Dear Dr Abbasi,

Thank you for submitting your manuscript entitled "Frequency-specific brain networks in continuous speaking and listening" for consideration as a Research Article by PLOS Biology and using our Portable Peer Review system. I have discussed your manuscript with the in-house editorial team and with an Academic Editor with relevant expertise and am writing to let you know we feel the study has potential for PLOS Biology. Your revision seems quite responsive to the questions and concerns that had been raised in the prior rounds of peer review at Nature Communications. We would therefore like to have the reviewers take a look at this new revision before making a final call. I have reached out to Nature Communications to see if we can obtain these reviewer identities. If we are able to get these, we will go back to the same reviewers. If we are unable to obtain these identities, we will approach a few new reviewers but will ask them to specifically comment on the revision in light of the reviews you've already received.

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Kris

Kris Dickson, Ph.D., (she/her)

Neurosciences Senior Editor/Section Manager

PLOS Biology

kdickson@plos.org

Dear Dr Abbasi,

Thank you for your patience while we considered your revised manuscript "Frequency-specific brain networks in continuous speaking and listening" 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 of the original Nature Communications reviewers. Unfortunately we were not able to obtain the identity of reviewer #1, but the Academic Editor has assessed your responses to that reviewer (as has reviewer #2). Please accept my apologies for the extreme delay incurred earlier in the process.

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

IMPORTANT - Please attend to the following:

a) Please could you make your Title more informative and accessible, preferably including an active verb? Based on your Abstract, I'd suggest something like "Brain networks involved in continuous speaking and listening are partially shared but dissociated in space, time and frequency," but you will likely be able to fashion something more accurate.

b) Please provide a blurb, according to the instructions in the submission form.

c) Please address the remaining concerns from the two reviewers.

c) Please address my Data Policy requests below; specifically, we need you to supply the numerical values underlying Figs 2, 3AB, 4, 5AB, 6ABCD, 7ABCD, 8AB, 9ABCDE, S1, S2AB, S3, S4AB, S5, S6AB, S7, S8AB, S9, either as a supplementary data file or as a permanent DOI’d deposition. We note that you have already deposited MatLab code and a small subset of the data in OSF; I will need to see the full deposition before accepting the paper for publication. I understand that the raw data are protected by privacy laws, and are therefore exempt from PLOS’ data policy.

d) Please cite the location of the data clearly in all relevant main and supplementary Figure legends, e.g. “The data underlying this Figure can be found in S1 Data” or “The data underlying this Figure can be found in https://osf.io/XXXXX"

e) We note that you state that "The study was approved by the local ethics committee" - please could you also include an approval number?

As you address these items, please take this last chance to review your reference list to ensure that it is complete and correct. If you have cited papers that have been retracted, please include the rationale for doing so in the manuscript text, or remove these references and replace them with relevant current references. Any changes to the reference list should be mentioned in the cover letter that accompanies your revised manuscript.

We expect to receive your revised manuscript within two weeks.

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

Sincerely,

Roli Roberts

Roland Roberts, PhD

Senior Editor,

rroberts@plos.org,

PLOS Biology

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DATA POLICY:

You may be aware of the PLOS Data Policy, which requires that all data be made available without restriction: http://journals.plos.org/plosbiology/s/data-availability. For more information, please also see this editorial: http://dx.doi.org/10.1371/journal.pbio.1001797

Note that we do not require all raw data. Rather, we ask that all individual quantitative observations that underlie the data summarized in the figures and results of your paper be made available in one of the following forms:

1) Supplementary files (e.g., excel). Please ensure that all data files are uploaded as 'Supporting Information' and are invariably referred to (in the manuscript, figure legends, and the Description field when uploading your files) using the following format verbatim: S1 Data, S2 Data, etc. Multiple panels of a single or even several figures can be included as multiple sheets in one excel file that is saved using exactly the following convention: S1_Data.xlsx (using an underscore).

2) Deposition in a publicly available repository. Please also provide the accession code or a reviewer link so that we may view your data before publication.

Regardless of the method selected, please ensure that you provide the individual numerical values that underlie the summary data displayed in the following figure panels as they are essential for readers to assess your analysis and to reproduce it: Figs 2, 3AB, 4, 5AB, 6ABCD, 7ABCD, 8AB, 9ABCDE, S1, S2AB, S3, S4AB, S5, S6AB, S7, S8AB, S9. NOTE: the numerical data provided should include all replicates AND the way in which the plotted mean and errors were derived (it should not present only the mean/average values).

IMPORTANT: Please also ensure that figure legends in your manuscript include information on where the underlying data can be found, and ensure your supplemental data file/s has a legend.

