Dear Dr Varlet,

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Dear Dr Varlet,

Thank you for your patience while your manuscript "Who is who? Behavioural and brain mechanisms underpinning the self-other distinction and action coordination through sound" was peer-reviewed at PLOS Biology. It has now been evaluated by the PLOS Biology editors, an Academic Editor with relevant expertise, and by several independent reviewers.

In light of the reviews, which you will find at the end of this email, we would like to invite you to revise the work to thoroughly address the reviewers' reports.

As you will see below, the reviewers overall think that your study is well executed and provides important insights. However, they also raised a number of concerns about a lack of clarity, the statistical analyses and some potential confounders, including the small number of trials in each condition (Reviewer 2, point 2).

Given the extent of revision needed, we cannot make a decision about publication until we have seen the revised manuscript and your response to the reviewers' comments. Your revised manuscript is likely to be sent for further evaluation by all or a subset of the reviewers.

In addition to these revisions, you will need to complete some formatting changes, which you will receive in a follow up email. A member of our team will be in touch with a set of requests shortly.

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Christian Schnell, PhD,

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------------------------------------

REVIEWS:

Reviewer #1: Thank you for the opportunity to read this very interesting and timely paper. I found it very well written and a valuable addition to research on social neuroscience of coordination. The findings have implications for broader work on communication and validate a novel paradigm that could be adapted to test these more ecologically-valid, complex interaction contexts. I have relatively minor comments below that I hope will add to the clarity of the manuscript and broaden its reach to multi-disciplinary audiences.

Introduction:

- It would be helpful for the reader to be more clearly guided through the Introduction, adding clear link from previous research to the focus of the current study. Can the research question be brought in earlier to help frame the background? The main study hypothesis is also presented in a long sentence of dense language- this would benefit from being split into two sentences with more concrete language: "We hypothesised that these mechanisms, together with those based on lower-level acoustic differences, constitute a powerful and flexible processing gradient enabling stream segregation at peripheral and central levels along the auditory pathway, key to maintaining the self-other distinction in everyday dynamic interactions characterised by temporally overlapping acoustic signals and uncertainty."

- The definition for 'brightness'- the key manipulation for participants in this study is buried in the Figure 1 legend - please define this key term in the main text.

Results:

- The pattern is results is described before (and independent of) stats, which makes hard to match up.

- Some effects are not mentioned at all (eg. leadership x frequency on interpersonal synchronisation, or main effect and interaction on sound brightness)- I assume these were all non-significant, but at the very least they should be acknowledged as such in the text with maximal p-value.

- Plots only show significant effects, rather than means for the fully crossed designs- non-significant factors are excluded. Please show complete plots of fully crossed data for transparency.

- Analysis of neural tracking- it's not clear why regressions are needed, they are presented as simply 'confirming' the ANOVA results?

- Currently behavioural and neural data are considered/analysed completed separately. Can additional analyses be run to test for links between these datasets, to support interpretation of the oscillations data?

Method:

- Please justify your sample size and comment on power. Did any of the dyads know each other before the experiment?

Reviewer #2: This innovative study combined dual-EEG and frequency-tagging techniques in a more ecological dyadic improvisation synchronization task to investigate a relevant and relatively understudied question in auditory perception: how individuals discriminate between self- and other-generated sounds in joint auditory-motor synchronization. By integrating behavioral and neural measures, the study shows that auditory self-other distinction is modulated by both stimulus characteristics (spectral information) and participant's control over one's own actions (leader-follower roles). These findings have important implications for our understanding of the mechanisms underlying social coordination in auditory interactions. There are several laudatory aspects to highlight, including the innovative integration of dual-EEG and frequency tagging techniques, as well as the relatively large sample size for hyperscanning research (25 dyads). However, some methodological and conceptual issues temper confidence in some of the authors' claims. Addressing the points outlined below would substantially strengthen the manuscript.

A. Method:

1.Participants: The manuscript does not report information on participants' musical training or rhythmic ability, which may represent a major confound for synchronization accuracy and sensitivity to changes in sound frequency (e.g., Scheurich et al., 2020). If these data were not collected, the authors should at least acknowledge this as a limitation and discuss the likely direction of its impact. Sample size justification should also be added to the manuscript.

2.Trial number: A critical limitation concerns the small number of trials in each condition. With only two trials per condition, the robustness of within-condition EEG estimates is likely reduced, despite the long trial duration (180 s each). This experimental choice requires justification. In addition, the authors should consider providing supplementary reliability analyses to increase confidence in the robustness of the data interpretation.

3.Another issue refers to the force-brightness mapping. The perceptual validity of the mapping between linear force and sound brightness is not demonstrated, leaving unclear whether "brightness" manipulations were perceived consistently across participants. More information is needed. For example, a pilot perceptual validation could demonstrate that participants reliably perceive increasing force as increased brightness.

4.I am also concerned with potential motor contamination of EEG signals. The finger force manipulation used to control sound brightness may introduce motor-related artifacts that could influence the EEG measures (see, for example, Timm et al., 2014). The authors should justify how motor contamination was addressed or mitigated.

5.The manuscript also lacks a description of hardware/software timing accuracy, timing alignment procedures, and drift correction - all essential components of hyperscanning methodology (e.g., see Zamm et al., 2024). Additional information on dual-EEG synchronization procedures is necessary.

B. Discussion:

6.In the discussion, certain claims extend beyond what the data directly support. For example, EEG-based source localization, particularly with a template head model, has limited anatomical precision, especially for hemispheric lateralization. The authors should temper these anatomical claims and discuss the limitations of template-based source localization more explicitly.

7.Interpretations involving participants' attentional expectations, "agency confusion," and hierarchical self-other processing along the auditory pathway should be presented more cautiously, as these mechanisms were not directly probed. Clearly distinguishing between empirically supported conclusions and more speculative theoretical interpretations would strengthen the discussion.

Cited references:

*Scheurich, R., Pfordresher, P. Q., & Palmer, C. Musical training enhances temporal adaptation of auditory-motor synchronization. Exp Brain Res 238, 81-92 (2020). https://doi.org/10.1007/s00221-019-05692-y

*Timm, J., SanMiguel, I., Keil, J., Schröger, E., & Schönwiesner, M. Motor intention determines sensory attenuation of brain responses to self-initiated sounds. J Cogn Neurosci 26(7), 1481-1489 (2014). https://doi.org/10.1162/jocn_a_00552

*Zamm, A., Loehr, J. D., Vesper, C., Konvalinka, I., Kappel, S. L., Heggli, O. A., Vuust, P., & Keller, P. E. A practical guide to EEG hyperscanning in joint action research: from motivation to implementation. Soc Cogn Affect Neurosci 19(1), nsae026 (2024). https://doi.org/10.1093/scan/nsae026

Reviewer #3: The study by Varlert et al. uses an auditory-motor synchronization paradigm to investigate how humans maintain a distinction between self- and other-generated sounds during joint action. The authors show that participants synchronize more effectively when their sounds are acoustically distinct and when leadership roles are assigned. I found the frequency-tagging approach to isolate neural responses to self- versus other-related stimuli particularly elegant, and the analyses and results are generally convincing. I have 3 major comments, as well as several smaller points, which I believe would strengthen the manuscript.

Major comments

Clarity of expression. The manuscript is very difficult to follow due to extremely long sentences and the use of technical terms before they are introduced or intuitively motivated. Early on, it is unclear what "inferring/coordinating from sound" refers to. The task description is especially hard to parse: even after repeated readings of the Fig. 1 caption, I struggled to understand (1) what the participant instructions actually were and (2) how exactly were the auditory signals experimentally manipulated. Several concepts appear abruptly (e.g., "brightness of sound", "frequency tagging"), though they are not widely familiar and require explanation for a general readership. In addition, some results are presented without even brief methodological context. Below, I point to a few particularly challenging passages; however, I would recommend that the manuscript be more broadly rewritten for clarity.

Contextualization and theoretical framing. In my opinion, the title (phrases like "self-other distinction from sound") strongly suggests a link to voice or speech processing, yet this literature is not addressed at all. Once it becomes clear that the study is not about voice or speech, the reader might then expect connections to other more typical auditory-motor coordination paradigms - such as tapping, joint rhythmic entrainment, or musical synchronization - but these are also largely neglected. Instead, the introduction moves rapidly to a very specific and technically complex paradigm without providing intuitive conceptual grounding. I recommend introducing the motivation more directly, clarifying what kind of "coordination from sound" the study targets (and why), and situating the work more clearly within both the vocal-agency and sensorimotor-synchronization literatures so that readers can understand how this paradigm extends or complements existing (more classical) approaches.

