Temporal dissociation of neural activity underlying synesthetic and perceptual colors.

Grapheme-color synesthetes experience color when seeing achromatic symbols. We examined whether similar neural mechanisms underlie color perception and synesthetic colors using magnetoencephalography. Classification models trained on neural activity from viewing colored stimuli could distinguish synesthetic color evoked by achromatic symbols after a delay of ∼100 ms. Our results provide an objective neural signature for synesthetic experience and temporal evidence consistent with higher-level processing in synesthesia.

Reduced perceptual narrowing in synesthesia.

Synesthesia is a neurologic trait in which specific inducers, such as sounds, automatically elicit additional idiosyncratic percepts, such as color (thus "colored hearing"). One explanation for this trait-and the one tested here-is that synesthesia results from unusually weak pruning of cortical synaptic hyperconnectivity during early perceptual development. We tested the prediction from this hypothesis that synesthetes would be superior at making discriminations from nonnative categories that are normally weakened by experience-dependent pruning during a critical period early in development-namely, discrimination among nonnative phonemes (Hindi retroflex /d̪a/ and dental /ɖa/), among chimpanzee faces, and among inverted human faces. Like the superiority of 6-mo-old infants over older infants, the synesthetic groups were significantly better than control groups at making all the nonnative discriminations across five samples and three testing sites. The consistent superiority of the synesthetic groups in making discriminations that are normally eliminated during infancy suggests that residual cortical connectivity in synesthesia supports changes in perception that extend beyond the specific synesthetic percepts, consistent with the incomplete pruning hypothesis.

Substantiating synesthesia: a novel aid in a case of grapheme-colour synesthesia and concomitant dyscalculia.

Here we present the case of SP, a 21-year-old female with life-long dyscalculia. SP was subsequently diagnosed with grapheme-color synesthesia, a diagnosis that serendipitously catalyzed our development of a novel aid:The digit-color calculator (DCC). The DCC substantiates SP's color concurrents, dramatically ameliorating her difficulties with basic calculations. We envisage the DCC and its analogues may assist others in educational settings, particularly if they experience difficulties with the acquisition of literacy and numeracy. Further devices that leverage synesthesia may also have the potential to improve the quality of life for others with trait synesthesia regardless of concomitant disorder.

Constructing calendars in the brain.

By studying an enigmatic condition called, "calendar synesthesia", we explored the elusive boundary between perception, visual imagery, and the manner in which we construct an internal mental calendar by mapping time-sequences onto spatial maps. We use a series of demonstrations to establish that these calendars act more like real objects activating sensory pathways rather than purely abstract symbolic descriptions that bear no resemblance to an actual calendar. We propose that the calendar is enshrined in acircuitry involving the hippocampal place-cells and entorhinal grid-cells, which are connected to the angular gyrus (involved with computing sequences) via the inferior longitudinal fasciculus.

Association between synesthetic colors and sensitivity to physical colors changed by type of synesthetic experience in grapheme-color synesthesia.

Grapheme-color synesthesia is a condition in which visual perception of letters induces simultaneous perception of a specific color. Previous studies indicate that grapheme-color synesthetes are more sensitive to physical colors than non-synesthetes. Synesthetic colors are found to be concentrated in multiple regions of the color space, forming "synesthetic color clusters". The present study investigated whether color sensitivity corresponding to synesthetic color clusters (clustered colors) is higher than color sensitivity that does not correspond to synesthetic color clusters (non-clustered colors). However, we found no difference in the color sensitivity for clustered and non-clustered colors. We also investigated whether the color sensitivity is dependent on the synesthetic experience (associators and projectors). We found that the greater the tendency toward associator characteristics, the greater the sensitivity for clustered colors compared to that for non-clustered colors. Our findings suggest an association between synesthetic colors and physical color sensitivity that is modulated by synesthetic experience.

Synesthesia in children with difficulties in written language learning.

