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').     

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.     

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.     

The colors of the alphabet: naturally-biased associations between shape and color

Many letters of the alphabet are consistently mapped to specific colors in English-speaking adults, both in the general population and in individuals with grapheme-color synaesthesia who perceive letters in color. Here, across six experiments, we tested the ubiquity of the color/letter associations with typically developing toddlers, literate children, and adults. We found that pre-literate children associate O with white and X with black and discovered that they also associate I and ameboid nonsense shapes with white; Z and jagged nonsense shapes with black; and C with yellow; but do not make a number of other associations (B blue; Y yellow; A red; G green) seen in literate children and adults. The toddlers' mappings were based on the shape and not the sound of the letter. The results suggest that sensory cortical organization initially binds specific colors to some specific shapes and that learning to read can induce additional associations, likely through the influence of higher order networks as letters take on meaning.     

The colour of Os: naturally biased associations between shape and colour

Many letters of the alphabet are consistently mapped to specific colours by English-speaking adults, both in the general population and in individuals with grapheme-colour synaesthesia who perceive letters in colour. Such associations may be naturally biased by intrinsic sensory cortical organisation, or may be based in literacy (eg 'A' is for 'apple', apples are red; therefore A is red). To distinguish these two hypotheses, we tested pre-literate children in three experiments and compared their results to those of literate children (aged 7-9 years) and adults. The results indicate that some colour letter mappings (O white, X black) are naturally biased by the shape of the letter, whereas others (A red, G green) may be based in literacy. They suggest that sensory cortical organisation initially binds colour to some shapes, and that learning to read can induce additional associations, likely through the influence of higher-order networks as letters take on meaning.     

The development of infant learning about specific face-voice relations

This study examined the development of infants' ability to perceive, learn, and remember the unique face-voice relations of unfamiliar adults. Infants of 2, 4, and 6 months were habituated to the faces and voices of 2 same-gender adults speaking and then received test trials where the faces and voices were synchronized yet mismatched. Results indicated that 4- and 6-month-olds, but not 2-month-olds, detected the change in face-voice pairings. Two-month-olds did, however, discriminate among the faces and voices in a control study. Results of a subsequent intermodal matching procedure indicated that only the 6-month-olds showed matching and memory for the face-voice relations. These findings suggest that infants' ability to detect the arbitrary relations between specific faces and voices of unfamiliar adults emerges between 2 and 4 months of age, whereas matching and memory for these relations emerges somewhat later, perhaps between 4 and 6 months of age.     

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.     

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.     

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.     

Grapheme–colour synaesthesia yields an ordinary rather than extraordinary memory advantage: Evidence from a group study

In synaesthesia, the input of one sensory modality automatically triggers an additional experience, not normally triggered by the input of that modality. Therefore, compared to non-synaesthetes, additional experiences exist and these may be used as retrieval cues when memory is tested. Previous case studies have suggested that synaesthesia may yield even extraordinary memory abilities. However, group studies found either a task-specific memory advantage or no performance advantage at all. The aim of the present study was to test whether grapheme–colour synaesthesia gives rise to a general memory benefit using a standardised memory test (Wechsler Memory Scale). The synaesthetes showed a performance advantage in episodic memory tests, but not in short-term memory tests. However, performance was still within the ordinary range. The results support the hypothesis that synaesthesia provides for a richer world of experience and as a consequence additional retrieval cues may be available and beneficial but not to the point of extraordinary memory ability.     

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.     

Training synaesthesia

Synaesthetic inducers such as graphemes are typically cultural artifacts. Thus, a learning component seems evident in synaesthesia (Simner et al, 2009 Brain 132 57-64). Normally, synaesthetes report to have their experiences since they can remember. Nevertheless, a recent training study suggests that synaesthesia can be mimicked in non-synaesthetes. To date, the role of learning during the development of synaesthesia is still debated. It is not clear whether synaesthesia can be learned or trained at all. To address this question, we compared a non-adaptive and an adaptive training for their effectiveness. We assessed their impact on two types of priming tasks, before and after the training. We found stronger priming in the adaptive training group suggesting that adaptive training is more efficient to mimic synaesthesia.     

疼痛共情的神经机制

疼痛是一种多维度的体验。情感成分和感觉成分是疼痛体验的主要成分,二者有着不同的神经机制。疼痛共情是痛觉"共鸣"的情绪反应,其神经机制的研究也围绕疼痛的情感成分和感觉成分展开。疼痛共情的情感成分的相关脑区主要有前扣带皮层和脑岛等脑区,与自身疼痛时相比,它们的激活位点及所处的神经网络存在差异;疼痛共情的感觉成分主要与初级感觉皮层以及一些其它的躯体感觉皮层有关,对它们的激活情况的研究受技术手段和实验范式的影响较大。未来的研究应从技术手段、分析方法及实验操作上予以关注。     

When does an ostrich become a bird? The role of typicality in early word comprehension.

