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PONE-D-22-34285Emergence and retention of a collective memory in cockroachesPLOS ONE

Dear Dr. Calvo Martín,

Thank you for submitting your manuscript to PLOS ONE. After careful consideration, we feel that it has merit but does not fully meet PLOS ONE’s publication criteria as it currently stands. Therefore, we invite you to submit a revised version of the manuscript that addresses the points raised during the review process.

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The experiments presented in this research have been conducted rigorously, with appropriate controls and replication. The sample sizes are large enough to produce robust results and methods and reagents are described in sufficient detail for another researcher to reproduce the experiments described. The data presented in the results section of the manuscript supports the conclusions drawn and the research meets all applicable standards for the ethics of experimentation and research integrity.

After a lengthy process of recruiting reviewers, we have obtained a single review, hence I highly recommend following the suggestions given by the reviewer and resubmitting your paper after major revision.

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Please submit your revised manuscript by Mar 19 2023 11:59PM. If you will need more time than this to complete your revisions, please reply to this message or contact the journal office at plosone@plos.org. When you're ready to submit your revision, log on to https://www.editorialmanager.com/pone/ and select the 'Submissions Needing Revision' folder to locate your manuscript file.

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We look forward to receiving your revised manuscript.

Kind regards,

Lemana Spahić

Academic Editor

PLOS ONE

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Reviewers' comments:

Reviewer's Responses to Questions

Comments to the Author

1. Is the manuscript technically sound, and do the data support the conclusions?

The manuscript must describe a technically sound piece of scientific research with data that supports the conclusions. Experiments must have been conducted rigorously, with appropriate controls, replication, and sample sizes. The conclusions must be drawn appropriately based on the data presented.

Reviewer #1: Yes

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2. Has the statistical analysis been performed appropriately and rigorously?

Reviewer #1: Yes

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3. Have the authors made all data underlying the findings in their manuscript fully available?

The PLOS Data policy requires authors to make all data underlying the findings described in their manuscript fully available without restriction, with rare exception (please refer to the Data Availability Statement in the manuscript PDF file). The data should be provided as part of the manuscript or its supporting information, or deposited to a public repository. For example, in addition to summary statistics, the data points behind means, medians and variance measures should be available. If there are restrictions on publicly sharing data—e.g. participant privacy or use of data from a third party—those must be specified.

Reviewer #1: No

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4. Is the manuscript presented in an intelligible fashion and written in standard English?

PLOS ONE does not copyedit accepted manuscripts, so the language in submitted articles must be clear, correct, and unambiguous. Any typographical or grammatical errors should be corrected at revision, so please note any specific errors here.

Reviewer #1: No

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5. Review Comments to the Author

Please use the space provided to explain your answers to the questions above. You may also include additional comments for the author, including concerns about dual publication, research ethics, or publication ethics. (Please upload your review as an attachment if it exceeds 20,000 characters)

Reviewer #1: In this study, the authors perform experimentation with cockroaches to discern their decision making processes when faced with two choices of light conditions. One of the choices (red light) is apparently favoured by cockroaches over the green light which is also what the authors find. However, when applied with an inversion of the light conditions, the findings differ between individual and group behaviours. The authors find that while the (isolated) individuals can transition from the less preferred zone to the preferred zone, the groups show inability to do that. Hence, existence of collective memory is suggested whereby group choice or ‘inertia’ prevents individuals to pursue their choices. The results are explained by a mathematical model (with two versions: deterministic and stochastic). The deterministic model is a differential equation model with the rate of change of number of individuals in a particular zone being modeled by the number of free or available individuals that can move to that zone, individual joining, and leaving rates. The leaving rate, in the model, is multiplied by an exponentially decreasing function of the number of individuals in that zone. This, supposedly, captures the trend observed in the experiments; for small group size the model shows flexibility in decision making, i.e. the proportion of individuals in a given zone decreases continuously if the individuals do not prefer that zone. On the other hand, for large group sizes, the proportion decreases, however, in a discontinuous manner or shows hysteresis. While I like the overall idea of both the experiments and the model, I have some criticisms and I state them as major comments.

Major comments.

• The choice of the exponentially decaying function with number of individuals in a zone that modulates the leaving rate is not justified enough. For a reader to understand the origin of this choice is crucial not only to appreciate the modelling part but to link it with the actual behaviours. Hence, I expect more justification by adding a paragraph on it. It took me some time and also I had to cross check the reference cited (Calvo

Mart´ın et al. 2019) to understand the motivation behind the choice. While, I mechanistically understand, how this choice can lead to the observed experimental results, it is yet unclear to me why this choice was made. For instance, a modulation of the joining rate with a similar function that increases exponentially with the number of individuals in that zone may also address the observation. (The authors claim that they find no evidence for attraction in their experiments, but it was unclear to me what is the basis of this claim.) Or, for that matter, both the modulation functions can depend on the number of individuals in a zone, however their relative strengths can be different. Hence, I think that this point could be addressed by making the model more general to modulate both the joining as well as leaving rate. More broadly, alternative models (at least one) and why they may as well or may not work should be checked and included in the text if possible. If the modelling is not possible, a small survey of literature of similar models should be incorporated.

