Modulating short-term auditory memory with focal transcranial direct current stimulation applied to the supramarginal gyrus.

Previous studies have shown that transcranial direct current stimulation (tDCS) can affect performance by decreasing regional excitability in a brain region that contributes to the task of interest. To our knowledge, no research to date has found both enhancing and diminishing effects on performance, depending upon which polarity of the current is applied. The supramarginal gyrus (SMG) is an ideal brain region for testing tDCS effects because it is easy to identify using the 10-20 electroencephalography coordinate system, and results of neuroimaging studies have implicated the left SMG in short-term memory for phonological and nonphonological sounds. In the present study, we found that applying tDCS to the left SMG affected pitch memory in a manner that depended upon the polarity of stimulation: cathodal tDCS had a negative impact on performance whereas anodal tDCS had a positive impact. These effects were significantly different from sham stimulation, which did not influence performance; they were also specific to the left hemisphere - no effect was found when applying cathodal stimulation to the right SMG - and were unique to pitch memory as opposed to memory for visual shapes. Our results provide further evidence that the left SMG is a nodal point for short-term auditory storage and demonstrate the potential of tDCS to influence cognitive performance and to causally examine hypotheses derived from neuroimaging studies.

Valproate reopens critical-period learning of absolute pitch.

Absolute pitch, the ability to identify or produce the pitch of a sound without a reference point, has a critical period, i.e., it can only be acquired early in life. However, research has shown that histone-deacetylase inhibitors (HDAC inhibitors) enable adult mice to establish perceptual preferences that are otherwise impossible to acquire after youth. In humans, we found that adult men who took valproate (VPA) (a HDAC inhibitor) learned to identify pitch significantly better than those taking placebo-evidence that VPA facilitated critical-period learning in the adult human brain. Importantly, this result was not due to a general change in cognitive function, but rather a specific effect on a sensory task associated with a critical-period.

Non-invasive brain stimulation enhances the effects of melodic intonation therapy.

Research has suggested that a fronto-temporal network in the right hemisphere may be responsible for mediating melodic intonation therapy's (MIT) positive effects on speech recovery. We investigated the potential for a non-invasive brain stimulation technique, transcranial direct current stimulation (tDCS), to augment the benefits of MIT in patients with non-fluent aphasia by modulating neural activity in the brain during treatment with MIT. The polarity of the current applied to the scalp determines the effects of tDCS on the underlying tissue: anodal-tDCS increases excitability, whereas cathodal tDCS decreases excitability. We applied anodal-tDCS to the posterior inferior frontal gyrus of the right hemisphere, an area that has been shown both to contribute to singing through the mapping of sounds to articulatory actions and to serve as a key region in the process of recovery from aphasia, particularly in patients with large left hemisphere lesions. The stimulation was applied while patients were treated with MIT by a trained therapist. Six patients with moderate to severe non-fluent aphasia underwent three consecutive days of anodal-tDCS + MIT, and an equivalent series of sham-tDCS + MIT. The two treatment series were separated by 1 week, and the order in which the treatments were administered was randomized. Compared to the effects of sham-tDCS + MIT, anodal-tDCS + MIT led to significant improvements in fluency of speech. These results support the hypothesis that, as the brain seeks to reorganize and compensate for damage to left hemisphere language centers, combining anodal-tDCS with MIT may further recovery from post-stroke aphasia by enhancing activity in a right hemisphere sensorimotor network for articulation.

Music to my eyes: cross-modal interactions in the perception of emotions in musical performance.

We investigate non-verbal communication through expressive body movement and musical sound, to reveal higher cognitive processes involved in the integration of emotion from multiple sensory modalities. Participants heard, saw, or both heard and saw recordings of a Stravinsky solo clarinet piece, performed with three distinct expressive styles: restrained, standard, and exaggerated intention. Participants used a 5-point Likert scale to rate each performance on 19 different emotional qualities. The data analysis revealed that variations in expressive intention had their greatest impact when the performances could be seen; the ratings from participants who could only hear the performances were the same across the three expressive styles. Evidence was also found for an interaction effect leading to an emergent property, intensity of positive emotion, when participants both heard and saw the musical performances. An exploratory factor analysis revealed orthogonal dimensions for positive and negative emotions, which may account for the subjective experience that many listeners report of having multi-valent or complex reactions to music, such as "bittersweet."