Which objects and animals are children willing to accept as referents for words they know? To answer this question, the authors assessed early word comprehension using the preferential looking task. Children were shown 2 stimuli side by side (a target and a distractor) and heard the target stimulus named. The target stimulus was either a typical or an atypical exemplar of the named category. It was predicted that children first connect typical examples with the target name and broaden the extension of the name as they get older to include less typical examples. Experiment 1 shows that when targets are named, 12-month-olds display an increase in target looking for typical but not atypical targets whereas 24-month-olds display an increase for both. Experiment 2 shows that 18-month-olds display a pattern similar to that of 24-month-olds. Implications for the early development of word comprehension are discussed.     

错误信息持续影响效应的神经基础

本研究通过分析任务态fMRI下相关脑区激活及功能连接的条件间差异以揭示CIEM神经基础并为心理模型更新及记忆提取失败假说提供更多证据。结果发现, 更正条件的推理分显著高于控制条件, 存在CIEM。编码阶段左颞中回在更正条件下的激活显著弱于控制条件, 提取阶段更正条件下额中回及前扣带回激活更弱、额中回与中央前回的功能连接更强。结果提示上述脑区可能参与了CIEM的形成, 并从神经层面提供了心理模型更新和记忆提取失败假说可能解释了CIEM形成的不同阶段的证据。     

Longitudinal stability of temperamental exuberance and social-emotional outcomes in early childhood

The goals of the current study were to investigate the stability of temperamental exuberance across infancy and toddlerhood and to examine the associations between exuberance and social-emotional outcomes in early childhood. The sample consisted of 291 4-month-olds followed at 9, 24, and 36 months and again at 5 years of age. Behavioral measures of exuberance were collected at 9, 24, and 36 months. At 36 months, frontal electroencephalogram (EEG) asymmetry was assessed. At 5 years, maternal reports of temperament and behavior problems were collected, as were observational measures of social behavior during an interaction with an unfamiliar peer in the laboratory. Latent profile analysis revealed a high, stable exuberance profile that was associated with greater ratings of 5-year externalizing behavior and surgency, as well as observed disruptive behavior and social competence with unfamiliar peers. These associations were particularly true for children who displayed left frontal EEG asymmetry. Multiple factors supported an approach bias for exuberant temperament but did not differentiate between adaptive and maladaptive social-emotional outcomes at 5 years of age.     

Toward a psychology of positive youth development

This article analyzes the development of initiative as an exemplar of one of many learning experiences that should be studied as part of positive youth development. The capacity for initiative is essential for adults in our society and will become more important in the 21st century, yet adolescents have few opportunities to learn it. Their typical experiences during schoolwork and unstructured leisure do not reflect conditions for learning initiative. The context best suited to the development of initiative appears to be that of structured voluntary activities, such as sports, arts, and participation in organizations, in which youths experience the rare combination of intrinsic motivation in combination with deep attention. An incomplete body of outcome research suggests that such activities are associated with positive development, but the developmental processes involved are only beginning to be understood. One promising approach has recorded language use and has found that adolescents participating in effective organizations acquire a new operating language that appears to correspond to the development of initiative.     

Development of the functional visual field

Andries Sanders' dissertation examined selective mechanisms in the functional visual field, and much of his work since has been concerned with the stages that underlie visual information processing particularly while making saccades. We argue that the study of orienting in the functional visual field is timely because it deals with the relation of covert attention shifts, eye movements and head movements to their underlying neurology. In our paper we develop a method to study learning of sequences at all ages from infants to adults. Our studies focus on how learning influences anticipatory eye movements. We examined the learning of unambiguous and context dependent sequences by 4-, 10-, and 18-month-old infants and undergraduates. We found clear learning of unambiguous sequences at 4 months, but learning of context dependent associations was found only in 18-month-olds and in adults. We hypothesize that the learning of unambiguous sequences by 4-month-olds reflects maturation of a basal ganglia-parietal circuit related to adult implicit learning, while the learning of context dependent sequences requires development of frontal structures underlying more general attentional abilities.     

Inhibition and exuberance in preschool classrooms: associations with peer social experiences and changes in cortisol across the preschool year

Associations between behavioral inhibition and activity of the hypothalamic-pituitary-adrenocortical (HPA) system, a stress-sensitive neuroendocrine system indexed by salivary cortisol, have varied widely across studies. In the current study, we examined the role of peer social experiences in moderating patterns of association between inhibition/risk-aversion and cortisol reactivity. As expected based on previous research, preschool children (N = 165, 78 boys, 87 girls, 3.0-5.0 years) had significantly different social experiences in their preschool classrooms depending on temperament. Highly inhibited/risk-averse children were less socially integrated, less dominant, and less involved in aggressive encounters than both average and highly exuberant/risk-seeking children, but they were no more likely to be peer rejected. Highly exuberant children were more dominant, exhibited anger more often, and had friendships characterized by higher conflict. Cortisol levels fell from fall to spring for average and highly exuberant children but not for highly inhibited children. Unexpectedly, for highly inhibited children, having friends and being more dominant and popular than other highly inhibited children was associated with increasing cortisol levels over the school year. In contrast, highly exuberant children who were less socially integrated than other highly exuberant children maintained higher cortisol levels. Results indicate that the types of social experiences that affect stress-responsive biological systems may differ markedly for highly inhibited and highly exuberant children.