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Annu. Rev. Psychol. 2013. 64:49–75 Keywords

First published online as a Review in Advance on color, synaesthesia, imagery, connectivity, multisensory, spatial
June 29, 2012

The Annual Review of Psychology is online at Abstract

[Link]
Although synesthesia has been known about for 200 years, it is only in
This article’s doi: the past decade or so that substantial progress has been made in study-
10.1146/annurev-psych-113011-143840
ing it empirically and in understanding the mechanisms that give rise to
Copyright  c 2013 by Annual Reviews. it. The first part of the review considers the characteristics of synesthe-
All rights reserved
sia: its elicited nature, automaticity, prevalence, and consistency, and
its perceptual and spatial phenomenology. The second part considers
the causes of synesthesia both in terms of candidate neural mechanisms
and the distal influences that shape this: genetic differences in devel-
opmental synesthesia and plasticity following sensory loss in acquired
synesthesia. The final part considers developmental synesthesia as an
individual difference in cognition and summarizes evidence of its influ-
ence on perception, imagery, memory, art/creativity, and numeracy.

49
 PS64CH03-Ward ARI 15 November 2012 14:35

In fact, the trigger itself may be an internal

Contents representation that is not physically presented.
A sum such as 5 + 2 may be sufficient to trigger
INTRODUCTION . . . . . . . . . . . . . . . . . . 50
the color associated with 7 (Dixon et al. 2000),
HISTORICAL PERSPECTIVES
and synesthetic experiences of taste can occur
ON SYNESTHESIA . . . . . . . . . . . . . . 52
even if the inducing word is inaccessible, as in a
THE CHARACTERISTICS
tip-of-the-tongue state (Simner & Ward 2006).
OF SYNESTHESIA . . . . . . . . . . . . . . . 53
Of course to those who do have synesthesia, it
Internal Consistency of
seems equally remarkable that these kinds of
Associations . . . . . . . . . . . . . . . . . . . . 53
experiences aren’t shared by everyone.
The Prevalence of Synesthesia . . . . . . 54
A precise definition of what synesthesia is
The Automaticity of Synesthesia . . . . 54
(and what it isn’t) remains a matter of some
The Perceptual Reality of
debate. An uncontroversial starting point
Synesthesia . . . . . . . . . . . . . . . . . . . . . 55
would be to say that synesthesia is elicited.
Annu. Rev. Psychol. 2013.64:49-75. Downloaded from [Link]

Extension in Space . . . . . . . . . . . . . . . . . 56
Grossenbacher & Lovelace (2001) introduced
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THE CAUSES OF SYNESTHESIA . . 58

the term “inducer” to describe the stimulus
Proximal Causes: The Neural
that elicits the synesthesia and the term “con-
Mechanisms of Synesthesia . . . . . . 58
current” to denote the experience itself. Thus,
Distal Causes of Developmental
every type of synesthesia can be minimally de-
Synesthesia . . . . . . . . . . . . . . . . . . . . . 61
scribed as an inducer-concurrent pairing. For
Distal Causes of Acquired
instance, music-taste and grapheme-color de-
Synesthesia . . . . . . . . . . . . . . . . . . . . . 62
note types of synesthesia in which music elicits
Where Do the Specific
taste, and graphemes (units of written language)
Associations Originate? . . . . . . . . . 63
elicit colors. It is important to note that the
THE CONSEQUENCES OF
inducer is not substituted by the concurrent, so
SYNESTHESIA . . . . . . . . . . . . . . . . . . . 65
music-taste synesthetes continue to hear music
Perception . . . . . . . . . . . . . . . . . . . . . . . . . 65
as well as tasting it (Beeli et al. 2005). This start-
Imagery . . . . . . . . . . . . . . . . . . . . . . . . . . . 66
ing position enables us to distinguish synesthe-
Memory . . . . . . . . . . . . . . . . . . . . . . . . . . . 66
sia from seemingly similar phenomena such as
Art and Creativity . . . . . . . . . . . . . . . . . . 67
illusions (in which the inducer is, in some sense,
Numeracy . . . . . . . . . . . . . . . . . . . . . . . . . 68
misperceived) and hallucinations (a concurrent
Summary . . . . . . . . . . . . . . . . . . . . . . . . . . 68
without an inducer). A further generally agreed
CONCLUDING REMARKS . . . . . . . . . 68
defining property is the notion of automaticity,
by which it is generally meant that the con-
current is a (virtually) inevitable consequence
of encountering the inducer. Synesthetes have
INTRODUCTION very little control over the onset or content of
Synesthesia is a remarkable way of perceiving their experiences. It is in this sense, among oth-
the world. One attribute of a stimulus (e.g., its ers, that we can say that synesthesia is more akin
sound, shape, or meaning) may inevitably lead to perception than mental imagery. For those
to the conscious experience of an additional without synesthesia (nonsynesthetes), if we see
attribute. For example, the word “Phillip” may a green 5 then we can neither switch the green
taste of sour oranges, the letter A may be lumi- off nor change it to pink; but if we imagine a 5
nous red, and a C# note on the violin may be it could take any color or none at all. For those
a brown fuzzy line extending from left to right with synesthesia, their experiences resemble the
in the lower left part of space. The attributes former more than the latter. However, in other
do not always lie in different senses (e.g., a important respects synesthetic experiences
visually presented letter may trigger a color). resemble mental images more than percepts;

50 Ward
 PS64CH03-Ward ARI 15 November 2012 14:35

for instance, synesthetic visual responses to are needed. More controversially, it has been
music aren’t affected by shutting or moving the suggested that synesthesia-like experiences can
eyes. be induced by hypnosis (Kadosh et al. 2009) or
One of the things that a theory of synesthe- by expertise with sensory substitution devices
sia must explain is the pattern of inducers and in the blind (Ward & Meijer 2010). Again, the
concurrents that are found. The most common main controversy here is whether this relates
synesthetic inducers are linguistic in nature: let- in any way to the same causal mechanisms that
ters, digits, and words—particularly words that underpin conventional forms of synesthesia.
form part of a series (Simner et al. 2006). The One of the reasons why synesthesia is of in-
most common synesthetic concurrents are vi- terest to psychology in general is the assump-
sual in nature. This includes color and colored tion that it could inform “normal” models of
textures. It also includes spatial forms in which perception and cognition. Synesthesia should
sequences such as numbers or years are visu- be explicable in terms of modifications to mod-
alized as linear or convoluted landscapes (Sagiv els of sensory and cognitive development and
Annu. Rev. Psychol. 2013.64:49-75. Downloaded from [Link]

et al. 2006). These are also called number forms, brain function just like other acquired and de-
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calendar forms, etc., or number-space, time- velopmental conditions. At the very least, the
space synesthesia, etc. However, the range of existence of synesthesia invites psychologists to
possible inducers and concurrents is very large, reconsider their notions of what “normal” is.
with new combinations still being discovered. Synesthesia could be regarded as an example
This also leads to a puzzle as to what combina- of an individual difference in cognition: It is
tions of inducer and concurrent could reason- not only associated with unusual experiences
ably be labeled as synesthesia. With a very broad but also impacts on many aspects of function-
definition, any automatic association could po- ing (memory, numeracy, perception, personal-
tentially meet the criteria for being classed as ity) (see sidebar Famous Synesthetes).
synesthesia. This review starts by considering the his-
A common solution to this problem is to re- tory of scientific enquiry into synesthesia. The
strict the definition of synesthesia to percept- remainder of the review is divided into three
like concurrent experiences. Another solution main sections. First, the characteristics of synes-
would be to go beyond the superficial char- thesia are considered such as its prevalence,
acteristics and consider the underlying causes
(Simner 2012). The assumption of most con-
temporary researchers is that there will be a set FAMOUS SYNESTHETES
of underlying causal mechanisms that give rise
to synesthesia, and these will probably differ Some famous synesthetes include the philosopher Ludwig
from those causal mechanisms that lead some Wittgenstein (1889–1951; “the vowel e is yellow,” etc.; see ter
people to have, say, vivid visual imagery (al- Hark 2009); the composer Olivier Messiaen (1908–1992; his
though the mechanisms could also overlap in compositions were influenced by the color of musical keys;
interesting ways). There could also be multi- see Bernard 1986); the actor Geoffrey Rush (born 1951; col-
ple causal pathways that give rise to synesthe- ored numbers and days; see Astle 2007); the psychiatrist Eugen
sia. Synesthesia exists as both a developmental Bleuler (1857–1939; he published an influential book on synes-
condition (i.e., existing throughout the lifespan thesia before turning to his more famous work on schizophrenia;
with no known precipitating event) and as an ac- see Bleuler & Lehmann 1881); the physicist Richard Feynman
quired condition (i.e., as the result of a precipi- (1918–1988; he saw his formulae in color and wondered what
tating event such as acquired sensory loss). It re- they looked like to his students; see Feynman 1988); and the au-
mains a matter of debate as to whether these two thor Vladimir Nabokov (1899–1977; who described the letters
different distal causes have anything in common kzspygv as creating the spectrum from red to violet; see Nabokov
in terms of the underlying neural mechanisms 1967).
or whether two completely separate accounts

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automaticity, consistency, and its similarity to the opposite of Daltonism (or color blindness),
veridical perception. The main focus of this part and gave synesthesia its first name, hyperchro-
of the review is on developmental synesthesia. matopsia. Although not explicitly articulated
However, the second section, which considers as such, we could regard this as implying ad-
the causes of synesthesia, discusses how both ditional forms of color receptors in the eye.
acquired and developmental synesthesia may Cornaz sparked the first significant interest in
arise. The final section considers how develop- synesthesia, leading to new cases being reported
mental synesthesia is related to other aspects of ( Jewanski et al. 2011). However, his eye-based
cognitive functioning. For instance, I consider explanation never gained widespread credence.
how it could be linked to enhanced memory Evidence against his account included obser-
and, in some cases, impaired numeracy. vations in new cases that colors can be trig-
gered by the idea of a letter alone (Chabalier
1864) and that synesthetic colors coexist with
HISTORICAL PERSPECTIVES the real text color and therefore resemble an
Annu. Rev. Psychol. 2013.64:49-75. Downloaded from [Link]