We tested whether the acquisition of grapheme-color synesthesia during childhood is related to difficulties in written language learning by measuring whether it is more frequent in 79 children receiving speech and language therapy for such difficulties than in the general population of children (1.3%). By using criteria as similar as possible to those used in the reference study (Simner et al., 2009), we did not identify any synesthete (Bayesian 95% credible interval [0, 4.5]% for a flat prior). The odds of the null model (no difference between 0/79 and 1.3%) over alternative models is 28 (Bayes Factor). A higher prevalence of grapheme-color synesthetes among children with learning difficulties is therefore very unlikely, questioning the hypothesis of a link between synesthesia and difficulties in language acquisition. We also describe the difficulty of diagnosing synesthesia in children and discuss the need for new approaches to do so.

A single system account of enhanced recognition memory in synaesthesia.

Researchers often adjudicate between models of memory according to the models' ability to explain impaired patterns of performance (e.g., in amnesia). In contrast, evidence from special groups with enhanced memory is very rarely considered. Here, we explored how people with unusual perceptual experiences (synaesthesia) perform on various measures of memory and test how computational models of memory may account for their enhanced performance. We contrasted direct and indirect measures of memory (i.e., recognition memory, repetition priming, and fluency) in grapheme-colour synaesthetes and controls using a continuous identification with recognition (CID-R) paradigm. Synaesthetes outperformed controls on recognition memory and showed a different reaction-time pattern for identification. The data were most parsimoniously accounted for by a single-system computational model of the relationship between recognition and identification. Overall, the findings speak in favour of enhanced processing as an explanation for the memory advantage in synaesthesia. In general, our results show how synaesthesia can be used as an effective tool to study how individual differences in perception affect cognitive functions.

Hearing through Your Eyes: Neural Basis of Audiovisual Cross-activation, Revealed by Transcranial Alternating Current Stimulation.

Some people experience auditory sensations when seeing visual flashes or movements. This prevalent synaesthesia-like visually evoked auditory response (vEAR) could result either from overexuberant cross-activation between brain areas and/or reduced inhibition of normally occurring cross-activation. We have used transcranial alternating current stimulation (tACS) to test these theories. We applied tACS at 10 Hz (alpha band frequency) or 40 Hz (gamma band), bilaterally either to temporal or occipital sites, while measuring same/different discrimination of paired auditory (A) versus visual (V) Morse code sequences. At debriefing, participants were classified as vEAR or non-vEAR, depending on whether they reported "hearing" the silent flashes. In non-vEAR participants, temporal 10-Hz tACS caused impairment of A performance, which correlated with improved V; conversely under occipital tACS, poorer V performance correlated with improved A. This reciprocal pattern suggests that sensory cortices are normally mutually inhibitory and that alpha-frequency tACS may bias the balance of competition between them. vEAR participants showed no tACS effects, consistent with reduced inhibition, or enhanced cooperation between modalities. In addition, temporal 40-Hz tACS impaired V performance, specifically in individuals who showed a performance advantage for V (relative to A). Gamma-frequency tACS may therefore modulate the ability of these individuals to benefit from recoding flashes into the auditory modality, possibly by disrupting cross-activation of auditory areas by visual stimulation. Our results support both theories, suggesting that vEAR may depend on disinhibition of normally occurring sensory cross-activation, which may be expressed more strongly in some individuals. Furthermore, endogenous alpha- and gamma-frequency oscillations may function respectively to inhibit or promote this cross-activation.

Implicit and explicit spatial-numerical representations diverge in number-form synesthetes.

