Colours sometimes count: awareness and bidirectionality in grapheme-colour synaesthesia

Three experiments were conducted with 10 grapheme-colour synaesthetes and 10 matched controls to investigate (a) whether awareness of the inducer grapheme is necessary for synaesthetic colour induction and (b) whether grapheme-colour synaesthesia may be bidirectional in the sense that not only do graphemes induce colours, but that colours influence the processing of graphemes. Using attentional blink and Stroop paradigms with digit targets, we found that some synaesthetes did report “seeing” synaesthetic colours even when they were not able to report the inducing digit. Moreover, congruency effects (effects of matching the colour of digit presentation with the synaesthetic colour associated with that digit) suggested that grapheme-colour synaesthesia can be bidirectional, at least for some synaesthetes.     

Not all synaesthetes are created equal: Projector versus associator synaesthetes

In synaesthesia, ordinary stimuli elicit extraordinary experiences. When grapheme-color synaesthetes view black text, each grapheme elicits a photism-a highly specific experience of color. Importantly, some synaesthetes (projectors) report experiencing their photisms in external space, whereas other synaesthetes (associators) report experiencing their photisms "in the mind's eye." We showed that projectors and associators can be differentiated not only by their subjective reports, but also by their performance on Stroop tasks. Digits were presented in colors that were either congruent or incongruent with the synaesthetes' photisms. The synaesthetes named either the video colors of the digits or the colors of the photisms elicited by the digits. The results revealed systematic differences in the patterns of Stroop interference between projectors and associators. Converging evidence from first-person reports and third-person objective measures of Stroop interference establish the projector/ associator distinction as an important individual difference in grapheme-color synaesthesia.     

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.     

Colours sometimes count: Awareness and bidirectionality in grapheme–colour synaesthesia

Three experiments were conducted with 10 grapheme–colour synaesthetes and 10 matched controls to investigate (a) whether awareness of the inducer grapheme is necessary for synaesthetic colour induction and (b) whether grapheme–colour synaesthesia may be bidirectional in the sense that not only do graphemes induce colours, but that colours influence the processing of graphemes. Using attentional blink and Stroop paradigms with digit targets, we found that some synaesthetes did report “seeing” synaesthetic colours even when they were not able to report the inducing digit. Moreover, congruency effects (effects of matching the colour of digit presentation with the synaesthetic colour associated with that digit) suggested that grapheme–colour synaesthesia can be bidirectional, at least for some synaesthetes.     

Out of sight,out of mind: The attentional blink can eliminate synaesthetic colours

Mechanisms of selective attention exert a powerful influence on visual perception. We examined whether attentional selection is necessary for generation of the vivid colours experienced by individuals with grapheme-colour synaesthesia. Twelve synaesthetes and matched controls viewed rapid serial displays of nonsense characters within which were embedded an oriented grating (T1) and a letter-prime (T2), forming a modified attentional blink (AB) task. At the end of the stream a coloured probe appeared that was either congruent or incongruent with the synaesthetic colour elicited by the letter-prime. When the prime was attended, synaesthetes showed a reliable effect of prime-probe congruency. In contrast, when the prime appeared at 350 ms following T1 (during the AB), the congruency effect was eliminated. Our findings suggest that focused attention is crucial for inducing letters to elicit colours in synaesthesia.     

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.     

Colour fluctuations in grapheme‐colour synaesthesia: The effect of clinical and non‐clinical mood changes

Synaesthesia is a condition that gives rise to unusual secondary sensations (e.g., colours are perceived when listening to music). These unusual sensations tend to be reported as being stable throughout adulthood (e.g., Simner & Logie, 2007, Neurocase, 13, 358) and the consistency of these experiences over time is taken as the behavioural hallmark of genuineness. Our study looked at the influence of mood states on synaesthetic colours. In Experiment 1, we recruited grapheme‐colour synaesthetes (who experience colours from letters/digits) and elicited their synaesthetic colours, as well as their mood and depression states, in two different testing sessions. In each session, participants completed the PANAS‐X (Watson & Clark, 1999) and the BDI‐II (Beck, Steer, & Brown, 1996, Manual for Beck Depression Inventory‐II), and chose their synaesthetic colours for letters A‐Z from an interactive colour palette. We found that negative mood significantly decreased the luminance of synaesthetic colours. In Experiment 2, we showed that synaesthetic colours were also less luminant for synaesthetes with anxiety disorder, versus those without. Additional evidence suggests that colour saturation, too, may inversely correlate with depressive symptoms. These results show that fluctuations in mood within both a normal and clinical range influence synaesthetic colours over time. This has implications for our understanding about the longitudinal stability of synaesthetic experiences, and of how mood may interact with the visual (imagery) systems.     

