swers from brain imaging and neurostimulation
techniques.
Evidence from Brain Imaging Studies
One early study conducted by Paulesu et al.
(1995) employed positron emission tomogra-
phy (PET) to examine cerebral activity in six
individuals who experienced colors when they
heard spoken phonemes or words. In the ex-
perimental condition the participants listened
Annals of the New York Academy of Sciences
to spoken words, whereas in a control task they
heard pure tones. In the spoken-word condi-
tion, synesthetes showed significantly greater
activation than did controls in several extras-
triate regions, including the superior occipital
lobe, bilateral parietal cortex, and the posterior
portion of the left inferior temporal cortex. This
pattern implies some contribution of higher
level associative brain areas, and perhaps lan-
guage centers, in phonemically driven synes-
thetic color experiences. Interestingly, there was
no differential activity within the human homo-
logue of area V4, located in the fusiform gyrus,
which in humans is assumed to be crucial for
color perception (Lueck et al. 1989; Zeki &
Marini 1998).
The results of the Paulesu et al. (1995) study
are intriguing because they imply a relatively
high level locus for synesthetic color experi-
ences. Unfortunately, it is difficult to draw firm
conclusions from the findings for two reasons.
First, the authors did not have participants en-
gage in a relevant behavioral task, so it is impos-
sible to determine what they actually perceived
during the scanning runs. Second, the baseline
task of listening to pure tones, while presum-
ably effective in not eliciting synesthetic colors,
did not control for the various linguistic and
cognitive processes engaged while listening to
words.
A similar limitation applies to the study
by Nunn et al. (2002), who used fMRI to
compare brain activity in synesthetes and
nonsynesthetic controls, using spoken words
versus pure tones as stimuli. In addition to
extensive activation of language areas, as
expected from their experimental and baseline
conditions, these authors found a significant
cluster of activity in the vicinity of area V4/V8
in the left hemisphere of synesthetes only. In-
terestingly, in a second experiment the authors
had participants view colored Mondrians and
found a similar left-sided V4/V8 cluster in
nonsynesthetic controls, but no such activity
for these chromatic stimuli in the synesthetes
(though the synesthetes did show a significant
color-related response in the right V4/V8
Mattingley: Attention, Automaticity and Awareness in Synesthesia
region). Based upon these findings, the authors
suggested that area V4/V8 in the left hemi-
sphere of synesthetes is no longer available for
processing real, chromatic stimuli because it
has been annexed for generating synesthetic
color experiences instead. Such modularity
of color processing has some appeal, since it
might explain why synesthetes rarely confuse
real and synesthetic colors, and why they
report experiencing both concurrently. On
the other hand, such radical “rewiring” of the
extrastriate visual system would be expected to
have measurable consequences for synesthetes’
ability to process real colors, but to date there
is no evidence to support this prediction.
A further feature of the fMRI study by Nunn
et al. (2002) is worthy of mention. It is often sug-
gested that the experiences of grapheme-color
synesthetes are akin to a vivid form of mental
imagery. Indeed, one recent study of a group
of 38 synesthetes found that self-rated visual
imagery, as indexed by the Vividness of Visual
Imagery Scale (VVIQ) (Marks 1973), was sig-
nificantly higher for the synesthetes than it was
for a group of nonsynesthetic controls (Barnett
& Newell 2008). Moreover, nonsynesthetes with
high scores on the VVIQ show increased ac-
tivation in visual cortex during imagery tasks
(Cui et al. 2007). It is therefore important to
know whether the patterns of brain activation
revealed in fMRI studies of synesthesia can be
distinguished from those one might expect to
see in nonsynesthetes who are instructed to
imagine colors. Nunn et al. (2002) addressed
this issue by training nonsynesthetes to asso-
ciate colors with words. Following training, the
participants underwent fMRI scans in which
they were instructed, in separate blocks, ei-
ther to imagine the color associated with each
spoken word presented over headphones or to
imagine the name of the color associated with
each word. The controls showed no differential
activity within V4/V8, unlike the synesthetes in
the word-versus-tone experiment, leading the
authors to conclude that the V4/V8 result in
synesthetes could not be due to color imagery.
Unfortunately, the null effect from the imagery
159
experiment is difficult to interpret, since the
nonsynesthetic controls received only a handful
of trials in which to learn the word−color asso-
ciations, whereas synesthetes presumably have
had a lifetime of such associations. Moreover, it
is impossible to rule out that the control partic-
ipants actually generated color images in both
the imagery task and the word-association con-
dition (the intended baseline), especially since
the training regime involved explicit pairings of
display colors and color names.
At least one other fMRI study examined
the neural correlates of color imagery in
synesthesia (Rich et al. 2006). During scan-
ning, synesthetes and nonsynesthetic controls
viewed pairs of grayscale photographs of fruits
and vegetables (Fig. 6). In the experimental
trials the participants’ task was to indicate
which of the two items, if viewed in their
natural state, would be darker in color (the
color−imagery condition). In the control trials
participants viewed the identical photographs
but now judged which of the pair, if viewed
in their natural state, would be larger in size
(the size−imagery condition). A comparison
between the two tasks revealed significant ac-
tivity in the right inferior occipital lobe, around
area V4, in both synesthetes and controls. This
occipital cluster was much larger in the synes-
thetes than the controls, perhaps implying a
greater capacity for color imagery in this group
(see also Barnett & Newell 2008). Interestingly,
in a separate task in which participants viewed
colored letters versus achromatic placeholders
that did not induce synesthesia, the synesthetes
showed significant activity in the left medial
lingual gyrus, but not area V4/V8.
Several other fMRI studies have examined
different aspects of synesthetic color experi-
ences, with varying results (Aleman et al. 2001;
Hubbard et al. 2005; Steven et al. 2006; Weiss
et al. 2001, 2005). Due to space constraints,
the findings of these studies will not be covered
here; instead the reader is referred to the excel-
lent review by Hubbard and Ramachandran
(2005). Before concluding this section, however,
it is worth mentioning two studies that have
160 Annals of the New York Academy of Sciences

Figure 6. Example stimuli and fMRI activations for the voluntary color imagery experiment
of Rich et al. (2006). (A) Example stimuli for the color imagery and size (baseline) tasks.
Participants were asked to judge which object would be darker in color or larger in size. (B)
Activity within the functionally defined ROIs for the color imagery versus baseline contrast.
Brain slices show regions of significantly greater activity for control participants (yellow) and
synesthetes (red). For controls significant activation was found in the right inferior occipital
lobe (x, y, z: 26, −90, −18; p = 0.016); for synesthetes significant activation occurred in
the right inferior occipital lobe (red; x, y, z : 34, −88, −20; p < 0.001) and right fusiform
gyrus (x, y, z: 30, −94, −26; p = 0.001). Coordinates in this and all subsequent figures
have been calculated using the MNI three-dimensional space.

provoked considerable interest for their novelty
and rigor.
Hubbard et al. (2005) conducted a study in
which six synesthetes and a group of matched,
nonsynesthetic controls completed two behav-
ioral tasks and a series of fMRI scans, with the
researchers’ aim being that of uncovering any
parametric changes in brain activity as a func-
tion of behavioral performance. The behav-
ioral tasks involved visual search for an embed-
ded figure composed of achromatic graphemes
(Hubbard et al. 2005) and identification of a pe-
ripheral grapheme under conditions of visual
crowding (Ramachandran & Hubbard 2001b).
The fMRI studies involved fine-grained retino-
topic mapping of striate and extrastriate ar-
eas, as well as a separate delineation of regions
that responded selectively to graphemic forms.
Mattingley: Attention, Automaticity and Awareness in Synesthesia
Hubbard et al. (2005) found that color-selective
area V4 was uniquely active in synesthetes rela-
tive to controls when participants viewed achro-
matic graphemes. Moreover, they showed that
V4 activity in the synesthetes tended to be
greater for those individuals who performed
better on the visual crowding task. The authors
interpreted their findings as consistent with the
notion of abnormal cross-activation between
grapheme-processing areas in the ventral tem-
poral lobe and the nearby color-selective area
V4, though fMRI data alone do not permit
firm conclusions regarding patterns of neural
connectivity.