In number-form (NF) synesthesia-a condition in which people report vivid, automatic and consistent mental layouts for numerical sequences-numbers and space are closely linked. These explicit associations are similar to the implicit associations demonstrated by the Spatial-Numerical Association of Response Codes (SNARC) effect. Thus, NF synesthesia offers a unique opportunity to investigate spatial-numerical associations. We tested implicit and explicit representations in NF synesthetes using a multiple case-study design. Over two sessions, synesthetes participated in a semi-structured interview focusing on the nature of their associations, as well as SNARC and number line estimation tasks. Contrary to our hypotheses, only one synesthete demonstrated SNARC effects congruent with her reported form, whereas two others exhibited SNARC effects that were the opposite of their explicit NFs. While this inconsistency between implicit and explicit representations may indicate separate underlying cognitive mechanisms, factors such as task-specific constraints and strategic variability must also be considered.

Automaticity of pitch class-color synesthesia as revealed by a Stroop-like effect.

Pitch classes (e.g., do, re, and mi) in music evoke color sensations in pitch class-color synesthesia, which is a recently described form of synesthesia in musicians. The synesthetic color sensations were confirmed to be consistent over an extended time interval, fulfilling a widely-accepted criterion for the authenticity of synesthesia. However, it remains unclear whether the color sensations occurred automatically (i.e., without voluntary effort), which is another defining property of synesthesia. We utilized the Stroop paradigm to investigate this issue in 10 pitch class-color synesthetes. Participants were visually presented with pitch class names in font colors that were either congruent or incongruent with the participants' own color sensations. The speed for reporting the font color was slower when it was incongruent with the synesthetic sensation than when it was congruent. The finding verifies the authenticity of pitch class-color synesthesia by demonstrating that the color sensations occur automatically, even when unnecessary.

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.

The role of synaesthesia in reading written musical key signatures.

This study is the first empirical demonstration of synaesthesia for reading written musical keys signatures. Nine music-color synaesthetes and 9 controls took part in 5 experiments that aimed to confirm the authenticity of synaesthesia for reading musical keys, and to demonstrate that this type of synaesthesia is linked to conceptual rather than to purely perceptual processing of the inducing stimulus. First, the existence of a synaesthetic association with written musical keys was validated in an objective manner by employing 2 measures of consistency as diagnostic criteria. Second, the automaticity of the synaesthetes' responses was tested by demonstrating the presence of interference when naming synaesthetic colors for incongruent pairings of color and musical key. To test whether a change in form altered the concept of the musical key, stimuli were randomly presented in 3 separate modes (words, treble clef, or bass clef). Last, the interference of synaesthetic colors with veridical colors was assessed in a task-irrelevant manner, that is, without the need for the explicit naming of synaesthetic color. Findings showed synaesthesia for written musical keys to be a genuine form of synaesthesia elicited from the concept, or the idea, of the key. (PsycInfo Database Record (c) 2022 APA, all rights reserved).

Synesthesia, hallucination, and autism.

Synesthesia literally means a "union of the senses" whereby two or more of the five senses that are normally experienced separately are involuntarily and automatically joined together in experience (1, 2, 3). For example, some synesthetes experience a color when they hear a sound, although many instances of synesthesia also occur entirely within the visual sense. In this paper, I first mainly engage critically with Sollberger's view that there is reason to think that at least some synesthetic experiences can be viewed as truly veridical perceptions, and not as illusions or hallucinations (4). Among other things, I explore the possibility that many forms of synesthesia can be understood as experiencing what I will call "second-order secondary properties," that is, experiences of properties of objects induced by the secondary qualities of those objects. In doing so, I shed some light on why synesthesia is typically one-directional and its relation to some psychopathologies such as autism.

Perceptual Gains and Losses in Synesthesia and Schizophrenia.