• The model does show hysteresis as a function of the individual choice for large group size. However, since this particular result hasn’t been shown in the experiments, the claim particularly about hysteresis should be toned down. A possible experimentation to show this could be inverting the light conditions in a more gradual or continuous manner. Two initial conditions for different group sizes might be used. One: keeping all individuals in the preferred zone at the start, and two: keeping all of them in the non preferred zone. A

plot of the proportion of individuals in either of the zone as a function of changing lighting conditions might actually be able to show hysteresis properly in experiments. However, this, I understand, is definitely out of the current scope of the work. Therefore, I suggest to avoid using hysteresis. The authors have right now shown the existence of collective memory or inertia but just for an abrupt switch of the conditions which I think is insufficient to prove hysteresis.

• The use of the stochastic version of the model was not clear to me. Basically, if the deterministic version is capable of capturing the observed trends, then the reasons to make it stochastic are not clear.

Minor comments.

I found several grammatical mistakes and also typos (especially in equations) that must be fixed. Some of the following points are also not really minor as they deal with the readability of the paper and are, therefore, very important for building an understanding while reading the text; While I think the experimentation and the ideas are not that difficult to grasp, overall, I found it very hard to follow them through the text. Therefore, the writing needs considerable work. I only state few of the points hoping that everything gets rectified through a thorough revision by the authors.

• Line 162: Definition of sheltered proportion missing.

• Line 168: Interpretation missing – What is meant by weak effect of time? Can it be written more simply? No figure is referred to highlight this effect.

• Line 191: Definition and unit of cumulative occupation?

• Line 192: Claim of difference between group choice and individual is unclear until the order parameter is defined clearly at the start.

• Line 195 - 198: Reference missing.

• Line 198: What thresholds are we talking about here? The authors cannot leave it for the readers to figure out themselves.

• In so many place use of comma is entirely missing. Just for an example, in line 205, the sentence should be – Indeed, if the quality of ... decreases, the decision ... (Note the use of comma).

• Line 218: Isn’t the interpretation simple: that the groups are unable to respond quickly or react to environ- mental changes?

• Fig 6 is referred before figure 5 in the main text. Hence, change their numbering order.

• Line 227: The plot seems unimodal to me even after 660 minutes.

• Definition of consensual fiedelity not clear

• Supplementary text line 121: isn’t it attraction of unsheltered to sheltered?

• Supplementary text line 134: It should be mentioned that the solution is for a steady state by putting derivatives equal to zero.

• Supplementary text line 206: Typos in equation.

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If you choose “no”, your identity will remain anonymous but your review may still be made public.

Do you want your identity to be public for this peer review? For information about this choice, including consent withdrawal, please see our Privacy Policy.

Reviewer #1: No

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[NOTE: If reviewer comments were submitted as an attachment file, they will be attached to this email and accessible via the submission site. Please log into your account, locate the manuscript record, and check for the action link "View Attachments". If this link does not appear, there are no attachment files.]

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PONE-D-22-34285R1Emergence and retention of a collective memory in cockroachesPLOS ONE

Dear Dr. Calvo Martín,

Thank you for submitting your manuscript to PLOS ONE. After careful consideration, we feel that it has merit but does not fully meet PLOS ONE’s publication criteria as it currently stands. Therefore, we invite you to submit a revised version of the manuscript that addresses the points raised during the review process.

The decision is “Minor Revision” to give you a chance to respond to the comments of Reviewer #1. If this is done adequately, I will most likely be able to accept the manuscript without involving reviewers again.

Please submit your revised manuscript by July 1st, 2023. If you will need more time than this to complete your revisions, please reply to this message or contact the journal office at plosone@plos.org. When you're ready to submit your revision, log on to https://www.editorialmanager.com/pone/ and select the 'Submissions Needing Revision' folder to locate your manuscript file.

Please include the following items when submitting your revised manuscript:

• A rebuttal letter that responds to each point raised by the academic editor and reviewer(s). You should upload this letter as a separate file labeled 'Response to Reviewers'.
• A marked-up copy of your manuscript that highlights changes made to the original version. You should upload this as a separate file labeled 'Revised Manuscript with Track Changes'.
• An unmarked version of your revised paper without tracked changes. You should upload this as a separate file labeled 'Manuscript'.

If you would like to make changes to your financial disclosure, please include your updated statement in your cover letter. Guidelines for resubmitting your figure files are available below the reviewer comments at the end of this letter.

If applicable, we recommend that you deposit your laboratory protocols in protocols.io to enhance the reproducibility of your results. Protocols.io assigns your protocol its own identifier (DOI) so that it can be cited independently in the future. For instructions see: https://journals.plos.org/plosone/s/submission-guidelines#loc-laboratory-protocols. 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.

We look forward to receiving your revised manuscript.

