Olympia Colizoli

Pseudo-Synesthesia through Reading Books with Colored Letters

Background Synesthesia is a phenomenon where a stimulus produces consistent extraordinary subjective experiences. A relatively common type of synesthesia involves perception of color when viewing letters (e.g. the letter ‘a’ always appears as light blue). In this study, we examine whether traits typically regarded as markers of synesthesia can be acquired by simply reading in color. Methodology/Principal Findings Non-synesthetes were given specially prepared colored books to read. A modified Stroop task was administered before and after reading. A perceptual crowding task was administered after reading. Reading one book (>49,000 words) was sufficient to induce effects regarded as behavioral markers for synesthesia. The results of the Stroop tasks indicate that it is possible to learn letter-color associations through reading in color (F(1, 14) = 5.85, p = .030). Furthermore, Stroop effects correlated with subjective reports about experiencing letters in color (r(13) = 0.51, p = .05). The frequency of viewing letters is related to the level of association as seen by the difference in the Stroop effect size between upper- and lower-case letters (t(14) = 2.79, p = .014) and in a subgroup of participants whose Stroop effects increased as they continued to read in color. Readers did not show significant performance advantages on the crowding task compared to controls. Acknowledging the many differences between trainees and synesthetes, results suggest that it may be possible to acquire a subset of synesthetic behavioral traits in adulthood through training. Conclusion/Significance To our knowledge, this is the first evidence of acquiring letter-color associations through reading in color. Reading in color appears to be a promising avenue in which we may explore the differences and similarities between synesthetes and non-synesthetes. Additionally, reading in color is a plausible method for a long-term ‘synesthetic’ training program.

Dynamic modulation of decision biases by Brainstem Arousal Systems

Decision-makers often arrive at different choices when faced with repeated presentations of the same evidence. Variability of behavior is commonly attributed to noise in the brain’s decision-making machinery. We hypothesized that phasic responses of brainstem arousal systems are a significant source of this variability. We tracked pupil responses (a proxy of phasic arousal) during sensory-motor decisions in humans, across different sensory modalities and task protocols. Large pupil responses generally predicted a reduction in decision bias. Using fMRI, we showed that the pupil-linked bias reduction was (i) accompanied by a modulation of choice-encoding pattern signals in parietal and prefrontal cortex and (ii) predicted by phasic, pupil-linked responses of a number of neuromodulatory brainstem centers involved in the control of cortical arousal state, including the noradrenergic locus coeruleus. We conclude that phasic arousal suppresses decision bias on a trial-by-trial basis, thus accounting for a significant component of the variability of choice behavior. DOI: http://dx.doi.org/10.7554/eLife.23232.001

A taste for words and sounds: a case of lexical-gustatory and sound-gustatory synesthesia

Gustatory forms of synesthesia involve the automatic and consistent experience of tastes that are triggered by non-taste related inducers. We present a case of lexical-gustatory and sound-gustatory synesthesia within one individual, SC. Most words and a subset of non-linguistic sounds induce the experience of taste, smell and physical sensations for SC. SCs lexical-gustatory associations were significantly more consistent than those of a group of controls. We tested for effects of presentation modality (visual vs. auditory), taste-related congruency, and synesthetic inducer-concurrent direction using a priming task. SCs performance did not differ significantly from a trained control group. We used functional magnetic resonance imaging to investigate the neural correlates of SCs synesthetic experiences by comparing her brain activation to the literature on brain networks related to language, music, and sound processing, in addition to synesthesia. Words that induced a strong taste were contrasted to words that induced weak-to-no tastes (“tasty” vs. “tasteless” words). Brain activation was also measured during passive listening to music and environmental sounds. Brain activation patterns showed evidence that two regions are implicated in SCs synesthetic experience of taste and smell: the left anterior insula and left superior parietal lobe. Anterior insula activation may reflect the synesthetic taste experience. The superior parietal lobe is proposed to be involved in binding sensory information across sub-types of synesthetes. We conclude that SCs synesthesia is genuine and reflected in her brain activation. The type of inducer (visual-lexical, auditory-lexical, and non-lexical auditory stimuli) could be differentiated based on patterns of brain activity.

Training Synesthetic Letter-color Associations by Reading in Color

Synesthesia is a rare condition in which a stimulus from one modality automatically and consistently triggers unusual sensations in the same and/or other modalities. A relatively common and well-studied type is grapheme-color synesthesia, defined as the consistent experience of color when viewing, hearing and thinking about letters, words and numbers. We describe our method for investigating to what extent synesthetic associations between letters and colors can be learned by reading in color in nonsynesthetes. Reading in color is a special method for training associations in the sense that the associations are learned implicitly while the reader reads text as he or she normally would and it does not require explicit computer-directed training methods. In this protocol, participants are given specially prepared books to read in which four high-frequency letters are paired with four high-frequency colors. Participants receive unique sets of letter-color pairs based on their pre-existing preferences for colored letters. A modified Stroop task is administered before and after reading in order to test for learned letter-color associations and changes in brain activation. In addition to objective testing, a reading experience questionnaire is administered that is designed to probe for differences in subjective experience. A subset of questions may predict how well an individual learned the associations from reading in color. Importantly, we are not claiming that this method will cause each individual to develop grapheme-color synesthesia, only that it is possible for certain individuals to form letter-color associations by reading in color and these associations are similar in some aspects to those seen in developmental grapheme-color synesthetes. The method is quite flexible and can be used to investigate different aspects and outcomes of training synesthetic associations, including learning-induced changes in brain function and structure.