Cross-modal influences of affect across social and non-social domains in individuals with Williams syndrome.

The Williams syndrome (WS) cognitive profile is characterized by relative strengths in face processing, an attentional bias towards social stimuli, and an increased affinity and emotional reactivity to music. An audio-visual integration study examined the effects of auditory emotion on visual (social/non-social) affect identification in individuals with WS and typically developing (TD) and developmentally delayed (DD) controls. The social bias in WS was hypothesized to manifest as an increased ability to process social than non-social affect, and a reduced auditory influence in social contexts. The control groups were hypothesized to perform similarly across conditions. The results showed that while participants with WS exhibited indistinguishable performance to TD controls in identifying facial affect, DD controls performed significantly more poorly. The TD group outperformed the WS and DD groups in identifying non-social affect. The results suggest that emotionally evocative music facilitated the ability of participants with WS to process emotional facial expressions. These surprisingly strong facial-processing skills in individuals with WS may have been due to the effects of combining social and music stimuli and to a reduction in anxiety due to the music in particular. Several directions for future research are suggested.

Modulating activity in the motor cortex affects performance for the two hands differently depending upon which hemisphere is stimulated.

We modulated neural excitability in the human motor cortex to investigate behavioral effects for both hands. In a previous study, we showed that decreasing excitability in the dominant motor cortex led to a decline in performance for the contralateral hand and an improvement for the ipsilateral hand; increasing excitability produced the opposite effects. Research suggests that the ipsilateral effects were mediated by interhemispheric inhibition. Physiological evidence points to an asymmetry in interhemispheric inhibition between the primary motor cortices, with stronger inhibitory projections coming from the dominant motor cortex. In the present study, we examined whether there is a hemispheric asymmetry in the effects on performance when modulating excitability in the motor cortex. Anodal and cathodal transcranial direct current stimulation were applied to the motor cortex of 17 participants, targeting the non-dominant hemisphere on one day and the dominant hemisphere on another day, along with one sham session. Participants performed a finger-sequence coordination task with each hand before and after stimulation. The dependent variable was calculated as the percentage of change in the number of correct keystrokes. We found that the effects of transcranial direct current stimulation depended upon which hemisphere was stimulated; modulating excitability in the dominant motor cortex significantly affected performance for the contralateral and ipsilateral hands, whereas modulating excitability in the non-dominant motor cortex only had a significant impact for the contralateral hand. These results provide evidence for a hemispheric asymmetry in the ipsilateral effects of modulating excitability in the motor cortex and may be important for clinical research on motor recovery.

Dual-hemisphere tDCS facilitates greater improvements for healthy subjects' non-dominant hand compared to uni-hemisphere stimulation.

BACKGROUND: Transcranial direct current stimulation (tDCS) is a non-invasive technique that has been found to modulate the excitability of neurons in the brain. The polarity of the current applied to the scalp determines the effects of tDCS on the underlying tissue: anodal tDCS increases excitability, whereas cathodal tDCS decreases excitability. Research has shown that applying anodal tDCS to the non-dominant motor cortex can improve motor performance for the non-dominant hand, presumably by means of changes in synaptic plasticity between neurons. Our previous studies also suggest that applying cathodal tDCS over the dominant motor cortex can improve performance for the non-dominant hand; this effect may result from modulating inhibitory projections (interhemispheric inhibition) between the motor cortices of the two hemispheres. We hypothesized that stimultaneously applying cathodal tDCS over the dominant motor cortex and anodal tDCS over the non-dominant motor cortex would have a greater effect on finger sequence performance for the non-dominant hand, compared to stimulating only the non-dominant motor cortex. Sixteen right-handed participants underwent three stimulation conditions: 1) dual-hemisphere - with anodal tDCS over the non-dominant motor cortex, and cathodal tDCS over the dominant motor cortex, 2) uni-hemisphere - with anodal tDCS over the non-dominant motor cortex, and 3) sham tDCS. Participants performed a finger-sequencing task with the non-dominant hand before and after each stimulation. The dependent variable was the percentage of change in performance, comparing pre- and post-tDCS scores. RESULTS: A repeated measures ANOVA yielded a significant effect of tDCS condition (F(2,30) = 4.468, p = .037). Post-hoc analyses revealed that dual-hemisphere stimulation improved performance significantly more than both uni-hemisphere (p = .021) and sham stimulation (p = .041). CONCLUSION: We propose that simultaneously applying cathodal tDCS over the dominant motor cortex and anodal tDCS over the non-dominant motor cortex produced an additive effect, which facilitated motor performance in the non-dominant hand. These findings are relevant to motor skill learning and to research studies of motor recovery after stroke.

