Wilson, Georgina. Paper and the Making of Early Modern Literature. University of Pennsylvania Press, 2025, https://www.pennpress.org/9781512827446/paper-and-the-making-of-early-modern-literature/.

Related to the idea of paper helping, as a technology, create modernity.

See also Roland Allen's The Notebook (2023)

emphasis: global sustainability

this seems like a super relavent study for my research!!!!!!!

1. Alan scruffling my hair whilst I relax on the sofa
2. Tin Tin sleeping on a bed of soft leaves and cotton in the jungle
3. A soft tissue which Lady Penelope holds close to my runny nose

1. Tin Tin using frogs and war paint to get her to sleep in the jungle
2. Tin Tin playing with a peacock, a red panda and a baby elephant
3. Tin Tin cuddling a red panda like it's a teddy bear

1. Splashing in the bath with Lady Penelope
2. Collapsing in a heap of giggles on the bed
3. Tickling Alan while I'm hiding in his red jumper

https://en.wikipedia.org/wiki/Why_the_Future_Doesn%27t_Need_Us

Brown, John Seely, and Paul Duguid. “A Response to Bill Joy and the Doom-and-Gloom Technofuturists.” 2000. Emerging Technologies: Ethics, Law and Governance, by Gary E. Marchant and Wendell Wallach, edited by Gary E. Marchant and Wendell Wallach, 1st ed., Routledge, 2020, pp. 65–71.

via: https://web.cs.ucdavis.edu/~koehl/Teaching/ECS188_W16/Reprints/Response_to_BillJoy.pdf

annotation URL: urn:x-pdf:1e8f84f1b5e3fb65dfe49ef6f173c79e

A reprint of: <br /> - “Re-Engineering the Future: A Response to Bill Joy and the doom-and-gloom technofuturists,” The Industry Standard, John Seely Brown and Paul Duguid. 24 April 2000, p.196. - “A Response to Bill Joy and the Doom-and-Gloom Technofuturists,” AAAS Science and Technology Policy Yearbook 2001, edited by Albert H. Teich, Stephen D. Nelson, Celia McEnaney and Stephen J. Lita, American Association for the Advancement of Science, 2001.

Cross reference: Bill Joy's paper and notes at urn:x-pdf:753822a812c861180bef23232a806ec0

Joy, Bill. “Why the Future Doesn’t Need Us.” Wired, April 1, 2000. https://www.wired.com/2000/04/joy-2/.

Annotation url: urn:x-pdf:753822a812c861180bef23232a806ec0

Annotations: https://jonudell.info/h/facet/?user=chrisaldrich&url=urn%3Ax-pdf%3A753822a812c861180bef23232a806ec0&max=100&exactTagSearch=true&expanded=true

Reprints available at: - Joy, Bill. “Why the Future Doesn’t Need Us.” 2000. AAAS Science and Technology Policy Yearbook 2001, edited by Albert H. Teich et al., Amer Assn for the Advancement of Science, 2002, pp. 47–75. Google Books, https://www.google.com/books/edition/Integrity_in_Scientific_Research/0X-1g8YElcsC.<br /> - Joy, Bill. “Why the Future Doesn’t Need Us.” 2000. Emerging Technologies: Ethics, Law and Governance, by Gary E. Marchant and Wendell Wallach, edited by Gary E. Marchant and Wendell Wallach, 1st ed., Routledge, 2020, pp. 65–71.

via eli

Not just the form / medium: also you learn bad habits. Coming in with your own clever thing instead of asking questions

B

D

G

F

C

D

اذا بدي اغير title

If you up the ante, you increase your demands or the risks in a situation in order to achieve a better result

DOI: 10.1016/j.isci.2025.113402

Resource: RRID:IMSR_JAX:000664

Curator: @areedewitt04

SciCrunch record: RRID:IMSR_JAX:000664

What is this?

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This mirrors our own struggles as Pacific Islanders, in particular, Native Hawaiians. We continue to thrive, evolve, and empower ourselves, yet we continue to face subversive efforts to disenfranchise our people. A recent example is the attacks on Kamehameha School's admission policies.

This exactly how I feel every time I have to fill up a form at the doctor's office or in other places. This the answer of my question before.

I have to mention that it is 2025 and I keep seeing in some of the important forms and paperwork options about race and there only appear the following: White, Caucasian, Black and what option Latinx can choose?

Do you miss the Torchy Gift Books?

What does this mean? carrying coals during this time period signified a person's low status, equaling little or giving no respect. Sampson was essentially saying that they will tolerate no disrespect from the house of Montague. Right out the gate, you certainly sense a high level of disregard from Sampson for house of Montague.

when capulet servants are looking for a fight montagues servants doesn’t entertain the idea

again Gregory trying to clarify if Sampson meant to be this rude

after a little google search I found out this is used be a punishment back in the days. Here Gregory tries to tell Sampson, if he tries to provoke a fight he might get fired

They set the stage in a way(armors) that viewer or readers are expected the scene to be dramatic.

Romeo speaks to him not being able to dance and being afraid of being frozen in front of all the guests.

the nurse is uplifting. making a joke about who Juliet got her wits and remarks from.

A language barrier is one thing, but having the drive and mentality to be capable of doing something is what really matters. It is important to not let things you struggle with take a tole on you.

This would be a very big struggle. Having limitations with your own english and having someone communicate for you would not be an easy thing to be able to do. It may feel like you are a failure in some ways which is not good for ones well being.

What a brilliant way to tie the beginning of her narrative into the end. Her mother, who the narrator spoke of poorly and partially is able to overcome her grievances about how she speaks and is able to celebrate her mother in a wholesome way.

This is what's wrong with the American healthcare system today. There are people who work countless hours providing care and medical assistance to those in need but then there's the 'bad apples' who deal with power struggles and superiority complexes based on their employed position.

What a powerful statement... it really gets me thinking about how speaking a fluent language does not mean you understand the culture of that language entirely.

This points out how this difference in language made her feel for most of her life, contrasting her view that she shares at the end of story.

I really like the way these beginning sentences relate to the rest of the story and help set up the theme.

I think that this is very underrated because many times that is not an area of consideration. If other countries don't have good relationships with a country that stops trade and commerce meaning the economy suffers and so does general business.

I definitely have noticed this first hand. During the ice raids in LA in my place of work business definitely dropped and there were less people willing to shop around in an area of high risk. As soon as there was less publicity on ice business improved and sales increased.

natural resources, labor (human resources), capital, and entrepreneurship, and knowledge

Communicating with others is not always easy. To have someone reach out to her to allow her to speak her mind helps a lot. It takes off the pressure of having to speak up for her own.

Fear is hidden when you choose to not speak, but emotions are not let go and are still kept to yourself, which may leave a negative impact on yourself.

I could not have written a better ending to this vulnerable and beautiful piece of literature. The narrator teaches readers a valuable lesson about closure and how overcoming adversity takes many different steps--acknowledging the problem, trying out a solution and letting that problem no longer phase you.

This language usage is phennomenal... she is weaving together a powerful statement of how women are tired of being treated a certain way and instead of 'trying' to be heard, using the word 'crying' brings a whole different tone and feel to her writing.

This is such a powerful statement. The narrator vulnerably states that even though she believes she is doing the right thing for herself and her family it is still challenging and she may be risking a lot of strength to overcome this obstacle.

Poetry can be used as a way to help us understand our own feelings better. Respecting your own feelings can be a struggle for a lot of people so having a way to write down your sad moments in a creative way can help people feel more validated and come out stronger and know it's ok to feel all the feelings.

I find this comment to be so fun. Women get to carry the ideals of this "keystone for survival" in means of poetry. It reminds me greatly of my love for the confusing and confounding poems, and my boyfriend's love for the simple and straightforward prose.

Here is definition of poetry as "distillation of experience." This definition doesn't provide a mention of the use of language. Perhaps poetry by this construal could be a dance, a picture, or architectural work.

At first glance I totally disagree. Poetry is definitely a luxury, as is the ability to read and write, the ability to share poetry freely (free speech), or even have the time to dedicate to poetry. Not saying it should be but unfortunately its how the world we live in works.

Wow I love this poem! This poem has so many holes in it to make you think and reflect on your own experiences. This sentence makes the metaphor as a drive to readers. Showing that us human have so many dreams and hopes, but sometimes we are too scared to act on those dreams and hopes because the fear overpowers it. This helps me realize and want to prove the reader wrong in my life.

This part of the sentence is powerful to me personally. Because everyone has battles and if we keep them silent they are going to take control of us. It encourages the reader to talk to loved ones so they don't lose the battle.

Thinking about it, I remember art projects and art included into lessons more than almost anything else. The creative parts of those lessons stayed with me, while regular worksheets or lectures were easier to forget because they weren't engaging. I think it’s because art made me get involved and use my own ideas, which helped the content we learned stick in my memory better.

This makes me think about how teachers typically reward silence and quiet, but arts integration rewards social interaction and conversation. It changes the classroom dynamic into a more active and engaging environment. I do worry about students discussing things that are off topic though.

Red Bull also markets in a way that makes the perceiver subconsciously believe that people who drink Red Bull can do insane stunts based on the stunts that people do while marketing Red Bull, from what I can tell. It makes it seem that it's healthy as well because these athletes who are performing the stunts are drinking Red Bull, when it's not healthy for the body.

Some demographics to consider might be that not everyone has had the experience of working for their parents or working at a food truck, which both apply to me, but not everyone can relate to that.

This makes sense for the speaker to go with dogs because it's a good, relatable subject to go into. People know what dogs are, and generally it is a topic most people can relate to. Sports, Freeganism, Wall Street, vinyl music and Hipster Culture are all not so common topics to go with

The ending of this will truly stick with me, as this notion of no one caring about Icarus' fall always makes me question the story. No one to care for you seems implausible.

I find this brings me the most joy when I have yet to fully understand the entire depth of poems and their meanings or knowledge.

Crazy statement. One other thing is that sometimes people dont recognize their own suffering. They go day in day out thinking that the pain they experience is normal just because they have been desensitized to it and then don't know that they need to advocate for themselves or that its even an issue. ei hoarders, people with abusive parents, addicts.

What I think is interesting is how in one of the first poems I read they stated that use commas to a minimum, but in this sentence there are multiple commas. Should the author used more descriptive words?

This explantion of formative and summative assessments gave me a slightly new perspective. Thinking of formative assessments like homework and quiz assignments and students learning not being in the final form yet stood out to me. By the time the summative evaluation or unit test is given, students should feel like they experience enough "draft" exercises and feel prepared to demonstrate a deep understanding of the topic without the need for revisions.

I have personally used both chatgpt and game based learning in my classroom. I don't use chatgpt to provide feedback or assess. I do believe that those aspects are primarily the educators responsibility. However, I have used chatgpt to create collaborative groups, design seating charts, create short stories using specific information and more! I have found It to be quite useful when feeling overwhelmed with the amount of tasks we face daily. As for game based learning, my students love playing blooket and gimkit. I use this as an opportunity to connect vocabulary to images for better memory and to do drill practice with topics like verb conjugations. This past school year, I had students create a trivia game by creating questions about the country of their year long portfolio.

It's very interesting to see how these predictions were being made in the early 80's and while many schools do have 1:1 devices, there are several that do not. It's rare to see a school without student laptops these days, I remember having to go to the computer lab, then it changed to the computer cart and finally, a device for each student. What will come next?

This is something educators are very familiar with today. We attend several mandatory trainings a year that are completed online using a training video to instruct us on topics such as hazardous materials,sexual harassment,etc.

I think in today's society, the radio acts as a tool for gathering information which can then be applied to other situations. I was shocked to hear about how bad the traffic at my school was yesterday morning on the radio while I was actively sitting in it! We continue to collect information from radio broadcasts,podcasts, audiobooks,etc.

While we are much less dependent on written content in books in 2025, Edison's belief was incorrect. We still watch films in school but they we also reread books and hope that between the written format, discussions and visual.audio media, students can engage in meaningful learning.

I find this connection between the process of bringing in actual items in the 1900's to now having a digital media center. In my opinion, the writer mentions the district wide media center to imply that learners now have even more extensive access to the outside world/history than they did when a few schools had a school museum.

I find this sentence extremely interesting- as many times stories do cause hardships to write. It almost feels endearing knowing that writers struggle as often as common-folk, and experience such normalities as writers block.

