Brain areas involved in synaesthesia: a review

Despite a recent upsurge of research, much remains unknown about the neurobiological mechanisms underlying synaesthesia. By integrating results obtained so far in Magnetic Resonance Imaging (MRI) studies, this contribution sheds light on the role of particular brain regions in synaesthetic experiences. First, in accordance with its sensory nature, it seems that the sensory brain areas corresponding to the type of synaesthetic experience are activated. Synaesthetic colour experiences can activate colour regions in occipito-temporal cortex, but this is not necessarily restricted to V4. Furthermore, sensory and motor brain regions have been obtained that extend beyond the particular type of synaesthesia studied. Second, differences in experimental setup, number and type of synaesthetes tested, and method to delineate regions of interest may help explain inconsistent results obtained in the BOLD-MRI (Blood Oxygen Level Dependent functional MRI) studies. Third, an overview of obtained results shows that a network of brain areas rather than a single brain region underlies synaesthesia. Six brain regions of overlapping results emerge, these regions are in sensory and motor regions as well as 'higher level' regions in parietal and frontal lobe. We propose that these regions are related to three different cognitive processes inherently part of synaesthesia; the sensory processes, the (attentional) 'binding' processes, and cognitive control processes. Finally, we discuss how these functional and structural brain properties might relate to the development of synaesthesia. In particular, we believe this relationship is better understood by separating the question what underlies the presence of synaesthesia ('trait') from what determines particular synaesthetic associations ('type').     

Psychophysiological evidence for the genuineness of swimming-style colour synaesthesia

Recently, swimming-style colour synaesthesia was introduced as a new form of synaesthesia. A synaesthetic Stroop test was used to establish its genuineness. Since Stroop interference can occur for any type of overlearned association, in the present study we used a modified Stroop test and psychophysiological synaesthetic conditioning to further establish the genuineness of this form of synaesthesia. We compared the performance of a swimming-style colour synaesthete and a control who was trained on swimming-style colour associations. Our results showed that behavioural aspects of swimming-style colour synaesthesia can be mimicked in a trained control. Importantly, however, our results showed a psychophysiological conditioning effect for the synaesthete only. We discuss the theoretical relevance of swimming-style colour synaesthesia according to different models of synaesthesia. We conclude that swimming-style colour synaesthesia is a genuine form of synaesthesia, can be mimicked behaviourally in non-synaesthetes, and is best explained by a re-entrant feedback model.     

Development of synaesthetic consistency: Repeated autonomous engagement with graphemes and colours leads to consistent associations

Synaesthetic consistency is the hallmark of synaesthesia and plays an important role in the definition and validation of synaesthesia. It has been hypothesised that the acquisition of initially unspecified synaesthetic associations is based on consolidation processes. Thus, we investigated in non-synaesthetes whether repeatedly engaging with grapheme-colour associations mimics the developmental trajectory of synaesthetic consistency in genuine grapheme-colour synaesthesia. This was the case for the two tested experimental groups, irrespective of whether they were instructed to memorize their chosen associations, but not for the passive control group. Moreover, consolidated associations of the experimental groups resembled those frequently found in genuine synaesthesia. Furthermore, the acquisition of consistent grapheme-colour associations resulted in a transfer of benefits to performance in recognition memory for abstract stimuli, as also found in genuine synaesthesia. Our findings suggest that consistent synaesthetic associations are based on consolidation processes due to repeated engagement with graphemes and colours.     

Searching for Shereshevskii: what is superior about the memory of synaesthetes?

Some individuals with superior memory, such as the mnemonist Shereshevskii (Luria, 1968), are known to have synaesthesia. However, the extent to which superior memory is a general characteristic of synaesthesia is unknown, as is the precise cognitive mechanism by which synaesthesia affects memory. This study demonstrates that synaesthetes tend to report subjectively better than average memory and that these reports are borne out with objective testing. Synaesthetes experiencing colours for words show better memory than matched controls for stimuli that induce synaesthesia (word lists) relative to stimuli that do not (an abstract figure). However, memory advantages are not limited to material that elicits synaesthesia because synaesthetes demonstrate enhanced memory for colour per se (which does not induce a synaesthetic response). Our results suggest that the memory enhancement found in synaesthetes is related to an enhanced retention of colour in both synaesthetic and nonsynaesthetic situations. Furthermore, this may account for the fact that synaesthetic associations, once formed, remain highly consistent.     

Searching for Shereshevskii: What is superior about the memory of synaesthetes?

Some individuals with superior memory, such as the mnemonist Shereshevskii (Luria, 1968), are known to have synaesthesia. However, the extent to which superior memory is a general characteristic of synaesthesia is unknown, as is the precise cognitive mechanism by which synaesthesia affects memory. This study demonstrates that synaesthetes tend to report subjectively better than average memory and that these reports are borne out with objective testing. Synaesthetes experiencing colours for words show better memory than matched controls for stimuli that induce synaesthesia (word lists) relative to stimuli that do not (an abstract figure). However, memory advantages are not limited to material that elicits synaesthesia because synaesthetes demonstrate enhanced memory for colour per se (which does not induce a synaesthetic response). Our results suggest that the memory enhancement found in synaesthetes is related to an enhanced retention of colour in both synaesthetic and nonsynaesthetic situations. Furthermore, this may account for the fact that synaesthetic associations, once formed, remain highly consistent.     

The cross-activation theory at 10

In 2001, Ramachandran and Hubbard introduced the cross-activation model of grapheme-colour synaesthesia. On the occasion of its 10-year anniversary, we review the evidence from experiments that have been conducted to test the model to assess how it has fared. We examine data from behavioural, functional magnetic resonance imaging (fMRI), anatomical studies (diffusion tensor imaging and voxel-based morphometry), and electroencephalography (EEG) and magnetoencephalography (MEG) studies of grapheme-colour synaesthesia. Although much of this evidence has supported the basic cross-activation hypothesis, our growing knowledge of the neural basis of synaesthesia, grapheme, and colour processing has necessitated two specific updates and modifications to the basic model: (1) our original model assumed that binding and parietal cortex functions were normal in synaesthesia; we now recognize that parietal cortex plays a key role in synaesthetic binding, as part of a two-stage model. (2) Based on MEG data we have recently collected demonstrating that synaesthetic responses begin within 140 ms of stimulus presentation, and an updated understanding of the neural mechanisms of reading as hierarchical feature extraction, we present a revised and updated version of the cross-activation model, the cascaded cross-tuning model. We then summarize data demonstrating that the cross-activation model may be extended to account for other forms of synaesthesia and discuss open questions about how learning, development, and cortical plasticity interact with genetic factors to lead to the full range of synaesthetic experiences. Finally, we outline a number of future directions needed to further test the cross-activation theory and to compare it with alternative theories.     

Color opponency in synaesthetic experiences

Grapheme-color synaesthesia is a rare condition in which perception of a letter or a digit is associated with concurrent perception of a color. Synaesthetes report that these color experiences are vivid and realistic. We used a Stroop task to show that synaesthetically induced color, like real color, is processed in color-opponent channels (red-green or blue-yellow). Synaesthetic color produced maximal interference with the perception and naming of the real color of a grapheme if the real color was opponent to the synaesthetic color. Interference was reduced considerably if the synaesthetic and real colors engaged different color channels (e.g., synaesthetic blue and real red). No dependence on color opponency was found for semantic conflicts between shape and color (e.g., a blue lemon). Thus, the neural representation of synaesthetic colors closely resembles that of real colors. This suggests involvement of early stages of visual processing in color synaesthesia and explains the vivid and realistic nature of synaesthetic experiences.     

Mental imagery and synaesthesia: Is synaesthesia from internally-generated stimuli possible?

Previous studies provide empirical support for the reported colour experience in grapheme-colour synaesthesia by measuring the synaesthetic experience from an externally presented grapheme. The current study explored the synaesthetic experience resulting from a visual mental image of a grapheme. Grapheme-colour synaesthetes (N=6) and matched controls (N=10 per synaesthete) completed a visual mental imagery task that involved visualising a letter and making a size-based decision about it. The background colour that the grapheme was visualised against was manipulated so that it was congruent or incongruent with the synaesthetic colour for the grapheme being visualised. Compared to matched controls, an effect of colour condition was found for four of the six synaesthetes, although importantly the direction of the effect varied between synaesthetes. In addition, a significant effect of group was found, as the synaesthetes were all faster than the matched controls at the imagery task, regardless of background colour. We conclude that there is some support for subjective reports of imagery-induced synaesthesia, but there are important individual differences. These findings are discussed in relation to both the visual imagery and synaesthesia literature.     

Familial patterns and the origins of individual differences in synaesthesia

The term synaesthesia has been applied to a range of different sensory-perceptual and cognitive experiences, yet how these experiences are related to each other is not well understood. Not only are there disparate types of synaesthesia, but even within types there are vast individual differences in the way that stimuli induce synaesthesia and in the subjective synaesthetic experience. An investigation of the inheritance patterns of different types of synaesthesia is likely to elucidate whether a single underlying mechanism can explain all types. This study is the first to systematically survey all types of synaesthesia within a familial framework. We recruited 53 synaesthetes and 42% of these probands reported a first-degree relative with synaesthesia. We then directly contacted as many first-degree relatives as possible and collected complete data on synaesthetic status for all family members for 17 families. We found that different types of synaesthesia can occur within the same family and that the qualitative nature of the experience can differ between family members. Our findings strongly indicate that various types of synaesthesia are fundamentally related at the genetic level, but that the explicit associations and the individual differences between synaesthetes are influenced by other factors. Synaesthesia thus provides a good model to explore the interplay of all these factors in the development of cognitive traits in general.     

Frequency correlates in grapheme-color synaesthesia

Individuals with synaesthesia experience certain stimuli in more than one sensory modality. Most common is the linkage of letters and digits (graphemes) to colors. Whereas synaesthesia might be partly genetically determined, the linkages to specific colors are assumed to be learned. We present a systematic statistical analysis of synaesthetic color perception based on subjects' reproduction of individual colors for each grapheme, instead of simple verbal categorizations. The statistical analysis revealed that the color perceptions, measured with the HSL (hue, saturation, and luminance) scale, varied systematically among the different digits and letters. The frequencies of the digits and letters (in the German language) partly explained these systematic variations. However, digit frequency was more strongly related to color perception in the synaesthetes than was letter frequency. The results for digit and letter frequency indicate that experience with graphemes may shape synaesthetic color perception.     

"That's not a real body": identifying stimulus qualities that modulate synaesthetic experiences of touch

Mirror-touch synaesthesia is a condition where observing touch to another’s body induces a subjective tactile sensation on the synaesthetes body. The present study explores which characteristics of the inducing stimulus modulate the synaesthetic touch experience. Fourteen mirror-touch synaesthetes watched videos depicting a touch event while indicating (i) whether the video induced a tactile sensation, (ii) on which side of their body they felt this sensation and (iii) the intensity of the experienced sensation. Results indicate that the synaesthetes experience stronger tactile sensations when observing touch to real bodies, whereas observing touch to dummy bodies, pictures of bodies and disconnected dummy body parts elicited weaker sensations. These results suggest that mirror-touch synaesthesia is not entirely bottom-up driven, but top-down information, such as knowledge about real and dummy body parts, also modulate the intensity of the experience.     

Varieties of grapheme-colour synaesthesia: a new theory of phenomenological and behavioural differences

Recent research has suggested that not all grapheme-colour synaesthetes are alike. One suggestion is that they can be divided, phenomenologically, in terms of whether the colours are experienced in external or internal space (projector–associator distinction). Another suggestion is that they can be divided according to whether it is the perceptual or conceptual attributes of a stimulus that is critical (higher–lower distinction). This study compares the behavioural performance of 7 projector and 7 associator synaesthetes. We demonstrate that this distinction does not map on to behavioural traits expected from the higher–lower distinction. We replicate previous research showing that projectors are faster at naming their synaesthetic colours than veridical colours, and that associators show the reverse profile. Synaesthetes who project colours into external space but not on to the surface of the grapheme behave like associators on this task. In a second task, graphemes presented briefly in the periphery are more likely to elicit reports of colour in projectors than associators, but the colours only tend to be accurate when the grapheme itself is also accurately identified. We propose an alternative model of individual differences in grapheme-colour synaesthesia that emphasises the role of different spatial reference frames in synaesthetic perception. In doing so, we attempt to bring the synaesthesia literature closer to current models of non-synaesthetic perception, attention and binding.     

Colored-Speech Synaesthesia Is Triggered by Multisensory, Not Unisensory, Perception

ABSTRACT— Although it is estimated that as many as 4% of people experience some form of enhanced cross talk between (or within) the senses, known as synaesthesia, very little is understood about the level of information processing required to induce a synaesthetic experience. In work presented here, we used a well-known multisensory illusion called the McGurk effect to show that synaesthesia is driven by late, perceptual processing, rather than early, unisensory processing. Specifically, we tested 9 linguistic-color synaesthetes and found that the colors induced by spoken words are related to what is perceived (i.e., the illusory combination of audio and visual inputs) and not to the auditory component alone. Our findings indicate that color-speech synaesthesia is triggered only when a significant amount of information processing has occurred and that early sensory activation is not directly linked to the synaesthetic experience.     

Synaesthesia,creativity and art: What is the link?

It has been suggested that individuals with synaesthesia may show heightened creativity as a result of being able to form meaningful associations between disparate stimuli (e.g. colour, sound). In this study, a large sample (N=82) of people with various kinds of synaesthesia were given two psychometric tests of creativity (Remote Associates Test, Alternate Uses Test) and were also asked about the amount of time engaged in creative arts (visual art, music). There was a significant tendency for synaesthetes to spend more time engaged in creative arts and this was, at least in part, dependent upon the type of synaesthesia experienced. For example, synaesthetes experiencing vision from music were far more likely to play an instrument than their other synaesthetic counterparts. There was no relationship between this tendency and the psychometric measures of creativity, but synaesthetes did outperform controls on one of the two psychometric measures (Remote Associates). We conclude that the tendency for synaesthetes to be more engaged in art is likely to have a different mechanism to psychometric measures of creativity, and that there is no direct link between them. Although synaesthetes may well perform better on some measures of creativity, we suggest that synaesthetes have better bottom‐up access to certain associations, but are not necessarily better able to use them flexibly (in divergent thinking).     