Individual differences in perception are widespread. Considering inter-individual variability, synesthetes experience stable additional sensations; schizophrenia patients suffer perceptual deficits in, eg, perceptual organization (alongside hallucinations and delusions). Is there a unifying principle explaining inter-individual variability in perception? There is good reason to believe perceptual experience results from inferential processes whereby sensory evidence is weighted by prior knowledge about the world. Perceptual variability may result from different precision weighting of sensory evidence and prior knowledge. We tested this hypothesis by comparing visibility thresholds in a perceptual hysteresis task across medicated schizophrenia patients (N = 20), synesthetes (N = 20), and controls (N = 26). Participants rated the subjective visibility of stimuli embedded in noise while we parametrically manipulated the availability of sensory evidence. Additionally, precise long-term priors in synesthetes were leveraged by presenting either synesthesia-inducing or neutral stimuli. Schizophrenia patients showed increased visibility thresholds, consistent with overreliance on sensory evidence. In contrast, synesthetes exhibited lowered thresholds exclusively for synesthesia-inducing stimuli suggesting high-precision long-term priors. Additionally, in both synesthetes and schizophrenia patients explicit, short-term priors-introduced during the hysteresis experiment-lowered thresholds but did not normalize perception. Our results imply that perceptual variability might result from differences in the precision afforded to prior beliefs and sensory evidence, respectively.

Language evolution: examining the link between cross-modality and aggression through the lens of disorders.

We demonstrate how two linguistic phenomena, figurative language (implicating cross-modality) and derogatory language (implicating aggression), both demand a precise degree of (dis)inhibition in the same cortico-subcortical brain circuits, in particular cortico-striatal networks, whose connectivity has been significantly enhanced in recent evolution. We examine four cognitive disorders/conditions that exhibit abnormal patterns of (dis)inhibition in these networks: schizophrenia (SZ), autism spectrum disorder (ASD), synaesthesia and Tourette's syndrome (TS), with the goal of understanding why the two phenomena altered reactive aggression and altered cross-modality cluster together in these disorders. Our proposal is that enhanced cross-modality (necessary to support language, in particular metaphoricity) was a result, partly a side-effect, of self-domestication (SD). SD targeted the taming of reactive aggression, but reactive impulses are controlled by the same cortico-subcortical networks that are implicated in cross-modality. We further add that this biological process of SD did not act alone, but was engaged in an intense feedback loop with the cultural emergence of early forms of language/grammar, whose high degree of raw metaphoricity and verbal aggression also contributed to increased brain connectivity and cortical control. Consequently, in conjunction with linguistic expressions serving as approximations/'fossils' of the earliest stages of language, these cognitive disorders/conditions serve as confident proxies of brain changes in language evolution, helping us reconstruct certain crucial aspects of early prehistoric languages and cognition, as well as shed new light on the nature of the disorders. This article is part of the theme issue 'Reconstructing prehistoric languages'.

Olfactory-colour crossmodal correspondences in art, science, and design.

The last few years have seen a rapid growth of interest amongst researchers in the crossmodal correspondences. One of the correspondences that has long intrigued artists is the putative association between colours and odours. While traditionally conceptualised in terms of synaesthesia, over the last quarter century or so, at least 20 published peer-reviewed articles have assessed the consistent, and non-random, nature of the colours that people intuitively associate with specific (both familiar and unfamiliar) odours in a non-food context. Having demonstrated such consistent mappings amongst the general (i.e. non-synaesthetic) population, researchers have now started to investigate whether they are shared cross-culturally, and to document their developmental acquisition. Over the years, several different explanations have been put forward by researchers for the existence of crossmodal correspondences, including the statistical, semantic, structural, and emotional-mediation accounts. While several of these approaches would appear to have some explanatory validity as far as the odour-colour correspondences are concerned, contemporary researchers have focussed on learned associations as the dominant explanatory framework. The nature of the colour-odour associations that have been reported to date appear to depend on the familiarity of the odour and the ease of source naming, and hence the kind of association/representation that is accessed. While the bidirectionality of odour-colour correspondences has not yet been rigorously assessed, many designers are nevertheless already starting to build on odour-colour crossmodal correspondences in their packaging/labelling/branding work.

On the Relative Nature of (Pitch-Based) Crossmodal Correspondences.