It is upsetting how women having control and being in power, even in times way past ours now, is considered feminist and inane. Women having the same rights and power stance as a man will seemingly always be feminist and not just something that should happen- something equal.

first time seeing this word a "philologist is a specialist in philology, which is the study of language in its historical, oral, and written contexts, focusing on literary texts, historical records, and the associated cultures"

Interesting to see that first author we have in the historical record was a woman. In so many ancient cultures all of priesthood, access to education, and public life were limited to men.

I like how the definitions of the words were included in the poetry. It gives more of learning aspect and better understanding of what each word means.

This is a great metaphor. More than likely a poem will not tell you word by word what they are trying to covey, instead the use mystery to help you think more on the poem and express your own thoughts with your own experiences.

Before reading this article I just assumed over time we as people figured out that to access clean and drinkable water, we have to have it purified just like anything else. I had no idea that cholera was the main threat that pushed the sanitation movement. Another thing I had never heard of was the sanitation movement. I had no clue that fighting against waterborne diseases came from a cholera root cause discovery by John Snow.

I find it interesting that even though this theory was wrong it should have also improved conditions where those with a strong enough immune system may not have been effected or effected as badly. It seems like a good start but impressive of Dr. Snow to conclude that the method of transmission was different which in turn completely changes the type of sanitation preformed / actions taken to prevent disease.

This is a classroom management technique that I think is underrated. How a teacher structures and designs their classroom can affect how a student pays attention. Which direction they are facing or learning content posters on the wall are examples.

I feel as though this could be fought against as untrue. Even the shortest poems have a beginning and end- and I think even an unfinished work still has the ending as the last word written.

Kind of like school or life as a whole. Even in Kindergarten we had a graduation. dressed up like college graduates and it probably felt like in that moment I could have titled it 'the end.' Of course we've all gone on passed that phase. Yet even now in college when we graduate it will feel similar even though millions graduate every year and countless others did in all the years prior. In a way it is child like to recognize achievements in our lives that have been done before but at the same time most peoples lives aren't that unique and most everyone goes through the same motions. What else would we get to celebrate

this is the key lesson

I think this definition implies a separation between instructional design and technology/media. Contrary to earlier definitions, this title acknowledges the inclusion of systemic processes used to design meaningful curriculum while also including the availability to include different modes of technology to follow these instructional procedures.

Answers my question, how do i think critically about potential sources?

I think this statement is a great way to start the article and definitely plays a role in why there have been so many young teacher burnt out and switching careers.

the two greatest dangers to American prosperity.

ver esto me recuerda a la clase de fundamentos de investigación y a como sufri para aprender a aplicar medianamente bien (no nos vamos a engañar) APA 7ma edición a mis trabajos

I have seen this countless times and I am not even a teacher yet!

I think that this is a great point!! Teachers have to put in the prep to see learning and growth as an outcome

It is nice to know the info is from a good source.

Argument about being more human.

Imporant

We have to prevent leaving things to the last minute.

This shows me effective strategic leaders are able to collectively understand and combine a variety of ideas to improve an organization.

Coastal trunk cable or something?

I always have a question that when some people have a theory ahead, then gather the data. They sometimes tend to pretend that the data perfectly proves their theory, maybe delete some irrelevant data, and then write a fraudulent paper. Would things like that happen? And how can academia prevent that?

Why is is I2 and not H2? @rebelford

En este foto, creo que las personas son contentos y queríamos celebrar los tiempos mejores durante la gran depresión. Que interesante las personas pueden usar la cine para escapar.

I can get behind 1-6 they seems like effective methods for understanding a poem. But all of 7 I think can actually be summarized to "does it have thought behind it?" Any one of those rules can be broken if the author intends for them to be broken or there is a greater purpose for including or omitting that technical detail. For example maybe the voice of the work is inconsistent to show the author's insanity or incorrectly using punctuation as a metaphor for something greater. I dont know, but I think that as long as there is thought and purpose behind the technical details its really doesn't matter how you write a poem

This is an excellent requirement for an A in this course. Being able to work well with others while standing out beyond the number of words on a page, and more importantly, how you convey and explain your writing, is a more effective way for a student to demonstrate their potential beyond the ideal requirements for a paper or worksheet. I believe that, in certain circumstances, where we have requirements for our writing, such as "500-word minimum essays" or limiting phrases, ultimately leads us to being more unfocused and uninteresting writers. This is an understandable requirement for an A, which is still achievable.

I agree with the standards that are required to get a C, B and an A. Being able to prioritize quality over quantity is an important skill, along with speaking. The ability to improve also shows proof of learning the material sufficiently. I've never experienced a class like this but I am very much looking forward to it!

I understand and agree with the requirements for a C, B, and A. When discussing the value of quality over quantity and the need to be able to lead is a reasonable standard for earning an A. Consistently improving is important as well, as it is proof of properly learning and absorbing the information.

I like this concept and i've seen it in other courses, I like how these reflect how much work you put in reflects the grade as well, Doing things like turning assignments on time are bare minimum when it comes to turning assignments in.

I like this section because it encourages students to go beyond the basics and really polish their work. The emphasis on listening as a skill is also valuable it shows that good participation is not just about talking but also paying attention to others.

I like how this section sets a clear baseline for students. It makes it very straightforward, if you keep up with readings, assignments, discussions, and feedback, you can pass. I think this is fair because it rewards consistent effort.

Is this meant more to make us laugh, or to leave us thinking?

**It's interesting how Singh admits to that there some truth to these stereotypes **

**This Suprises me because of how students can even hold judgement towards their peers in the same major as them. **

This shows a rhetorical appeal (logos) because it explains the real reason behind stereotyping in a logical way.

Ends with playfully to remind readers stereotypes aren’t serious and to make the article memorable. Setting the tone of sarcastic and funny

The author is addressing this specific text because the audience it is addressing is directly to college students who are choosing or defending their majors.

*This could be an example of pathos because it sets a tone of being balanced, fair-minded. It softens the stereotype argument by recognizing difficulty across fields. *

One of the main points to the key claim, it explains why stereotypes happen.

This is a large downside to seeing and getting knowledge from the internet, because it's so easy to see a biased perspective when its the only one you've seen.

Influencers on Tik Tok showing days in the life videos in different parts of the country.

I chose this title because I have learned most of my knoledge about Africa from the internet and promotion videos.

This doesn't make as much sense to me; I feel like the whole point (in my mind) is to be able to walk the path, to feel it under your feet. However, this might also be inaccessible for certain people and age groups.

feeling a bit like "every square is a rectangle but not every rectangle is a square," these are separate things but not unattached

These examples offer practical solutions and show that restrictions can work when well-implemented. These policies, while sometimes controversial, have led to positive feedback from educators regarding student engagement and focus.

This section emphasizes the grass-roots nature of many policies, showing how teacher advocacy and involvement can be a powerful catalyst for institutional change

Educators are increasingly concerned about the impact of cellphones not only on academic focus but also on students’ mental health, particularly due to social media’s effect.

Schools have struggled with how to regulate cellphones for over 15 years. Teens should not be on their phones all the time.

The narrator connects her own struggle to these imagined figures. Fantasy blends with reality, showing how women use myth to cope with hardship.

The women’s lives are a performance where they must “awe” others while hiding their own pain. This shows the theme of survival through illusion.

“Ghost women” symbolize women who live in the margins ignored, invisible, exhausted, but still carrying beauty (stars). The wave imagery shows instability and struggle, while brushing stars suggests both hope and weariness.

nodule

add space

increasing the expression of sodium channels, potassium channels, and the Na+/K+ ATPase in the distal tubule of the kidney.

binding

(DOC to corticosterone to 18-hydroxycorticosterone to aldosterone)

precursor

precursor

increase in sympathetic activity, and a decrease in NaCl passing through the renal nephron, as detected by the salt-sensing macula densa cells. (if "interesting" mechanistic text on a slide is not explained a little bit, it is very annoying for the students. But they can do without all the things that angiotensin II does for now.)

and starts with secretion of the protease renin from the juxtaglomerular cells of the kidney. Renin catalyzes the conversion of angiotensinogen, made in the liver, to angiotensin I that is converted to angiotensin II by the protease angiotensin-converting enzyme (ACE). (Students are often confused by and miss the fact that renin and ACE are proteases, so better to explain it here, as it is also illustrated in figure 4.)

as well as other peptides.

I honestly agree. I love to learn, I do, but sometimes my fear of failing gets so overwhelming. I think this highlights alot of the anxiety students feel about getting good grades and passing.

Sounds like the "flattening your voice" argument

thats crazy

Seeing this as one of the requirements for a passing grade is refreshing to see listed. We sometimes forget, as students, when we are navigating new courses, job opportunities, and everyday life, that it is vital to feed our minds with ideas that don't initially come to mind. Thinking deeply and forming connections with past readings and courses has helped me tremendously as a student when it comes to passing a class. I believe it's excellent that you brought this to students' attention in hopes of them succeeding within your course.

The idea of Gorbachev working in a field where there are benefits and security is clearly demonstrated in this paragraph. We see more of the good and effectiveness that Gorbachev has begun to enhance with China. Increasing trade and the opening of borders definitely give Gorbachev some credit; however, what caught my attention was that, despite some basic things flourishing, the Soviet Union's relationship with China remains somewhat rocky and distant. This is not to say they can't work together (which they undoubtedly have and can), but it's the question asked as to why tensions can't decline and relationships grow stronger for the benefit of resources. One of my biggest takeaways from this paragraph is how, even though things are not perfect within their feelings and restrictions towards each other, there is still a balance that is acceptable to maintain consistency and keep things working for both sides.

Due to Gorbachev's extensive pride and loyalty towards the Soviet Union, I sense that the author fears that Gorbachev would rather keep and enforce based on previous Soviet Union orders to fix the problematic pressures there are economically rather than bringing in new possible solutions that would better fix the issues but may be to new or skeptical or rely to much on the people. I sense that the author subjectively feels that Gorbachev would rather respect and preserve the power of the Soviet Union and not risk the idea of giving the people too much power to make executive decisions.

This paragraph answers the question "Why is your participation important?". Participation is important because there must be engagement and effort put in to learn what liberal education teaches.

I never knew what DALN was until I read this. It seems like it could be something I'd be interested in. Seems to be something purposefully comfortable and down to earth

This I feel is super important to the reader/audience. Wether we like or dislike a character especially the main antagonist Because that could end up being the whole point.

Language is the way we speak to one another about any sort of our ideas, feelings and information.

Basically is finding people with your same goals and learning together on how to accomplish that goal.

This is a really interesting method and was a super fun read! I like that not only can it clearly accelerate binder studies, but it can be done in any lab. I'm looking for a use case to try it out already!

Can you interrogate the effect of these different features on the signal output in this method?

This is interesting to me because geographers construct models when they are not able to study or experiment with the real thing. If they are not able to study or experiment on the real object, how do they know their model is accurate?

This way of backing out recombination rate from Ner makes some strict assumptions, particularly that pi reflect mutation-drift equilibrium, and that mu is invariant across the genome. This is unlikely to be true in most datasets. It would be very helpful if more region specific comparisons could be made to pedigree based estimates to evaluate how well this approximation works in this dataset.

Again I think the influence of phylogenetic signal is important to consider here. A phylogenetically corrected regression (e.g. phylogenetic generalized least squares) would take into account the non-independence of species observations and provide a more accurate view of the data.

It would be nice to get an idea of how stable this porting over of the estimated recombination map is across phylogenetic distance. There is considerable variation in the correlation among species pairs which would be good to understand. Also would be valuable to show the reader how the phylogenetic distance of these comparisons relates to the distances used for comparisons of the main analysis.

The message of this figure get's a little muddled. One would expect that as as Ne increases, that the proportion of mutations experiencing a larger (more negative) Nes would grow (as in the case of increasing r) but the opposite is shown. Unless the figure implies that fewer segregating mutations will have large Nes? But then this conflicts with the sequence for recombination.

Aerial photographs provide us with a real-world view of the earth's surface, whereas maps are just a representation of the real world.

Remote sensing is just one of the many tools geographers use to observe the earth's physical features. Remote sensing probably helps make maps more accurate.

There are various types of data that allow for diverse maps. Each map is designed to tell a story using the data.

It's interesting to see how different maps were made to represent different types of data.

This is something that I've struggle with but this year I am trying to change it.

It's a balancing act, if you don't know or understand something, it okay to ask your instructor for help, but overloading them with so many questions can make it hard for them to help other students. If you have trouble understanding the material, you may need a tutor.

The best writers are the ones who seek help from those around them.

Some professors require a specific subject line and this is the example for English.

I purposely don't have an exception set on what will be required of me during my time at CNM in general, not just this class. No matter how unconfident I was before this semester, this a new opportunity to prove myself.

Writing with different styles, tones and circumstances is going to help in every area of life. Whether it be a job application, a university application, or just an important letter/email. Knowing the intended audience and level of professionalism is a learned skill.

Darwinism-in other words socialism; many laborers believed it would lead to better wages/more rights for laborers.

robber barons- financial and industrial titans

J.P. Morgan- the company which controlled steel & had a lot of profit from competition melting away and companies rising and falling

mass production- the production of many machines and items with cheap production costs McCormick- the person associated with making the machines

Taylorism-the idea that with the new machine age, firms needed scientific organization and to subdivide tasks to speed up production.

Using militia showed more force towards the strikers and showed their lack of understanding towards them. Using militia just worsened the situation by showing they didn't care. Innocent people died by using extreme force.

The author mentions to love one another and yet the native americans showed kindness and the colonizers showed hate and basically laughed at the gods that the native americans worshipped and told them there is only one god and that is my God.

It seems kinda ironic for the author to say that he is spreading God's word to cleanse the word of evil when history has shown that they had caused so much harms to other people i.e. the native americans. Not only that but they mention on how both the rich and the poor should respect each other while the rich are hoarding money, jewelry, and power. All while the poor are in famine and dying from dieseas cause they are so poor.

it would also be valuable to see which Raman wavenumbers drive the LDA axes to see if they make conceptual sense and are not just noisy artifacts. by eye, the representative spectra in fig 1c are very similar, so it would be good to see that the differences between the strains are driven by real differences in peaks

were the transcriptomes (or at least a subset) validated in-house? some more details about how the transcriptome data was used would be very beneficial to the reader.

I like seeing the different ways Geographic Information Systems can be used. It's so interesting to see how multiple professions can use them effectively.

writing is such a boring, mundane task to many, so it is important to make the prompts interesting

the education system needs to alter itself in order to prevent the complete takeover of AI in academic environments

taking the long route gives us necessary skills to progress in life

I like this use of this word because it emphasizes the process rather than the result

This really highlights the heart of teaching; it’s not just about academics but helping students grow into curious, capable, and purposeful individuals.

I like how this connects to building strong teacher-student relationships. It shows that understanding the law helps create a safe, fair environment where those relationships can grow.

This is an important reminder that court rulings can vary within a state and may even conflict with each other. It shows why educators need to stay updated on legal decisions, since what’s true in one area might not apply statewide

This reminds teachers that laws heavily shape how schools operate, but legal cases can have different outcomes for the same issue.

If someone’s rights are violated, they must be given a fair and unbiased hearing before any decisions or punishments are made.

eLife Assessment

This valuable model-based study seeks to mimic bat echolocation behavior and flight under conditions of high interference, such as when large numbers of bats leave their roost together. Although some of the assumptions made in the model may be questioned, the simulations convincingly suggest that the problem of acoustic jamming in these situations may be less severe than previously thought. This finding will be of broad interest to scientists working in the fields of bat biology and collective behaviour.

Reviewer #1 (Public review):

Summary:

Mazer & Yovel 2025 dissect the inverse problem of how echolocators in groups manage to navigate their surroundings despite intense jamming using computational simulations.

The authors show that despite the 'noisy' sensory environments that echolocating groups present, agents can still access some amount of echo-related information and use it to navigate their local environment. It is known that echolocating bats have strong small and large-scale spatial memory that plays an important role for individuals. The results from this paper also point to the potential importance of an even lower-level, short-term role of memory in the form of echo 'integration' across multiple calls, despite the unpredictability of echo detection in groups. The paper generates a useful basis to think about the mechanisms in echolocating groups for experimental investigations too.

Strengths:

* The paper builds on biologically well-motivated and parametrised 2D acoustics and sensory simulation setup to investigate the various key parameters of interest

* The 'null-model' of echolocators not being able to tell apart objects & conspecifics while echolocating still shows agents succesfully emerge from groups - even though the probability of emergence drops severely in comparison to cognitively more 'capable' agents. This is nonetheless an important result showing the direction-of-arrival of a sound itself is the 'minimum' set of ingredients needed for echolocators navigating their environment.

* The results generate an important basis in unraveling how agents may navigate in sensorially noisy environments with a lot of irrelevant and very few relevant cues.

* The 2D simulation framework is simple and computationally tractable enough to perform multiple runs to investigate many variables - while also remaining true to the aim of the investigation.

Weaknesses:

* Authors have not yet provided convincing justification for the use of different echolocation phases during emergence and in cave behaviour. In the previous modelling paper cited for the details - here the bat-agents are performing a foraging task, and so the switch in echolocation phases is understandable. While flying with conspecifics, the lab's previous paper has shown what they call a 'clutter response' - but this is not necessarily the same as going into a 'buzz'-type call behaviour. As pointed out by another reviewer - the results of the simulations may hinge on the fact that bats are showing this echolocation phase-switching, and thus improving their echo-detection. This is not necessarily a major flaw - but something for readers to consider in light of the sparse experimental evidence at hand currently.

* The decision to model direction-of-arrival with such high angular resolution (1-2 degrees) is not entirely justifiable - and the authors may wish to do simulation runs with lower angular resolution. Past experimental paradigms haven't really separated out target-strength as a confounding factor for angular resolution (e.g. see the cited Simmons et al. 1983 paper). Moreover, to this reviewer's reading of the cited paper - it is not entirely clear how this experiment provides source-data to support the DoA-SNR parametrisation in this manuscript. The cited paper has two array-configurations, both of which are measured to have similar received levels upon ensonification. A relationship between angular resolution and signal-to-noise ratio is understandable perhaps - and one can formulate such a relationship, but here the reviewer asks that the origin/justification be made clear. On an independent line, also see the recent contrasting results of Geberl, Kugler, Wiegrebe 2019 (Curr. Biol.) - who suggest even poorer angular resolution in echolocation.

Reviewer #2 (Public review):

This manuscript describes a detailed model for bats flying together through a fixed geometry. The model considers elements which are faithful to both bat biosonar production and reception and the acoustics governing how sound moves in air and interacts with obstacles. The model also incorporates behavioral patterns observed in bats, like one-dimensional feature following and temporal integration of cognitive maps. From a simulation study of the model and comparison of the results with the literature, the authors gain insight into how often bats may experience destructive interference of their acoustic signals and those of their peers, and how much such interference may actually negatively effect the groups' ability to navigate effectively. The authors use generalized linear models to test the significance of the effects they observe.

The work relies on a thoughtful and detailed model which faithfully incorporates salient features, such as acoustic elements like the filter for a biological receiver and temporal aggregation as a kind of memory in the system. At the same time, the authors abstract features that are complicating without being expected to give additional insights, as can be seen in the choice of a two-dimensional rather than three-dimensional system. I thought that the level of abstraction in the model was perfect, enough to demonstrate their results without needless details. The results are compelling and interesting, and the authors do a great job discussing them in the context of the biological literature.

With respect to the first version of the manuscript, the authors have remedied all my outstanding questions or concerns in the current version. The new supplementary figure 5 is especially helpful in understanding the geometry.

Author response:

The following is the authors’ response to the original reviews.

Reviewer #1 (Public review):

We thank the reviewer for his valuable input and careful assessment, which have significantly improved the clarity and rigor of our manuscript.

Summary:

Mazer & Yovel 2025 dissect the inverse problem of how echolocators in groups manage to navigate their surroundings despite intense jamming using computational simulations.

The authors show that despite the 'noisy' sensory environments that echolocating groups present, agents can still access some amount of echo-related information and use it to navigate their local environment. It is known that echolocating bats have strong small and large-scale spatial memory that plays an important role for individuals. The results from this paper also point to the potential importance of an even lower-level, short-term role of memory in the form of echo 'integration' across multiple calls, despite the unpredictability of echo detection in groups. The paper generates a useful basis to think about the mechanisms in echolocating groups for experimental investigations too.

Strengths:

(1) The paper builds on biologically well-motivated and parametrised 2D acoustics and sensory simulation setup to investigate the various key parameters of interest

(2) The 'null-model' of echolocators not being able to tell apart objects & conspecifics while echolocating still shows agents successfully emerge from groups - even though the probability of emergence drops severely in comparison to cognitively more 'capable' agents. This is nonetheless an important result showing the directionof-arrival of a sound itself is the 'minimum' set of ingredients needed for echolocators navigating their environment.

(3) The results generate an important basis in unraveling how agents may navigate in sensorially noisy environments with a lot of irrelevant and very few relevant cues.

(4) The 2D simulation framework is simple and computationally tractable enough to perform multiple runs to investigate many variables - while also remaining true to the aim of the investigation.

Weaknesses:

There are a few places in the paper that can be misunderstood or don't provide complete details. Here is a selection:

(1) Line 61: '... studies have focused on movement algorithms while overlooking the sensory challenges involved' : This statement does not match the recent state of the literature. While the previous models may have had the assumption that all neighbours can be detected, there are models that specifically study the role of limited interaction arising from a potential inability to track all neighbours due to occlusion, and the effect of responding to only one/few neighbours at a time e.g. Bode et al. 2011 R. Soc. Interface, Rosenthal et al. 2015 PNAS, Jhawar et al. 2020 Nature Physics.

We appreciate the reviewer's comment and the relevant references. We have revised the manuscript accordingly to clarify the distinction between studies that incorporate limited interactions and those that explicitly analyze sensory constraints and interference. We have refined our statement to acknowledge these contributions while maintaining our focus on sensory challenges beyond limited neighbor detection, such as signal degradation, occlusion effects, and multimodal sensory integration (see lines 58-64):

(2) The word 'interference' is used loosely places (Line 89: '...took all interference signals...', Line 319: 'spatial interference') - this is confusing as it is not clear whether the authors refer to interference in the physics/acoustics sense, or broadly speaking as a synonym for reflections and/or jamming.

To improve clarity, we have revised the manuscript to distinguish between different types of interference:

• Acoustic interference (jamming): Overlapping calls that completely obscure echo detection, preventing bats from perceiving necessary environmental cues.

• Acoustic interference (masking): Partial reduction in signal clarity due to competing calls.

• Spatial interference: Physical obstruction by conspecifics affecting movement and navigation.

We have updated the manuscript to use these terms consistently and explicitly define them in relevant sections (see lines 84-85, 119-120). This distinction ensures that the reader can differentiate between interference as an acoustic phenomenon and its broader implications in navigation.

(3) The paper discusses original results without reference to how they were obtained or what was done. The lack of detail here must be considered while interpreting the Discussion e.g. Line 302 ('our model suggests...increasing the call-rate..' - no clear mention of how/where call-rate was varied) & Line 323 '..no benefit beyond a certain level..' - also no clear mention of how/where call-level was manipulated in the simulations.

All tested parameters, including call rate dynamics and call intensity variations, are detailed in the Methods section and Tables 1 and 2. Specifically:

• Call Rate Variation: The Inter-Pulse Interval (IPI) was modeled based on documented echolocation behavior, decreasing from 100 msec during the search phase to 35 msec (~28 calls per second) at the end of the approach phase, and to 5 msec (200 calls per second) during the final buzz (see Table 2). This natural variation in call rate was not manually manipulated in the model but emerged from the simulated bat behavior.

• Call Intensity Variation: The tested call intensity levels (100, 110, 120, 130 dB SPL) are presented in Table 1 under the “Call Level” parameter. The effect of increasing call intensity was analyzed in relation to exit probability, jamming probability, and collision rate. This is now explicitly referenced in the Discussion. We have revised the manuscript to explicitly reference these aspects in the Results and Discussion sections – see lines 346-349, 372-375.

Reviewer #2 (Public review):

We are grateful for the reviewer’s insightful feedback, which has helped us clarify key aspects of our research and strengthen our conclusions.

This manuscript describes a detailed model of bats flying together through a fixed geometry. The model considers elements that are faithful to both bat biosonar production and reception and the acoustics governing how sound moves in the air and interacts with obstacles. The model also incorporates behavioral patterns observed in bats, like one-dimensional feature following and temporal integration of cognitive maps. From a simulation study of the model and comparison of the results with the literature, the authors gain insight into how often bats may experience destructive interference of their acoustic signals and those of their peers, and how much such interference may actually negatively affect the groups' ability to navigate effectively. The authors use generalized linear models to test the significance of the effects they observe.