Direct evidence for the cross-wiring hypoth-
esis has proven elusive, but a recent study by
Rouw and Scholte (2007) has gone some way
toward addressing this issue. They used diffu-
sion tensor imaging (DTI) to measure fractional
anisotropy in the brains of 18 grapheme-color
synesthetes and a group of nonsynesthetic con-
trols. In DTI magnetic resonance imaging is
used to measure the diffusion properties of wa-
ter molecules. Regions in which white matter
tracts are organized in a common direction will
tend to have higher fractional anisotropy val-
ues than those in which the white matter is
arranged more heterogeneously with respect
to direction. DTI thus provides a means of
assessing the relative coherence of white mat-
ter connectivity within and between brain ar-
eas. Rouw and Scholte (2007) performed DTI
scans in their synesthetes and controls, as well
as an fMRI study in which the same individuals
viewed achromatic graphemes selected to elicit
strong, intermediate, or weak color experiences
for the synesthetes. Significantly higher values
of fractional anisotropy, indicating a greater
strength of connectivity within white matter
tracts, were found for synesthetes versus con-
trols in two distinct areas of interest: the left
superior parietal cortex and the right inferior
temporal cortex, immediately adjacent to the
fusiform gyrus. Moreover, the authors observed
a significant correlation between the strength of
connectivity within the right inferior temporal
site and blood oxygen level-dependent (BOLD)
161
responses to the color-inducing graphemes in
the group of synesthetes. These data provide
the first evidence in favor of the hypothesis that
abnormal cross-activation of grapheme- and
color-processing areas gives rise to synesthetic
experiences of color.
Evidence from Event-Related
Potential Studies
Brain imaging techniques such as PET and
fMRI provide a reasonably good estimate of
the location of brain activity corresponding
with perceptual, cognitive, and motor states,
but are less sensitive to subtle changes in the
timing of involvement of brain regions. The
significant BOLD activity observed within ex-
trastriate cortex (including the color selective
area V4) in association with synesthetic color
experiences might be interpreted as evidence
for the involvement of this area in the initial
feedforward sweep of input from the primary
visual cortex. In principle, however, such acti-
vation might be due to later feedback signals
arising from temporal, parietal, or frontal as-
sociation areas. To address this interpretative
limitation, a small number of studies have used
scalp-recorded, event-related potentials to ex-
amine moment to moment changes in neural
activity associated with synesthetic experiences.
An early study by Schiltz et al. (1999) had 17
grapheme-color synesthetes view achromatic
letters and digits that appeared briefly on a
computer display. The researchers asked sub-
jects to respond to occasional predefined targets
(a subset of the letters and digits). They found a
significant enhancement in the P300 response
over frontal electrode sites in the synesthetes
compared with a group of nonsynesthetic con-
trols. This result is somewhat surprising in light
of the early perceptual effects that synesthetic
colors appear to have on behavioral perfor-
mance and considering the unique brain ac-
tivity associated with synesthetic experiences in
color-selective area V4.
A more recent ERP study was conducted
by Sagiv et al. (2006). Their grapheme-color
162
synesthete was shown letters colored congru-
ently or incongruently with her synesthetic ex-
periences. During the experiment she was in-
structed to monitor for the appearance of an
occasional target letter (an “I,” which was ex-
perienced as white) and to report the number
of times this target appeared at the comple-
tion of each block. Averaged ERPs showed a
significantly larger negative amplitude in the
N1/N170 component for congruent relative to
incongruent trials. This difference was appar-
ent for posterior parietal electrodes, and was
larger over the right hemisphere than the left.
These findings suggest a much earlier emer-
gence of synesthetic color experiences than sug-
gested by Schiltz et al. (1999), or at least a
much earlier onset for the congruency effect
that arises from such experiences.
Broadly similar findings were reported by
Beeli et al. (2008), who measured ERPs in a
group of 13 synesthetes and 13 nonsynesthetic
controls in response to spoken words and let-
ters presented over headphones. Their partic-
ipants performed a cued, one-back task to en-
sure they maintained attention on the stimuli
(though performance for the group was at ceil-
ing so it is impossible to know to what extent
the task actually taxed attentional resources). A
direct comparison of the averaged ERP data
for synesthetes revealed longer latencies for the
N1 and P2 components and a significantly re-
duced amplitude for the P2 component relative
to those of controls. Source localization sug-
gested stronger activity for synesthetes within
the posterior region of the left inferior tempo-
ral cortex, including area V4, and in the lat-
eral and medial prefrontal cortex, and these ef-
fects emerged as early as 122 ms after stimulus
presentation.