Results interpretation: top-down vs. bottom-up mechanisms. Several paragraphs in the introduction talk about hierarchical processes that might contribute to self-other distinction, but the argument remains difficult to follow, and the connection between these proposed top-down mechanisms and the present paradigm is not clearly articulated. For example, after the brief mention of "anticipation, adaptation, and ongoing planning" I was expecting that such processes would be manipulated or discussed, yet the manuscript never returns to them. In contrast, the broader auditory self-other literature often targets specific processes possibly guiding the hierarchical predictions: efference-copy mechanisms in self-voice monitoring (e.g., Whitford et al., 2019, Biological Psychiatry: CNN), memory-based self-voice representations (e.g., Iannotti et al., 2021, Cerebral Cortex), attentional and emotional biases (e.g., Pinheiro et al., 2022, Cortex), and even biases from concurrent sensory stimuli such as bone conduction (e.g., Orepic et al., 2023, RSOS). These lines of research provide a clearer theoretical grounding for understanding how self-related auditory information is processed. Moreover, many of self-other studies also converge on right-hemisphere specialization for self-related processing - both the sense of self more broadly (Northoff et al., 2006, NeuroImage) but also within auditory self-representation (e.g., Candini et al., 2018, NeuroImage: Clinical; Iannotti et al., 2021, Cerebral Cortex; Kaplan et al., 2008, SCAN). I therefore recommend that the authors relate their own right-hemisphere findings to this wider self-other literature, both in the Introduction (for context) and more importantly in the Discussion.

Minor

Line numbers would be helpful for the review

Title & abstract

I find "action coordination through sound" quite vague

The first part of abstract is also rather unclear. It is hard to understand the scope of the study: are people speaking in synchrony or tapping in rhythm, or something else? I am also not convinced by the statement "such interaction is challenging in real-life". I'd argue that the brain does do this quite well in real life.

Other vague expressions: "leader-follower roles", "stronger facilitation", "modality-specific" (which modalities other than auditory are manipulated?)

Introduction

Page 4:

"impossible in principle" - exclusive and too strong, not tested explicitly

"growing evidence point to a complementary and necessary involvement of selective attentional and predictive mechanisms (…)" - such as? References?

"Such top-down brain mechanisms (…)" - my major comments

Page 5:

"Here we combined dual-EEG and frequency-tagging techniques in a dyadic improvisation synchronization" - terms not clear for general readership

"an auditory-motor analogue of the mirror game paradigm" - please briefly mention what is the paradigm. The reader does not want to open another paper to properly understand this one

"Illustration of the improvisation synchronisation task in which (…)" and "This frequency tagging of the sound envelope enabled neural processing (…)" - long sentences very hard to follow

Fig 1: What instructions were given to participants? Upper panel - I find that the 2 rows are rather confusing than helping the interpretation - the only difference is the color, but then the color is introduced only below. The brightness aspect is not clear and not obvious if it is (and even should be) the same in both rows. Is this auditory manipulation evolving in time (the arrow)? Is this happening at each trial? Very hard to understand the figure without reading the methods.

"improvisation" suggests freedom, but "synchronisation" implies alignment. Not obvious how they are merged together in the paradigm.

The manuscript states that participants produced "original and synchronized fluctuations in brightness", but does not explain what these fluctuations should look like, whether they followed an external cue, or how participants knew they were on time with one another.

The manipulation of brightness through finger pressure is highly unconventional and is not sufficiently explained in intuitive terms.

Why 33 and 47 Hz exactly?

Isn't it critical to mention that these amplitude fluctuations cannot be consciously perceived?

Intermodulation frequencies - not obvious why they would indicate integration. Does their processing necessarily reflects integration? Could low-level auditory processing be really decoupled from higher-level mechanisms (e.g. paying attention to self/other identity)?

Results

Coherence - briefly mention how it is computed

ANOVAs - was there an interaction term and why yes or why not (in all analyses)?

Fig 2 - "1 × 95% CI of the mean computed for within-subject designs" - not clear

Why up to 5 Hz?

Page 9: Why is Leadership variable now modeled differently (3 levels instead of 2)? Why is this not consistent across analyses (or both reported for all)?

Fig 3 - it would be helpful to indicate what is significant. The same for some other plots.

Why averaging across all EEG channels?

Fig 4 - What is the take-home message of this plot? Left: There are a few other prominent peaks than the 4 indicated ones. What could they represent? Is it then justified to only use 14 and 80 for further analyses? Can there be other markers of integration (e.g. some other nonlinear combination of 33 and 47)? Right: The 80 peak does not seem to stand out from many others. Can it therefore be used reliably in further analyses?

"suggesting an increase in the neural tracking (…) not only originating from peripheral changes in sound brightness but also centrally from selective enhancement at the brain level" Do we even know that there are any peripheral changes (I assume here you mean the cochlea or auditory nerve?) Not sure this statement is justified if this was not manipulated.

Fig 5 - what are the units on the x and y axes?

"These analyses also show that it is the right hemisphere that benefits the most from designated leadership roles, which could reflect the spectral superiority of this hemisphere" What is "spectral superiority of a brain hemisphere"?