In terms of its strengths, the work relies on a thoughtful and detailed model that faithfully incorporates salient features, such as acoustic elements like the filter for a biological receiver and temporal aggregation as a kind of memory in the system. At the same time, the authors' abstract features are complicating without being expected to give additional insights, as can be seen in the choice of a twodimensional rather than three-dimensional system. I thought that the level of abstraction in the model was perfect, enough to demonstrate their results without needless details. The results are compelling and interesting, and the authors do a great job discussing them in the context of the biological literature. 

The most notable weakness I found in this work was that some aspects of the model were not entirely clear to me. 

For example, the directionality of the bat's sonar call in relation to its velocity. Are these the same?

For simplicity, in our model, the head is aligned with the body, therefore the direction of the echolocation beam is the same as the direction of the flight. 

Moreover, call directionality (directivity) is not directly influenced by velocity. Instead, directionality is estimated using the piston model, as described in the Methods section. The directionality is based on the emission frequency and is thus primarily linked to the behavioral phases of the bat, with frequency shifts occurring as the bat transitions from search to approach to buzz phases. During the approach phase, the bat emits calls with higher frequencies, resulting in increased directionality. This is supported by the literature (Jakobsen and Surlykke, 2010; Jakobsen, Brinkløv and Surlykke, 2013). This phase is also associated with a natural reduction in flight speed, which is a well-documented behavioral adaptation in echolocating bats(Jakobsen et al., 2024).

To clarify this in the manuscript, we have updated the text to explicitly state that directionality follows phase-dependent frequency changes rather than being a direct function of velocity, see lines 543-545. 

If so, what is the difference between phi_target and phi_tx in the model equations? 

𝝓_{𝒕𝒂𝒓𝒈𝒆𝒕} represents the angle between the bat and the reflected object (target).

𝝓_𝑻𝒙 the angle [rad], between the masking bat and target (from the transmitter’s perspective)

𝝓_{𝑻𝒙𝑹𝒙} refers to the angle between the transmitting conspecific and the receiving focal bat, from the transmitter’s point of view.

𝝓_{𝑹𝒙𝑻𝒙} represents the angle between the receiving bat and the transmitting bat, from the receiver’s point of view.

These definitions have been explicitly stated in the revised manuscript to prevent any ambiguity (lines 525-530). Additionally, a Supplementary figure demonstrating the geometrical relations has been added to the manuscript.

What is a bat's response to colliding with a conspecific (rather than a wall)? 

In nature, minor collisions between bats are common and typically do not result in significant disruptions to flight (Boerma et al., 2019; Roy et al., 2019; Goldshtein et al., 2025). Given this, our model does not explicitly simulate the physical impact of a collision event. Instead, during the collision event the bat keeps decreasing its velocity and changing its flight direction until the distance between bats is above the threshold (0.4 m). We assume that the primary cost of such interactions arises from the effort required to avoid collisions, rather than from the collision itself. This assumption aligns with observations of bat behavior in dense flight environments, where individuals prioritize collision avoidance rather than modeling post-collision dynamics. See lines 479-484.

From the statistical side, it was not clear if replicate simulations were performed. If they were, which I believe is the right way due to stochasticity in the model, how many replicates were used, and are the standard errors referred to throughout the paper between individuals in the same simulation or between independent simulations, or both? 

The number of repetitions for each scenario is detailed in Table 1, but we included it in a more prominent location in the text for clarity. Specifically, we now state (Lines 110-111):

"The number of repetitions for each scenario was as follows: 1 bat: 240; 2 bats: 120; 5 bats: 48; 10 bats: 24; 20 bats: 12; 40 bats: 12; 100 bats: 6."

Regarding the reported standard errors, they are calculated across all individuals within each scenario, without distinguishing between different simulation trials. 

We clarified in the revised text (Lines 627-628 in Statistical Analysis) 

Overall, I found these weaknesses to be superficial and easily remedied by the authors. The authors presented well-reasoned arguments that were supported by their results, and which were used to demonstrate how call interference impacts the collective's roost exit as measured by several variables. As the authors highlight, I think this work is valuable to individuals interested in bat biology and behavior, as well as to applications in engineered multi-agent systems like robotic swarms.

Reviewer #3 (Public review):

We sincerely appreciate the reviewer’s thoughtful comments and the time invested in evaluating our work, which have greatly contributed to refining our study.

We would like to note that in general, our model often simplifies some of the bats’ abilities, under the assumption that if the simulated bats manage to perform this difficult task with simpler mechanisms, real better adapted bats will probably perform even better. This thought strategy will be repeated in several of the s below.

Summary:

The authors describe a model to mimic bat echolocation behavior and flight under high-density conditions and conclude that the problem of acoustic jamming is less severe than previously thought, conflating the success of their simulations (as described in the manuscript) with hard evidence for what real bats are actually doing. The authors base their model on two species of bats that fly at "high densities" (defined by the authors as colony sizes from tens to tens of thousands of individuals and densities of up to 33.3 bats/m2), Pipistrellus kuhli and Rhinopoma microphyllum. This work fits into the broader discussion of bat sensorimotor strategies during collective flight, and simulations are important to try to understand bat behavior, especially given a lack of empirical data. However, I have major concerns about the assumptions of the parameters used for the simulation, which significantly impact both the results of the simulation and the conclusions that can be made from the data. These details are elaborated upon below, along with key recommendations the authors should consider to guide the refinement of the model.

Strengths:

This paper carries out a simulation of bat behavior in dense swarms as a way to explain how jamming does not pose a problem in dense groups. Simulations are important when we lack empirical data. The simulation aims to model two different species with different echolocation signals, which is very important when trying to model echolocation behavior. The analyses are fairly systematic in testing all ranges of parameters used and discussing the differential results.

Weaknesses:

The justification for how the different foraging phase call types were chosen for different object detection distances in the simulation is unclear. Do these distances match those recorded from empirical studies, and if so, are they identical for both species used in the simulation? 

The distances at which bats transition between echolocation phases are identical for both species in our model (see Table 2). These distances are based on welldocumented empirical studies of bat hunting and obstacle avoidance behavior (Griffin, Webster and Michael, 1958; Simmons and Kick, 1983; Schnitzler et al., 1987; Kalko, 1995; Hiryu et al., 2008; Vanderelst and Peremans, 2018). These references provide extensive evidence that insectivorous bats systematically adjust their echolocation calls in response to object proximity, following the characteristic phases of search, approach, and buzz.

To improve clarity, we have updated the text to explicitly state that the phase transition distances are empirically grounded and apply equally to both modeled species (lines 499-508).

What reasoning do the authors have for a bat using the same call characteristics to detect a cave wall as they would for detecting a small insect? 

In echolocating bats, call parameters are primarily shaped by the target distance and echo strength. Accordingly, there is little difference in call structure between prey capture and obstacles-related maneuvers, aside from intensity adjustments based on target strength (Hagino et al., 2007; Hiryu et al., 2008; Surlykke, Ghose and Moss, 2009; Kothari et al., 2014). In our study, due to the dense cave environment, the bats are found to operate in the approach phase most of the time, which is consistent with natural cave emergence, where they are navigating through a cluttered environment rather than engaging in open-space search. For one of the species (Rhinopoma), we also have empirical recordings of individuals flying under similar conditions (Goldshtein et al., 2025). Our model was designed to remain as simple as possible while relying on conservative assumptions that may underestimate bat performance. If, in reality, bats fine-tune their echolocation calls even earlier or more precisely during navigation than assumed, our model would still conservatively reflect their actual capabilities. See lines 500-508.

The two species modeled have different calls. In particular, the bandwidth varies by a factor of 10, meaning the species' sonars will have different spatial resolutions. Range resolution is about 10x better for PK compared to RM, but the authors appear to use the same thresholds for "correct detection" for both, which doesn't seem appropriate.

The detection process in our model is based on Saillant’s method using a filterbank, as detailed in the paper (Saillant et al., 1993; Neretti et al., 2003; Sanderson et al., 2003). This approach inherently incorporates the advantages of a wider bandwidth, meaning that the differences in range resolution between the species are already accounted for within the signal-processing framework. Thus, there is no need to explicitly adjust the model parameters for bandwidth variations, as these effects emerge from the applied method.

Also, the authors did not mention incorporating/correcting for/exploiting Doppler, which leads me to assume they did not model it.

The reviewer is correct. To maintain model simplicity, we did not incorporate the Doppler effect or its impact on echolocation. The exclusion of Doppler effects was based on the assumption that while Doppler shifts can influence frequency perception, their impact on jamming and overall navigation performance is minor within the modelled context.

The maximal Doppler shifts expected for the bats in this scenario are of ~ 1kHz. These shifts would be applied variably across signals due to the semi-random relative velocities between bats, leading to a mixed effect on frequency changes. This variability would likely result in an overall reduction in jamming rather than exacerbating it, aligning with our previous statement that our model may overestimate the severity of acoustic interference. Such Doppler shifts would result in errors of 2-4 cm in localization (i.e., 200-400 micro-seconds) (Boonman, Parsons and Jones, 2003).

We have now explicitly highlighted this in the revised version (see 548-581).

The success of the simulation may very well be due to variation in the calls of the bats, which ironically enough demonstrates the importance of a jamming avoidance response in dense flight. This explains why the performance of the simulation falls when bats are not able to distinguish their own echoes from other signals. For example, in Figure C2, there are calls that are labeled as conspecific calls and have markedly shorter durations and wider bandwidths than others. These three phases for call types used by the authors may be responsible for some (or most) of the performance of the model since the correlation between different call types is unlikely to exceed the detection threshold. But it turns out this variation in and of itself is what a jamming avoidance response may consist of. So, in essence, the authors are incorporating a jamming avoidance response into their simulation. 

We fully agree that the natural variations in call design between the phases contribute significantly to interference reduction (see our discussion in a previous paper in Mazar & Yovel, 2020). However, we emphasize that this cannot be classified as a Jamming Avoidance Response (JAR). In our model, bats respond only to the physical presence of objects and not to the acoustic environment or interference itself. There is no active or adaptive adjustment of call design to minimize jamming beyond the natural phase-dependent variations in call structure. Therefore, while variation in call types does inherently reduce interference, this effect emerges passively from the modeled behavior rather than as an intentional strategy to avoid jamming. 

The authors claim that integration over multiple pings (though I was not able to determine the specifics of this integration algorithm) reduces the masking problem. Indeed, it should: if you have two chances at detection, you've effectively increased your SNR by 3dB.  

The reviewer is correct. Indeed, integration over multiple calls improves signal-tonoise ratio (SNR), effectively increasing it by approximately 3 dB per doubling of observations. The specifics of the integration algorithm are detailed in the Methods section, where we describe how sensory information is aggregated across multiple time steps to enhance detection reliability.

They also claim - although it is almost an afterthought - that integration dramatically reduces the degradation caused by false echoes. This also makes sense: from one ping to the next, the bat's own echo delays will correlate extremely well with the bat's flight path. Echo delays due to conspecifics will jump around kind of randomly. However, the main concern is regarding the time interval and number of pings of the integration, especially in the context of the bat's flight speed. The authors say that a 1s integration interval (5-10 pings) dramatically reduces jamming probability and echo confusion. This number of pings isn't very high, and it occurs over a time interval during which the bat has moved 5-10m. This distance is large compared to the 0.4m distance-to-obstacle that triggers an evasive maneuver from the bat, so integration should produce a latency in navigation that significantly hinders the ability to avoid obstacles. Can the authors provide statistics that describe this latency, and discussion about why it doesn't seem to be a problem? 

As described in the Methods section, the bat’s collision avoidance response does not solely rely on the integration process. Instead, the model incorporates real-time echoes from the last calls, which are used independently of the integration process for immediate obstacle avoidance maneuvers. This ensures that bats can react to nearby obstacles without being hindered by the integration latency. The slower integration on the other hand is used for clustering, outlier removal and estimation wall directions to support the pathfinding process, as illustrated in Supplementary Figure 1.

Additionally, our model assumes that bats store the physical positions of echoes in an allocentric coordinate system (x-y). The integration occurs after transforming these detections from a local relative reference frame to a global spatial representation. This allows for stable environmental mapping while maintaining responsiveness to immediate changes in the bat’s surroundings.

See lines 600-616 in the revised version.