The neural basis of illusory gustatory sensations: two rare cases of lexical-gustatory synaesthesia

Lexical-gustatory synaesthesia is a rare phenomenon in which the individual experiences flavour sensations when they read, hear, or imagine words. In this study, we provide insight into the neural basis of this form of synaesthesia using functional neuroimaging. Words known to evoke pleasant, neutral, and unpleasant synaesthetic tastes and synaesthetically tasteless words were presented to two lexical-gustatory synaesthetes, during fMRI scanning. Ten non-synaesthetic participants were also scanned on the same list of words. The synaesthetic brain displayed a different pattern of activity to words when compared to the non-synaesthetes, with insula activation related to viewing words that elicited tastes that have an associated emotional valence (i.e., pleasant or unpleasant tastes). The subjective intensity of the synaesthesia was correlated with activity in the medial parietal lobes (precuneus/retrosplenial cortex), which are implicated in polymodal imagery and self-directed thought. This region has also previously been activated in studies of lexical-colour synaesthesia, suggesting its role may not be limited to the type of synaesthesia explored here.     

Synaesthetic perception of colour and visual space in a blind subject: an fMRI case study

In spatial sequence synaesthesia (SSS) ordinal stimuli are perceived as arranged in peripersonal space. Using fMRI, we examined the neural bases of SSS and colour synaesthesia for spoken words in a late-blind synaesthete, JF. He reported days of the week and months of the year as both coloured and spatially ordered in peripersonal space; parts of the days and festivities of the year were spatially ordered but uncoloured. Words that denote time-units and triggered no concurrents were used in a control condition. Both conditions inducing SSS activated the occipito-parietal, infero-frontal and insular cortex. The colour area hOC4v was engaged when the synaesthetic experience included colour. These results confirm the continued recruitment of visual colour cortex in this late-blind synaesthetes. Synaesthesia also involved activation in inferior frontal cortex, which may be related to spatial memory and detection, and in the insula, which might contribute to audiovisual integration related to the processing of inducers and concurrents.     

What is the relationship between synaesthesia and visuo-spatial number forms?

This study compares the tendency for numerals to elicit spontaneous perceptions of colour or taste (synaesthesia) with the tendency to visualise numbers as occupying particular visuo-spatial configurations (number forms). The prevalence of number forms was found to be significantly higher in synaesthetes experiencing colour compared both to synaesthetes experiencing taste and to control participants lacking any synaesthetic experience. This suggests that the presence of synaesthetic colour sensations enhances the tendency to explicitly represent numbers in a visuo-spatial format although the two symptoms may nevertheless be logically independent (i.e. it is possible to have number forms without colour, and coloured numbers without forms). Number forms are equally common in men and women, unlike previous reports of synaesthesia that have suggested a strong female bias. Individuals who possess a number form are also likely to possess visuo-spatial forms for other ordinal sequences (e.g. days, months, letters) which suggests that it is the ordinal nature of numbers rather than numerical quantity that gives rise to this particular mode of representation. Finally, we also describe some consequences of number forms for performance in a number comparison task.     

Synaesthesia: learned or lost?

The question why synaesthesia, an atypical binding within or between modalities, occurs is both enduring and important. Two explanations have been provided: (1) a congenital explanation: we are all born as synaesthetes but most of us subsequently lose the experience due to brain development; (2) a learning explanation: synaesthesia is related to some learning process during childhood. Three recent studies provide conflicting support for these explanations. Two studies supported the idea that synaesthesia is learned by showing that the frequency of everyday language implicitly modulates the synaesthetic experience. Another study argued that synaesthesia reflects basic, innate magnitude representations. In this paper we reassess these points of view, and show that it is possible for both to be valid. These findings are integrated into an interactive specialization account of development in order to explain the neuronal mechanism underlying synaesthesia.     

Synaesthetic associations decrease during infancy

Early development is characterized by a period of exuberant neural connectivity followed by a retraction and reweighting of connections over the course of development. It has been proposed that this connectivity may facilitate arbitrary sensory experiences in infants that are unlike anything experienced by typical adults but are similar to the sensory experiences of adults with synaesthesia, a rare sensory phenomenon that has been associated with exuberant neural connectivity and that is characterized by strong arbitrary associations between different sensations. We provide the first evidence for this infant-synaesthesia hypothesis by showing that the presence of particular shapes influences color preferences in typical 2- and 3-month-olds, but not in 8-month-olds or adults. These results are consistent with the possibility that exuberant neural connectivity facilitates synaesthetic associations during infancy that are typically eliminated during development, but that a failure of the retraction process leads in rare cases to synaesthesia in adults.