What is the relationship between Aphantasia,Synaesthesia and Autism?

For people with aphantasia, visual imagery is absent or markedly impaired. Here, we investigated the relationship between aphantasia and two other neurodevelopmental conditions also linked to imagery differences: synaesthesia, and autism. In Experiment 1a and 1b, we asked whether aphantasia and synaesthesia can co-occur, an important question given that synaesthesia is linked to strong imagery. Taking grapheme-colour synaesthesia as a test case, we found that synaesthesia can be objectively diagnosed in aphantasics, suggesting visual imagery is not necessary for synaesthesia to occur. However, aphantasia influenced the type of synaesthesia experienced (favouring ‘associator’ over ‘projector’ synaesthesia - a distinction tied to the phenomenology of the synaesthetic experience). In Experiment 2, we asked whether aphantasics have traits associated with autism, an important question given that autism – like aphantasia – is linked to weak imagery. We found that aphantasics reported more autistic traits than controls, with weaknesses in imagination and social skills.     

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.     

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.     

Type-based associations in grapheme-color synaesthesia revealed by response time distribution analyses

Determining the nature of binding in grapheme-color synaesthesia has consequences for understanding the neural basis of synaesthesia and visual awareness in general. We evaluated type- and token-based letter-color binding using a synaesthetic version of the object-reviewing paradigm. Although mean response times failed to reveal any significant differences between synaesthetes and control participants, RT analyses with ex-Gaussian distributions revealed that the response facilitation in the synaesthesia group reflected type representations exclusively, while response facilitation in the control group, who learned letter-color associations, was dominated by token representations. Thus, letter-color associations in associator synaesthetes are type-based, and do not involve binding to object tokens, consistent with their subjective reports. Contrary to recent studies that failed to find differences between synaesthetes and non-synaesthetes with behavioral measures, response time distribution analyses indicate that color sensations in synaesthetes are not simply the extreme form of normal letter-color associations, and cannot be attributed to demand characteristics.     

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.     

Is synaesthesia an X-linked dominant trait with lethality in males?

In previous research the inheritance patterns of synaesthesia (eg experiencing colours from graphemes) has been studied and it was concluded that synaesthesia is most likely to be the outcome of a single gene passed on the X chromosome in a dominant fashion. In addition, it has been reported that the female-male ratio of synaesthetes is as high as 6:1 and the families of synaesthetes contain more female than male members. This raises the possibility that the gene may be associated with lethality in males. In this study we replicate and extend previous research by investigating the female-male ratio and inheritance patterns in a large sample of synaesthetic families (N = 85). We were able to verify the authenticity of grapheme-colour associates in at least one proband from each family using internal consistency. As before, our results show a female-male bias and are broadly consistent with an X-linked dominant mode of inheritance. However, there was no evidence of male lethality (eg synaesthetes are just as likely to give birth to sons as to daughters). Moreover, our female-male ratio of synaesthetes within families was 2:1--considerably lower than previous estimates. We speculate that men may be more reluctant to disclose synaesthesia than women (indeed, our female-male ratio based on self-referral was 3.7: 1). Finally, we discuss how the genotype may give rise to the phenotype in terms of changes in synaptogenesis or plasticity extending into childhood, to be subsequently shaped by the environment.     

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.     

Biases and regularities of grapheme–colour associations in Japanese nonsynaesthetic population

Associations between graphemes and colours in a nonsynaesthetic Japanese population were investigated. Participants chose the most suitable colour from 11 basic colour terms for each of 40 graphemes from the four categories of graphemes used in the Japanese language (kana characters, English alphabet letters, and Arabic and kanji numerals). This test was repeated after a three-week interval. In their responses, which were not as temporally consistent as those of grapheme–colour synaesthetes, participants showed biases and regularities that were comparable to those of synaesthetes reported in past studies. Although it has been believed that only synaesthetes, and not nonsynaesthetes, tended to associate graphemes with colours based on grapheme frequency, Berlin and Kay's colour typology, and colour word frequency, participants in this study tended in part to associate graphemes with colours based on the above factors. Moreover, participants that were nonsynaesthetes tended to associate different graphemes that shared sounds and/or meanings (e.g., Arabic and kanji numerals representing the same number) with the same colours, which was analogous to the findings in Japanese synaesthetes. These results support the view that grapheme–colour synaesthesia might have its origins in cross-modal association processes that are shared with the general population.     