The authors are using a 2D simulation, but this very much simplifies the challenge of a 3D navigation task, and there is an explanation as to why this is appropriate. Bat densities and bat behavior are discussed per unit area when realistically it should be per unit volume. In fact, the authors reference studies to justify the densities used in the simulation, but these studies were done in a 3D world. If the authors have justification for why it is realistic to model a 3D world in a 2D simulation, I encourage them to provide references justifying this approach. 

We acknowledge that this is a simplification; however, from an echolocation perspective, a 2D framework represents a worst-case scenario in terms of bat densities and maneuverability:

• Higher Effective Density: A 2D model forces all bats into a single plane rather than distributing them through a 3D volume, increasing the likelihood of overlap in calls and echoes and making jamming more severe. As described in the text: the average distance to the nearest bat in our simulation is 0.27m (with 100 bats), whereas reported distances in very dense colonies are 0.5m (Fujioka et al., 2021), as observed in Myotis grisescens (Sabol and Hudson, 1995) and Tadarida brasiliensis (Theriault et al., no date; Betke et al., 2008; Gillam et al., 2010)

• Reduced Maneuverability: In 3D space, bats can use vertical movement to avoid obstacles and conspecifics. A 2D constraint eliminates this degree of freedom, increasing collision risk and limiting escape options.

Thus, our 2D model provides a conservative difficult test case, ensuring that our findings are valid under conditions where jamming and collision risks are maximized. Additionally, the 2D framework is computationally efficient, allowing us to perform multiple simulation runs to explore a broad parameter space and systematically test the impact of different variables.

To address the reviewer’s concern, we have clarified this justification in the revised text and will provide supporting references where applicable (see Methods lines 450455).

The focus on "masking" (which appears to be just in-band noise), especially relative to the problem of misassigned echoes, is concerning. If the bat calls are all the same waveform (downsweep linear FM of some duration, I assume - it's not clear from the text), false echoes would be a major problem. Masking, as the authors define it, just reduces SNR. This reduction is something like sqrt(N), where N is the number of conspecifics whose echoes are audible to the bat, so this allows the detection threshold to be set lower, increasing the probability that a bat's echo will exceed a detection threshold. False echoes present a very different problem. They do not reduce SNR per se, but rather they cause spurious threshold excursions (N of them!) that the bat cannot help but interpret as obstacle detection. I would argue that in dense groups the mis-assignment problem is much more important than the SNR problem. 

There is substantial literature supporting the assumption that bats can recognize their own echoes and distinguish them from conspecific signals (Schnitzler, Bioscience and 2001, no date; Kazial, Burnett and Masters, 2001; Burnett and Masters, 2002; Kazial, Kenny and Burnett, 2008; Chili, Xian and Moss, 2009; Yovel et al., 2009; Beetz and Hechavarría, 2022)). However, we acknowledge that false echoes may present a major challenge in dense groups. To address this, we explicitly tested the impact of the self-echo identification assumption in our study see Results Figure 1: The impact of confusion on performance, and lines 399-404 in the Discussion.

Furthermore, we examined a full confusion scenario, where all reflected echoes from conspecifics were misinterpreted as obstacle reflections (i.e., 100% confusion). Our results show that this significantly degrades navigation performance, supporting the argument that echo misassignment is a critical issue. However, we also explored a simple mitigation strategy based on temporal integration with outlier rejection, which provided some improvement in performance. This suggests that real bats may possess additional mechanisms to enhance self-echo identification and reduce false detections. See lines 411-420 in the manuscript for further discussion. 

We actually used logarithmically frequency modulated (FM) chirps, generated using the MATLAB built-in function chirp(t, f0, t1, f1, 'logarithmic'). This method aligns with the nonlinear FM characteristics of Pipistrellus kuhlii (PK) and Rhinopoma microphyllum (RM) and provides a realistic approximation of their echolocation signals. We acknowledge that this was not sufficiently emphasized in the original text, and we have now explicitly highlighted this in the revised version to ensure clarity (see Lines 509-512 in Methods).

The criteria set for flight behavior (lines 393-406) are not justified with any empirical evidence of the flight behavior of wild bats in collective flight. How did the authors determine the avoidance distances? Also, what is the justification for the time limit of 15 seconds to emerge from the opening? Instead of an exit probability, why not instead use a time criterion, similar to "How long does it take X% of bats to exit?"  :

While we acknowledge that wild bats may employ more complex behaviors for collision avoidance, we chose to implement a simplified decision-making rule in our model to maintain computational tractability.

The avoidance distances (1.5 m from walls and 0.4 m from other bats) were selected as internal parameters to support stable and realistic flight trajectories while maintaining a reasonable collision rate. These values reflect a trade-off between maneuverability and behavioral coherence under crowding. To address this point, we added a sensitivity analysis to the revised manuscript. Specifically, we tested the effect of varying the conspecific avoidance distance from 0.2 to 1.6 meters at bat densities of 2 to 40 bats/3m². The only statistically significant impact was at the highest density (40 bats/3m²), where exit probability increased slightly from 82% to 88% (p = 0.024, t = 2.25, DF = 958). No significant changes were observed in exit time, collision rate, or jamming probability across other densities or conditions (GLM, see revised Methods). These results suggest that the selected avoidance distances are robust and not a major driver of model performance, see lines 469-47.

The 15-second exit limit was determined as described in the text (Lines 489-491): “A 15-second window was chosen because it is approximately twice the average exit time for 40 bats and allows for a second corrective maneuver if needed.” In other words, it allowed each bat to circle the ‘cave’ twice to exit even in the most crowded environment. This threshold was set to keep simulation time reasonable while allowing sufficient time for most bats to exit successfully.

We acknowledge that the alternative approach suggested by the reviewer— measuring the time taken for a certain percentage of bats to exit—is also valid. However, in our model, some outlier bats fail to exit and continue flying for many minutes, such simulations would lead to excessive simulation times making it difficult to generate repetitions and not teaching us much – they usually resulted from the bat slightly missing the opening (see video S1. Our chosen approach ensures practical runtime constraints while still capturing relevant performance metrics.

What is the empirical justification for the 1-10 calls used for integration?  

The "average exit time for 40 bats" is also confusing and not well explained. Was this determined empirically? From the simulation? If the latter, what are the conditions?

Does it include masking, no masking, or which species? 

Previous studies have demonstrated that bats integrate acoustic information received sequentially over several echolocation calls (2-15), effectively constructing an auditory scene in complex environments (Ulanovsky and Moss, 2008; Chili, Xian and Moss, 2009; Moss and Surlykke, 2010; Yovel and Ulanovsky, 2017; Salles, Diebold and Moss, 2020). Additionally, bats are known to produce echolocation sound groups when spatiotemporal localization demands are high (Kothari et al., 2014). Studies have documented call sequences ranging from 2 to 15 grouped calls (Moss and Surlykke, 2010), and it has been hypothesized that grouping facilitates echo segregation.

We did not use a single integration window - we tested integration sizes between 1 and 10 calls and presented the results in Figure 3A. This range was chosen based on prior empirical findings and to explore how different levels of temporal aggregation impact navigation performance. Indeed, the results showed that the performance levels between 5-10 calls integration window (Figure 3A)

Regarding the average exit time for 40 bats, this value was determined from our simulations, where it represents the mean time for successful exits under standard conditions with masking. We have revised the text to clarify these details see, lines 489-491.

Reviewer #1 (Recommendations for the authors):

(1) Data Availability:

As it stands now, this reviewer cannot vouch for the uploaded code as it wasn't accessible according to F.A.I.R principles. The link to the code/data points to a private company's file-hosting account that requires logging in or account creation to see its contents, and thus cannot be accessed.

This reviewer urges the authors to consider uploading the code onto an academic data repository from the many on offer (e.g. Dryad, Zenodo, OSF). Some repositories offer an option to share a private link (e.g. Zenodo) to the folder that can then be shared only with reviewers so it is not completely public.

This is a computational paper, and the credibility of the results is based on the code used to generate them.

The code is available at GitHub as required:

https://github.com/omermazar/Colony-Exit-Bat-Simulation

(2) Abstract:

Line 22: 'To explore whether..' - replace 'whether' with 'how'?

The sentence was rephrased as suggested by the reviewer.

(2) Main text:

Line 43: '...which may share...' - correct to '...which share...', as elegantly framed in the authors' previous work - jamming avoidance is unavoidable because all FM bats of a species still share >90% of spectral bandwidth despite a few kHz shift here and there.

The sentence was rephrased as suggested by the reviewer.

Line 49: The authors may wish to additionally cite the work of Fawcett et al. 2015 (J. Comp. Phys A & Biology Open)

Thank you for the suggestion. We have included a citation to the work of Fawcett et al. (2015) in the revised manuscript.

Line 61: This statement does not match the recent state of the literature. While the previous models may have assumed that all neighbours can be detected, there are models that specifically study the role of limited interaction arising from the potential inability to track all neighbours, and the effect of responding to only one/few neighbours at a time e.g. Bode et al. 2011 R. Soc. Interface, Jhawar et al. 2020 Nature Physics.

We have added citations to the important studies suggested by the reviewer, as detailed in the Public Review above.

Line 89: '..took all interference signals into account...' - what is meant by 'interference signals' - are the authors referring to reflections, unclear.

We have revised the sentence and detailed the acoustic signals involved in the process: self-generated echoes, calls from conspecifics, and echoes from cave walls and other bats evoked by those calls, see lines 99-106.

Figure 1A: The colour scheme with overlapping points makes the figure very hard to understand what is happening. The legend has colours from subfigures B-D, adding to the confusion.

What does the yellow colour represent? This is not clear. Also, in general, the color schemes in the simulation trajectories and the legend are not the same, creating some amount of confusion for the reader. It would be good to make the colour schemes consistent and visually separable (e.g. consp. call direct is very similar to consp. echo from consp. call), and perhaps also if possible add a higher resolution simulation visualisation. Maybe it is best to separate out the colour legends for each sub-figure.

The updated figure now includes clearer, more visually separable colors, and consistent color coding across all sub-panels. The yellow trajectory representing the focal bat’s flight path is now explicitly labeled, and we adjusted the color mapping of acoustic signals (e.g., conspecific calls vs. echoes) to improve distinction. We also revised the figure caption accordingly and ensured that the legend is aligned with the updated visuals. These modifications aim to enhance interpretability and reduce ambiguity for the reader.

Figure C3: What is 'FB Channel', this is not explained in the legend.

FB Channel’ stands for ‘Filter Bank Channel’. This clarification has been added to the caption of Figure 1. 

Figure 3: Visually noticing that the colour legend is placed only on sub-figure A is tricky and readers may be left searching for the colour legend. Maybe lay out the legend horizontally on top of the entire figure, so it stands out?

We have adjusted the placement of the color legend in Figure 3 to improve visibility and consistency.

Line 141: '..the probability of exiting..' - how is this probability calculated - not clear.

We have clarified in the revised text that the probability of exiting the cave within 15 seconds is defined as the number of bats that exited the cave within that time divided by the total number of bats in each scenario, see lines 159160.

Line 142: What are the sample sizes here - i.e. how many simulation replicates were performed?

We have clarified the number of repetitions in each scenario the revised text, as detailed in the Public Review above.

Line 151: 'The jamming probability,...number of jammed echoes divided by the total number of reflected echoes' - it seems like these are referring to 'own' echoes or first-order reflections, it is important to clarify this.

The reviewer is right. We have clarified it in the revised text, see lines 173175.

Line 153: '..with a maximum difference of ...' - how is this difference calculated? What two quantities are being compared - not clear.

We have revised the text to clarify that the 14.3% value reflects the maximum difference in jamming probability between the RM and PK models, which occurred at a density of 10 bats. The values at each density are shown in Figure 2D, see lines 175-177.

Line 221: '..temporal aggregation helps..' - I'm assuming the authors meant temporal integration? However, I would caution against using the exact term 'temporal integration' as it is used in the field of audition to mean something different. Perhaps something like 'sensory integration' , or 'multi-call integration'

To avoid ambiguity and better reflect the process modeled in our work, we have replaced the term "temporal aggregation" with "multi-call integration" throughout the revised manuscript. This term more accurately conveys the idea of combining information from multiple echolocation calls without conflicting with existing terminology.

(4) Discussion

Lines 302: 'Our model suggests...increasing the call-rate..' - not clear where this is explicitly tested or referred to in this manuscript. Can't see what was done to measure/quantify the effect of this variable in the Methods or anywhere else.

We have rephrased this paragraph as detailed in the Public Review above, see lines 346-349.

