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Spatial for RTs or hit rates HR , suggesting that the baseline difficulty levels of each task were consistent. TMS experiments were conducted in a dimly lit sound-attenuated low-reverberation chamber.

TMS coil orientation was perpendicular to the individual local curvature of Sylvian fissure, i. Bilateral TMS was chosen because sounds are processed simultaneously in both hemispheres, and because previous studies 4 suggest more consistent effects after bilateral than unilateral AC deactivations. Bilateral TMS, presumably, also helped avoid lateralized distraction effects during Spatial task. Based on previous studies 3 , 46 , the TMS pulse pairs were applied randomly at four different latencies 55, 85, , ms after the probe-sound onset, covering the typical N1 m activity in ACs.

The experiment was divided into eight runs, each containing 80 trials and lasting on average 7 min 18 s. Each session, thus, included ppTMS events.

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On average, ppTMS was delivered about every 10—13 s. To avoid making the experiment excessively long, all conditions in one TMS location were always conducted in a row coil re-targeting may require extra time when two coils are used. The order of TMS locations and task conditions was counterbalanced across subjects.

The TMS target locations were selected based on a sample of previously published maximally-activated fMRI 10 , 11 , 13 — 16 and positron emission tomography 51 voxels during audiospatial or sound-object identity processing. More specifically, the posterior AC targets were defined by averaging the peak-voxel Talairach coordinates of contrasts reflecting sound movement vs. When needed, MNI coordinates were converted to Talairach coordinates. The anterior AC targets were defined similarly, based on peak voxels in contrasts reflecting varying vs. For both targets, average loci were calculated first within, then across studies.

Finally, before the TMS experiment, an optimized entry point closer to the inner skull was selected, as guided by the E-field estimates produced by the navigation system, to allow the TMS effects to be maximally focused to the target area. Electromagnetic modeling analyses utilizing realistic anatomy were conducted to estimate the actual cortically induced E-field distributions in each subject.

The intracranial space was considered as a homogenous isotropic volume conductor. TMS-induced E-field amplitudes were computed at the white matter surfaces extracted using Freesurfer according to well-established physical principles, as described by Nummenmaa and colleagues A full factorial design was utilized to control for potential biases caused by the TMS side effects acoustic clicks, muscle stimulation.

Behavioral data, including RTs correct responses only and HRs, were recorded separately for each task condition. Statistical analyses were conducted using the R packages lme4, languageR, and lsmeans 58 — To further mitigate within-session fluctuations e. The RT lag time series were, finally, entered into a linear mixed model: The random effects included the subject and trial type sound direction nested within Spatial task; AM frequency nested within Identity task ; The fixed effects included the task Identity vs.

Spatial , TMS target location anterior vs.

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We controlled for potential biases caused by non-normality and homogeneity by examining the model residuals against fitted values. Multicollinearity was addressed by residualizing predictors correlating with each other. Our main hypothesis regarded the interaction between the task and TMS target location: We hypothesized that RT lags are significantly larger with TMS targeted to the anterior vs.

A priori comparisons were computed based on the main models using the R lsmeans package Finally, HR results were analyzed with a linear mixed model with the task, target location, treatment TMS vs. Author I. The authors declare no competing financial interests. The content is solely the responsibility of the authors and does not necessarily represent the official views of the funding agencies. National Center for Biotechnology Information , U. Nat Commun. Author manuscript; available in PMC Feb John W.

Iiro P. Josef P. Author information Copyright and License information Disclaimer. Corresponding Author: Jyrki Ahveninen, Ph. Copyright notice. The publisher's final edited version of this article is available at Nat Commun. See other articles in PMC that cite the published article. Abstract Neurophysiological animal models suggest that anterior auditory cortex AC areas process sound-identity information, whereas posterior ACs specialize in sound location processing.

Open in a separate window. Figure 1. Figure 2. Table 1 Group-average RTs in milliseconds. Figure 3. Table 2 Group-average HR percentages. Verifying TMS targeting by cortical E-field estimates The results of electromagnetic modeling analyses, which were conducted to verify the foci of E-fields induced by TMS in each subject, are shown in Figure 4.

Figure 4. Discussion Here, we studied the effects of transient focal AC deactivations induced by TMS 44 on processing of object-identity vs. TMS target post hoc confirmation Electromagnetic modeling analyses utilizing realistic anatomy were conducted to estimate the actual cortically induced E-field distributions in each subject. Data analysis A full factorial design was utilized to control for potential biases caused by the TMS side effects acoustic clicks, muscle stimulation.


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Footnotes The authors declare no competing financial interests. Author Contributions J. References 1. Rauschecker JP, Tian B. Functional specialization in rhesus monkey auditory cortex. Ahveninen J, et al. Lomber SG, Malhotra S. Nat Neurosci. Romanski LM, et al. Dual streams of auditory afferents target multiple domains in the primate prefrontal cortex. Auditory agnosia and auditory spatial deficits following left hemispheric lesions: evidence for distinct processing pathways. Maeder PP, et al.

Distinct pathways involved in sound recognition and localization: a human fMRI study. Bushara KO, et al. Modality-specific frontal and parietal areas for auditory and visual spatial localization in humans. Perception of sound-source motion by the human brain.


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Brunetti M, et al. Hum Brain Mapp. Cerebral responses to change in spatial location of unattended sounds.

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Kopco N, et al. Neuronal representations of distance in human auditory cortex. The processing of temporal pitch and melody information in auditory cortex. Binder JR, et al. Human temporal lobe activation by speech and nonspeech sounds. Cereb Cortex. Identification of a pathway for intelligible speech in the left temporal lobe. Obleser J, et al.

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