These findings are broadly consistent with
those of fMRI studies in which inferior tem-
poral regions were found to be active during
synesthetic color experiences (Hubbard et al.
2005; Nunn et al. 2002; Paulesu et al. 1995;
Rich et al. 2006). Moreover, the ERP data of
Beeli et al. (2008) imply that such color experi-
ences can emerge quite early in cortical process-
Annals of the New York Academy of Sciences
ing, perhaps after the initial feedforward sweep
of inputs from the primary visual cortex. A fur-
ther recent ERP study used contextual priming,
in which sentences used to invoke a particular
color concept (e.g., “The sky is. . .”) ended with
a grapheme (e.g., the number “7”) that elicited
a congruent or incongruent synesthetic color
(Brang et al. 2008). Averaged waveforms for
the synesthetes showed a significant congru-
ency effect for early ERP components, includ-
ing the N1 and P2, consistent with other rele-
vant findings (Beeli et al. 2008; Sagiv & Ward
2006), plus a significant congruency effect for
the N400 component, which is widely recog-
nized as providing an index of the extent to
which a meaningful stimulus matches a primed
context.
To summarize, data from ERP studies sug-
gest that differences attributable to synesthetic
colors can arise as early as 120 ms after the pre-
sentation of an appropriate auditory or visual
inducer. Later components, perhaps reflecting
a more conceptual level of analysis, have also
been found as a distinguishing feature of ERPs
in synesthetes versus controls. Future studies
will need to focus carefully on source localiza-
tion, which is a perennial problem with ERP
research. This could be achieved by combining
ERP and fMRI, as has been done in a recent
case study of an individual with digit−color
synesthesia (Kadosh et al. 2007). It will also be
important to characterize the influence of at-
tention on ERPs, since attention clearly exerts
a profound influence both on subjective reports
of synesthetic experiences, and on objective be-
havioral measures of the phenomenon.
Evidence from Transcranial Magnetic
Stimulation Studies
Despite numerous insights into the neural
underpinnings of synesthesia offered by fMRI
and ERP studies, these two approaches suffer
from a fundamental limitation, namely, that
they can only identify patterns of neural ac-
tivity that are correlated with behavior, rather
than regions that have played a causal role in
Mattingley: Attention, Automaticity and Awareness in Synesthesia
the phenomenon. To address this limitation two
recent studies have used transcranial magnetic
stimulation (TMS) to transiently disrupt corti-
cal areas thought to be involved in synesthe-
sia (see Chambers & Mattingley 2005). Both
studies examined the potential contributions of
subregions of the inferior parietal cortex, since
this area has been implicated, both directly and
indirectly, in attentional binding of visual form
and color in normals and neuropsychological
patients (Robertson 2003).
Esterman et al. (2006) tested two grapheme-
color synesthetes on a standard version of the
synesthetic Stroop task, in which participants
named the colors of congruent, incongruent,
and neutral letters. Participants received repet-
itive TMS (rTMS) for 8 min at 115% of motor
threshold, with the stimulating coil positioned
over the left or right angular gyrus (near the
junction of the intraparietal sulcus and trans-
verse occipital sulcus). Baseline performance
was measured in two control conditions: a pari-
etal “sham” condition, in which the coil was
oriented at 90◦ from the scalp so that the mag-
netic pulse was directed away from the brain;
and an active control, in which the coil was
positioned over the occipital pole, close to the
primary visual cortex (V1). Both synesthetes
showed a significant reduction in the magni-
tude of Stroop interference following rTMS
over the right angular gyrus relative to the size
of the effect following the sham treatment. This
reduction was not apparent following stimula-
tion of the left angular gyrus, or for stimulation
of area V1. The findings suggest that right pari-
etal stimulation transiently disrupts the neural
binding of form and color, thereby reducing
any perceptual or cognitive conflict between
the synesthetic and real colors on incongruent
trials.