Grapheme-color synaesthesia benefits rule-based Category learning

Researchers have long suspected that grapheme-color synaesthesia is useful, but research on its utility has so far focused primarily on episodic memory and perceptual discrimination. Here we ask whether it can be harnessed during rule-based Category learning. Participants learned through trial and error to classify grapheme pairs that were organized into categories on the basis of their associated synaesthetic colors. The performance of synaesthetes was similar to non-synaesthetes viewing graphemes that were physically colored in the same way. Specifically, synaesthetes learned to categorize stimuli effectively, they were able to transfer this learning to novel stimuli, and they falsely recognized grapheme-pair foils, all like non-synaesthetes viewing colored graphemes. These findings demonstrate that synaesthesia can be exploited when learning the kind of material taught in many classroom settings.     

A systematic, large-scale study of synaesthesia: implications for the role of early experience in lexical-colour associations

For individuals with synaesthesia, stimuli in one sensory modality elicit anomalous experiences in another modality. For example, the sound of a particular piano note may be 'seen' as a unique colour, or the taste of a familiar food may be 'felt' as a distinct bodily sensation. We report a study of 192 adult synaesthetes, in which we administered a structured questionnaire to determine the relative frequency and characteristics of different types of synaesthetic experience. Our data suggest the prevalence of synaesthesia in the adult population is approximately 1 in 1150 females and 1 in 7150 males. The incidence of left-handedness in our sample was within the normal range, contrary to previous claims. We did, however, find that synaesthetes are more likely to be involved in artistic pursuits, consistent with anecdotal reports. We also examined responses from a subset of 150 synaesthetes for whom letters, digits and words induce colour experiences ('lexical-colour' synaesthesia). There was a striking consistency in the colours induced by certain letters and digits in these individuals. For example, 'R' elicited red for 36% of the sample, 'Y' elicited yellow for 45%, and 'D' elicited brown for 47%. Similar trends were apparent for a group of non-synaesthetic controls who were asked to associate colours with letters and digits. Based on these findings, we suggest that the development of lexical-colour synaesthesia in many cases incorporates early learning experiences common to all individuals. Moreover, many of our synaesthetes experienced colours only for days of the week, letters or digits, suggesting that inducers that are part of a conventional sequence (e.g. Monday, Tuesday, Wednesday...; A, B, C...; 1, 2, 3...) may be particularly important in the development of synaesthetic inducer-colour pairs. We speculate that the learning of such sequences during an early critical period determines the particular pattern of lexical-colour links, and that this pattern then generalises to other words.     

Isolating automatic photism generation from strategic photism use in grapheme-colour synaesthesia

Grapheme-colour synaesthesia is a phenomenon in which ordinary black numbers and letters (graphemes) trigger the experience of highly specific colours (photisms). The Synaesthetic Stroop task has been used to demonstrate that graphemes trigger photisms automatically. In the standard Stroop task, congruent trial probability (CTP) has been manipulated to isolate effects of automaticity from higher-order strategic effects, with larger Stroop effects at high CTP attributed to participants strategically attending to the stimulus word to facilitate responding, and smaller Stroop effects at low CTP reflecting automatic word processing. Here we apply this logic for the first time to the Synaesthetic Stroop task. At high CTP we showed larger Stroop effects due to synaesthetes using their synaesthetic colours strategically. At low CTP Stroop effects were reduced but were still significant. We directly isolate automatic processing of graphemes from strategic effects and conclusively show that, in synaesthesia, viewing black graphemes automatically triggers colour experiences.     

An independent component analysis of fMRI data of grapheme-colour synaesthesia

The independent component analysis (ICA) method was applied to fMRI data from two synaesthetes and their matched controls while they performed the coloured-word Stroop task and the single-letter (synaesthetic) Stroop task. ICA identified an 'attention' network, a 'perceptual' network as well as a 'conflict monitoring' network. Increased activity was observed in right V4 during the single-letter Stroop task for synaesthetes only. The finding confirms that the same neural substrate that is known to support the experience of physical colours also supports the experience of synaesthetic colours.