Nina Bien

The Brain's Intention to Imitate: The Neurobiology of Intentional versus Automatic Imitation

Whenever we observe a movement of a conspecific, our mirror neuron system becomes activated, urging us to imitate the observed movement. However, because such automatic imitation is not always appropriate, an inhibitive component keeping us from imitating everything we see seems crucial for an effective social behavior. This becomes evident from neuropsychological conditions like echopraxia, in which this suppression is absent. Here, we unraveled the neurodynamics underlying this proposed inhibition of automatic imitation by measuring and manipulating brain activity during the execution of a stimulus-response compatibility paradigm. Within the identified connectivity network, right middle/inferior frontal cortex sends neural input concerning general response inhibition to right premotor cortex, which is involved in automatic imitation. Subsequently, the fully prepared imitative response is sent to left opercular cortex that functions as a final gating mechanism for intentional imitation. We propose an informed neurocognitive model of inhibition of automatic imitation, suggesting a functional dissociation between automatic and intentional imitation.

The sound of size: Crossmodal binding in pitch-size synesthesia: A combined TMS, EEG and psychophysics study

Crossmodal binding usually relies on bottom-up stimulus characteristics such as spatial and temporal correspondence. However, in case of ambiguity the brain has to decide whether to combine or segregate sensory inputs. We hypothesise that widespread, subtle forms of synesthesia provide crossmodal mapping patterns which underlie and influence multisensory perception. Our aim was to investigate if such a mechanism plays a role in the case of pitch-size stimulus combinations. Using a combination of psychophysics and ERPs, we could show that despite violations of spatial correspondence, the brain specifically integrates certain stimulus combinations which are congruent with respect to our hypothesis of pitch-size synesthesia, thereby impairing performance on an auditory spatial localisation task (Ventriloquist effect). Subsequently, we perturbed this process by functionally disrupting a brain area known for its role in multisensory processes, the right intraparietal sulcus, and observed how the Ventriloquist effect was abolished, thereby increasing behavioural performance. Correlating behavioural, TMS and ERP results, we could retrace the origin of the synesthestic pitch-size mappings to a right intraparietal involvement around 250 ms. The results of this combined psychophysics, TMS and ERP study provide evidence for shifting the current viewpoint on synesthesia more towards synesthesia being at the extremity of a spectrum of normal, adaptive perceptual processes, entailing close interplay between the different sensory systems. Our results support this spectrum view of synesthesia by demonstrating that its neural basis crucially depends on normal multisensory processes.

Automatic and intentional number processing both rely on intact right parietal cortex: A combined fMRI and neuronavigated TMS study

Practice and training usually lead to performance increase in a given task. In addition, a shift from intentional toward more automatic processing mechanisms is often observed. It is currently debated whether automatic and intentional processing is subserved by the same or by different mechanism(s), and whether the same or different regions in the brain are recruited. Previous correlational evidence provided by behavioral, neuroimaging, modeling, and neuropsychological studies addressing this question yielded conflicting results. Here we used transcranial magnetic stimulation (TMS) to compare the causal influence of disrupting either left or right parietal cortex during automatic and intentional numerical processing, as reflected by the size congruity effect and the numerical distance effect, respectively. We found a functional hemispheric asymmetry within parietal cortex with only the TMS-induced right parietal disruption impairing both automatic and intentional numerical processing. In contrast, disrupting the left parietal lobe with TMS, or applying sham stimulation, did not affect performance during automatic or intentional numerical processing. The current results provide causal evidence for the functional relevance of right, but not left, parietal cortex for intentional, and automatic numerical processing, implying that at least within the parietal cortices, automatic, and intentional numerical processing rely on the same underlying hemispheric lateralization.

Audio-visual onset differences are used to determine syllable identity for ambiguous audio-visual stimulus pairs

Content and temporal cues have been shown to interact during audio-visual (AV) speech identification. Typically, the most reliable unimodal cue is used more strongly to identify specific speech features; however, visual cues are only used if the AV stimuli are presented within a certain temporal window of integration (TWI). This suggests that temporal cues denote whether unimodal stimuli belong together, that is, whether they should be integrated. It is not known whether temporal cues also provide information about the identity of a syllable. Since spoken syllables have naturally varying AV onset asynchronies, we hypothesize that for suboptimal AV cues presented within the TWI, information about the natural AV onset differences can aid in speech identification. To test this, we presented low-intensity auditory syllables concurrently with visual speech signals, and varied the stimulus onset asynchronies (SOA) of the AV pair, while participants were instructed to identify the auditory syllables. We revealed that specific speech features (e.g., voicing) were identified by relying primarily on one modality (e.g., auditory). Additionally, we showed a wide window in which visual information influenced auditory perception, that seemed even wider for congruent stimulus pairs. Finally, we found a specific response pattern across the SOA range for syllables that were not reliably identified by the unimodal cues, which we explained as the result of the use of natural onset differences between AV speech signals. This indicates that temporal cues not only provide information about the temporal integration of AV stimuli, but additionally convey information about the identity of AV pairs. These results provide a detailed behavioral basis for further neuro-imaging and stimulation studies to unravel the neurofunctional mechanisms of the audio-visual-temporal interplay within speech perception.