Please ensure that your Data Statement in the submission system accurately describes where your data can be found.

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DATA NOT SHOWN?

- Please note that per journal policy, we do not allow the mention of "data not shown", "personal communication", "manuscript in preparation" or other references to data that is not publicly available or contained within this manuscript. Please either remove mention of these data or provide figures presenting the results and the data underlying the figure(s).

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REVIEWERS' COMMENTS:

Reviewer #1:

[identifies himself as Benjamin Morillon]

I reviewed a previous version of the manuscript for another journal. My current review addresses the comments raised by all the previous reviewers. As I previously wrote, this work is innovative, methodologically and conceptually, the methods are sound, and the results are important to further our understanding of the brain as being a dynamical system.

Concerning the previous concerns of Reviewer #1, I tend to agree with their, and appreciate that the authors have deemphasized references to predictive coding from their manuscript.

However, I would suggest removing any mention of predictive coding in the abstract (where it is present in two instances). Indeed, none of the results investigate predictive processing per se, as is for instance done with the estimation of surprisal or entropy variables that are then correlated to brain dynamics and connectivity patterns. Here predictive coding is only one of the possible interpretations of their result (and not the most straightforward).

The authors also added a fifth question in the introduction, but again I would use caution regarding this point (and actually, I suggest to simply remove it), for a specific set of related reasons:

- First, beta (∼20 Hz) activity is the default oscillatory mode of the motor system.

- Second, the framework championed by A. Bastos and colleagues (that the authors rely on here) has recently been heavily put into question, notably by the work of M. Vinck and colleagues (see, e.g. Schneider et al., Neuron 2021, or Vinck Neuron 2023 - but also by the group of C.E. Schroeder or A. Thiele). They basically show that inter-areal directed coherence can be predicted by the dominant power amongst the two explored regions.

- Third, regrouping these two information results in the hypothesis that beta activity will be necessarily be directed from motor to auditory regions. Imho, this reflects the most parsimonious hypothesis, and there is no need to rely on predictive coding assumptions of dedicated frequency channels.

- Finally, as I said before, the authors entirely rely on the proposal by Bastos et al. (which was actually never empirically validated!) but do not show any evidence that some predictions or prediction error signals are transmitted between motor and auditory regions in the beta band. It is simply an interpretation of their results that they can of course mention in the discussion. But I would use caution (and mention alternative interpretations too, such as the one described above, for instance).

Regarding the negative-lag result, I would also first emphasize the most accepted interpretations, notably that the motor cortex is involved in motor preparation, then in the processing of auditory feedback (see, e.g. Ozker…Flinker, PloS Biol 2022 for a relevant example), and then in predictive processing.

I have no further comments and congratulate the authors for their excellent work.

Reviewer #3:

The study's main contribution is a comprehensive description of the brain tracking of the speech envelope during continuous speaking and listening. Moreover, it also describes connectivity patterns that could be interpreted in the light of the predictive coding framework.

Since a substantial part is dedicated to the understanding of the neural substrate of speaking, the inclusion of relevant related bibliography could strengthen the arguments and provide additional context to the findings:

Pérez, A. et al. Timing of brain entrainment to the speech envelope during speaking, listening and self-listening. Cognition (2022).

Ozker M. et al. A cortical network processes auditory error signals during human speech production to maintain fluency. Plos Biology (2022).

Page 5 of the document: the narrative ends abruptly with the disconnected paragraph "Recently, we characterised …".

To improve clarity, the captions of Figures 3 and 4 should include "during speaking".

Dear Dr Abbasi,

Thank you for the submission of your revised Research Article "Spatio-temporal dynamics characterise spectral connectivity profiles of continuous speaking and listening" for publication in PLOS Biology. On behalf of my colleagues and the Academic Editor, David Poeppel, I'm 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.

Please take a minute to log into Editorial Manager at http://www.editorialmanager.com/pbiology/, click the "Update My Information" link at the top of the page, and update your user information to ensure an efficient production process.

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We also ask that you take this opportunity to read our Embargo Policy regarding the discussion, promotion and media coverage of work that is yet to be published by PLOS. As your manuscript is not yet published, it is bound by the conditions of our Embargo Policy. Please be aware that this policy is in place both to ensure that any press coverage of your article is fully substantiated and to provide a direct link between such coverage and the published work. For full details of our Embargo Policy, please visit http://www.plos.org/about/media-inquiries/embargo-policy/.

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

Sincerely, 

Roli Roberts

Roland G Roberts, PhD, PhD

Senior Editor

PLOS Biology

rroberts@plos.org