ON SYNESTHESIA associated perception rather than a mispercep-

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The first documented case of synesthesia, that is tion/illusion (Perroud 1863). The term pseu-
generally accepted as such, dates back to 1812. dochromesthesia was coined to reflect this new
Georg Sachs was an Austrian doctor who, for his understanding (Chabalier 1864), and more de-
medical dissertation, submitted an account of tailed neuroscientific accounts were proposed
himself and his sister on the topic of albinism—a during the 1870s. One suggestion was that
condition that affected both of them. It is within two different sensory nerves might stimulate
this thesis that he describes his own synesthe- each other if they cross or are near each other
sia, giving detailed examples of colors for num- (Nussbaumer 1873), or, in more phrenologi-
bers and letters and other series (for an English cal terms, that the cerebral “organs” of music,
translation, see Jewanski et al. 2009). color, and language lie next to each other and
Although we know of no earlier docu- may interconnect (Lusanna 1873). Another in-
mented case, the idea that color may not solely teresting account is that the sensory specializa-
be derived from light/vision was treated with tion of the nerves is not present from birth but
open-mindedness in the intellectual circles is learned via experience, and synesthetes may
of the Enlightenment period. The scientist fail to make this developmental differentiation
Robert Boyle (1627–1691) mentions the case of of the senses (Ranke 1875).
a congenital blind man who could tell the color During the mid-nineteenth century there
of cloth by touch (see Larner 2006). Although was general acceptance that synesthesia was
unlikely to be a true case of touch-color not pathological despite consensus that it
synesthesia ( Jewanski et al. 2009), the case reflected an alteration in underlying biology
illustrates the fact that eminent figures were ( Jewanski et al. 2011). Although not considered
willing to entertain such ideas. Isaac Newton dysfunctional, nor was it typically considered
(1642–1727) himself believed that there was functional. Perhaps the first systematic attempt
a law of physics that connected the colors of to describe synesthesia as a functional entity
the spectrum with seven tonal intervals of the came from the British polymath, Francis
musical octave (Pesic 2006). Galton (1822–1911). In 1878, Galton read an
By the nineteenth century, color associa- obituary by the son of a famous calculating
tions were recognized as a phenomenon of bi- prodigy and engineer, George Bidder (1806–
ology rather than physics. However, one early 1878), in which both father and son were
debate centered on whether synesthesia orig- described as seeing “mental pictures of figures
inated from the eyes or brain. Cornaz (1848) and diagrams” when dealing with numbers.
rediscovered the Sachs case, interpreted it as Bidder Jr. provided Galton with a drawing

52 Ward
 PS64CH03-Ward ARI 15 November 2012 14:35

of his own number form, sparking Galton’s The twentieth century took synesthesia on
interest in mental imagery and its possible role a rather different journey away from its earlier
in intellectual ability (Burbridge 1994). For medical heritage. The notion of synesthesia
Galton, phenomena such as number forms and became incorporated in cultural movements
color associations were regarded as an individ- of the time in the visual arts (e.g., the work of
ual difference in mental imagery, and he did Kandinsky), literature, music, and spirituality
not seek to attach a special name to it. While (see Dann 1998). As such, synesthesia began
he asserted its biological and hereditary nature, to coexist as both a biological entity and
he also acknowledged that number forms are as a cultural metaphor for the expression
in some sense constructed during childhood of intersensory correspondence. Moreover,
(he likens it to nest building in animals) and synesthesia was not an area of research that fit-
only retained into adulthood if they are useful ted well with the new psychological movement
to the user (Galton 1883). Examples of spatial of the time, namely behaviorism. In the behav-
forms are shown in Figure 1. iorist tradition, synesthesia was little more than
Annu. Rev. Psychol. 2013.64:49-75. Downloaded from [Link]

In the last two decades of the nineteenth a learned association (e.g., Howells 1944), and
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century, there was a rapid growth in the num- so by the mid-twentieth century synesthesia
ber of different varieties of synesthesia and the had stopped being a puzzle for many psychol-
number of new cases that were reported. For ogists and had become trivial. It would take
instance, Bleuler & Lehmann (1881) document several more decades to reclaim some scientific
76 synesthetes, including the first examples credibility.
of pain-color, taste-color, and smell-color.
Nor were the reports limited to color as the
concurrent, and nor were the concurrent expe- THE CHARACTERISTICS
riences necessarily percept-like in nature. For OF SYNESTHESIA
instance, personifications (associating genders Although the general characteristics of synes-
and personalities to, say, graphemes) were thesia were described well over 100 years
typically grouped alongside color associations ago, there has been a strong emphasis in the
(Galton 1883). As such, it was clear that the contemporary literature on developing testable
most popular terminology of the day, namely criteria that might distinguish synesthesia from
“colored hearing,” did not do justice to what phenomena such as imagery or cross-sensory
was being documented. The term “sunaisthe- metaphor (such as “sharp cheese”). The sec-
sis” (from which synesthesia derives) had been tions below consider the prevalence of synes-
used since antiquity. For instance, Aristotle thesia, its internal consistency and automaticity,
used it to refer to rather different phenomena and its perceptual and spatial characteristics.
such as shared feelings—similar to modern no-
tions of empathy (Flakne 2005). It had also been
used in medical contexts to denote things such Internal Consistency of Associations
as referred sensations, the photic sneeze, and Efforts in the 1980s to study synesthesia ob-
bodily sensations from hearing nails scratching jectively, rather than just as case descriptions,
a blackboard (e.g., Vulpian 1866), and it was concentrated on measures of test-retest con-
from this context that the word synesthesia was sistency (e.g., Baron-Cohen et al. 1987). In a
coined as an umbrella term to accommodate typical test, synesthetes would be given a list of
the recent proliferation of cases that had been inducers and asked to generate their concurrent
reported (e.g., Calkins 1895). The 1890s notion experience, whereas controls would be asked to
of synesthesia is broadly similar to that used freely associate a concurrent. This procedure is
today, although the links to referred sensations then repeated, often weeks or months later, and
(and so on) have been largely dropped. test-retest consistency is calculated. Using this

[Link] • Synesthesia 53
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method, grapheme-color synesthetes are often Table 1 Current estimates of the prevalence
found to be around 80% to 100% consistent and of different types of synesthesia
controls around 30% to 50% consistent (e.g., Type of synesthesia Prevalence (%)
Mattingley et al. 2001). The method can be Spatial forms (e.g., 2.2–20.0
adapted for other concurrent experiences such calendar→space)a,b
as taste/flavor (Ward & Simner 2003). In re- Days→colorc 2.8
cent years, there has been a shift away from us- Vision→touch (mirror 1.6
ing verbal descriptions of color concurrents to touch)d
using visual color pickers and having the test Grapheme (letter + 1.4
and retest occurring within the same session number)→colorc
(Eagleman et al. 2007). Month→colorc 1.0
Although measures of consistency have been People→colorc 0.4
extremely influential, it goes without saying Music→colorc 0.2
that having a very high consistency isn’t nec- Taste→shapec 0.2
Annu. Rev. Psychol. 2013.64:49-75. Downloaded from [Link]

essarily uniquely diagnostic, and some element

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of first-person report of synesthesia is used in a

Brang et al. (2010), b Sagiv et al. (2006), c Banissy et al.
conjunction with this measure. In line with (2009a), d Simner et al. (2006).
this, controls trained to have highly consistent
word-color associations do not activate color- types of developmental synesthesia have a
sensitive regions of the brain (area V4) as much common cause. For instance, spatial forms are
as synesthetes in response to those same words five times more prevalent in grapheme-color
(Nunn et al. 2002). synesthetes than in those lacking color concur-
rents, suggesting that these types of synesthesia
are not independent (Sagiv et al. 2006). The
The Prevalence of Synesthesia
most prevalent type of nonvisual synesthesia
One method of assessing prevalence has been appears to be so-called mirror touch or vision-
to place an advertisement in a newspaper touch synesthesia, in which observing touch
or magazine and divide the number of re- to another person triggers tactile sensations on
spondents who both report synesthesia and the body of the perceiver (Banissy et al. 2009a).
pass an internal consistency measure by the
circulation figure. These studies have tended
to find a prevalence of around 1 in 2,000, The Automaticity of Synesthesia
with a preponderance of female synesthetes as Aside from consistency, the most common way
high as 5:1 (Barnett et al. 2008a, Baron-Cohen of establishing the authenticity of synesthesia
et al. 1996, Rich et al. 2005). However, this has been to use a synesthetic version of the
should be construed as a very conservative Stroop test. The original version of this test
prevalence rate given that one can’t draw firm uses color names printed in congruent or
conclusions about people who fail to respond incongruent colors (e.g., RED printed in
to the advertisement. A more reliable method either red or green ink; see MacLeod 1991). In
is to screen a large sample of the population synesthetic versions, color names are replaced
for reports of synesthesia and supplement this by the inducer. If a synesthete perceives the
with an objective measure (typically internal letter “A” as red, then he/she will be faster at
consistency). Table 1 shows the prevalence naming the printed color of a grapheme if it is
of various types of synesthesia obtained using congruent with the synesthesia (“A” printed in
this method. This method produced higher red) relative to when it is incongruent with it
prevalence rates and a female:male ratio close (“A” printed in green; Mattingley et al. 2001).
to 1:1. Most of these synesthetes have multiple The effect is found for other types of synesthe-
types of synesthesia, suggesting that different sia, such as music-taste (Beeli et al. 2005) and