This review deals with the question of the relative vs absolute nature of crossmodal correspondences, with a specific focus on those correspondences involving the auditory dimension of pitch. Crossmodal correspondences have been defined as the often-surprising crossmodal associations that people experience between features, attributes, or dimensions of experience in different sensory modalities, when either physically present, or else merely imagined. In the literature, crossmodal correspondences have often been contrasted with synaesthesia in that the former are frequently said to be relative phenomena (e.g., it is the higher-pitched of two sounds that is matched with the smaller of two visual stimuli, say, rather than there being a specific one-to-one crossmodal mapping between a particular pitch of sound and size of object). By contrast, in the case of synaesthesia, the idiosyncratic mapping between inducer and concurrent tends to be absolute (e.g., it is a particular sonic inducer that elicits a specific colour concurrent). However, a closer analysis of the literature soon reveals that the distinction between relative and absolute in the case of crossmodal correspondences may not be as clear-cut as some commentators would have us believe. Furthermore, it is important to note that the relative vs absolute question may receive different answers depending on the particular (class of) correspondence under empirical investigation.

A colorful advantage in iconic memory.

Synesthesia is a benign neurodevelopmental condition in which stimulation of one sensory modality evokes experiences in a second, unstimulated modality (Simner and Hubbard, 2013). In grapheme-color synesthesia (GCS), which is experienced by 1-2% of adults, synesthetes reliably and involuntarily experience specific colors when viewing blackand-white graphemes. Previous case-studies have identified synesthetes with spectacular memory (Luria, 1968; Smilek, Dixon, Cudahy, & Merikle, 2001) and group studies have found advantages for synesthetes compared to nonsynesthetes in long-term memory (Rothen, Meier, & Ward, 2012). Here, we tested whether similar advantages are also present in earlier stages of memory. We tested visual iconic memory-the sensory store for vision-which has a very large capacity, but decays approximately 1000 ms after stimulus offset (Chow, 1985; Sergent et al., 2013; Sperling, 1960). We tested 20 synesthetes and 20 nonsynesthetes in a direct replication of the Sperling (1960) Partial Report Paradigm using letters (Experiment 1) and non-alphanumeric symbols (Experiment 2) as stimuli. Overall, synesthetes had a greater iconic memory capacity than nonsynesthetes when presented with synesthesia-inducing letter stimuli. This advantage was reduced when they were presented with non-synesthesia inducing symbol stimuli. Critically this advantage was most prominent when memory was stressed by asking participants to remember large arrays. Our results demonstrate that synesthetic memory advantages extend to the earliest stages of memory, and suggest that advantages in later stages of memory may arise from these earlier advantages.

Double-blind study of visual imagery in grapheme-color synesthesia.

Synesthesia is an atypical perceptual phenomenon that has been associated with generalized differences in other cognitive and perceptual domains. Given similarities in the qualitative nature of synesthetic experiences to visual imagery perceptions, several studies have sought to examine whether synesthetes demonstrate increased visual imagery abilities. Using subjective imagery questionnaires, some studies have identified superior imaging abilities in synesthetes, while others have not. However, because most research on synesthesia uses un-blinded group membership prior to data collection, such methods for studying group differences may be prone to participant and experimenter biases (e.g., a motivated synesthete may rate themselves as having stronger visual imagery abilities due to their own bias and perceived experimenter expectations). To address this issue, we demonstrate the feasibility of double-blind designs in synesthesia research, applied here to examine differences in subjectively reported levels of imagery usage and intensity. Prior to identifying synesthetes' and non-synesthetes' group membership (in order to eliminate the potential for bias), subjects completed two common measures of visual imagery experiences. Using this approach, we replicated findings of greater visual imagery usage in synesthetes on the Spontaneous Use of Imagery Scale (SUIS) measure, but not of enhanced imagery abilities on the standardized Vividness of Visual Imagery Questionnaire (VVIQ) measure. The present study strengthens prior evidence that synesthesia is associated with heightened visual imagery and demonstrates the utility of double-blind designs in order to limit biases and promote further replicability of other findings in research on synesthesia.