Line 319: 'spatial interference' - unclear what this means. This reviewer would strongly caution against creating new terms unless there is an absolute need for it. What is meant by 'interference' in this paper is hard to assess given that the word seems to be used as a synonym for jamming and also for actual physical wave-based interference.

We have rephrased this paragraph as detailed in the Public Review above, see line 119-120, 366-367.

Line 323: '..no benefit beyond a certain level...' - also not clear where this is explicitly tested. It seems like there was a set of simulations run for a variety of parameters but this is not written anywhere explicitly. What type of parameter search was done, was it all possible parameter combinations - or only a subset? This is not clear.

We have rephrased this paragraph as detailed in the Public Review above, see lines 372-375.

Line 324: '..ca. 110 dB-SPL.' - what reference distance?

All call levels were simulated and reported in dB-SPL, referenced at 0.1 meters from the emitting bat. We have clarified it in the revised text in the relevant contexts and specifically in line 529.

(5) Methods

Line 389 : '...over a 2 x 1.5 m2 area..' It took a while to understand this statement and put it in context. Since there is no previous description of the entire L-arena, the reviewer took it to mean the simulations happened over the space of a 2 x 1.5 m2 area. Include a top-down description of the simulation's spatial setup and rephrase this sentence.

To address the confusion, we revised the text to clarify that the full simulation environment represents a corridor-shaped cave measuring 14.5 × 2.5 meters, with a right-angle turn located 5.5 meters before the exit, as shown in Figure 1A. The 2 × 1.5 m area refers specifically to the small zone at the far end of the cave where bats begin their flight. The revised description now includes a clearer spatial overview to prevent ambiguity, see lines 456-460.

Line 398: Replace 'High proximity' with 'Close proximity'

Replaced.

Line 427: 'uniform target strength of -23 dB' - at what distance is this target strength defined? Given the reference distance can vary by echolocation convention (0.1 or 1 m), one can't assess if this is a reasonable value or not.

The reference distance for the reported target strength is 1 meter, in line with standard acoustic conventions. We have revised the text to clarify this explicitly (line 531).

Also, independent of the reference distance, particularly with reference to bats, the target strength is geometry-dependent, based on whether the wings are open or not. Using the entire wingspan of a bat to parametrise the target strength is an overestimate of the available reflective area. The effective reflective area is likely to be somewhere closer to the surface area of the body and a fraction of the wingspan together. This is important to note and/or mention explicitly since the value is not experimentally parametrised.

For comparison, experimentally based measurements used in Goetze et al. 2016 are -40 dB (presumably at 1 m since the source level is also defined at 1 m?), and Beleyur & Goerlitz 2019 show a range between -43 to -34 dB at 1 m.

We agree with the reviewer that target strength in bats is strongly influenced by their geometry, particularly wing posture during flight. In our model, we simplified this aspect by using a constant target strength, as the detailed temporal variation in body and wing geometry is pseudo-random and not explicitly modeled. We acknowledge that this is a simplification, and have now stated this limitation clearly in the revised manuscript. We chose a fixed value of –23 dB at 1 meter to reflect a plausible mid-range estimate, informed by anatomical data and consistent with values reported for similarly sized species (Beleyur and Goerlitz, 2019). To support this, we directly measured the target strength of a 3D-printed RM bat model, obtaining –32dB. 

Moreover, a sensitivity analysis across a wide range (–49 to –23 dB) confirmed that performance metrics remain largely stable, indicating that our conclusions are not sensitive to this parameter, and suggesting that our results hold for different-sized bats. See lines 384-390, 533-538, and Supplementary Figures 3 and 4 in the revised article. 

Line 434: 'To model the bat's cochlea...'. Bats have two cochleas. This model only describes one, while the agents are also endowed with the ability to detect sound direction - which requires two ears/cochleas.... There is missing information about the steps in between that needs to be provided.

We appreciate the reviewer’s observation. Indeed, our model is monaural, and simulates detection using a single cochlear-like filter bank receiver. We have clarified this in the revised text to avoid confusion. This paragraph specifically describes the detection stage of the auditory processing pipeline. The localization process, which builds on detection and includes directional estimation, is described in the following paragraph (see line 583 onward), as discussed in the next comment and response.

Line 457: 'After detection, the bat estimates the range and Direction of Arrival...' This paragraph describes the overall idea, but not the implementation. What were the inputs and outputs for the range and DOA calculation performed by the agent? Or was this information 'fed' in by the simulation framework? If there was no explicit DOA step that the agent performed, but it was assumed that agents can detect DOA, then this needs to be stated.

In the current simulation, the Direction of Arrival (DOA) was not modeled via an explicit binaural processing mechanism. Instead, based on experimental studies (Simmons et al., 1983; Popper and Fay, 1995).  we assumed that bats can estimate the direction of an echo with an angular error that depends on the signal-to-noise ratio (SNR). Accordingly, the inputs to the DOA estimation were the peak level of the desired echo, noise level, and the level of acoustic interference. The output was an estimated direction of arrival that included a random angular error, drawn from a normal distribution whose standard deviation varied with the SNR. We have revised the relevant paragraph (Lines 583-592) to clarify this implementation.

Line 464: 'To evaluate the impact of the assumption...' - the 'self' and 'non-self' echoes can be distinguished perhaps using pragmatic time-delay cues, but also using spectro-temporal differences in individual calls/echoes. Do the agents have individual call structures, or do all the agents have the same call 'shape'? The echolocation parameters for the two modelled species are given, but whether there is call parameter variation implemented in the agents is not mentioned.

In our relatively simple model, all individuals emit the same type of chirp call, with parameters adapted only based on the distance to the nearest detected object. However, individual variation is introduced by assigning each bat a terminal frequency drawn from a normal distribution with a standard deviation of 1 kHz, as described in the revised version -lines 519-520. This small variation is not used explicitly as a spectro-temporal cue for echo discrimination.

In our model, all spectro-temporal variations—whether due to call structure or variations resulting from overlapping echoes from nearby reflectors—are processed through the filter bank, which compares the received echoes to the transmitted call during the detection stage. As such, the detection process itself can act as a discriminative filter, to some extent, based on similarity to the emitted call.

We acknowledge that real bats likely rely on a variety of spectro-temporal features for distinguishing self from non-self-echoes—such as call duration, received level, multi-harmonic structure, or amplitude modulation. In our simulation, we focus on comparing two limiting conditions: full recognition of self-generated echoes versus full confusion. Implementing a more nuanced self-recognition mechanism based on temporal or spectral cues would be a valuable extension for future work.

(6) References

Reference 22: Formatting error - and extra '4' in the reference.

The error has been fixed.

(7) Thoughts/comments

Even without 'recogntion' of walls & conspecifics, bats may be able to avoid obstacles - this is a neat result. Also, using their framework the authors show that successful 'blind' object-agnostic obstacle avoidance can occur only when supported by some sort of memory. In some sense, this is a nice intermediate step showing the role of memory in bat navigation. We know that bats have good long-term and long-spatial scale memory, and here the authors show that short-term spatial memory is important in situations where immediate sensory information is unreliable or unavailable.

We appreciate the reviewer’s thoughtful summary. Indeed, one of the main takeaways of our study is that successful obstacle avoidance can occur even without explicit recognition of walls or conspecifics—provided that a clustered multi-call integration is in place. Our model shows that when immediate sensory information is unreliable, integrating detections over time becomes essential for effective navigation. This supports the broader view that memory, even on short timescales, plays an important role in bat behavior.

(8) Reporting GLM results

The p-value, t-statistic, and degrees of freedom are reported consistently across multiple GLM results. However, the most important part which is the effect size is not consistently reported - and this needs to be included in all results, and even in the table. The effect size provides an indicator of the parameter's magnitude, and thus scientific context.

We agree that the effect size provides essential scientific context. In fact, we already include the effect size explicitly in Table 1, as shown in the “Effect Size” column for each tested parameter. These values describe the magnitude of each parameter’s effect on exit probability, jamming probability, and collision rate. In the main text, effect sizes are presented as concrete changes in performance metrics (e.g., “exit probability increased from 20% to 87%,” or “with a decrease of 3.5%±8% to 5.5%±5% (mean ± s.e.)”), which we believe improves interpretability and scientific relevance.  

To further clarify this in the main text, we have reviewed the reported results and ensured that effect sizes are mentioned more consistently wherever GLM outcomes are discussed. Additionally, we have added a brief note in the table caption to emphasize that effect sizes are provided for all tested parameters.

The 'tStat' appears multiple times and seems to be the output of the MATLAB GLM function. This acronym is specific to the MATLAB implementation and needs to be replaced with a conventionally used acronym such as 't', or the full form 't-statistic' too. This step is to keep the results independent of the programming language used.

We have replaced all instances of tStat with the more conventional term ‘t’ throughout the manuscript to maintain consistency with standard reporting practices.

Reviewer #2 (Recommendations for the authors):

In addition to my public review, I had a few minor points that the authors may want to consider when revising their paper.

(1) Figures 2, 3, and 4 may benefit from using different marker styles, in addition to different colors, to show the different cases.

Thank you for the suggestion. In Figures 2–4, the markers represent means with standard error bars. To maintain clarity and consistency across all conditions, we have chosen to keep a standardized marker style – and we clarify this in the legend. We found that varying only the colors is sufficient for distinguishing between conditions without introducing visual clutter.

(2) The text "PK" in the inset for Figure 2A is very difficult to read. I would suggest using grey as with "RM" in the other inset.

We have updated the insert in Figure 2A to improve legibility.

(3) Are the error bars in Figure 3 very small? I wasn't able to see them. If that is the case, the authors may want to mention this in the caption.

You are correct—the error bars are present in all plots but appear very small due to the large number of simulation repetitions and low variability. We have revised the caption to explicitly mention this.

(4) The species name of PK is spelled inconsistently (kuhli, khulli, and kuhlii).

We have corrected the species name throughout the manuscript.

(5) Table 1 is a great condensation of all the results, but the time to exit is missing. It may be helpful if summary statistics on that were here as well.

We have added time-to-exit to the effect size column in Table 1, alongside the other performance metrics, to provide a more complete summary of the simulation results.

(6) I may have missed it, but why are there two values for the exit probability when nominal flight speed is varied?

The exit probability was not monotonic with flight speed, but rather showed a parabolic trend with a clear optimum. Therefore, we reported two values representing the effect before and after the peak. We have clarified this in the revised table and updated the caption accordingly.

(7) Table 2 has an extra header after the page break on page 18.

The extra header in Table 2 after the page break has been removed in the revised manuscript.

(8) The G functions have 2 arguments in their definitions and Equation 1, but only one argument in Equations 2 and 3. I wasn't able to see why.

Thank you for pointing this out. You are correct—this was a typographical error. We have corrected the argument notation in Equations 2 and 3 and explicitly included the frequency dependence of the gain (G) functions in both equations.

(9) D_txrx was not defined but it was used in Equation 2.

The variable D_txrx is defined in the equation notation section as: D_₍ₜₓr_ₓ – the distance [m] between the transmitting conspecific and the receiving focal bat, from the transmitter’s perspective. We have now ensured that this definition is clearly linked to Equation 2 in the revised text. Moreover, we have added a supplementary figure that illustrates the geometric configuration defined by the equations to further support clarity, as described in the Public Review above.

(10) It was hard for me to understand what was meant by phi_rx and phi_tx. These were described as angles between the rx or tx bats and the target, but I couldn't tell what the point defining the angle was. Perhaps a diagram would help, or more precise definitions.

We have revised the caption to provide clearer and more precise definitions Additionally, we have included a geometric diagram as a supplementary figure, as noted in the Public Review above, to visually clarify the spatial relationships and angle definitions used in the equations, see lines 498-499.

(11) Was the hearing threshold the same for both species?

Yes. We have clarified it in the revised version.

(12) Collision avoidance is described as turning to the "opposite direction" in the supplemental figure explaining the model. Is this 90 degrees or 180 degrees? If 90 degrees, how do these turns decide between right and left?

In our model, the bat does not perform a fixed 90° or 180° turn. Instead, the avoidance behavior is implemented by setting the maximum angular velocity in the direction opposite to the detected echo. For example, if the obstacle or conspecific is detected on the bat’s right side, the bat begins turning left, and vice versa.

This turning direction is re-evaluated at each decision step, which occurs after every echolocation pulse. The bat continues turning in the same direction if the obstacle remains in front, otherwise it resumes regular pathfinding. We have clarified this behavior in the updated figure caption and model description, see lines 478-493.