54 Ward
 PS64CH03-Ward ARI 15 November 2012 14:35

music-color (Ward et al. 2006a). Related tasks this larger group as associators. As such, it is
have also been used for spatial forms by show- possible that different types of synesthetes may
ing a slower response time when the spatial make somewhat different demands on the per-
arrangement of externally presented stimuli ception/attention network, and there is some
(such as months, numbers) is incompatible with evidence to support this idea.
the spatial arrangement of the form (e.g., Sagiv In terms of psychophysical studies, synes-
et al. 2006, Smilek et al. 2006). The synesthetic thetic and real colors interact under binocular
Stroop effect does not prove that synesthetic rivalry and can bias the direction of apparent
colors are truly seen rather than just known (as motion (Kim et al. 2006). Thus, synesthetic col-
an overlearned association). Indeed, controls ors can affect other aspects of perception (which
can be trained to show comparable interference suggests that they are not just verbal associa-
effects, but trained controls still differ from tions). Another line of evidence has come from
synesthetes on measures such as functional visual search tasks. In normal perception, the
magnetic resonance imaging (fMRI) (Elias time taken to detect a target is fast, and indepen-
Annu. Rev. Psychol. 2013.64:49-75. Downloaded from [Link]

et al. 2003) and conditioned physiological dent of the number of items to search, if the tar-
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responses to color (Meier & Rothen 2009). get can be discriminated from distractors on the
When attention to the grapheme is reduced, basis of color—a phenomenon known as pop-
the synesthetic Stroop effect is also reduced out (e.g., Treisman 1988). Would the same hold
(Mattingley et al. 2006), and the Stroop effect true for a color that is not physically present,
is abolished if the grapheme is presented such as synesthetically induced colors? Some
subliminally followed by a supraliminal color case reports have suggested that this might be
patch (Mattingley et al. 2001). This suggests so (e.g., Palmeri et al. 2002, Smilek et al. 2001),
that the degree of automaticity of synesthesia but this is not replicated at the group level (e.g.,
is a function of attention to (and awareness of ) Edquist et al. 2006). In those cases that did re-
the inducer. This issue is returned to in the port an effect, it was qualitatively different from
next section. pop-out: for instance, by being dependent on
the number of distractors (Palmeri et al. 2002)
or dependent on initial fixation location (Laeng
The Perceptual Reality of Synesthesia et al. 2004). This is consistent with the view
Synesthesia can be construed as a conscious ex- that attention to a grapheme is needed to elicit
perience in the absence of the normal precon- a synesthetic color, whereas veridical colors can
scious sensory stimulation and therefore has guide visual search preattentively.
the potential to inform theories about con- A task conceptually related to visual search
scious perception. Psychophysical and func- is the embedded figures task of Ramachandran
tional imaging studies have attempted to es- & Hubbard (2001). They presented synesthetes
tablish which perceptual processes are involved with visual arrays of graphemes (for one second)
in the synesthetic experience. Most of the ev- in which some graphemes could be grouped to-
idence here has come from grapheme-color gether into shapes (e.g., a triangle). The task was
synesthesia. It should be noted at the outset to report the shape from four alternatives, and
that the experiences of synesthetes are het- their two synesthetes outperformed controls on
erogeneous. For some grapheme-color synes- this task. Hubbard et al. (2005a) and Ward et al.
thetes, the colors are seen on the surface of (2010) replicated the finding but noted that the
the grapheme itself, and these individuals have effect was variable across synesthetes and fell
been referred to as projectors (Dixon et al. short of true pop-out levels based on perfor-
2004). Other grapheme-color synesthetes claim mance with real color. Importantly, Ward et al.
to see the colors elsewhere (e.g., on an exter- (2010) obtained trial-by-trial phenomenologi-
nal or internal screen) or just to know the col- cal reports and were able to link these to per-
ors, and Dixon et al. (2004) have referred to formance. For these briefly presented stimuli,

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many synesthetes reported no color at all. For undermined by this research). With regard to
synesthetes who did report colors (and these the visual ventral stream, it would be reason-
were more likely to be projectors), only some able to conclude that color concurrents involve
of the letters were perceived as colored, and higher regions (V4 and beyond), with little ev-
the more letters that were colored, the better idence behaviorally (Hong & Blake 2008) or
they performed. Thus, synesthetic colors may neurally (Rouw et al. 2011) that lower regions
facilitate perceptual grouping, but not indepen- such as V1 are implicated in the color expe-
dently of attention and varying according to the rience itself (although they could still be im-
spatial location of the colors. plicated in synesthesia in other ways; Barnett
A more comprehensive account of the neu- et al. 2008b). In other modalities, there is func-
ral basis of synesthesia is given below in the tional imaging evidence that concurrent experi-
context of the causes of synesthesia. However, ences of synesthetic touch are linked to activity
in light of the present discussion, it is worth- within somatosensory cortices (Beauchamp &
while considering activity in one particular re- Ro 2008, Blakemore et al. 2005). The evidence
Annu. Rev. Psychol. 2013.64:49-75. Downloaded from [Link]

gion, namely the color-sensitive region in the that synesthetic flavor experiences are linked to
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fusiform cortex (V4), and its possible link to gustatory cortex is more equivocal ( Jones et al.
concurrent color experiences. Typically, stud- 2011).
ies that have investigated this have attempted
to locate the region functionally by compar-
ing veridical colors against grey-scale equiva- Extension in Space
lents with fMRI and then assessing whether the In veridical perception, linking spatial informa-
same region responds during the presence of tion from different senses requires a series of
an achromatic inducer that elicits synesthetic computations that take into account informa-
color. Area V4 has been shown to be activated tion about the orientation of the sensory recep-
by synesthetic color when the inducer consists tors in space (e.g., Morgan 2003). For example,
of speech (Nunn et al. 2002) or an achromatic in order to decide whether a seen object is lo-
grapheme (e.g., Hubbard et al. 2005a). How- cated to the left, front, or right of space, one
ever, not all studies have reported this (e.g., cannot simply read off the location of the ob-
Hupé et al. 2012). Areas more anterior to V4 ject on the retina. One would also need to take
have been shown by various studies to be more into account whether the eyes (and the head)
active during synesthetic experiences of color, are themselves oriented to the front, the left, or
and this has been linked to other aspects of the right. This suggests a hierarchy of spatial
processing such as color retrieval or cascaded processes starting from the sensory receptors
feedback (for a review, see Rouw et al. 2011). themselves and progressing ultimately to a spa-
Van Leeuwen et al. (2011) provide evidence tial representation of the external world that is
that different patterns of functional connectiv- independent of the observer.
ity between these more anterior regions and Synesthetic concurrents tap into these same
V4 may discriminate between projectors and spatial maps. Consider this quote from Galton
associators. (1883): “The diagram of the numerals has
In summary, with regard to the perceptual roughly the shape of a horseshoe, lying on a
reality of the synesthetic concurrent, most of slightly inclined plane with the open end to-
the evidence has come from experiences of wards me; zero is in front of my left eye. When
color. The empirical evidence suggests that I move my eyes without moving my head, the
synesthetic colors are not just like real colors diagram remains fixed in space and does not
in terms of either behavior or neural basis, al- follow the movement of my eye. When I move
though the evidence shows that synesthetes do the head, the diagram unconsciously follows
indeed differ from controls on a number of mea- the movement.” What Galton’s participant
sures (i.e., the existence of synesthesia is not is describing is a number form that lies in a

56 Ward
 PS64CH03-Ward ARI 15 November 2012 14:35

head-centered spatial reference frame. Not of spatial reference frames provides one exam-
all spatial forms are like this: Some are fixed ple of how characteristics of synesthesia can
relative to the body rather than head, and still be understood in terms of differences within a
others are reported to lie only on an “inner normative model of perception. Other object-
screen” or “mind’s eye” that is not defined by centered examples include projecting colored
egocentric coordinates (Smilek et al. 2006). auras onto faces or projecting colors onto the
Some synesthetes can vary the perspective from fingers when counting (see Ward 2008). Body-
which the form is viewed ( Jarick et al. 2009). and head-centered reference frames would in-
With respect to spatial forms, one can say that clude seeing colors inside the mouth when eat-
there are two levels of spatial relationship: an ing (Downey 1911) or projecting synesthetic
internal structure that determines the spatial tastes into the mouth (Ward & Simner 2003).
relationship of the elements (e.g., numbers) to Some sound-vision synesthetes experience the
each other, and a second level that specifies the vision as emanating from the sound source (ob-
spatial relationship(s) between the form itself ject centered), others in front of their body
Annu. Rev. Psychol. 2013.64:49-75. Downloaded from [Link]