Extinguishing extinction: Hemispheric differences in the modulation of tms-induced visual extinction by directing covert spatial attention

The topic of spatial attention is of great relevance for researchers in various fields, including neuropsychology, cognitive neuroscience, and cognitive psychology, as well as for clinical practice. Deficits of spatial attentional arising from parietal brain damage remain largely confined to the left visual field. The mechanisms underlying this hemispheric asymmetry are still elusive. We mimicked the neuropsychological syndrome of contralesional extinction by temporarily inducing a spatial attentional bias in healthy volunteers with TMS. We investigated whether directing covert spatial attention could enhance or, more importantly, counteract the resulting behavioral deficits. Although both the left and right parietal TMS induced contralateral extinction, only left hemifield extinction following right parietal TMS was severely aggravated by a competing stimulus in the ipsilesional (right) hemifield. We put forward the hypothesis that an asymmetry with respect to the ability of detaching attention from a distractor is contributing to the right hemispheric lateralization with regard to extinction. On a broader level, we suggest that “virtual patients” might be used for evaluating neuropsychological treatment in an early stage of development, reducing the burden on actual patients.

Dissecting hemisphere-specific contributions to visual spatial imagery using parametric brain mapping

In the current study we aimed to empirically test previously proposed accounts of a division of labour between the left and right posterior parietal cortices during visuospatial mental imagery. The representation of mental images in the brain has been a topic of debate for several decades. Although the posterior parietal cortex is involved bilaterally, previous studies have postulated that hemispheric specialisation might result in a division of labour between the left and right parietal cortices. In the current fMRI study, we used an elaborated version of a behaviourally-controlled spatial imagery paradigm, the mental clock task, which involves mental image generation and a subsequent spatial comparison between two angles. By systematically varying the difference between the two angles that are mentally compared, we induced a symbolic distance effect: smaller differences between the two angles result in higher task difficulty. We employed parametrically weighed brain imaging to reveal brain areas showing a graded activation pattern in accordance with the induced distance effect. The parametric difficulty manipulation influenced behavioural data and brain activation patterns in a similar matter. Moreover, since this difficulty manipulation only starts to play a role from the angle comparison phase onwards, it allows for a top-down dissociation between the initial mental image formation, and the subsequent angle comparison phase of the spatial imagery task. Employing parametrically weighed fMRI analysis enabled us to top-down disentangle brain activation related to mental image formation, and activation reflecting spatial angle comparison. The results provide first empirical evidence for the repeatedly proposed division of labour between the left and right posterior parietal cortices during spatial imagery.

Moving beyond attentional biases: Shifting the interhemispheric balance between left and right posterior parietal cortex modulates attentional control processes

The concept of interhemispheric competition has been very influential in attention research, and the occurrence of biased attention due to an imbalance in posterior parietal cortex (PPC) is well documented. In this context, the vast majority of studies have assessed attentional performance with tasks that did not include an explicit experimental manipulation of attention, and, as a consequence, it remains largely unknown how these findings relate to core attentional constructs such as endogenous and exogenous control and spatial orienting and reorienting. We here addressed this open question by creating an imbalance between left and right PPC with transcranial direct current stimulation, resulting in right-hemispheric dominance, and assessed performance on three experimental paradigms that isolate distinct attentional processes. The comparison between active and sham transcranial direct current stimulations revealed a highly informative pattern of results with differential effects across tasks. Our results demonstrate the functional necessity of PPC for endogenous and exogenous attentional control and, importantly, link the concept of interhemispheric competition to core attentional processes, thus moving beyond the notion of biased attention after noninvasive brain stimulation over PPC.

Corrigendum to ‘The sound of size’ crossmodal binding in pitch-size synesthesia: A combined TMS, EEG and psychophysics study [NeuroImage 59/1(2012) 663–672]

The authors regret that on page 664 of the Introduction it states that “To this end, we employed the Ventriloquist paradigm (Driver, 1996)”, and in the Discussion on page 668 the article states: “To this end we employed a pitch-size variant of the spatial Ventriloquist illusion”. Although in this context we also cited the work by Parise and Spence (2009), we would like to explicitly clarify that we here in fact employed the behavioral task described by Parise and Spence (2009) in their psychophysical experiment 3. Also, on page 668 of our Discussion it states that “Recent studies (Parise and Spence, 2008, 2009) reported some initial proof of the existence of pitch-size synesthesia, using a similar approach.” However, this should rather read that: “Parise and Spence (2009) were the first to provide psychophysical evidence that synesthetic congruency (i.e., crossmodal correspondences) can promote multisensory integration.”.

Audiovisual onset differences are used to determine the identity of ambiguous syllables

Content and temporal cues have been shown to interact during audiovisual (AV) speech identification. Typically, the most reliable unimodal cue is used to identify specific speech features; however, visual cues are only used if the audiovisual stimuli are presented within a certain temporal integration window (TWI). This suggests that temporal cues denote whether unimodal stimuli belong together and should be integrated. It is unknown whether temporal cues also provide information about speech content. Since spoken syllables have naturally varying audiovisual onset asynchronies, we hypothesize that for suboptimal AV cues presented within the TWI, these natural AV onset differences can aid in speech identification. To test this, we presented low-intensity auditory syllables concurrently with visual speech signals, and varied the stimulus onset asynchronies (SOA) of the audiovisual pair, while participants identified the syllables. We revealed that the most reliable cues of the audiovisual input were used to identify specific speech features (e.g., voicing). Additionally, we showed that the TWI was wider for congruent stimulus pairs. Finally, we found a specific response pattern across the SOA-range for syllables that were not reliably identified by the unimodal cues, which we explained by the use of natural onset differences between audiovisual speech signals. This indicates that temporal cues not only determine whether or not different inputs belong together, but additionally convey identity information of audiovisual pairs. These results provide a detailed behavioral basis for further neuro-imaging and stimulation studies to unravel the neurofunctional mechanisms of the audiovisual–temporal interplay within speech perception.