Reviewer #3 (Recommendations for the authors):

(1) Lines 27-31: These sentences mischaracterize the results. This claim appears to equate "the model works" with "this is what bats actually do." Also, the model does not indicate that bats' echolocation strategies are robust enough to mitigate the effects of jamming - this is self-evident from the fact that bats navigate successfully via echolocation in dense groups.

Thank you for the comment. Our aim was not to claim that the model confirms actual bat behavior, but rather to demonstrate that simple and biologically plausible strategies—such as signal redundancy and basic pathfinding—are sufficient to explain how bats might cope with acoustic interference in dense settings. We have revised the wording to better reflect this goal and to avoid overinterpreting the model's implications.

See abstract in the revised version.  

(2) Line 37: This number underestimates the number of bats that form some of the largest aggregations of individuals worldwide - the free-tailed bats can form aggregations exceeding several million bats.

We have revised the text to reflect that some bat species, such as free-tailed bats, are known to form colonies of several million individuals, which exceed the typical range. The updated sentence accounts for these extreme cases, see lines 36-37.

(3) The flight densities explained in the introduction and chosen references are not representative of the literature - without providing additional justification for the chosen species, it can be interpreted that the selection of the species for the simulation is somewhat arbitrary. If the goal is to model dense emergence flight, why not use a species that has been studied in terms of acoustic and flight behavior during dense emergence flights---such as Tadarida brasiliensis?

Our goal was to develop a general model applicable to a broad class of FMecholocating bat species. The two species we selected—Pipistrellus kuhlii (PK) and Rhinopoma microphyllum (RM)—span a wide range of signal characteristics: from wideband (PK) to narrowband (RM), providing a representative contrast in call structure. 

Although we did not include Tadarida brasiliensis (TB) specifically, its echolocation calls are acoustically similar to RM in terminal frequency and fall between PK and RM in bandwidth. Therefore, we believe our findings are likely to generalize to TB and other FM-bats.

Moreover, as noted in a previous response, the average inter-bat distance in our highest-density simulations (0.27 m) is still smaller than those reported for Tadarida brasiliensis during dense emergences—further supporting the relevance of our model to such scenarios.

To support broader applicability, we also provide a supplementary graphical user interface (GUI) that allows users to modify key echolocation parameters and explore their impact on behavior—making the framework adaptable to additional species, including TB.

(4) Line 78: It is not clear how (or even if) the simulated bats estimate the direction of obstacles. The explanation given in lines 457-463 is quite confusing. What is the acoustic/neurological mechanism that enables this direction estimation? If there is some mechanism (such as binaural processing), how does this extrapolate to 3D?

This comment echoes a similar concern raised by a previous reviewer. As explained earlier, in the current simulation, the Direction of Arrival (DOA) was not modeled via an explicit binaural processing mechanism. The complete  is detailed in  to Reviewer #1, Line 457. This implementation is now clarified in the revised text, and a detailed description of the localization process is also provided in the Methods section (lines 583-592).

(5) The authors propose they are modeling the dynamic echolocation of bats in the simulation (line 79), but it appears (whether this is due to a lack of information in the manuscript or true lack in the simulation) that the authors only modeled a flight response. How did the authors account for bats dynamically changing their echolocation? This is unclear and from what I can tell may just mean that the bats can switch between foraging phase call types depending on the distance to a detected obstacle. Can the authors elaborate more on this?

The echolocation behavior of the bats—including dynamic call adjustments— was implemented in the simulation and is described in detail in the Methods section (lines 498-520 and Table 2). To avoid redundancy, the Results chapter originally referred to this section, but we have now added a brief explanation in the Results to clarify that the bats’ call parameters (IPI, duration, and frequency range) adapt based on the distance to detected objects, following empirically documented echolocation phases ("search," "approach," "buzz"). These dynamics are consistent with established bat behavior during navigation in cluttered environments such as caves.

(6) Figure 1 C3: "Detection threshold": what is this and how was it derived?

The caption also mentions yellow arrows, but they are absent from the figure. C4: Each threshold excursion is marked with an asterisk, but there are many more excursions than asterisks. Why are only some marked? Unclear.

C3: The detection threshold is determined dynamically. It is set to the greater of either 7 dB above the noise level (0 dB-SPL)(Kick, 1982; Saillant et al., 1993; Sanderson et al., 2003; Boonman et al., 2013) or the maximal received level minus 70 dB, effectively applying a dynamic range of 70 dB. This clarification has been added to the Methods section. The yellow arrow has been added.

C4: Thank you for this important observation. Only peaks marked with asterisks represent successful detections—those that were identified in both the interference-free and full detection conditions, as explained in the Methods. Other visible peaks result from masking signals or overlapping echoes from nearby reflectors, but they do not meet the detection criteria. To keep the figure caption concise, we have elaborated on this process more clearly in the revised Methods section. We added this information to the legend

(7) Figure 2: A line indicating RM, No Masking is absent

Thank you for pointing this out. The missing line for RM, No Masking has now been added in the revised version of Figure 2.

(8) Line 121: "reflected off conspecifics". Does this mean echoes due to conspecifics?

The phrase "reflected off conspecifics" refers to echoes originating from the bat’s own call and reflected off the bodies of nearby conspecifics. We have clarified the wording in the revised text to avoid confusion

(9) Line 125: Why are low-frequency channels stimulated by higher frequencies? This needs further clarification.

The cochlear filter bank in our model is implemented using gammatone filters, each modeled as an 8th-order Butterworth filter. Due to the non-ideal filter response and relatively broad bandwidths—especially in the lower-frequency channels—strong energy from the beginning of the downward FM chirp (at higher frequencies) can still produce residual activation in lower-frequency channels. While these stimulations are usually below the detection threshold, they may still be visible as early sub-threshold responses. Given the technical nature of this explanation (a property of the filter implementation) and it does not influence the detection outcomes, we have chosen not to elaborate on it in the figure caption or Methods.

(10) Lines 146-150: This is an interesting finding. Is there a theoretical justification for it?

This outcome arises directly from the simulation results. As noted in the Discussion (lines 359-365), although Pipistrellus kuhlii (PK) shows a modest advantage in jamming resistance due to its broader bandwidth, the redundancy in sensory information across calls—enabled by frequent echolocation—appears to compensate for these signal differences. As a result, the small variations in echo quality between species do not translate into significant differences in performance. We speculate that if the difference in jamming probability had been larger, performance disparities would likely have emerged.

(11) Line 151: The authors define a jammed echo as an echo entirely missed due to masking. Is this appropriate? Doesn't echo mis-assignment also constitute jamming?

We agree that echo mis-assignment can also degrade performance; however, in our model, we distinguish between two outcomes: (1) complete masking (echo not detected), and (2) detection with a localization error. As explained in the Methods (lines 500–507), we run the detection analysis twice—once with only desired echoes (“interference-free detection”) and once including masking signals (“full detection”). If a previously detected echo is no longer detected, it is classified as a jammed echo. If the echo is still detected but the delay shifts by more than 100 µs compared to the interference-free condition, it is also considered jammed. If the delay shift is smaller, it is treated as a detection with localization error rather than full jamming. We have clarified this distinction in the revised Methods section.

(12) Figure 2-E: Detection probability statistics are of limited usefulness without accompanying false alarm rate (FAR) statistics. Do the authors have FAR numbers?

We understand FAR to refer to instances where masking signals or other acoustic phenomena are mistakenly interpreted as real echoes from physical objects. As explained in the manuscript, we implemented two model versions: one without confusion, and one with full confusion.

Figure 2E reports detection performance under the non-confusion model, in which only echoes from actual physical reflectors are used, and no false detections occur—hence, the false alarm rate is effectively zero in this condition. In the full-confusion model, all detected echoes—including those originating from masking signals or conspecific calls—are treated as valid detections, which may include false alarms. However, we did not explicitly quantify the false alarm rate as a separate metric in this simulation.

We agree that tracking FAR could be informative and will consider incorporating it into future versions of the model.

(13) Line 161: RM bats suffered from a significantly higher probability of the "desired conspecific's echoes" being jammed. What does "desired conspecific's echoes" mean? This is unclear.

The term “desired conspecific's echoes” refers to echoes originating from the bat’s own call, reflected off nearby conspecifics, which are treated as relevant reflectors for collision avoidance. We have revised the wording in the text for clarity.

(14) Line 188: Why didn't the size of the integration window affect jamming probability? I couldn't find this explained in the discussion.

The jamming probability in our analysis is computed at the individual-echo level, prior to any temporal integration. Since the integration window is applied after the detection step, it does not influence whether a specific echo is masked (i.e., jammed) or not. Therefore, as expected, we did not observe a significant effect of integration window size on jamming probability.

(15) Line 217-218: Why do the authors think this would be?

Thank you for the thoughtful question. We agree that, in theory, increasing call intensity should raise the levels of both desired echoes and masking signals proportionally. However, in our model, the environmental noise floor and detection threshold remain constant, meaning that higher call intensities increase the signal-to-noise ratio (SNR) more effectively for weaker echoes, especially those at longer distances or with low reflectivity. This could lead to a higher likelihood of those echoes crossing the detection threshold, resulting in a small but measurable reduction in jamming probability.

Additionally, the non-linear behavior of the filter-bank receiver—including such as thresholding at multiple stages—can introduce asymmetries in how increased signal levels affect the detection of target versus masking signals.

That said, the effect size was small, and the improvement in jamming probability did not translate into any significant gain in behavioral performance (e.g., exit probability or collision rate), as shown in Figure 3C.

(16) Line 233: I'm not sure I understand how a slightly improved aggregation model that clustered detected reflectors over one-second periods is different. Doesn't this just lead to on average more calls integrated into memory?

While increasing the memory duration does lead to more detections being available, the enhanced aggregation model (we now refer to as multi-call clustering) differs fundamentally from the simpler one. As detailed in the Methods, it includes additional processing steps: clustering spatially close detections, removing outliers, and estimating wall directions based on the spatial structure of clustered echoes. In contrast, the simpler model treats each detection as an isolated point without estimating obstacle orientation. These additional steps allow for more robust environmental interpretation and significantly improve performance under high-confusion conditions. We have clarified it in revised text (lines 606-616) and added a Supplementary Figure 2B.

(17) Table 1: What about conspecific target strength?

We have now added the conspecific target strength as a tested parameter in Table 1, along with its tested range, default value, and measured effect sizes. A detailed sensitivity analysis is also presented in Supplementary Figure 4, demonstrating that variations in conspecific target strength had relatively minor effects on performance metrics.  

(18) Figure 3-A: The x-axis is the number of calls in the integration window. But the leftmost sample on each curve is at 0 calls. Shouldn't this be 1?

“0 calls” refers to the case where only the most recent call is used for pathfinding—without integrating any information from prior calls. The x-axis reflects the number of previous calls stored in memory, so a value of 0 still includes the current call. We’ve clarified this terminology in the figure caption.

(19) Lines 282-283: This statement needs to be clarified that it is with the constraints of using a 2D simulation with at most 33 bats/m^2. It also should be clarified that it is assumed the bat can reliably distinguish between its own echoes and conspecific echoes, which is a very important caveat.

We have revised the text to clarify that the results are based on a 2D simulation with a maximum tested density of 33 bats/m². We also now explicitly state that the model assumes bats can distinguish between their own echoes and those generated by conspecifics—an assumption we recognize as a simplification. These clarifications help place the results within the scope and constraints of the simulation. Moreover, as described in the text (and noted in previous response): the average distance to the nearest bat in our simulation is 0.27m (with 100 bats), whereas reported distances in very dense colonies are 0.5m

(20) Line 294: What is this sentence referring to?

The sentence refers to the finding that, even under high bat densities, a substantial portion of the echoes—particularly those reflected from nearby obstacles (e.g., 1 m away)—were jammed due to masking. Nevertheless, the bats in the simulation were still able to navigate successfully using partial sensory input. We have clarified the sentence in the revised text to make this point more explicit, see line 333-336.

(21) Line 302: Was jamming less likely when IPI was higher or lower? I could not find this demonstrated anywhere in the manuscript.

We agree that the original text was not sufficiently clear on this point. While we did not explicitly test fixed IPI values as a parameter, the model does simulate the natural behavior of decreasing IPI as bats approach obstacles. This behavior is supported by empirical observations and is incorporated into the echolocation dynamics of the simulation. We have clarified this point in the revised text (see Lines 346-351) and explained that while lower IPI introduces more acoustic overlap, it also increases redundancy and improves detection through temporal integration.