and the observer. In terms of neural mech- (body centered), and still others on an inner
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anisms, Eagleman (2009) speculates that the screen (Goller et al. 2009). When touch is
former may be part of the (right-lateralized) the concurrent (e.g., mirror-touch synesthesia),
ventral visual stream of object perception; then it is necessarily experienced on the body,
the latter may involve regions around the but even here there are two spatial subtypes
intraparietal sulcus known to be involved in (Banissy et al. 2009a). For some synesthetes,
coding egocentric space (Tang et al. 2008). seeing touch to a left cheek may elicit touch
These kinds of differences are by no means on their left cheek (an anatomical spatial rela-
limited to spatial forms. Consider grapheme- tionship), whereas for others it may be felt on
color synesthesia. So-called projectors can be the right cheek (a reflected or specular spatial
understood in terms of the concurrent experi- relationship).
ence lying in an object-centered frame of ref- In acquired synesthesia, Armel &
erence (the object, in this case, is the printed Ramachandran (1999) report a case of touch-
word). Other grapheme-color synesthetes de- vision synesthesia arising after blindness.
scribe seeing the colors on a body-centered Synesthetic visual sensations were projected
frame of reference (e.g., the colors are seen on onto the spatial location of the touched body
a screen that is a fixed location relative to the part (e.g., hands) irrespective of where the body
body, sometimes even inside the body itself ), part was located in space. They measured the
and still others see it on a nonegocentric imag- intensity of tactile stimulation required to in-
ined screen (Ward et al. 2007). These differ- duce visual photisms and noted that they were
ences are not well captured by the umbrella easier to induce (i.e., lower threshold) when
term “associator,” and there is evidence that the hand was in front of the person rather than
these finer spatial distinctions manifest them- behind the head. This suggests a body-based
selves in terms of both behavior (Ward et al. spatial reference frame that additionally incor-
2007) and neural substrates (van Leeuwen et al. porates information about gaze and/or head
2010). orientation. Acquired audio-visual synesthesia
At least five different spatial reference frames arising from partial loss of vision due to optic
can be observed across the range of synesthe- nerve injury is linked to photisms that appear
sias: object centered, head centered, body cen- in the blind scotoma of the synesthete’s visual
tered, imaginal centered (or nonegocentric), field, that is, an eye-centered spatial reference
and eye centered. All of these are known to ex- frame ( Jacobs et al. 1981). There are no known
ist in normal cognition from single-cell record- examples of an eye-centered frame of reference
ings (Morgan 2003) or spatial impairments such in developmental synesthesia consistent with
as neglect (Halligan et al. 2003), and the use this primarily involving higher visual processes.

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Finally, there are some synesthetes who these differences are typically assumed to lie at
claim to know the color (or other concurrent) the cortical level, reflecting the complex nature
but deny perceiving it. These individuals could of the inducer/concurrent. Beyond this starting
be classed as having a nonspatial (and indeed position, there is far less agreement as to how
a nonperceptual) form of synesthesia. The dis- these differences in connectivity might be real-
tinction between knowing and perceiving has ized. Bargary & Mitchell (2008) provide a use-
parallels with Block’s (e.g., 2005) notion of ac- ful taxonomy of candidate neural mechanisms
cess versus phenomenal consciousness (briefly, in terms of a 2 × 2 space of theories. One factor
the former is reportable and the latter experi- is whether the inducer and concurrent are con-
ential). It could also reflect a resolved synesthe- nected directly or indirectly. This is reminis-
sia in which the perceptual components have cent of the debate in the multisensory process-
faded over time but associative components are ing literature as to whether unimodal sensory
retained. regions influence each other directly (feedfor-
ward only) or indirectly by converging (feed-
Annu. Rev. Psychol. 2013.64:49-75. Downloaded from [Link]

forward) onto some common hub that then in-

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THE CAUSES OF SYNESTHESIA fluences the input regions via feedback activity.
When considering the causes of synesthesia, it Both types of direct and indirect mechanism are
can be helpful to make a distinction between known to exist in the normal brain during mul-
proximal and distal (or ultimate) causes. For tisensory processing, and the dominance of one
instance, in developmental synesthesia we can type of mechanism over the other may reflect
say that the unusual experiences are caused the timing, the task, or the nature of the stim-
by functional and structural differences within uli (e.g., Driver & Noesselt 2008). The second
the brain. This is the proximal cause. The orthogonal factor is the nature of the connec-
distal cause, in this instance, would be the role tivity differences themselves: structural or func-
of genetic (and environmental?) differences tional. Structural differences are assumed to re-
in creating these brain-level differences.1 flect extra connections in synesthetes relative to
The challenge for research in this area is to controls, whereas functional differences are as-
join up these different levels of explanation: sumed to reflect pathways that are common to
from genes, to brain structure/function, to synesthetes and nonsynesthetes alike but such
cognition and unusual conscious experiences. that these routes are normally inhibited in the
This section begins by considering the neural typical brain yet are active (i.e., disinhibited) in
basis of synesthesia. It then examines the distal synesthetes.
causes of both developmental and acquired Structural differences are found in the brains
synesthesia. Finally, it considers the origins of of people with developmental synesthesia. Dif-
the particular associations themselves. fusion tensor imaging shows greater white mat-
ter organization in synesthetes relative to con-
trols in several brain regions (Rouw & Scholte
Proximal Causes: The Neural 2007). There are also differences in gray matter
Mechanisms of Synesthesia density (both cortical thickening and thinning)
Candidate neural mechanisms of synesthesia all in synesthetes relative to controls (Banissy et al.
have something in common insofar as they are 2012, Weiss & Fink 2009). One current chal-
believed to reflect differences in connectivity lenge for researchers is to link these differences
relative to the neurotypical brain. Moreover, to the actual characteristics of synesthesia be-
cause many of the structural changes are not in
the regions that process the inducer and/or con-
1
current (the affected regions are considered in
One could even push the levels of explanation further back
to the evolutionary origins of these genetic differences (see more detail below). As such, these studies have
Brang & Ramachandran 2011). not struck a definitive blow against functional

58 Ward
 PS64CH03-Ward ARI 15 November 2012 14:35

accounts. The functional versus structural di- may bias the pairing of inducers and concur-
chotomy can also be construed as two extreme rents in synesthesia. Ramachandran & Hubbard
end points, and it is possible that both are in- (2001) proposed that grapheme-color synesthe-
volved to different degrees. Faster-acting func- sia may be particularly prevalent because re-
tional changes (e.g., disinhibition) may initi- gions involved in color perception (V4) and let-
ate structural changes, as has been documented ter/digit recognition lie adjacent to each other
in the neuroplastic changes following blind- in the fusiform cortex (and this affords oppor-
ness (Pascual-Leone et al. 2005). Functional ac- tunities for connectivity). This basic idea has
counts also offer a way of explaining both de- been extended to other types of synesthesia (for
velopmental and acquired synesthesia within a a review, see Hubbard et al. 2011). Consider
single mechanism (Grossenbacher & Lovelace the difference between grapheme-color synes-
2001). But there is no necessary reason why de- thesia and lexical-gustatory synesthesia (Ward
velopmental and acquired synesthesia should be et al. 2005). When grapheme-color synesthetes
explicable by the same neural mechanism. see (or hear) a word, the color depends on its
Annu. Rev. Psychol. 2013.64:49-75. Downloaded from [Link]

With regard to the issue of whether inducer- spelling: A word such as PHOTO may be col-
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concurrent connectivity is direct or indirect, ored by the first letter (P), or sometimes the
Bargary & Mitchell (2008) argue that the uni- vowel (O), or, less frequently, as a sequence
directional nature of synesthesia (graphemes of five colors. (Even in Chinese, the colors of
trigger colors but not vice versa) favors the words depend on their constituent characters
direct, feedforward-only models. However, and radicals; Simner et al. 2011.) But when a
there is evidence of some implicit bidirection- word is experienced as a taste/flavor, the taste is
ality; for instance, judging the size of colored unrelated to the spelling; rather, similar sound-
lines is biased by the presence of number-color ing words tend to taste the same (Ward et al.
associations, suggesting that the color can 2005). There is no cognitive reason why the
activate numerical representations (e.g., Cohen color of words should depend on written lan-
Kadosh et al. 2007). It is also conceivable that guage but the taste of words should depend on
the directness of the mapping varies according spoken language. These differences do make
to the type of synesthesia or even between sense when one considers the neural architec-
individuals with putatively the same type of ture of the brain.
synesthesia. In mirror-touch synesthesia, there The parietal lobes are involved in number
is a rather complex spatial transformation processing (and possibly other ordinal se-
between the inducer (seeing a body part quences) and spatial processing, and it has been
touched) and concurrent (feeling touch) that is suggested that differences in connectivity here
suggestive of an indirect inducer-to-concurrent may underpin the spatial form of synesthesia
route (Banissy et al. 2009a). Grapheme-color (Hubbard et al. 2005b). Not all types of synes-
synesthesia would be a stronger candidate for thesia can be readily accounted for by adjacency
a direct route, given that they are anatomically between inducer and concurrent, but nor is ad-
adjacent (Ramachandran & Hubbard 2001). jacency the sole mode of connectivity. A more
But there is also some evidence that this could serious limitation of the adjacency principle is
be variable between synesthetes depending on that it offers no account as to why spatial forms
the spatial reference frame, which may depend should be more prevalent than grapheme-
on indirect connectivity via parietal regions color synesthesia, or why grapheme-color
(van Leeuwen et al. 2011). synesthesia should be more prevalent than
It has been shown that dense connectiv- lexical-gustatory synesthesia. One needs to
ity between adjacent and nearby regions (with make further assumptions to account for this:
sparser long-distance connectivity) is an opti- for instance, by relating to regional difference
mal configuration for information transfer and in gene expression or by assuming there are
minimizing wiring (Sporns et al. 2004), and this other differences in visual/spatial perception

[Link] • Synesthesia 59
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a that lead to vision being more prevalent as a

Intra-parietal sulcus (IPS) concurrent (for which there is some evidence).
Motor/premotor and parieto-occipital area
60 Finally, it was noted above that many of
the structural and functional brain differences
z-axis (Talairach coordinates)

40 between synesthetes and controls do not lie

in regions implicated in the processing of the
20 inducer or concurrent. In particular, regions
in the frontal and parietal lobes have been con-
0 sistently implicated. Activation maxima on the
lateral surface of the frontal and parietal lobes
–20 are depicted in Figure 2a,b. Within the pari-
etal lobes, two areas have been highlighted: the
–40 Left ventro-lateral Left intraparietal sulcus (IPS) on the lateral surface
prefrontal cortex
(Broca’s area) and the precuneus/retrosplenial region on the
Annu. Rev. Psychol. 2013.64:49-75. Downloaded from [Link]

medial surface (these regions themselves can

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60 40 20 0 –20 –40 –60 –80 –100

be subdivided, but possible subdivisions aren’t
y-axis (Talairach coordinates)
considered here). The IPS region is associated
with increased gray matter in grapheme-color
b
Intra-parietal sulcus(IPS) synesthesia (Weiss & Fink 2009) and is acti-
and parieto-occipital area Motor/premotor
60 vated in fMRI (relative to control participants)
z-axis (Talairach coordinates)

when graphemes are presented (Weiss et al.

