Dear Dr Turner,

Happy New Year!

Thank you for submitting your manuscript entitled "Progressive multi-stage extrapolation of predictable motion in human visual cortex" for consideration as a Research Article by PLOS Biology.

Your manuscript has now been evaluated by the PLOS Biology editorial staff as well as by an academic editor with relevant expertise and I am writing to let you know that we would like to send your submission back to external peer review, despite some reservations whether the reviewers' concerns have been fully addressed.

However, before we can send your manuscript to reviewers, we need you to complete your submission by providing the metadata that is required for full assessment. To this end, please login to Editorial Manager where you will find the paper in the 'Submissions Needing Revisions' folder on your homepage. Please click 'Revise Submission' from the Action Links and complete all additional questions in the submission questionnaire.

Once your full submission is complete, your paper will undergo a series of checks in preparation for peer review. After your manuscript has passed the checks it will be sent out for review. To provide the metadata for your submission, please Login to Editorial Manager (https://www.editorialmanager.com/pbiology) within two working days, i.e. by Jan 11 2025 11:59PM.

If your manuscript has been previously peer-reviewed at another journal, PLOS Biology is willing to work with those reviews in order to avoid re-starting the process. Submission of the previous reviews is entirely optional and our ability to use them effectively will depend on the willingness of the previous journal to confirm the content of the reports and share the reviewer identities. Please note that we reserve the right to invite additional reviewers if we consider that additional/independent reviewers are needed, although we aim to avoid this as far as possible. In our experience, working with previous reviews does save time.

If you would like us to consider previous reviewer reports, please edit your cover letter to let us know and include the name of the journal where the work was previously considered and the manuscript ID it was given. In addition, please upload a response to the reviews as a 'Prior Peer Review' file type, which should include the reports in full and a point-by-point reply detailing how you have or plan to address the reviewers' concerns.

During the process of completing your manuscript submission, you will be invited to opt-in to posting your pre-review manuscript as a bioRxiv preprint. Visit http://journals.plos.org/plosbiology/s/preprints for full details. If you consent to posting your current manuscript as a preprint, please upload a single Preprint PDF.

Feel free to email us at plosbiology@plos.org if you have any queries relating to your submission.

Kind regards,

Christian

Christian Schnell, PhD

Senior Editor

PLOS Biology

cschnell@plos.org

Dear Dr Turner,

Thank you for your patience while we considered your revised manuscript "Progressive multi-stage extrapolation of predictable motion in human visual cortex" for consideration as a Research Article at PLOS Biology. Please allow me first to apologize for the long delay in sending our decision. Unfortunately, one of the original reviewers was not able to submit their report after agreeing to re-review your manuscript, so we had to find a replacement reviewer. Your revised study has now been evaluated by the PLOS Biology editors, the Academic Editor, one the original reviewers and one new reviewer. Please note that Laurent was Reviewer 2 of the original submission, so Reviewer 2 of this submission is the replacement reviewer for Reviewer 1 of the original submission.

In light of the reviews, which you will find at the end of this email, we are pleased to offer you the opportunity to address the remaining points and comments from the reviewers in a revision that we anticipate should not take you very long. We will then assess your revised manuscript and your response to the reviewers' comments with our Academic Editor aiming to avoid further rounds of peer-review, although we might need to consult with the reviewers, depending on the nature of the revisions.

You will see that Reviewer 1 only has a few minor concerns and that Reviewer 2 is mostly satisfied with your response to Reviewer 1's original concerns, but there are a few items that need further addressing. We also encourage you to address the additional points that are raised by Reviewer 2. Please do not hesitate to get in touch if you have any questions or need a bit more time.

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.

We expect to receive your revised manuscript within 1 month. Please email us (plosbiology@plos.org) if you have any questions or concerns, or would like to request an extension.

At this stage, your manuscript remains formally under active consideration at our journal; please notify us by email if you do not intend to submit a revision so that we withdraw the manuscript.

**IMPORTANT - SUBMITTING YOUR REVISION**

Your revisions should address the specific points made by each reviewer. Please submit the following files along with your revised manuscript:

1. A 'Response to Reviewers' file - this should detail your responses to the editorial requests, present a point-by-point response to all of the reviewers' comments, and indicate the changes made to the manuscript.

*NOTE: In your point-by-point response to the reviewers, please provide the full context of each review. Do not selectively quote paragraphs or sentences to reply to. The entire set of reviewer comments should be present in full and each specific point should be responded to individually.

You should also cite any additional relevant literature that has been published since the original submission and mention any additional citations in your response.

2. In addition to a clean copy of the manuscript, please also upload a 'track-changes' version of your manuscript that specifies the edits made. This should be uploaded as a "Revised Article with Changes Highlighted " file type.

*Resubmission Checklist*

When you are ready to resubmit your revised manuscript, please refer to this resubmission checklist: https://plos.io/Biology_Checklist

To submit a revised version of your manuscript, please go to https://www.editorialmanager.com/pbiology/ and log in as an Author. Click the link labelled 'Submissions Needing Revision' where you will find your submission record.