(22) Lines 313-314: This is an interesting assumption, but it is not evident that is substantiated by the references.

The claim is based on well-established principles in signal processing and bioacoustics. Wideband signals—such as those emitted by PK bats— distribute their energy over a broader frequency range, which makes them inherently more resistant to narrowband interference and masking. This concept is commonly applied in both biological and artificial sonar systems and is supported by empirical studies in bats and theory in acoustic sensing.

For example, Beleyur & Goerlitz (2019) demonstrate that broader bandwidth calls improve detection in cluttered and jamming-prone environments. Similarly, Ulanovsky et al. (2004) and Schnitzler & Kalko (200) discuss how FM bats' wideband calls enhance temporal and spatial resolution, helping to reduce the impact of overlapping signals from conspecifics. These findings align with communication theory where spread-spectrum techniques improve robustness in noisy environments.

We agree with the reviewer that this is an important point and we have updated the manuscript to clarify this rationale and cite the relevant literature accordingly – lines 631-363,

(23) Lines 318-319: What is the justification for "probably"? Isn't this just a supposition?

We agree with the reviewer’s point and have rephrased the sentence

(24) Line 320: How does this 63% performance match the sentence in line 295?

The sentence in Line 295 refers to the overall ability of the bats to navigate successfully despite high jamming levels, highlighting the robustness of the strategy under challenging conditions. The figure in Line 320 (63%) quantifies this performance under the most extreme simulated scenario (100 bats / 3 m²), where both spatial and acoustic interferences are maximal. We have rephrased the text in the revised version (lines 324-327).

(25) Lines 341-345: It seems like this is more likely to be the main takeaway of the paper.

As noted in the Public Review above, there is substantial literature supporting the assumption that bats can recognize their own echoes and distinguish them from those of conspecifics (e.g., Schnitzler, Bioscience, 2001; Kazial et al., 2001, 2008; Burnett & Masters, 2002; Chiu et al., 2009; Yovel et al., 2009; Beetz & Hechavarría, 2022). Therefore, we consider our assumption of selfrecognition to be well-supported, at least under typical conditions. That said, we agree that the impact of echo confusion on performance is significant and highlights a critical challenge in dense environments.

To our knowledge, this is the first computational model to explicitly simulate both self-recognition and full echo confusion under high-density conditions. We believe that the combination of modeled constraints and the demonstrated robustness of simple sensorimotor strategies, even under worst-case assumptions, is what makes this contribution both novel and meaningful.

(26) Lines 349-350: What is the aggregation model? What is meant by "integration"?

We have revised the text to clarify that the “aggregation model” refers to a multi-call clustering process that includes clustering of detections, removal of outliers, and estimation of wall orientation, as described in detail in the revised Methods and Results sections.

(27) Line 354: Again, why isn't this the assumption we're working under?

As addressed in our response to Comment 25, our primary model assumes that bats can recognize their own echoes—an assumption supported by substantial empirical evidence. The alternative "full confusion" model was included to explore a worst-case scenario and highlight the behavioral consequences of failing to distinguish self from conspecific echoes. We assume that real bats may experience some degree of echo misidentification; however, our assumption of full confusion represents a worst-case scenario.

(28) Line 382: "Under the assumption that..." I agree that bats probably can, but if we assume they can differentiate them all, where's the jamming problem?

The assumption that bats can theoretically distinguish between different signal sources applies after successful detection. However, the jamming problem arises during the detection and localization stages, where acoustic interference can prevent echoes from crossing the detection threshold or distort their timing.

(29) Lines 386-387: The paper referenced focused on JAR in the context of foraging. What changes were made to the simulation to switch to obstacle avoidance?

While the simulation framework in Mazar & Yovel (2020) was developed to study jamming avoidance during foraging, the core components—such as the acoustic calculations, receiver model, and echolocation behavior—remain applicable. For the current study, we adapted the simulation extensively to address colony-exit behavior. These modifications include modeling cave walls as acoustic reflectors, implementing a pathfinding algorithm, integrating obstacle-avoidance maneuvers, and adapting the integration window and integration processes. These updates are detailed throughout the Methods section.

(30) Line 400-402: Something doesn't add up with the statement: each decision relies on an integration window that records estimated locations of detected reflectors from the last five echolocation calls, with the parameter being tested between 1 and 10 calls. Can the authors reword this to make it less confusing?

We have reworded the sentence to clarify that the default integration window includes five calls, while we systematically tested the effect of using 1 to 10 calls, see lines 486-487.

(31) Line 393: "30 deg/sec" why was this value chosen?

The turning rate of 30 deg/sec was manually selected to approximate the curvature of natural foraging flight paths observed in Rhinopoma microphyllum using on-board tags. Moreover, in Mazar & Yovel (2020), we showed that the flight dynamics of simulated bats in a closed room closely matched those of Pipistrellus kuhlii flying in a room of similar dimensions. However, in the current simulation, bats rarely follow a random-walk trajectory due to the structured environment and frequent obstacle detection. As a result, this parameter has no meaningful impact on the simulation outcomes.

(32) Line 412: "Harmony" --- do you mean harmonic? And what is the empirical evidence that RM bats use the 2nd harmonic compared to the 1st?

Perhaps showing a spectrogram of a real RM signal would be helpful.

The typo-error was corrected. For reference See (Goldshtein et al., 2025)

(33) Table 2: Something is incorrect with the table. The first row on the next page is the wrong species name. Also, where are the citations for these parameter values?

The table header has been corrected in the revised version. The parameter values for flight and echolocation behavior were derived from existing literature and empirical data: Pipistrellus kuhlii parameters were based on Kalko (1995), and Rhinopoma microphyllum parameters were extracted from our own recordings using on-board tags, as described in Goldstein et al. (2025). We have added the appropriate citations to Table 2.

(34) Line 442: How was the threshold level chosen?

The detection threshold in each level is set to the greater of either 7 dB above the noise level (0 dB-SPL) or the maximal received level minus 70 dB, effectively applying a dynamic range of 70 dB.

(35) Line 445: 100 micros: This is about 3cm. The resolution of PK is about 1cm. For RM it's about 10cm. So, this window is generous for PK, but too strict for RM.

To keep the model simple and avoid introducing species-specific detection thresholds, we selected a biologically plausible compromise that could reasonably apply to both species. This simplification ensures consistency across simulations while remaining within the known behavioral range.

(36) Line 448: What is the spectrum of the Gaussian noise, and did it change between PK and RM?

We used the same white Gaussian noise with a flat spectrum across the relevant frequency range (10–80 kHz) for both species. We have clarified this in the revised text in lines 570-572.

(37) Line 451: 4 milliseconds is 1.3m. Is this appropriate?

The 4 milliseconds window was selected based on established auditory masking thresholds described in Mazar & Yovel (2020), and supported by (Popper and Fay, 1995) ch. 2.4.5, ((Blauert, 1997),  ch. 3.1 and (Mohl and Surlykke, 1989). These values provide conservative lower bounds on bats’ ability to cope with masking (Beleyur and Goerlitz, 2019). For simplicity, we used constant thresholds within each window, see lines 574-576.  

(38) Line 452: Citation for the forward and backward masking durations?

See the  to the previous comment.

(39) Lines 460-461: This is unclear. How does the bat get directional information? The authors claim to be able to measure direction-of-arrival for each detection, but it is not clear how this is done

As noted in our response to Reviewer 1 (Comment on Line 457), directional information is not computed via an explicit binaural model. Instead, we assume the bat estimates the direction of arrival with an angular error that depends on the SNR, based on established studies (e.g., Simmons et al., 1983; Popper & Fay, 1995). We have clarified this in the revised text in lines 583-592.

(40) Line 467: It seems like the authors are modeling pulse-echo ambiguity, at least in this one alternative model, which is good! However the alternative model doesn't get much attention in the paper. Is there a reason for this?

We would like to clarify that we did not model pulse-echo. In our confusion model, all echoes received within the IPI are attributed to the bat’s most recent call. This includes echoes that may in fact originate from conspecific calls, but the model does not assign self-echoes to earlier pulses or span multiple IPIs. Therefore, while the model captures echo confusion, it does not include true pulse-echo ambiguity. We have clarified this point in the revised text in lines 551-553.

(41) Line 41: "continuous" is more appropriate than "constant".

Thank you, we have rephrased the text accordingly.

(42) Line 69: "band width" should be one word.

Thank you, we have corrected it to “bandwidth”.

(43) Line 79: "bats" should be in the possessive.

Thank you, the text has been rephrased.

(44) Line 128: "convoluted" don't you mean "convolved"?

We have replaced “convoluted” with the correct term “convolved” in the revised text.

(45) Please check your references, as there are some incomplete citations and typos.

Thank you, we have reviewed and corrected all references for completeness and consistency.

References

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Academic Year: A school year that usually lasts 2 semesters

Registration: Applying and signing up for the college and classes.

Enrollment: The act of officially joining a college.

Admission: How much money is due to attend college.

Student Number: The ID number assigned to you.

Probation: The last attempt to get your grade up.

Credit Hour/Unit: A measure of workload and learning time for a course.

Term: Academic year when classes are held.

Tuition: Full cost for a whole school year.

General Eduction/Gen Ed: Basic courses taken as freshmen and sophomores.

Elective: A course that a student can choose to take based on interest.

Degree: A certificate awarded for completing a specific course.

Certificate: stating you completed a course.

Career Pathway: A look at education and training.

Financial Aid: Money to help pay for college.

FAFSA: An application you fill out to apply for financial aid.

Stafford Loan: A type of federal loan that helps students pay for college.

Scholarship: Financial support awarded to a student.

Grant: Financial aid that does not need to be repaid.

Federal Work Study: Part-time employment for undergraduate and graduate students to help them earn money.

Transcript: An academic record.

Non-Credit/Continuing Education: Short term courses or programs.

Audit: To attend and participate in a class without receiving academic credit or grade.

Grade Options: The choice between receiving a letter for grade (A,B,C,D,F).

Course Number: Indicate’s a course subject, level, and sometime its credit hours.

College Level Course: Academic instruction that requires critical thinking,etc.

Pre-College Level Course: A college-level course taken by a high school student.

Lower Division Course: A introductory class taken during first two years of a bachelor’s program.

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Prerequisite: A course or condition you have to do before doing something else.

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Learning Community: Group of students who share common academic goals.

Major: A primary field of study you specialize in to earn a degree.

A widely used reading process that involves surveying the text and forming questions before reading; reading to answer questions, predict test material, and form new questions and predictions; reciting or recording the main points of the text; and reviewing and reflecting upon the material.

called Abstract also includes electronic databases include summaries of articles

When you begin reading a section, try to identify two to three questions you should be able to answer after you finish it. Write down your questions and use them to test yourself on the reading. If you cannot answer a question, try to determine why. Is the answer buried in that section of reading but just not coming across to you? Or do you expect to find the answer in another part of the reading?

At the end of each section, pause to summarize the main points in a few sentences. If you have trouble doing so, revisit that section.

good way to understand what the author is getting at

Details, facts and explanations that develop and clarify a writer's main point

This is the challenge when it comes to comprehension

Some of your reading assignments will be fairly straightforward. Others, however, will be longer or more complex, so you will need a plan for how to handle them.

Measuring gene similarity within each Chromosome (the traditional method of detecting relatedness) would be a very strong supplemental figure to establish a baseline of comparison to GeneCompare. How does the previously biased approach compare to this new unbiased approach?

Because of the high number of match queries, it unlikely that the percent ratios would change much with subsequent runs of GenomeCompare. However to alleviate concerns of algorithm stability, it may be worth running GeneCompare at these three granularities multiple times to add confidence intervals to these ratios.

As mentioned, changing the granularity of chromosome comparisons does not perfectly preserve the rank order of relatedness (eg chromosome 16 going from third lowest at 32bp to lowest at 200bp, while chromosomes 1,6,11 remain ranked third, first and second respectively). However, this isn't necessarily "more" indicative of relatedness. Applying CompareGenome to more species (especially with varied evolutionary histories, genetic architectures, mutation rates, etc), seems like the next logical step (as suggested in the discussion section) towards providing evidence for this claim.

Observation: Hard workers won't have to worry about food. Interpretation: Benjamin signifies the importance of hard-working people compared to those who laze around. My observation indicates that there was struggle and Franklin shows that the people can't only blame taxes but can also blame the laziness of others. Context: People complained about the struggle of working with harsh taxes to the government. Franklin addresses their concerns but many dismiss him and walk away.