A conceptually similar study was reported
by Muggleton et al. (2007), who tested five
grapheme-color synesthetes on the synesthetic
Stroop task. Their participants received rTMS
at 10Hz for 500 ms following the onset of
each target grapheme. Stimulation was deliv-
ered over one of two parietal sites in both the
163
left and right hemispheres: a region neighbor-
ing the parieto−occipital junction and a region
just anterior to this, roughly in the vicinity of the
angular gyrus location targeted by Esterman
et al. (2006). The cost of incongruently colored
graphemes was significantly reduced following
rTMS of the right parieto−occipital area, again
implicating the right inferior parietal lobe in the
anomalous binding of form and color in synes-
thesia. Closer inspection of the results, however,
indicates that stimulation of the right angular
gyrus and left parieto−occipital area also re-
sulted in a substantially reduced Stroop effect,
suggesting that the effect might be mediated via
a fairly widespread, bilateral network of cortical
areas.
Future studies using TMS should focus on
early visual areas, including V1, and elsewhere
within ventral and dorsal extrastriate pathways
to help determine where in the cortical process-
ing hierarchy synesthetic colors arise. It will also
be important to use time-locked TMS protocols
to reveal the fine-grained temporal dynamics
involved in the integration of visual form and
color.
Conclusion
In this article I have provided a selective
overview of recent research on the role of atten-
tion, automaticity, and awareness in the behav-
ioral and neural manifestations of grapheme-
color synesthesia. The picture that emerges is
that synesthetic color experiences are typically
triggered automatically, in the absence of vol-
untary effort. Synesthetic colors might help to
guide attention during visual search of achro-
matic displays, but there is no convincing evi-
dence to suggest they “pop out” like unique col-
ors in chromatic displays. On the other hand,
synesthetic colors can be strongly attenuated
by demanding secondary tasks or when induc-
ers must compete with one another for selec-
tion. Indeed, it seems likely that mechanisms
of attention play a causal role in synesthesia by
acting to bind, in an obligatory fashion, visual
164
representations of graphemic form and color.
Whether synesthetic colors can be triggered
without explicit awareness of the inducing stim-
ulus is still debated, but currently the evidence
suggests that inducers must be consciously per-
ceived to exert a measurable influence on be-
havior. These behavioral findings are mirrored
by data from brain imaging studies, which have
shown modulations of neural activity associated
with synesthetic color experiences within ex-
trastriate regions, including the color-selective
area V4. Transient disruption of the inferior
parietal lobule, a region widely recognized as
a key node in the attention network, can selec-
tively reduce the otherwise automatic binding
of form and color in synesthesia.
Clearly there are many questions yet to be
answered. For example, are all grapheme-color
synesthetes alike, or are there important indi-
vidual differences? In this article I considered
evidence for the proposed distinction between
“projector” synesthetes, for whom colors are
said to be experienced “out in the world,” and
“associator” synesthetes, who see colors in their
mind’s eye (Dixon et al. 2004). Although the
dichotomy has been widely used in the liter-
ature, recent evidence suggests that the man-
ner in which individuals use language to de-
scribe their color experiences can change over
time (Edquist et al. 2006). It has also been sug-
gested that the projector−associator distinction
reflects the engagement of different spatial ref-
erence frames for synesthetic color experiences,
rather than fundamentally different underlying
neural processes. The same caveats apply to
the proposed distinction between “lower” and
“higher” synesthetes (Ramachandran & Hub-
bard 2001a).
The danger faced by synesthesia researchers
is that we will be reduced to cataloguing new
and unusual variants of the condition, rather
than constructing scientific accounts for the
phenomena based upon currently accepted
principles of brain structure and function.
What is desperately needed is a model of synes-
thesia that can generate testable predictions
with respect to brain activity and behavior, and
Annals of the New York Academy of Sciences
that can be falsified by suitably designed exper-
iments. Arguably, no such model currently ex-
ists. Nevertheless, given the enormous progress
made in the last decade, there is reason for
optimism.
Acknowledgments
My sincere thanks to Anina Rich for stimu-
lating my interest in synesthesia and for her sub-
stantial contributions to much of the research
reviewed in this article. Thanks also to Mike
Dixon for his helpful comments on an earlier
version of the manuscript.
Conflicts of Interest
The author declares no conflicts of interest.
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