Please make sure to read the following important policies and guidelines while preparing your revision:

*Published Peer Review*

Please note while forming your response, if your article is accepted, you may have the opportunity to make the peer review history publicly available. The record will include editor decision letters (with reviews) and your responses to reviewer comments. If eligible, we will contact you to opt in or out. Please see here for more details:

https://blogs.plos.org/plos/2019/05/plos-journals-now-open-for-published-peer-review/

*PLOS Data Policy*

Please note that as a condition of publication PLOS' data policy (http://journals.plos.org/plosbiology/s/data-availability) requires that you make available all data used to draw the conclusions arrived at in your manuscript. If you have not already done so, you must include any data used in your manuscript either in appropriate repositories, within the body of the manuscript, or as supporting information (N.B. this includes any numerical values that were used to generate graphs, histograms etc.). For an example see here: http://www.plosbiology.org/article/info%3Adoi%2F10.1371%2Fjournal.pbio.1001908#s5

*Blot and Gel Data Policy*

We require the original, uncropped and minimally adjusted images supporting all blot and gel results reported in an article's figures or Supporting Information files. We will require these files before a manuscript can be accepted so please prepare them now, if you have not already uploaded them. Please carefully read our guidelines for how to prepare and upload this data: https://journals.plos.org/plosbiology/s/figures#loc-blot-and-gel-reporting-requirements

*Protocols deposition*

To enhance the reproducibility of your results, we recommend that if applicable you deposit your laboratory protocols in protocols.io, where a protocol can be assigned its own identifier (DOI) such that it can be cited independently in the future. Additionally, PLOS ONE offers an option for publishing peer-reviewed Lab Protocol articles, which describe protocols hosted on protocols.io. Read more information on sharing protocols at https://plos.org/protocols?utm_medium=editorial-email&utm_source=authorletters&utm_campaign=protocols

Thank you again for your submission to our journal. We hope that our editorial process has been constructive thus far, and we welcome your feedback at any time. Please don't hesitate to contact us if you have any questions or comments.

Sincerely,

Christian

Christian Schnell, PhD

Senior Editor

PLOS Biology

cschnell@plos.org

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

REVIEWS:

Reviewer #1 (Laurent U Perrinet): I would like to thank the authors for the new version of their manuscript, as well as for the detailed response to our comments. These two elements allow me to judge that this article is now acceptable for publication in this journal.

I have just a few minor points that could be included in the final version.

- The choice of decoding score (point R2C12) can be simplified by giving it a physical meaning. By using a value between -1 and 1, we are in fact calculating the cosine of the difference between the predicted angle and the actual angle. It may be simpler simply to give this angle difference.

- In Fig 2A, you could mention that the "hockey club curve" is similar to that observed in the Frohlich effect (see for instance work from Jancke) - this would also allow you to justify the fact that the temporal resolution here is not enough to conclude for a "jump" at reversal in panel C

- in point R2C7 evidence is defined as the posteriori probability however position eviidence (eg in Fig2 or 3, 4) can be negative. please provide a clear definition in the methods

Reviewer #2: I stepped in for R1 (as a new reviewer) after a first round of revision.

In my reading, the main concern of R1 is that reported effects may have oculomotor origins rather than visual-processing properties inherent in the visual system. I do agree with the original R1 that it would have been more convincing if the authors had included careful eye-tracking when collecting their data (simply assuming participants follow instructions to fixate may be too optimistic).

In response to this main first concern of R1, the authors now make the case that their findings are not driven by muscle artifacts by the eyes because their decoding localizes to posterior electrodes. While I agree this rules out the contribution of muscle artefacts associated with eye movements, it does not necessarily rule out potential secondary consequences of eye movements (moving the visual stimuli over the retina) that may also contribute to the decoding from visual areas. Thus, just because the decoding is posterior, it may still reflect a secondary consequence of eye movements (or micro eye movements), rather than reflecting a computation inherent to the visual system. This could possibly be acknowledged (or provided further evidence against). That said, I find the additional arguments convincing, such as how their decoder was trained on a case with static stimuli, and tested on a case with predictably moving stimuli.

I also agree with R1 that the data presentation could benefit from further statistical substantiation. For example, while figure 3 is extensively described on pages 7-8, there appears no statistical quantification to back things up for this point (and the same appears to apply for figure 2). This remains a bit odd, also to me.

In addition to the above points, R1 made several other good (and critical) comments, but I do not see these as major breaking points, and the authors' responses appear reasonable to me.

As a new reviewer brought in at this stage, reading the work did inevitably prompt some new points that I would like to share. However, I wish to refrain from listing these as major concerns, also because I see as my primarily role to evaluate the revisions done in response to original R1, as I reflected on above. I nonetheless hope my additional comments may be perceived as useful.