Testing for causality with transcranial direct current stimulation: pitch memory and the left supramarginal gyrus.

Neuroimaging studies have implicated the left supramarginal gyrus in short-term auditory memory processing, including memory for pitch. The present study investigated the causal role of the left supramarginal gyrus in short-term pitch memory by comparing the effects of cathodal transcranial direct current stimulation when applied over the left or right supramarginal gyrus with sham transcranial direct current stimulation. Only cathodal transcranial direct current stimulation over the left supramarginal gyrus had a detrimental effect on short-term pitch-memory performance in 11 adult participants. These results provide support for the important role of the left supramarginal gyrus in short-term memory for pitch information, and they further demonstrate the potential of transcranial direct current stimulation to modulate the functional contribution of a brain area to a particular cognitive process.

Contralateral and ipsilateral motor effects after transcranial direct current stimulation.

Transcranial direct current stimulation over the left motor area influenced both contralateral and ipsilateral finger sequence movements in seven healthy adults. Effects for the two hands were reversed: anodal stimulation improved right-hand performance significantly more than cathodal stimulation, whereas cathodal stimulation improved left-hand performance significantly more than anodal stimulation. The results show that stimulating a motor region directly, or indirectly by modulating activity in the homologous region on the opposite hemisphere, can affect motor skill acquisition, presumably by facilitating effective synaptic connectivity. This outcome provides evidence for the role of interhemispheric inhibition in corticomotor functioning, and also has implications for treatment methods aimed at facilitating motor recovery after stroke.

Cross-modal interactions in the perception of musical performance.

We investigate the dynamics of sensory integration for perceiving musical performance, a complex natural behavior. Thirty musically trained participants saw, heard, or both saw and heard, performances by two clarinetists. All participants used a sliding potentiometer to make continuous judgments of tension (a measure correlated with emotional response) and continuous judgments of phrasing (a measure correlated with perceived musical structure) as performances were presented. The data analysis sought to reveal relations between the sensory modalities (vision and audition) and to quantify the effect of seeing the performances on participants' overall subjective experience of the music. In addition to traditional statistics, functional data analysis techniques were employed to analyze time-varying aspects of the data. The auditory and visual channels were found to convey similar experiences of phrasing but different experiences of tension through much of the performances. We found that visual information served both to augment and to reduce the experience of tension at different points in the musical piece (as revealed by functional linear modeling and functional significance testing). In addition, the musicians' movements served to extend the sense of phrasing, to cue the beginning of new phrases, to indicate musical interpretation, and to anticipate changes in emotional content. Evidence for an interaction effect suggests that there may exist an emergent quality when musical performances are both seen and heard. The investigation augments knowledge of human communicative processes spanning language and music, and involving multiple modalities of emotion and information transfer.

Dimensions of emotion in expressive musical performance.

This paper explores the dimensions of emotion conveyed by music. Participants rated emotion terms after seeing and/or hearing recordings of clarinet performances that varied in expressive content. A factor analysis revealed four independent dimensions of emotion. Changes to the clarinetists' expressive intentions did not significantly affect emotions conveyed by sound. It was largely through the visual modality that expressive intentions influenced the experience for observers.