Fig 6 - Could the response be modulated by the absolute value of the 2 chosen frequencies? Why the choice of 200 and 500?

What about the effect of Leadership in the "Self-Other neural integration/segregation" part? Do you observe similar effects for other prominent peaks from Fig 4?

The dual-EEG setup is interesting, but only within-participant analyses are reported. Have the authors explored inter-brain synchrony or leader-follower dynamics? Measures such as cross-brain coherence or correlation of frequency-tagged responses could reveal how neural coordination supports behavioral synchrony. Such findings would largely strengthen the manuscript.

Discussion

"Our work (…) demonstrates that the processes underlying interpersonal coordination are modality-specific." - How does this study show that? What modality other than auditory is tested?

Broader scope - why this work is important - could be strengthened. Specifically, there is a large body of literature demonstrating that auditory self-other confusion is related to auditory-verbal hallucinations, both in schizophrenia and in healthy individuals (e.g. Duggirala et al., 2024, Schizophrenia Bulletin; Waters et al., 2012, Schizophrenia Bulletin) and in other clinical changes in the sense of self (e.g. Voruz et al., 2024, Heliyon; Candini et al., 2018, NeuroImage: Clinical)

Methods

Sill not clear to me what exactly the instruction for the participants were. How was brightness of sound explained? How exactly is one told to lead?

Dear Dr Varlet,

Thank you for your patience while we considered your revised manuscript "Mechanisms underpinning the self-other distinction during interpersonal auditory-motor coordination" for publication as a Research Article at PLOS Biology. This revised version of your manuscript has been evaluated by the PLOS Biology editors, the Academic Editor and two the original reviewers.

Based on the reviews and on our Academic Editor's assessment of your revision, we are likely to accept this manuscript for publication, provided you satisfactorily address the following data and other policy-related requests:

* We would like to suggest a different title to improve its accessibility for our broad audience:

Right-hemisphere-dominant neural mechanisms facilitate auditory self-other distinction and coordination in human dyads

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Christian Schnell, PhD

Senior Editor

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------------------------------------------------------------------------

Reviewer remarks:

Reviewer #2: The authors have responded carefully and comprehensively to the issues raised in a previous version of the manuscript. The manuscript has been substantially improved in terms of clarity, methodological transparency, theoretical framing, and interpretation of the findings. While some issues (e.g., formal validation of the brightness manipulation and empirical assessment of reliability) remain only partially resolved, these limitations are now adequately acknowledged and do not undermine the main conclusions of the study. I therefore support publication of the manuscript in its revised form.

Reviewer #3: I thank the authors for addressing my comments and applaud them for such a thorough revision.

Dear Dr Varlet,

Thank you for your patience while we considered your revised manuscript "Lateralised neural mechanisms facilitate auditory self-other distinction and coordination in human dyads " for publication as a Research Article at PLOS Biology.

Thank you also for largely addressing the editorial requests. There is one point, however, which needs further changes.

The references in the supplementary information file need to be moved to the main manuscript files. The reason for this is that the literature databases do not process references from supplementary information files, so the authors of those references would not receive any credit for these citations. You can then use the reference numbers from the main manuscript file to cite the papers in the supplementary information file.

Everything else looked and we'll be able to formally accept the your paper for publication once this is sorted.

In addition to these revisions, you may need to complete some formatting changes, which you will receive in a follow up email. A member of our team will be in touch with a set of requests shortly. If you do not receive a separate email within a few days, please assume that checks have been completed, and no additional changes are required.

We expect to receive your revised manuscript within one week.

To submit your revision, please go to https://www.editorialmanager.com/pbiology/ and log in as an Author. Click the link labelled 'Submissions Needing Revision' to find your submission record. Your revised submission must include the following:

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Please do not hesitate to contact me should you have any questions.

Sincerely,

Christian

Christian Schnell, PhD

Senior Editor

cschnell@plos.org

PLOS Biology

Dear Dr Varlet,

Thank you for the submission of your revised Research Article "Lateralised neural mechanisms facilitate auditory self-other distinction and coordination in human dyads " for publication in PLOS Biology. On behalf of my colleagues and the Academic Editor, Jennifer Bizley, I am pleased to say that we can in principle accept your manuscript for publication, provided you address any remaining formatting and reporting issues. These will be detailed in an email you should receive within 2-3 business days from our colleagues in the journal operations team; no action is required from you until then. Please note that we will not be able to formally accept your manuscript and schedule it for publication until you have completed any requested changes.

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Thank you again for choosing PLOS Biology for publication and supporting Open Access publishing. We look forward to publishing your study.

Sincerely,

Christian

Christian Schnell, PhD

Senior Editor

PLOS Biology

cschnell@plos.org