I was wondering the following about the data in figure 3. Given models were trained in response to localizer stimuli with sudden onsets, is it not trivial that the decoded data differ when applying the model to the onset of the motion (similar to the onset of the localizer) versus mid-motion (that is more distinct from the localizer, given single localizer stimuli were never presented for this long)? This difference (in figure 3) could thus either reflect a change in how predictable motion is represented (as currently interpreted) but may also be driven by an increasing mismatch between localizer and test data (when comparing onset vs. mid-motion) in terms of time from stimulation onset? (It appears to me that to rule this out, the authors would have had to also include long-duration localizer stimuli, and train the decoding model to both early-onset and mid-way points in the localizer data). I also note how the reported difference appears to be largely driven by a change in magnitude (more so than in reconstructed position; given the peaks appears to remain largely unaffected).

Also, if the primary findings reflect those in figure 4, why do we only see the data for mid-motion in this figure? Would it not be of central interest to also show a temporal profile starting from motion onset to see the development not only with respect to the time after localizer onset (in training data), but also the time after motion onset (in testing data)?

Finally, at various key instances, such as in the title and the abstract, the authors speak of "processing stages". At these key moments, it is not always sufficiently clear whether the authors mean 'brain area in the visual hierarchy' (V1 vs. MT), 'time from motion onset' (onset vs. mid-way), or something else. Upon further reading, it appears these terms actually reflect time after localizer onset and how this time varying localiser location signal predicts the location of a moving stimulus. Translating this operationalization to processing stages and hierarchy appears less trivial than the initial intuition I had when these terms were introduced. I wonder whether this could/should be better positioned up front, to avoid readers potentially feeling misled by the promise set up at the outset of this paper.

Dear Dr Turner,

Thank you for your patience while we considered your revised manuscript "Progressive multi-stage extrapolation of predictable motion in human visual cortex" 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 one of 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:

Predictable motion is progressively extrapolated across temporally distinct processing stages in the human visual cortex

* Please include information in the Methods section whether the study has been conducted according to the principles expressed in the Declaration of Helsinki.

* 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

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

* CODE POLICY

Per journal policy, if you have generated any custom code during the course of this investigation, please make it available without restrictions. Please ensure that the code is sufficiently well documented and reusable, and that your Data Statement in the Editorial Manager submission system accurately describes where your code can be found.

Please note that we cannot accept sole deposition of code in GitHub, as this could be changed after publication. However, you can archive this version of your publicly available GitHub code to Zenodo. Once you do this, it will generate a DOI number, which you will need to provide in the Data Accessibility Statement (you are welcome to also provide the GitHub access information). See the process for doing this here: https://docs.github.com/en/repositories/archiving-a-github-repository/referencing-and-citing-content

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.

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.

We expect to receive your revised manuscript within two weeks.

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:

- a cover letter that should detail your responses to any editorial requests, if applicable, and whether changes have been made to the reference list

- a Response to Reviewers file that provides a detailed response to the reviewers' comments (if applicable, if not applicable please do not delete your existing 'Response to Reviewers' file.)

- a track-changes file indicating any changes that you have made to the manuscript.

NOTE: If Supporting Information files are included with your article, note that these are not copyedited and will be published as they are submitted. Please ensure that these files are legible and of high quality (at least 300 dpi) in an easily accessible file format. For this reason, please be aware that any references listed in an SI file will not be indexed. For more information, see our Supporting Information guidelines:

https://journals.plos.org/plosbiology/s/supporting-information

*Published Peer Review History*

Please note that you may have the opportunity to make the peer review history publicly available. The record will include editor decision letters (with reviews) and your responses to reviewer comments. If eligible, we will contact you to opt in or out. Please see here for more details:

https://plos.org/published-peer-review-history/

*Press*

Should you, your institution's press office or the journal office choose to press release your paper, please ensure you have opted out of Early Article Posting on the submission form. We ask that you notify us as soon as possible if you or your institution is planning to press release the article.

*Protocols deposition*

To enhance the reproducibility of your results, we recommend that if applicable you deposit your laboratory protocols in protocols.io, where a protocol can be assigned its own identifier (DOI) such that it can be cited independently in the future. Additionally, PLOS ONE offers an option for publishing peer-reviewed Lab Protocol articles, which describe protocols hosted on protocols.io. Read more information on sharing protocols at https://plos.org/protocols?utm_medium=editorial-email&utm_source=authorletters&utm_campaign=protocols

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

Sincerely,

Christian

Christian Schnell, PhD

Senior Editor

cschnell@plos.org

PLOS Biology

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

Reviewer remarks:

Reviewer #2: The authors have done a great job in responding to my comments. I have no further comments.

Dear Dr Turner,

Thank you for the submission of your revised Research Article "Predictable motion is progressively extrapolated across temporally distinct processing stages in the human visual cortex" for publication in PLOS Biology. On behalf of my colleagues and the Academic Editor, Christopher Pack, 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.

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.

PRESS

We frequently collaborate with press offices. If your institution or institutions have a press office, please notify them about your upcoming paper at this point, to enable them to help maximise its impact. If the press office is planning to promote your findings, we would be grateful if they could coordinate with biologypress@plos.org. If you have previously opted in to the early version process, we ask that you notify us immediately of any press plans so that we may opt out on your behalf.

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, 

Christian

Christian Schnell, PhD

Senior Editor

PLOS Biology

cschnell@plos.org