Defining (trained) grapheme-color synesthesia

There is a current debate over the possibility and validity of synesthesia training experiments (Deroy and Spence, 2013; Rothen and Meier, 2014). In order to test whether a trainee should be considered to have acquired a trained form of synesthesia, a precise definition and specific diagnostic criteria of synesthesia are necessary. There is currently not one specific checklist available including all specific diagnostic methods and criteria, exacerbating the determination and interpretation of differences between (potentially) trained and genuine synesthesia. In order to facilitate communication surrounding these issues, we propose a practical guideline for diagnosing the specific characteristics that are typical of grapheme-color synesthesia (GCS). These guidelines can be applied to developmental cases of GCS, cases of trainees who may show synesthetic traits, and other types of acquired forms of GCS at the level of a single individual.

Task-evoked pupil responses reflect internal belief states

Perceptual decisions about the state of the environment are often made in the face of uncertain evidence. Internal uncertainty signals are considered important regulators of learning and decision-making. A growing body of work has implicated the brain’s arousal systems in uncertainty signaling. Here, we found that two specific computational variables, postulated by recent theoretical work, evoke boosts of arousal at different times during a perceptual decision: decision confidence (the observer’s internally estimated probability that a choice was correct given the evidence) before feedback, and prediction errors (deviations from expected reward) after feedback. We monitored pupil diameter, a peripheral marker of central arousal state, while subjects performed a challenging perceptual choice task with a delayed monetary reward. We quantified evoked pupil responses during decision formation and after reward-linked feedback. During both intervals, decision difficulty and accuracy had interacting effects on pupil responses. Pupil responses negatively scaled with decision confidence prior to feedback and scaled with uncertainty-dependent prediction errors after feedback. This pattern of pupil responses during both intervals was in line with a model using the observer’s graded belief about choice accuracy to anticipate rewards and compute prediction errors. We conclude that pupil-linked arousal systems are modulated by internal belief states.

Author Correction: Task-evoked pupil responses reflect internal belief states

A correction to this article has been published and is linked from the HTML and PDF versions of this paper. The error has not been fixed in the paper.

Creating Colored Letters: Familial Markers of Grapheme–Color Synesthesia in Parietal Lobe Activation and Structure

Perception is inherently subjective, and individual differences in phenomenology are well illustrated by the phenomenon of synesthesia (highly specific, consistent, and automatic cross-modal experiences, in which the external stimulus corresponding to the additional sensation is absent). It is unknown why some people develop synesthesia and others do not. In the current study, we tested whether neural markers related to having synesthesia in the family were evident in brain function and structure. Relatives of synesthetes (who did not have any type of synesthesia themselves) and matched controls read specially prepared books with colored letters for several weeks and were scanned before and after reading using magnetic resonance imaging. Effects of acquired letter–color associations were evident in brain activation. Training-related activation (while viewing black letters) in the right angular gyrus of the parietal lobe was directly related to the strength of the learned letter–color associations (behavioral Stroop effect). Within this obtained angular gyrus ROI, the familial trait of synesthesia related to brain activation differences while participants viewed both black and colored letters. Finally, we compared brain structure using voxel-based morphometry and diffusion tensor imaging to test for group differences and training effects. One cluster in the left superior parietal lobe had significantly more coherent white matter in the relatives compared with controls. No evidence for experience-dependent plasticity was obtained. For the first time, we present evidence suggesting that the (nonsynesthete) relatives of grapheme–color synesthetes show atypical grapheme processing as well as increased brain connectivity.

Pseudo-synesthesia through reading books with colored letters and experience-dependent plasticity of the visual system

Grapheme–color synesthesia is defined as the experience of color in relation to letters, words and numbers. This form of synesthesia is a relatively common type and has received the most attention in the research field in terms of the neural basis of the synesthetic experience and behavioral features of synesthesia such as attention and memory. Grapheme–color and linguistic–color synesthesia have a genetic component. Still, language (the most common synesthetic inducer) is acquired through an interaction between genes and the environment. In order to probe this interaction, we compared relatives of grapheme–color synesthetes to a group of matched controls in a reading-in-color training paradigm. All participants read specially prepared books in which four high-frequency letters were paired with four high-frequency colors. Magnetic resonance images (MRI) were acquired before and after training, including diffusion tensor imaging (DTI). In this sample, there was no evidence of increased acquisition of the letter–color associations by the relatives of synesthetes. No group differences were found in behavior, while there are differences in functional activation on the Stroop task. Imaging results showed significant differences in brain activation in regions known to be involved in grapheme–color synesthesia across both groups. These regions include temporal–occipital cortex, inferior and superior parietal lobe, precentral gyrus, and frontal pole. We conclude that the neural basis of the effects of reading in color relies primarily on the same brain regions as the experience of (authentic) grapheme–color synesthesia.