40 2005). Transcranial magnetic stimulation over
a nearby region disrupts the synesthetic Stroop
20 effect (Esterman et al. 2006, Muggleton et al.
2007). The functional role of this region in
0 synesthesia is assumed to be in binding, that
is, in linking the grapheme and color into a
–20
spatially localized perceptual whole. It may
also be implicated in attentional aspects of
synesthesia. The functional role of the pre-
–40 Right Right dorso-lateral
prefrontal cortex cuneus/retrosplenial region in synesthesia is
unclear, although the region itself is implicated
–100 –80 –60 –40 –20 0 20 40 60 in imagery across a variety of sensory domains
y-axis (Talairach coordinates) as well as self-directed and internal thought
Figure 2 more generally (Cavanna & Trimble 2006).
(a,b) The filled circles (•) show loci of maximal activity from nine functional Increased white matter in this region has been
magnetic resonance imaging (fMRI) studies reviewed by Rouw et al. (2011) on found in grapheme-color synesthesia (e.g.,
the lateral frontal and parietal cortex (activity elsewhere, including the ventral Hupé et al. 2012), and functional activity
visual stream, is not shown here but is discussed in the text). All studies in this region (relative to controls) has been
involved color as the concurrent and linguistic inducers (letters, numbers,
found in a diverse range of types of synesthesia,
words) presented either visually or as speech. The open circles (o) are from an
fMRI study of number forms (Tang et al. 2008). All contrasts compare including lexical-gustatory (contrasting synes-
synesthetes against a nonsynesthetic baseline (either control participants thetes against controls when presented with
and/or stimuli that do not induce synesthesia). The “+” indicates locations in inducers; Jones et al. 2011) and associating
which grapheme-color synesthesia has been shown to be disrupted by genders to graphemes (contrasting inducing
transcranial magnetic stimulation (Esterman et al. 2006, Muggleton et al. 2007,
versus noninducing stimuli; Amin et al. 2011)
Rothen et al. 2010). All coordinates are based on the Talairach system
[converted from the Montreal Neurological Institute (MNI) system if in addition to color (contrasting inducing
necessary], and the locations of the sulci are only approximate. versus noninducing stimuli; Nunn et al. 2002).
Finally, regions in the lateral frontal cortex

60 Ward
 PS64CH03-Ward ARI 15 November 2012 14:35

(ventral, dorsal, and motor) have been found to absence of axons between the regions. Bor-
be activated by synesthetes relative to controls ders are determined developmentally by dif-
(Rouw et al. 2011), but this may reflect differ- ferences in gene expression, and a breakdown
ences in task demands for synesthetes relative in border formation could be one explanation
to controls (who have an “extra” dimension to of synesthesia. Finally, there may be a failure
process). There is little evidence of structural to prune synaptic connections between regions
changes in prefrontal regions (Rouw et al. that are transiently connected in early devel-
2011), consistent with this suggestion. opment. This mechanism may be partly un-
der genetic control. Further progress in this
important area will depend on isolating can-
Distal Causes of Developmental didate genes in synesthetes and then exploring
Synesthesia the normal role of these genes in mouse models
Developmental synesthesia has long been of brain development.
known to run in families ( Jewanski et al. 2011). Failure to prune normally transient synaptic
Annu. Rev. Psychol. 2013.64:49-75. Downloaded from [Link]

However, the particular associations do not run connections has been a particularly influential
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in families. Family members don’t agree on the idea in the synesthesia literature. This derives
colors of graphemes any more than do unre- from the suggestion that synesthesia is a normal
lated synesthetes (Barnett et al. 2008a). Nor phase of early development that, in most peo-
do the patterns of inducers and concurrent ple, drops out during childhood (e.g., Maurer
appear to be strongly familial. For instance, & Mondloch 2006). Infants are born with about
synesthetes with lexical-gustatory synesthesia (a 150% the level of adult synapses, and synap-
rare type) tend to have relatives with the more tic elimination is a normal part of learning and
common color-based synesthesia (Ward et al. development (Huttenlocher & Dabholkar
2005). Genetic differences appear to convey a 1997). In animal studies, for instance, there are
susceptibility to synesthesia but don’t specify transient connections from auditory to visual
the presence/absence of synesthesia in a de- areas that diminish developmentally (e.g., In-
terministic way. One early suggested genetic nocenti et al. 1988), but one possibility is that
mechanism was X-linked dominant inheritance such pathways may be retained in synesthesia.
(Baron-Cohen et al. 1996). However, this has Infants show far less evidence of functional spe-
not been supported by further research on in- cialization and show somewhat different pat-
heritance patterns and genetic linkage studies terns of multisensory influence relative to older
(Asher et al. 2009, Tomson et al. 2011). These children (e.g., Maurer & Mondloch 2006).
studies confirm genetic differences between af- Moreover, many of the same associations found
fected and unaffected members of the same in adult synesthetes are found in infants (see dis-
families, and the current evidence suggests the cussion below). However, so far there is no di-
involvement of multiple genes, but with no can- rect evidence that links early differences in mul-
didates on the X chromosome. tisensory processing with later development
Bargary & Mitchell (2008) discuss three pos- of synesthesia, and virtually nothing is known
sible kinds of molecular mechanisms that could about the characteristics of synesthesia itself
mediate between gene level and connectivity prior to the age of 6 years. Given that synesthe-
levels of explanation. First, there are genes sia runs in families, future research could com-
involved in axon guidance. Genetic polymor- pare the infants/children of synesthetes against
phisms in molecules mediating axon guidance those lacking a familial susceptibility.
could result in different patterns of connectiv- Considering children ages 6 years and up,
ity. Second, there might be disruption of bor- grapheme-color synesthesia appears to be no
der formation. Borders are regions of the brain more prevalent in the early school years than
in which activity is not propagated between in adulthood (Simner et al. 2006, 2009a).
adjacent regions due to either inhibition or the However, there is evidence of developmental

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changes during this time. Child synesthetes are Acquired loss of sensory input can some-
far less consistent in their grapheme-color asso- times lead to symptoms of synesthesia. In all
ciations than are adult synesthetes—fewer than the examples considered below there is a clear
half of the letters and numbers are consistently pattern, namely, that the impaired modality
colored (Simner et al. 2009a). (Note: This level acts as the synesthetic concurrent. Induced vi-
of consistency is still significantly higher than sual experiences following blindness are the
that of the average child asked to freely asso- most frequently documented, particularly ac-
ciate colors for graphemes.) But when tested a quired auditory-visual synesthesia (see Afra
year later, the number of consistent associations et al. 2009), although the prevalence of these
in the child synesthetes (but not the controls) experiences is unknown. Acquired touch-vision
increased. This suggests that synesthesia is not synesthesia following blindness has also been
just a residual form of infantile perception, but documented (Armel & Ramachandran 1999).
rather continues to develop during childhood. Loss of somatosensory input through amputa-
However, it is unclear what is developing (the tion or paralysis is also commonly associated
Annu. Rev. Psychol. 2013.64:49-75. Downloaded from [Link]

concurrent experiences themselves or the sta- with phantom pain, and acquired synesthetic
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bility of the inducer-concurrent associations), sensations of touch may be induced through vi-
and it is unclear what precedes it developmen- sual observation of touch in the location of the
tally. Even in adulthood, new synesthetic as- phantom limb (e.g., Ramachandran et al. 1995)
sociations can be formed. For instance, if a or even by observing touch on another person
synesthete learns musical notation (Ward et al. (e.g., Goller et al. 2011). In the case of hear-
2006b) or learns a second written language that ing and deafness, tinnitus (phantom sounds) can
uses different printed characters (e.g., Witthoft arise after damage to the peripheral auditory
& Winawer 2006), the new associations are de- system, and it has been suggested that tinnitus
rived from old ones (e.g., on the basis of having is evoked by somatosensory inputs (Cacace et al.
similar shape, meaning, or phonology). Need- 1999), that is, acquired somatosensory-auditory
less to say, this also points to an important role synesthesia. After having cochlear implants fit-
for the environment in synesthesia insofar as ex- ted, prelingually deaf people report tactile ex-
posure to literacy is culturally dependent. We periences to sounds (McFeely et al. 1998), that
know nothing about how synesthesia is mani- is, acquired auditory-somatosensory synesthe-
fested in illiterate cultures. sia. In addition to sensory loss following damage
to the sensory organs and peripheral nerves, ac-
quired synesthesia has been reported after dam-
age to the thalamus (e.g., Ro et al. 2007). Ro
Distal Causes of Acquired Synesthesia et al. (2007) document a patient with left-sided
Synesthesia can be acquired in two general somatosensory weakness following a thalamic
ways: by sensory loss (e.g., blindness) or phar- lesion who developed auditory-somatosensory
macologically [e.g., lysergic acid diethylamide synesthesia.
(LSD)]. Superficially, acquired synesthesia has In terms of a general mechanism for ac-
different characteristics from developmental quired synesthesia arising from sensory loss, the
forms of synesthesia. The inducers tend to be sensory cortex associated with the concurrent
simple sensory stimuli rather than learned and experience remains functional but is denied
meaningful stimuli (e.g., graphemes, days of its previous primary inputs. It may therefore
the week). This could reflect the participation come to rely more on existing multisensory
of different brain regions or pathways in ac- inputs (cortico-cortical or from subcortical
quired versus developmental synesthesia, and projections), which then take on the role of the
it could also be related to the different times inducer. The onset of synesthesia can be as soon
of onset that afford different opportunities for as days or as long as one to two years (see Afra
reorganization. et al. 2009). This suggests the operation of two

62 Ward
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different kinds of neural plasticity (Pascual- origins in cross-sensory correspondences that

Leone et al. 2005): fast-acting removal of are found in typical (i.e., nonsynesthetic) brains.
inhibition from existing pathways and slower-
acting formation of new connections. One Audio-visual associations. Despite a signifi-
thing that needs to be accounted for in acquired cant amount of individual difference in the pre-
synesthesia is why some patients with sensory cise way in which music and other sounds are
loss experience it, but most (apparently) do “seen” by synesthetes, there are some common
not. This raises the possibility of individual underlying principles. High-pitch sounds, rela-
differences in reorganization or differences tive to low-pitch sounds, are perceived by synes-
in the way that transient phenomena (e.g., thetes as being brighter, smaller, and higher
spontaneous hallucinations) stabilize into in space (e.g., Fernay et al. 2012, Ward et al.
synesthesia. It would be interesting to know 2006a). Temporal aspects of music and sounds
whether there is any relationship between tend to be mapped on to visual space such that,
the genes that give rise to developmental at least in Western synesthetes, music is seen to
Annu. Rev. Psychol. 2013.64:49-75. Downloaded from [Link]

synesthesia and the tendency for some (but progress from left to right (Ward et al. 2008a).
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not others) to acquire synesthesia following a Changes in timbre (for a given pitch) tend to af-
coincidental sensory loss. fect saturation more than lightness (Ward et al.
Finally, the temporary induction of synes- 2006a).
thetic experiences following consumption of Studies with the general population show
hallucinogenic drugs (LSD, mescaline, psilocy- cross-modal biases in speed of responding to
bin) offers the clearest example of a fast-acting one dimension while ignoring an irrelevant di-
mechanism (minutes to hours; e.g., Hartman mension in the other modality (for a review, see
& Hollister 1963) that presumably arises via Marks 2004). This has been reported for pitch
unmasking of pre-existing pathways. Marek & and lightness (high is lighter), pitch and size
Aghajanian (1998) propose a neurobiological (high is smaller), and pitch and vertical position
mechanism of disinhibition leading to synes- (high pitch is spatially higher). Similar results
thesia based on LSD’s known action on sero- have been found in infants and young children
tonergic receptors on inhibitory interneurons (Walker et al. 2010), and luminance-pitch asso-
in the cortex. ciations are also found in chimpanzees (Ludwig
et al. 2011).

Where Do the Specific Letter-color associations. Synesthetes tend

Associations Originate? to agree on letter-color associations more than
Most synesthetes have little or no insight into would be expected by chance; thus A tends to
where their particular associations originate. be red, B tends to be blue, C tends be yellow,
They don’t tend to come from alphabet books O tends to be white, and so on (Barnett et al.
(Rich et al. 2005), although a few cases are 2008a, Rich et al. 2005, Simner et al. 2005).
known to have been biased by these (e.g., There does not appear to be a single factor that
Witthoft & Winawer 2006). If the associa- determines this, but rather at least three con-
tions reflected chance contingencies in the straining factors: grapheme frequency, color
environment (e.g., having a friend named names, and letter shape. With regard to the lin-
Anne who likes to wear red), then one would guistic frequency of the grapheme, Simner et al.
expect minimal agreement among synesthetes. (2005) considered color categories and found
But synesthetic associations are not chance a correlation between grapheme frequency and
occurrences. The subsections below consider color term frequency (i.e., common letters tend
evidence from various types of synesthesia that to have common colors such as red and yellow,
the associations are nonrandom. It is claimed, whereas uncommon letters have colors such as
instead, that synesthetic associations have their purple or brown). Beeli et al. (2007) considered

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RGB space (rather than color categories) and If asked to decide which of two numbers is
found inverse correlations between grapheme larger (e.g., 3 or 6), performance is facilitated
frequency and saturation and lightness (i.e., un- if the smaller number is lighter and the larger
common graphemes are darker and less satu- number is darker than when it is vice versa
rated). With regard to color names, it is found (Cohen Kadosh & Henik 2006), and these
that R = red, Y = yellow, B = blue, and G = number-luminance associations are mediated
green more than would be expected by chance by the parietal lobes (Kadosh et al. 2008).
(e.g., Simner et al. 2005). Finally, it has been
noted that similarly shaped graphemes (as de-
termined by letter confusability matrix) tend to Spatial forms. Spatial forms, at least from
evoke similar colors (Brang et al. 2011a). synesthetes in the West, tend to run in a left-
When nonsynesthetes are asked to freely to-right direction (e.g., Eagleman 2009, Jonas
associate color terms to graphemes, they don’t et al. 2011, Sagiv et al. 2006). This is assumed
show an effect of grapheme frequency, but to reflect a cultural bias from reading direc-
Annu. Rev. Psychol. 2013.64:49-75. Downloaded from [Link]

they are biased by color names such that tion. Associations between space and number
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G = green, etc. (Simner et al. 2005). There are found in nonsynesthetes too, with a general
are also shape-based influences. Spector & left-to-right association with magnitude on re-
Maurer (2011) asked children of different ages sponse time measures (Fias & Fischer 2005).
to associate colors to letters. In one condition In addition to direction, the shape of spa-
they were shown a Perspex shape (e.g., O) and tial forms reflects the internal structure of the
asked which colored box they would open to sequence. Number forms often bend or break
find another one (given two choices such as at decade boundaries—10, 20, etc. (Sagiv et al.
white/black or green/red). In another condi- 2006). Months are more likely to be repre-
tion they were told the letter name (e.g., O) sented in a circular arrangement than num-
and asked which box it might be in. Preliterate bers, reflecting their cyclical nature (Brang
children (ages 3 to 4) chose nonrandomly et al. 2010). Handedness may further influ-
when given a shape (e.g., O = white, X = ence the direction of rotation, with left-handers
black, C = yellow) but randomly when given a arranging the months counterclockwise and
letter name. More recently, it has been shown right-handers clockwise (Brang et al. 2011b).
that infants have nonrandom shape-color Spatial forms involving the alphabet tend to
associations (Wagner & Dobkins 2011). bend/break around phrases in the alphabet song
( Jonas et al. 2011).
Number-color associations. For number- Makioka (2009) provides a computational
color synesthetes, single-digit numbers tend model of number forms that could also be
to progress from light to dark and saturated extended to other sequences. Variations in an
to unsaturated as the number increases in input layer (corresponding to number size) are
magnitude (Beeli et al. 2007).2 Associations mapped onto a two-dimensional output layer
between hue and specific numbers have not (corresponding to space) using a self-organizing
been noted. Beyond 10, numbers tend to be map. Similarities in the inputs (e.g., the number
colored by the constituent digits (so 12 would 5 is more similar to 6 than it is 8) give rise to its
be some combination of the colors for 1 and sequential structure (i.e., 5 is spatially nearer to
2). The association between magnitude and lu- 6 than 8 in the output layer). Bends/breaks in
minance is also found in adult nonsynesthetes. the form reflect a combination of noise in the
model and a decomposition of the input layer
into separate representations of tens and units.
2
In light of the discussion above it is also worth noting Circular arrangements of months are predicted
that number size and linguistic frequency of numbers are
highly correlated, so it is unclear which of these dimensions is by this model if the input representation of
relevant. January and December are similar to each other

64 Ward
 PS64CH03-Ward ARI 15 November 2012 14:35

(despite being at opposite ends of the verbal

sequence). Model 1: synesthesia as directly causal
In summary, synesthetic associations are not
random, and nor are they likely to be based Structural and Cognitive
functional brain Synesthesia
on chance occurrences within the environment. differences
differences
Instead they have a rich structure that is based
on the multisensory correspondences found in
the typically developing brain from a young Model 2: synesthesia as a colorful sideshow
age. Different causal mechanisms, discussed
above, would then act on these associations so
that for individuals with synesthesia they are Synesthesia
consciously perceived (internally stable, etc.),
Structural and
whereas in most others they exert indirect functional brain
effects. differences
Annu. Rev. Psychol. 2013.64:49-75. Downloaded from [Link]
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Cognitive
differences
THE CONSEQUENCES OF
SYNESTHESIA
Figure 3
What are the consequences of having synesthe-
How might synesthesia be related to individual differences in cognition? Two
sia for other aspects of cognition such as mem- different models are outlined here, in which synesthesia either has a direct role
ory, creativity, or numeracy? The answer to this or is a co-occurring symptom linked to a mediating variable (such as
question is of general interest to psychologists structural/functional brain differences).
for a number of reasons. First, the question has
obvious relevance to those with an interest in both synesthesia and to a propensity towards
individual differences. Second, the question is linking seemingly unrelated concepts and
relevant for the broader theoretical issue as to ideas—in short, creativity. This would explain
how perception interfaces with (or underpins) why the apparently useless synesthesia gene
other aspects of cognition. has survived in the population.” Disentangling
In thinking about how synesthesia may these two scenarios is not straightforward be-
be linked to other cognitive differences (both cause the chain of causality is not amenable to
weaknesses and strengths), one could entertain experimental manipulation. However, one pre-
a number of possibilities. These are sum- diction is that if synesthesia is directly involved,
marized in Figure 3. The first possibility is then the affected domain should be relatively
that synesthesia exerts a direct influence on circumscribed to synesthesia-related material.
cognition. That is, the synesthetic associa-
tions themselves (e.g., 5 = red, 6 = salty)
participate in other cognitive domains and Perception
affect cognitive performance in those domains. Synesthetes with grapheme-color synesthesia
The second possibility is that changes in perform better on tests of color perception
cognitive performance are an outcome of the relative to controls (Banissy et al. 2009b, Yaro
more general structural/functional differences & Ward 2007). Moreover, there does appear
known to exist in the brains of synesthetes. to be a relationship between the modality of
For example, this position is advocated by the synesthetic concurrent and the modal-
Ramachandran & Hubbard (2003, p. 58) with ity in which perception is enhanced. Thus,
respect to creativity: “[S]ynesthesia causes mirror-touch synesthetes have enhanced tactile
excess communication among brain maps. acuity (but normal color perception), whereas
Depending on where and how widely in the grapheme-color synesthetes show the opposite
brain the trait was expressed, it could lead to profile (Banissy et al. 2009b). This suggests

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a direct relationship between synesthesia and verbalizing strategies in daily life in synesthetes
differences in perceptual ability: It is not simply with number forms (verbalizing weaker) rela-
the case that synesthetes are better perceivers in tive to controls.
some general sense. However, it is unclear from
these data whether we can conclude that the
differences in perception are a consequence or Memory
a cause. Barnett et al. (2008b) speculate that the Some notable cases of exceptional memory abil-
differences in perception may predate the emer- ities have been linked to synesthesia. Luria
gence of synesthesia because differences are (1968) describes the case of S (now known to be
found in early visual pathways (early in terms Solomon Shereshevskii), who performed feats
of developmental maturity and in terms of of extraordinary memory to entertain public au-
visual hierarchy). From electroencephalogram diences. S also reported having multiple forms
recordings, they suggest that grapheme-color of synesthesia in which, for instance, a voice
synesthetes may have a perceptual enhance- may elicit colors, textures, and tastes. Daniel
Annu. Rev. Psychol. 2013.64:49-75. Downloaded from [Link]

ment of the parvocellular system (sensitive to Tammet (2006) gives an autobiographical ac-
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color, high contrast, high spatial frequency) count of both his synesthesia and his prodigious
relative to magnocellular system (sensitive to memory. For instance, he was able to memo-
motion, low contrast, low spatial frequency). rize pi to over 20,000 decimal places and re-
Interestingly, synesthetes show both increased ported doing so by using his synesthesia: Each
cortical thickness in a color-related region (V4) digit had a characteristic color, shape, and tex-
and reduced cortical thickness in a region re- ture that, when placed together as a sequence,
lated to motion perception (V5/MT) (Banissy created an undulating landscape that he could
et al. 2012). As such, the existing evidence read off from his mind’s eye. In formal testing,
suggests that perceptual changes in visual Tammet appeared to rely less on conventional
synesthesia aren’t limited to the concurrent ways of chunking stimuli that were found to
(color), nor are they as broad as the sense itself benefit controls (Bor et al. 2007). This suggests
(vision), but rather may affect an intermediate that he relies on his own strategies for encod-
systems level. ing material mnemonically (e.g., based on his
synesthesia). Tammet also has Asperger’s syn-
drome, and one suggestion is that the (presum-
Imagery ably coincidental) co-occurrence of synesthesia
People with grapheme-color synesthesia re- and Asperger’s leads to a particularly beneficial
port more vivid visual imagery than matched synergy for memorizing.
controls (Barnett & Newell 2007). Synesthetes Both Tammet and S came to the attention
with spatial forms also report greater use and of researchers because of their abilities, but is
vividness of visual imagery, but not spatial im- synesthesia associated with enhanced mem-
agery (Price 2009). The latter may imply that ory in unselected cases? The answer to this
forms are more like visual objects than maps question is yes, but not all aspects of memory
(as suggested by Eagleman 2009). There is also are affected equally (e.g., Rothen & Meier
evidence that grapheme-color synesthetes are 2010a, Yaro & Ward 2007). In grapheme-color
faster at visual imagery, at least when it involves synesthesia, one of the most consistent findings
letters (Spiller & Jansari 2008). It is possible is that long-term memory for verbal material
that enhancements in imagery closely parallel (such as free recall of word lists) is better
enhancements in perception, given the close than for matched controls (e.g., Radvansky
relationship between perception, imagery, and et al. 2011, Yaro & Ward 2007). This may be
visual memory (e.g., Slotnick et al. 2012). Con- expected on the basis that, for these individuals,
sistent with this, Seron et al. (1992) report words are encoded visually (e.g., as a pattern
a greater imbalance between visualizing and of colors) in addition to verbally/semantically.

66 Ward
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Indeed, Radvansky et al. (2011) showed that the creative industries, relative to national es-
synesthetes were less influenced by semantic timates of numbers employed in that sector, in
factors than were controls in free recall, sug- both Australia (Rich et al. 2005) and the United
gesting that they may rely more on perceptual Kingdom (Ward et al. 2008b). Moreover, the
encoding. Synesthetes with calendar forms prevalence of grapheme-color synesthesia is
(i.e., experiencing months and years spatially) higher in arts students compared to those
are better at retrieving autobiographical events from other subjects (Rothen & Meier 2010b),
from date cues, and they are more accurate at and synesthetes (as a mixed group) are more
dating news events (Simner et al. 2009b). This likely to be engaged in visual arts as a hobby
suggests that they can use their internal synes- (Rich et al. 2005, Ward et al. 2008b). This is
thetic representations to give an organized particularly true of auditory-visual synesthetes,
structure to certain memoranda. for whom music and other sounds trigger a
The evidence above suggests a close link very rich experience of colors, shapes, texture,
between the nature of the synesthesia and the and movement. These individuals are far more
Annu. Rev. Psychol. 2013.64:49-75. Downloaded from [Link]

nature of memory enhancement. However, likely than other synesthetes to play musical
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the evidence is not quite as clear-cut as this. instruments and be engaged in visual art, and
First, not all verbal memory tasks show an Ward et al. (2008b) suggest that the synesthesia
enhancement. Grapheme-color synesthetes provides a particularly vivid source of artistic
do not show enhanced performance on digit inspiration for these individuals (i.e., Model 1
span (e.g., Rothen & Meier 2010a) or on in Figure 3). In addition, an enhanced color
remembering the position of digits in a matrix sense could lead to a more general interest in
(e.g., Yaro & Ward 2007). This suggests that it the visual arts in synesthetes with color as the
is not just the material (and whether it triggers concurrent.
synesthesia) but how one has to process it Artistic engagement is only one way in
mnestically. Associating an item to a position which creativity can be expressed. Creativity as
(in a sequence or matrix) requires forming a a cognitive ability can be assessed by tests that
specific contextual association, whereas free tap an ability to form novel and adaptive (i.e.,
recall can be done at the item level itself. Thus, useful/meaningful) associations. Ramachan-
spatial and sequential memory may not be dran & Hubbard (2003) suggest that the kinds
enhanced in grapheme-color synesthesia even of brain changes that give rise to synesthesia
if the participating stimuli induce synesthesia. may also give rise to creativity as a cognitive
Second, the memory enhancement is also found ability. Ward et al. (2008b) gave two tests
for some visual material. Synesthetes have bet- of creativity to a group of synesthetes and
ter memory for colors (Yaro & Ward 2007) and controls: a test of divergent thinking (gener-
do particularly well on tasks such as learning ating alternative uses for objects) and a test of
to pair colors with abstract shapes (Rothen & convergent thinking (finding a conceptual link
Meier 2010a). Gaining a better understanding between three words). The synesthetes did
of precisely which type of visual stimuli do outperform the controls, but only on the test
and do not give rise to enhanced memory is an of convergent thinking (even after entering
important challenge for future research. It is education as a regressor). This provides some
possible that the changes in perceptual ability support for Ramachandran & Hubbard’s
(described above) also affect the ability to (2003) suggestion, but not complete support
encode certain types of material into memory. (as their theory predicts that all aspects of
creativity are enhanced). It would be important
to extend and replicate the result using other
Art and Creativity tests and measures (including in the nonver-
It has been observed that both the hobbies and bal domain) and taking into account other
occupations of synesthetes are skewed toward potential confounds (e.g., vocabulary level).

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Numeracy Summary
Rich et al. (2005) asked a large sample of synes- At the start of this section, I outlined two broad
thetes (n = 192) to report their strengths and ways in which synesthesia could be linked to
weaknesses in an open-ended way and then individual differences in cognition. Synesthesia
compared this to a control group. The list of could be directly related to individual differ-
strengths included memory, languages, written ences in cognition, or the relationship may
and verbal communications, and art. The list of be more indirect (e.g., both synesthesia and
weaknesses included sense of direction and co- individual differences in cognition coemerge
ordination/balance. Interestingly, mathematics from a common mediating factor). One way in
was described both more often as a strength and which to explore this is to determine how spe-
more often as a weakness (relative to controls). cific or general the profile of differences is. In
Daniel Tammet (2006), described above, would the domain of perception and imagery there is
be an example of a synesthete who is able to use evidence of enhanced abilities, and the benefits
his synesthesia to perform extraordinary calcu- are found to be relatively specific to domains
Annu. Rev. Psychol. 2013.64:49-75. Downloaded from [Link]
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lations (e.g., 375 ). As with his general mem- related to the synesthesia (e.g., to color per-
orizing ability, he may use his synesthesia to ception and visual imagery in grapheme-color
store a large number of numerical facts to aid in synesthesia) but are not limited to stimuli that
calculation. act as inducers/concurrents. For memory, it is
In what ways could synesthesia also be dis- undoubtedly the case that synesthetic associ-
advantageous to numerical cognition? There ations (to words, etc.) can be used to directly
is some evidence of numerical difficulties in develop mnemonics and may bias the way
synesthetes with number forms (although not that information is encoded. However, there
necessarily in grapheme-color synesthesia). is also evidence of a broader influence (e.g.,
Specifically, this group is significantly slower grapheme-color synesthetes have enhanced
with single-digit multiplication but performed visual memory for certain achromatic stimuli;
normally on subtraction (Ward et al. 2009). Rothen & Meier 2010a). For psychomet-
Different arithmetical operations rely on ric measures of creativity, further evidence is
different cognitive mechanisms, with multipli- needed to determine how and when synesthesia
cation normally relying more on retrieval of affects performance on these tests. However,
verbal facts (e.g., 5 × 5 = 25) and subtraction the evidence that synesthesia (in general) is
relying on online calculation (e.g., Dehaene linked to artistic inclinations is more robust
et al. 2003). Ward et al. (2009) argued that and may be even more common in those who
synesthetes with number forms might tend have very rich synesthetic experiences (such as
to be biased toward a slower visuo-spatial those linked to visualized music; i.e., a direct
strategy (rather than faster verbal retrieval) and link between synesthesia and artistic tenden-
that this impacts multiplication most because cies). Although cognitive difficulties have been
the strategy is less suited to that operation. self-reported in synesthesia, these have been
Other studies have shown that synesthetes with under-researched. So far, the clearest evidence
number forms have worse performance on comes from a link between number forms and
numerical tasks when the numbers are spatially some difficulty on certain tests of numerical
incompatible with their internal representation cognition (i.e., a direct link between synesthetic
(e.g., Gertner et al. 2009, Sagiv et al. 2006). phenomenology and cognitive outcome).
Although spatial processes have a role to play in
nonsynesthetic numerical cognition, the pres- CONCLUDING REMARKS
ence of a number form may impair the ability Research into synesthesia is at a crossroads. The
to represent numbers in a flexible manner ac- onus is no longer on researchers to establish
cording to task demands (Gertner et al. 2009). its existence—this is beyond doubt. However,

68 Ward
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the nature of synesthesia remains harder to pin argued in this review that we can use synes-
down. I suggest that it straddles perception, thesia as a tool to understand cognition more
imagery, and memory, and this is broadly generally. Synesthesia provides a window into,
consistent with contemporary theories that among other things, how conscious experiences
acknowledge shared mechanisms across these are created; how genetic differences affect brain
domains. A greater challenge for researchers development and cognition; and how different
is to place synesthesia within the bigger ways of perceiving the world may impact
picture. Are we studying synesthesia in order upon memory, numeracy, artistic inclinations,
to understand synesthesia itself ? This is not an and so on. It is likely to be on these fronts
unreasonable research agenda given how many that the value of synesthesia research will be
people are estimated to have it. However, I have realized.
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SUMMARY POINTS
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1. Empirical research into synesthesia shows it to be a genuine phenomenon that cannot

be reduced to associative learning, vivid imagination, or metaphorical thinking. It can be
thought of as an alternative mode of perceiving (and conceptualizing) the world.
2. Synesthetic experiences are elicited (i.e., triggered by a stimulus) and automatic (at least
when the inducer is attended), and many definitions also incorporate the notion that they
are percept-like in nature. They tend to be internally consistent (at least in adults) and
spatially extended (utilizing various spatial reference frames).
3. It is generally believed that the patterns of inducer-concurrent pairings found in synesthe-
sia are derived from the same principles that determine cortical connectivity in general;
for instance, favoring connectivity between adjacent regions.
4. There is evidence of structural differences in the brains of developmental forms of synes-
thesia (both in white matter and gray matter), but these differences are by no means
limited to regions of the brain that process the inducer or concurrent.
5. Acquired synesthesia may occur as a result of sensory loss (such that the lost modality
acts as the synesthetic concurrent), and this may reflect both disinhibition of existing
pathways and slower-acting structural changes in connectivity.
6. The particular associations themselves are not random but structured. They may reflect
a universal tendency to associate certain attributes together (e.g., pitch with luminance,
shape with color) that is present from a very early age. This suggests a link between
synesthesia and normal modes of multisensory processing, but whereas synesthetes
directly experience these associations (e.g., 3 is bright yellow), they tend to exist as
declarative knowledge or implicit rules in other adults (e.g., as revealed in response time
measures).
7. Synesthesia conveys various cognitive benefits in perception, imagery, and memory.
These benefits tend to be relatively circumscribed (e.g., not all aspects of percep-
tion are improved) and bear some relation to the type of synesthesia experienced.
They also lead to possible difficulties (e.g., in numeracy), although these are less
explored.

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FUTURE ISSUES
1. How does synesthesia manifest itself in cultures that are very different from our own?
To what extent is synesthesia culturally situated despite having a biological basis?
2. What is the role of the hypothetical synesthesia genes in normal brain development, and
how is their action altered in people with synesthesia?
3. Why is it that only some individuals with sensory loss (blindness, deafness, amputation)
go on to acquire synesthesia? What is different about their brains or their environment?
4. What insights can synesthesia give us into consciousness—its function, its neural basis,
or the hard problem of why things feel the way they do?
5. What specific cognitive problems are associated with synesthesia, and in what ways are
they directly related to the presence of synesthesia itself ?
Annu. Rev. Psychol. 2013.64:49-75. Downloaded from [Link]
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DISCLOSURE STATEMENT
The author is not aware of any affiliations, memberships, funding, or financial holdings that might
be perceived as affecting the objectivity of this review.

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RELATED RESOURCES
• The author’s webpage is located at [Link]
• The Synesthesia Battery is an online resource for synesthetes to test themselves and for re-
searchers to obtain color consistency measures ([Link]
• The U.K. Synaesthesia Association ([Link] and the American
Synesthesia Association ([Link] organize annual meetings for re-
searchers, synesthetes, and others with a personal or professional interest.
• An email discussion list (the Synesthesia List) is moderated by Sean Day of Trident Techni-
cal College (Charleston, South Carolina). Details can be found at [Link]
[Link]

[Link] • Synesthesia 75
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Annu. Rev. Psychol. 2013.64:49-75. Downloaded from [Link] ARI 15 November 2012 14:35
Access provided by Monash University on 11/25/20. For personal use only.

Figure 1
Spatial forms for numbers and time may either be colored or not. The days of the week were produced by
Tessa Verrecchia (with permission). The number form is taken from Richard E. Cytowic, from Cytowic &
Eagleman (2009), Wednesday is Indigo Blue: Discovering the Brain of Synesthesia, Cambridge, MA: MIT Press.
(Permission is granted to copy, distribute, and/or modify this document under the terms of the GNU Free
Documentation License.)

[Link] • Synesthesia C-1

 PS64-FrontMatter ARI 15 November 2012 14:20

Annual Review of
Psychology

Volume 64, 2013 Contents

Prefatory
Shifting Gears: Seeking New Approaches for Mind/Brain Mechanisms
Michael S. Gazzaniga p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 1
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Biological Bases of Behavior

The Endocannabinoid System and the Brain
Raphael Mechoulam and Linda A. Parker p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p21

Vision
Synesthesia
Jamie Ward p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p49

Scene Perception, Event Perception, Object Recognition

Visual Aesthetics and Human Preference
Stephen E. Palmer, Karen B. Schloss, and Jonathan Sammartino p p p p p p p p p p p p p p p p p p p p p p p p p77

Attention and Performance

Detecting Consciousness: A Unique Role for Neuroimaging
Adrian M. Owen p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 109
Executive Functions
Adele Diamond p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 135

Animal Learning and Behavior

The Neuroscience of Learning: Beyond the Hebbian Synapse
C.R. Gallistel and Louis D. Matzel p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 169

Evolutionary Psychology
Evolutionary Psychology: New Perspectives on Cognition
and Motivation
Leda Cosmides and John Tooby p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 201
Origins of Human Cooperation and Morality
Michael Tomasello and Amrisha Vaish p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 231

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Language and Communication

Gesture’s Role in Speaking, Learning, and Creating Language
Susan Goldin-Meadow and Martha Wagner Alibali p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 257

Nonverbal and Verbal Communication

The Antecedents and Consequences of Human Behavioral Mimicry
Tanya L. Chartrand and Jessica L. Lakin p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 285

Intergroup Relations, Stigma, Stereotyping, Prejudice, Discrimination

Sexual Prejudice
Gregory M. Herek and Kevin A. McLemore p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 309
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Social Neuroscience
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A Cultural Neuroscience Approach to the Biosocial Nature

of the Human Brain
Shihui Han, Georg Northoff, Kai Vogeley, Bruce E. Wexler,
Shinobu Kitayama, and Michael E.W. Varnum p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 335

Organizational Climate/Culture
Organizational Climate and Culture
Benjamin Schneider, Mark G. Ehrhart, and William H. Macey p p p p p p p p p p p p p p p p p p p p p p p p 361

Industrial Psychology/Human Resource Management

Employee Recruitment
James A. Breaugh p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 389

Learning and Performance in Educational Settings

Self-Regulated Learning: Beliefs, Techniques, and Illusions
Robert A. Bjork, John Dunlosky, and Nate Kornell p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 417

Teaching of Subject Matter

Student Learning: What Has Instruction Got to Do With It?
Hee Seung Lee and John R. Anderson p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 445

Health Psychology
Bringing the Laboratory and Clinic to the Community: Mobile
Technologies for Health Promotion and Disease Prevention
Robert M. Kaplan and Arthur A. Stone p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 471

Research Methodology
Multivariate Statistical Analyses for Neuroimaging Data
Anthony R. McIntosh and Bratislav Mišić p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 499

Contents vii
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Social Network Analysis: Foundations and Frontiers on Advantage

Ronald S. Burt, Martin Kilduff, and Stefano Tasselli p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 527

Indexes

Cumulative Index of Contributing Authors, Volumes 54–64 p p p p p p p p p p p p p p p p p p p p p p p p p p p 549

Cumulative Index of Chapter Titles, Volumes 54–64 p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 554

Errata

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viii Contents