Kind regards,

Wolfgang Blenau

Academic Editor

PLOS ONE

Journal Requirements:

Please 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 rebuttal letter that accompanies your revised manuscript. If you need to cite a retracted article, indicate the article’s retracted status in the References list and also include a citation and full reference for the retraction notice.

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Reviewers' comments:

Reviewer's Responses to Questions

Comments to the Author

1. If the authors have adequately addressed your comments raised in a previous round of review and you feel that this manuscript is now acceptable for publication, you may indicate that here to bypass the “Comments to the Author” section, enter your conflict of interest statement in the “Confidential to Editor” section, and submit your "Accept" recommendation.

Reviewer #1: All comments have been addressed

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2. Is the manuscript technically sound, and do the data support the conclusions?

The manuscript must describe a technically sound piece of scientific research with data that supports the conclusions. Experiments must have been conducted rigorously, with appropriate controls, replication, and sample sizes. The conclusions must be drawn appropriately based on the data presented.

Reviewer #1: Yes

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3. Has the statistical analysis been performed appropriately and rigorously?

Reviewer #1: I Don't Know

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4. Have the authors made all data underlying the findings in their manuscript fully available?

The PLOS Data policy requires authors to make all data underlying the findings described in their manuscript fully available without restriction, with rare exception (please refer to the Data Availability Statement in the manuscript PDF file). The data should be provided as part of the manuscript or its supporting information, or deposited to a public repository. For example, in addition to summary statistics, the data points behind means, medians and variance measures should be available. If there are restrictions on publicly sharing data—e.g. participant privacy or use of data from a third party—those must be specified.

Reviewer #1: No

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5. Is the manuscript presented in an intelligible fashion and written in standard English?

PLOS ONE does not copyedit accepted manuscripts, so the language in submitted articles must be clear, correct, and unambiguous. Any typographical or grammatical errors should be corrected at revision, so please note any specific errors here.

Reviewer #1: Yes

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6. Review Comments to the Author

Please use the space provided to explain your answers to the questions above. You may also include additional comments for the author, including concerns about dual publication, research ethics, or publication ethics. (Please upload your review as an attachment if it exceeds 20,000 characters)

Reviewer #1: I am satisfied with the author's response and changes made to the manuscipt and the supplemental material. However, minor grammatical errors and typos still remain. I would lke to suggest that the captions of Table 1 and 2 in the main text can be expanded that will help the readers to understand the technical analysis performed there. Finally, I could hardly understand Figure 6 of the main text due to its poor quality. Apart from these concerns, tha manuscript seems good for a publication. The conclusions about collective memory inferred through intelligent experiments and a rigorous mathematical model are very interesting and should be welcomed by the community of researchers in this area.

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7. PLOS authors have the option to publish the peer review history of their article (what does this mean?). If published, this will include your full peer review and any attached files.

If you choose “no”, your identity will remain anonymous but your review may still be made public.

Do you want your identity to be public for this peer review? For information about this choice, including consent withdrawal, please see our Privacy Policy.

Reviewer #1: No

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[NOTE: If reviewer comments were submitted as an attachment file, they will be attached to this email and accessible via the submission site. Please log into your account, locate the manuscript record, and check for the action link "View Attachments". If this link does not appear, there are no attachment files.]

While revising your submission, please upload your figure files to the Preflight Analysis and Conversion Engine (PACE) digital diagnostic tool, https://pacev2.apexcovantage.com/. PACE helps ensure that figures meet PLOS requirements. To use PACE, you must first register as a user. Registration is free. Then, login and navigate to the UPLOAD tab, where you will find detailed instructions on how to use the tool. If you encounter any issues or have any questions when using PACE, please email PLOS at figures@plos.org. Please note that Supporting Information files do not need this step.

Emergence and retention of a collective memory in cockroaches

PONE-D-22-34285R2

Dear Dr. Calvo Martín,

We’re pleased to inform you that your manuscript has been judged scientifically suitable for publication and will be formally accepted for publication once it meets all outstanding technical requirements.

Within one week, you’ll receive an e-mail detailing the required amendments. When these have been addressed, you’ll receive a formal acceptance letter and your manuscript will be scheduled for publication.

An invoice for payment will follow shortly after the formal acceptance. To ensure an efficient process, please log into Editorial Manager at http://www.editorialmanager.com/pone/, click the 'Update My Information' link at the top of the page, and double check that your user information is up-to-date. If you have any billing related questions, please contact our Author Billing department directly at authorbilling@plos.org.

If your institution or institutions have a press office, please notify them about your upcoming paper to help maximize its impact. If they’ll be preparing press materials, please inform our press team as soon as possible -- no later than 48 hours after receiving the formal acceptance. Your manuscript will remain under strict press embargo until 2 pm Eastern Time on the date of publication. For more information, please contact onepress@plos.org.

Kind regards,

Wolfgang Blenau

Academic Editor

PLOS ONE

Additional Editor Comments (optional):

Reviewers' comments: