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Writing an Abstract for Your Research Paper

Definition and Purpose of Abstracts

An abstract is a short summary of your (published or unpublished) research paper, usually about a paragraph (c. 6-7 sentences, 150-250 words) long. A well-written abstract serves multiple purposes:

• an abstract lets readers get the gist or essence of your paper or article quickly, in order to decide whether to read the full paper;
• an abstract prepares readers to follow the detailed information, analyses, and arguments in your full paper;
• and, later, an abstract helps readers remember key points from your paper.

It’s also worth remembering that search engines and bibliographic databases use abstracts, as well as the title, to identify key terms for indexing your published paper. So what you include in your abstract and in your title are crucial for helping other researchers find your paper or article.

If you are writing an abstract for a course paper, your professor may give you specific guidelines for what to include and how to organize your abstract. Similarly, academic journals often have specific requirements for abstracts. So in addition to following the advice on this page, you should be sure to look for and follow any guidelines from the course or journal you’re writing for.

The Contents of an Abstract

Abstracts contain most of the following kinds of information in brief form. The body of your paper will, of course, develop and explain these ideas much more fully. As you will see in the samples below, the proportion of your abstract that you devote to each kind of information—and the sequence of that information—will vary, depending on the nature and genre of the paper that you are summarizing in your abstract. And in some cases, some of this information is implied, rather than stated explicitly. The Publication Manual of the American Psychological Association , which is widely used in the social sciences, gives specific guidelines for what to include in the abstract for different kinds of papers—for empirical studies, literature reviews or meta-analyses, theoretical papers, methodological papers, and case studies.

Here are the typical kinds of information found in most abstracts:

• the context or background information for your research; the general topic under study; the specific topic of your research
• the central questions or statement of the problem your research addresses
• what’s already known about this question, what previous research has done or shown
• the main reason(s) , the exigency, the rationale , the goals for your research—Why is it important to address these questions? Are you, for example, examining a new topic? Why is that topic worth examining? Are you filling a gap in previous research? Applying new methods to take a fresh look at existing ideas or data? Resolving a dispute within the literature in your field? . . .
• your research and/or analytical methods
• your main findings , results , or arguments
• the significance or implications of your findings or arguments.

Your abstract should be intelligible on its own, without a reader’s having to read your entire paper. And in an abstract, you usually do not cite references—most of your abstract will describe what you have studied in your research and what you have found and what you argue in your paper. In the body of your paper, you will cite the specific literature that informs your research.

When to Write Your Abstract

Although you might be tempted to write your abstract first because it will appear as the very first part of your paper, it’s a good idea to wait to write your abstract until after you’ve drafted your full paper, so that you know what you’re summarizing.

What follows are some sample abstracts in published papers or articles, all written by faculty at UW-Madison who come from a variety of disciplines. We have annotated these samples to help you see the work that these authors are doing within their abstracts.

Choosing Verb Tenses within Your Abstract

The social science sample (Sample 1) below uses the present tense to describe general facts and interpretations that have been and are currently true, including the prevailing explanation for the social phenomenon under study. That abstract also uses the present tense to describe the methods, the findings, the arguments, and the implications of the findings from their new research study. The authors use the past tense to describe previous research.

The humanities sample (Sample 2) below uses the past tense to describe completed events in the past (the texts created in the pulp fiction industry in the 1970s and 80s) and uses the present tense to describe what is happening in those texts, to explain the significance or meaning of those texts, and to describe the arguments presented in the article.

The science samples (Samples 3 and 4) below use the past tense to describe what previous research studies have done and the research the authors have conducted, the methods they have followed, and what they have found. In their rationale or justification for their research (what remains to be done), they use the present tense. They also use the present tense to introduce their study (in Sample 3, “Here we report . . .”) and to explain the significance of their study (In Sample 3, This reprogramming . . . “provides a scalable cell source for. . .”).

Sample Abstract 1

From the social sciences.

Reporting new findings about the reasons for increasing economic homogamy among spouses

Gonalons-Pons, Pilar, and Christine R. Schwartz. “Trends in Economic Homogamy: Changes in Assortative Mating or the Division of Labor in Marriage?” Demography , vol. 54, no. 3, 2017, pp. 985-1005.

Sample Abstract 2

From the humanities.

Analyzing underground pulp fiction publications in Tanzania, this article makes an argument about the cultural significance of those publications

Emily Callaci. “Street Textuality: Socialism, Masculinity, and Urban Belonging in Tanzania’s Pulp Fiction Publishing Industry, 1975-1985.” Comparative Studies in Society and History , vol. 59, no. 1, 2017, pp. 183-210.

Sample Abstract/Summary 3

From the sciences.

Reporting a new method for reprogramming adult mouse fibroblasts into induced cardiac progenitor cells

Lalit, Pratik A., Max R. Salick, Daryl O. Nelson, Jayne M. Squirrell, Christina M. Shafer, Neel G. Patel, Imaan Saeed, Eric G. Schmuck, Yogananda S. Markandeya, Rachel Wong, Martin R. Lea, Kevin W. Eliceiri, Timothy A. Hacker, Wendy C. Crone, Michael Kyba, Daniel J. Garry, Ron Stewart, James A. Thomson, Karen M. Downs, Gary E. Lyons, and Timothy J. Kamp. “Lineage Reprogramming of Fibroblasts into Proliferative Induced Cardiac Progenitor Cells by Defined Factors.” Cell Stem Cell , vol. 18, 2016, pp. 354-367.

Sample Abstract 4, a Structured Abstract

Reporting results about the effectiveness of antibiotic therapy in managing acute bacterial sinusitis, from a rigorously controlled study

Note: This journal requires authors to organize their abstract into four specific sections, with strict word limits. Because the headings for this structured abstract are self-explanatory, we have chosen not to add annotations to this sample abstract.

Wald, Ellen R., David Nash, and Jens Eickhoff. “Effectiveness of Amoxicillin/Clavulanate Potassium in the Treatment of Acute Bacterial Sinusitis in Children.” Pediatrics , vol. 124, no. 1, 2009, pp. 9-15.

“OBJECTIVE: The role of antibiotic therapy in managing acute bacterial sinusitis (ABS) in children is controversial. The purpose of this study was to determine the effectiveness of high-dose amoxicillin/potassium clavulanate in the treatment of children diagnosed with ABS.

METHODS : This was a randomized, double-blind, placebo-controlled study. Children 1 to 10 years of age with a clinical presentation compatible with ABS were eligible for participation. Patients were stratified according to age (<6 or ≥6 years) and clinical severity and randomly assigned to receive either amoxicillin (90 mg/kg) with potassium clavulanate (6.4 mg/kg) or placebo. A symptom survey was performed on days 0, 1, 2, 3, 5, 7, 10, 20, and 30. Patients were examined on day 14. Children’s conditions were rated as cured, improved, or failed according to scoring rules.

RESULTS: Two thousand one hundred thirty-five children with respiratory complaints were screened for enrollment; 139 (6.5%) had ABS. Fifty-eight patients were enrolled, and 56 were randomly assigned. The mean age was 6630 months. Fifty (89%) patients presented with persistent symptoms, and 6 (11%) presented with nonpersistent symptoms. In 24 (43%) children, the illness was classified as mild, whereas in the remaining 32 (57%) children it was severe. Of the 28 children who received the antibiotic, 14 (50%) were cured, 4 (14%) were improved, 4(14%) experienced treatment failure, and 6 (21%) withdrew. Of the 28children who received placebo, 4 (14%) were cured, 5 (18%) improved, and 19 (68%) experienced treatment failure. Children receiving the antibiotic were more likely to be cured (50% vs 14%) and less likely to have treatment failure (14% vs 68%) than children receiving the placebo.

CONCLUSIONS : ABS is a common complication of viral upper respiratory infections. Amoxicillin/potassium clavulanate results in significantly more cures and fewer failures than placebo, according to parental report of time to resolution.” (9)

Some Excellent Advice about Writing Abstracts for Basic Science Research Papers, by Professor Adriano Aguzzi from the Institute of Neuropathology at the University of Zurich:

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Writing a manuscript and mastering abstracts: a guide for authors

Author: guest contributor.

How you structure your article can affect how much it gets read—and cited. Each section serves a purpose, and does so in a particular order.

In this blog, we’ll take a look at the overall structure that you should follow in your article. And then we’ll focus specifically on the abstract—one of the most important sections that can help your article get attention.

Organizing your article: IMRaD structure

Structuring your article well helps readers find the information they’re looking for, and helps them easily follow your methodologies and arguments. To help do this, most articles follow a common pattern and structure—after the title and abstract and before the references—called IMRaD.

IMRaD stands for I ntroduction, M aterials and M ethods, R esults, (a)nd D iscussion and Conclusions. And although this is how you should structure your manuscript (in most cases), it’s not the best order for writing it.

You would actually want to start your writing with the Materials and Methods sections, followed by the Results. And you can work on these sections while running your experiments or doing your research as they describe what you’re doing.

Once you have results—and you’ve analyzed what they mean—you can work on the sections that describe that. So that’s the Discussion, Conclusion, and Introduction.

Only after that would you turn to the title and the abstract.

So let’s look next at how to write the best abstract.

Abstracts: Selecting the most important information

The first part of your article—after the title—that readers will see is your abstract. Readers will often only see your title and abstract, among lists of other articles on similar topics. This means that really well-done abstracts can have a big impact on how much your article gets read—and cited.

The abstract must outline the most important aspects of the study while providing only a limited amount of detail on its background, methodology and results. So you need to critically assess the different aspects of the manuscript and choose those that are sufficiently important to deserve inclusion in the abstract.

Once the abstract is ready it can be helpful to ask a colleague who is not involved in the research to go through it to ensure that the descriptions are clear. After you have drafted the manuscript, you should go back to the abstract to check that it agrees with the contents of the final manuscript.

Abstracts should have a structured format, serving several purposes: it helps authors summarize the different aspects of their work; it makes the abstract more immediately clear; and it helps peer reviewers and readers assess the contents of the manuscript.

The abstract structure varies between journals and between types of articles. You should check that the abstract of their manuscript is consistent with the requirements of the article type and journal to which the manuscript will be submitted. 

TIP: Journals often set a maximum word count for Abstracts, often 250 words, and no citations. This is to ensure that the full Abstract appears in indexing services.

These tips and suggestions should help you get started.

But to learn more about how to prepare for your next article submission, you can take Springe Nature’s free online tutorial (registration required), “ Writing a journal manuscript. ” You can also explore more resources at Nature Masterclasses and at AJE .

Best of luck with your next manuscript!

Guest Contributors include Springer Nature staff and authors, industry experts, society partners, and many others. If you are interested in being a Guest Contributor, please contact us via email: [email protected] .

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• How to Write an Abstract

Expedite peer review, increase search-ability, and set the tone for your study

The abstract is your chance to let your readers know what they can expect from your article. Learn how to write a clear, and concise abstract that will keep your audience reading.

How your abstract impacts editorial evaluation and future readership

After the title , the abstract is the second-most-read part of your article. A good abstract can help to expedite peer review and, if your article is accepted for publication, it’s an important tool for readers to find and evaluate your work. Editors use your abstract when they first assess your article. Prospective reviewers see it when they decide whether to accept an invitation to review. Once published, the abstract gets indexed in PubMed and Google Scholar , as well as library systems and other popular databases. Like the title, your abstract influences keyword search results. Readers will use it to decide whether to read the rest of your article. Other researchers will use it to evaluate your work for inclusion in systematic reviews and meta-analysis. It should be a concise standalone piece that accurately represents your research. 

What to include in an abstract

The main challenge you’ll face when writing your abstract is keeping it concise AND fitting in all the information you need. Depending on your subject area the journal may require a structured abstract following specific headings. A structured abstract helps your readers understand your study more easily. If your journal doesn’t require a structured abstract it’s still a good idea to follow a similar format, just present the abstract as one paragraph without headings. 

Background or Introduction – What is currently known? Start with a brief, 2 or 3 sentence, introduction to the research area. 

Objectives or Aims – What is the study and why did you do it? Clearly state the research question you’re trying to answer.

Methods – What did you do? Explain what you did and how you did it. Include important information about your methods, but avoid the low-level specifics. Some disciplines have specific requirements for abstract methods. 

• CONSORT for randomized trials.
• STROBE for observational studies
• PRISMA for systematic reviews and meta-analyses

Results – What did you find? Briefly give the key findings of your study. Include key numeric data (including confidence intervals or p values), where possible.

Conclusions – What did you conclude? Tell the reader why your findings matter, and what this could mean for the ‘bigger picture’ of this area of research. 

Writing tips

The main challenge you may find when writing your abstract is keeping it concise AND convering all the information you need to.

• Keep it concise and to the point. Most journals have a maximum word count, so check guidelines before you write the abstract to save time editing it later.
• Write for your audience. Are they specialists in your specific field? Are they cross-disciplinary? Are they non-specialists? If you’re writing for a general audience, or your research could be of interest to the public keep your language as straightforward as possible. If you’re writing in English, do remember that not all of your readers will necessarily be native English speakers.
• Focus on key results, conclusions and take home messages.
• Write your paper first, then create the abstract as a summary.
• Check the journal requirements before you write your abstract, eg. required subheadings.
• Include keywords or phrases to help readers search for your work in indexing databases like PubMed or Google Scholar.
• Double and triple check your abstract for spelling and grammar errors. These kind of errors can give potential reviewers the impression that your research isn’t sound, and can make it easier to find reviewers who accept the invitation to review your manuscript. Your abstract should be a taste of what is to come in the rest of your article.

Don’t

• Sensationalize your research.
• Speculate about where this research might lead in the future.
• Use abbreviations or acronyms (unless absolutely necessary or unless they’re widely known, eg. DNA).
• Repeat yourself unnecessarily, eg. “Methods: We used X technique. Results: Using X technique, we found…”
• Contradict anything in the rest of your manuscript.
• Include content that isn’t also covered in the main manuscript.
• Include citations or references.

Tip: How to edit your work

Editing is challenging, especially if you are acting as both a writer and an editor. Read our guidelines for advice on how to refine your work, including useful tips for setting your intentions, re-review, and consultation with colleagues.

• How to Write a Great Title
• How to Write Your Methods
• How to Report Statistics
• How to Write Discussions and Conclusions
• How to Edit Your Work

The contents of the Peer Review Center are also available as a live, interactive training session, complete with slides, talking points, and activities. …

The contents of the Writing Center are also available as a live, interactive training session, complete with slides, talking points, and activities. …

There’s a lot to consider when deciding where to submit your work. Learn how to choose a journal that will help your study reach its audience, while reflecting your values as a researcher…

Journal Article: Abstract

When to write the abstract.

• Introduction

Writing an abstract can be difficult because you are tasked with condensing tons of work into such a small amount of space. To make things easier, write your abstract last. Read through your entire paper and distill each section down to its main points. Sometimes it can be helpful to answer this question through a subtractive process. For example, if you are trying to distill down your results, simply list all your findings and then go through that list and start crossing off or consolidating each finding until you are left with a only the most crucial results.

Your title and abstract are the primary medium through which interested readers will find your work amidst the deluge of scientific publications, posters, or conference talks. When a fellow scientist happens upon your abstract they will quickly skim it to determine if it is worth their time to dive into the main body of the paper. The main purpose of an abstract, therefore, is to contextualize and describe your work in a concise and easily-understood manner. This will ensure that your scientific work is found and read by your intended audience.

Abstract Formula

Clarity is achieved by providing information in a predictable order: successful abstracts therefore are composed of 6 ordered components which are referred to as the “abstract formula”.

General and   Specific Background (~1 sentence each). Introduce the area of science that you will be speaking about and the state of knowledge in that area. Start broad in the general background, then narrow in on the relevant topic that will be pursued in the paper. I f you use jargon, be sure to very briefly define it.

Knowledge Gap (~1 sentence). Now that you’ve stated what is already known, state what is not known. W hat specific question is your work attempting to answer?

“Here we show…” (~1 sentence). State your general experimental approach and the answer to the question which you just posed in the “Knowledge Gap” section.

Experimental Approach & Results (~1-3 sentences). Provide a high-level description of your most important methods and results. How did you get to the conclusion that you stated in the “Here we show…” section?

Implications (~1 sentence).  Describe how your findings influence our understanding of the relevant field and/or their implications for future studies.

This content was adapted from from an article originally created by the  MIT Biological Engineering Communication Lab .

Resources and Annotated Examples

Annotated example 1.

Annotated abstract of a microbiology paper published in Nature . 4 MB

Annotated Example 2

Annotated abstract of a paper published in Nature . 2 MB

How to Write an Abstract?

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• First Online: 24 October 2021

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• Samiran Nundy 4 ,
• Atul Kakar 5 &
• Zulfiqar A. Bhutta 6  

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An abstract is a crisp, short, powerful, and self-contained summary of a research manuscript used to help the reader swiftly determine the paper’s purpose. Although the abstract is the first paragraph of the manuscript it should be written last when all the other sections have been addressed.

Research is formalized curiosity. It is poking and prying with a purpose. — Zora Neale Hurston, American Author, Anthropologist and Filmmaker (1891–1960)

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Writing the Abstract

Abstract and Keywords

Additional Commentaries

1 what is an abstract.

An abstract is usually a standalone document that informs the reader about the details of the manuscript to follow. It is like a trailer to a movie, if the trailer is good, it stimulates the audience to watch the movie. The abstract should be written from scratch and not ‘cut –and-pasted’ [ 1 ].

2 What is the History of the Abstract?

An abstract, in the form of a single paragraph, was first published in the Canadian Medical Association Journal in 1960 with the idea that the readers may not have enough time to go through the whole paper, and the first abstract with a defined structure was published in 1991 [ 2 ]. The idea sold and now most original articles and reviews are required to have a structured abstract. The abstract attracts the reader to read the full manuscript [ 3 ].

3 What are the Qualities of a Good Abstract?

The quality of information in an abstract can be summarized by four ‘C’s. It should be:

C: Condensed

C: Critical

4 What are the Types of Abstract?

Before writing the abstract, you need to check with the journal website about which type of abstract it requires, with its length and style in the ‘Instructions to Authors’ section.

The abstract types can be divided into:

Descriptive: Usually written for psychology, social science, and humanities papers. It is about 50–100 words long. No conclusions can be drawn from this abstract as it describes the major points in the paper.

Informative: The majority of abstracts for science-related manuscripts are informative and are surrogates for the research done. They are single paragraphs that provide the reader an overview of the research paper and are about 100–150 words in length. Conclusions can be drawn from the abstracts and in the recommendations written in the last line.

Critical: This type of abstract is lengthy and about 400–500 words. In this, the authors’ own research is discussed for reliability, judgement, and validation. A comparison is also made with similar studies done earlier.

Highlighting: This is rarely used in scientific writing. The style of the abstract is to attract more readers. It is not a balanced or complete overview of the article with which it is published.

Structured: A structured abstract contains information under subheadings like background, aims, material and methods, results, conclusion, and recommendations (Fig. 15.1 ). Most leading journals now carry these.

Example of a structured abstract (with permission editor CMRP)

5 What is the Purpose of an Abstract?

An abstract is written to educate the reader about the study that follows and provide an overview of the science behind it. If written well it also attracts more readers to the article. It also helps the article getting indexed. The fate of a paper both before and after publication often depends upon its abstract. Most readers decide if a paper is worth reading on the basis of the abstract. Additionally, the selection of papers in systematic reviews is often dependent upon the abstract.

6 What are the Steps of Writing an Abstract?

An abstract should be written last after all the other sections of an article have been addressed. A poor abstract may turn off the reader and they may cause indexing errors as well. The abstract should state the purpose of the study, the methodology used, and summarize the results and important conclusions. It is usually written in the IMRAD format and is called a structured abstract [ 4 , 5 ].

I: The introduction in the opening line should state the problem you are addressing.

M: Methodology—what method was chosen to finish the experiment?

R: Results—state the important findings of your study.

D: Discussion—discuss why your study is important.

Mention the following information:

Important results with the statistical information ( p values, confidence intervals, standard/mean deviation).

Arrange all information in a chronological order.

Do not repeat any information.

The last line should state the recommendations from your study.

The abstract should be written in the past tense.

7 What are the Things to Be Avoided While Writing an Abstract?

Cut and paste information from the main text

Hold back important information

Use abbreviations

Tables or Figures

Generalized statements

Arguments about the study

8 What are Key Words?

These are important words that are repeated throughout the manuscript and which help in the indexing of a paper. Depending upon the journal 3–10 key words may be required which are indexed with the help of MESH (Medical Subject Heading).

9 How is an Abstract Written for a Conference Different from a Journal Paper?

The basic concept for writing abstracts is the same. However, in a conference abstract occasionally a table or figure is allowed. A word limit is important in both of them. Many of the abstracts which are presented in conferences are never published in fact one study found that only 27% of the abstracts presented in conferences were published in the next five years [ 6 ].

Table 15.1 gives a template for writing an abstract.

10 What are the Important Recommendations of the International Committees of Medical Journal of Editors?

The recommendations are [ 7 ]:

An abstract is required for original articles, metanalysis, and systematic reviews.

A structured abstract is preferred.

The abstract should mention the purpose of the scientific study, how the procedure was carried out, the analysis used, and principal conclusion.

Clinical trials should be reported according to the CONSORT guidelines.

The trials should also mention the funding and the trial number.

The abstract should be accurate as many readers have access only to the abstract.

11 Conclusions

An Abstract should be written last after all the other sections of the manuscript have been completed and with due care and attention to the details.

It should be structured and written in the IMRAD format.

For many readers, the abstract attracts them to go through the complete content of the article.

The abstract is usually followed by key words that help to index the paper.

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Preparing a manuscript for submission to a medical journal. Available on http://www.icmje.org/recommendations/browse/manuscript-preparation/preparing-for-submission.html . Accessed 10 May 2020.

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Institute for Global Health and Development, The Aga Khan University, South Central Asia, East Africa and United Kingdom, Karachi, Pakistan

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Nundy, S., Kakar, A., Bhutta, Z.A. (2022). How to Write an Abstract?. In: How to Practice Academic Medicine and Publish from Developing Countries?. Springer, Singapore. https://doi.org/10.1007/978-981-16-5248-6_15

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How to write a good abstract for a scientific paper or conference presentation

Chittaranjan andrade.

Department of Psychopharmacology, National Institute of Mental Health and Neurosciences, Bangalore, Karnataka, India

Abstracts of scientific papers are sometimes poorly written, often lack important information, and occasionally convey a biased picture. This paper provides detailed suggestions, with examples, for writing the background, methods, results, and conclusions sections of a good abstract. The primary target of this paper is the young researcher; however, authors with all levels of experience may find useful ideas in the paper.

INTRODUCTION

This paper is the third in a series on manuscript writing skills, published in the Indian Journal of Psychiatry . Earlier articles offered suggestions on how to write a good case report,[ 1 ] and how to read, write, or review a paper on randomized controlled trials.[ 2 , 3 ] The present paper examines how authors may write a good abstract when preparing their manuscript for a scientific journal or conference presentation. Although the primary target of this paper is the young researcher, it is likely that authors with all levels of experience will find at least a few ideas that may be useful in their future efforts.

The abstract of a paper is the only part of the paper that is published in conference proceedings. The abstract is the only part of the paper that a potential referee sees when he is invited by an editor to review a manuscript. The abstract is the only part of the paper that readers see when they search through electronic databases such as PubMed. Finally, most readers will acknowledge, with a chuckle, that when they leaf through the hard copy of a journal, they look at only the titles of the contained papers. If a title interests them, they glance through the abstract of that paper. Only a dedicated reader will peruse the contents of the paper, and then, most often only the introduction and discussion sections. Only a reader with a very specific interest in the subject of the paper, and a need to understand it thoroughly, will read the entire paper.

Thus, for the vast majority of readers, the paper does not exist beyond its abstract. For the referees, and the few readers who wish to read beyond the abstract, the abstract sets the tone for the rest of the paper. It is therefore the duty of the author to ensure that the abstract is properly representative of the entire paper. For this, the abstract must have some general qualities. These are listed in Table 1 .

General qualities of a good abstract

SECTIONS OF AN ABSTRACT

Although some journals still publish abstracts that are written as free-flowing paragraphs, most journals require abstracts to conform to a formal structure within a word count of, usually, 200–250 words. The usual sections defined in a structured abstract are the Background, Methods, Results, and Conclusions; other headings with similar meanings may be used (eg, Introduction in place of Background or Findings in place of Results). Some journals include additional sections, such as Objectives (between Background and Methods) and Limitations (at the end of the abstract). In the rest of this paper, issues related to the contents of each section will be examined in turn.

This section should be the shortest part of the abstract and should very briefly outline the following information:

• What is already known about the subject, related to the paper in question
• What is not known about the subject and hence what the study intended to examine (or what the paper seeks to present)

In most cases, the background can be framed in just 2–3 sentences, with each sentence describing a different aspect of the information referred to above; sometimes, even a single sentence may suffice. The purpose of the background, as the word itself indicates, is to provide the reader with a background to the study, and hence to smoothly lead into a description of the methods employed in the investigation.

Some authors publish papers the abstracts of which contain a lengthy background section. There are some situations, perhaps, where this may be justified. In most cases, however, a longer background section means that less space remains for the presentation of the results. This is unfortunate because the reader is interested in the paper because of its findings, and not because of its background.

A wide variety of acceptably composed backgrounds is provided in Table 2 ; most of these have been adapted from actual papers.[ 4 – 9 ] Readers may wish to compare the content in Table 2 with the original abstracts to see how the adaptations possibly improve on the originals. Note that, in the interest of brevity, unnecessary content is avoided. For instance, in Example 1 there is no need to state “The antidepressant efficacy of desvenlafaxine (DV), a dual-acting antidepressant drug , has been established…” (the unnecessary content is italicized).

Examples of the background section of an abstract

The methods section is usually the second-longest section in the abstract. It should contain enough information to enable the reader to understand what was done, and how. Table 3 lists important questions to which the methods section should provide brief answers.

Questions regarding which information should ideally be available in the methods section of an abstract

Carelessly written methods sections lack information about important issues such as sample size, numbers of patients in different groups, doses of medications, and duration of the study. Readers have only to flip through the pages of a randomly selected journal to realize how common such carelessness is.

Table 4 presents examples of the contents of accept-ably written methods sections, modified from actual publications.[ 10 , 11 ] Readers are invited to take special note of the first sentence of each example in Table 4 ; each is packed with detail, illustrating how to convey the maximum quantity of information with maximum economy of word count.

Examples of the methods section of an abstract

The results section is the most important part of the abstract and nothing should compromise its range and quality. This is because readers who peruse an abstract do so to learn about the findings of the study. The results section should therefore be the longest part of the abstract and should contain as much detail about the findings as the journal word count permits. For example, it is bad writing to state “Response rates differed significantly between diabetic and nondiabetic patients.” A better sentence is “The response rate was higher in nondiabetic than in diabetic patients (49% vs 30%, respectively; P <0.01).”

Important information that the results should present is indicated in Table 5 . Examples of acceptably written abstracts are presented in Table 6 ; one of these has been modified from an actual publication.[ 11 ] Note that the first example is rather narrative in style, whereas the second example is packed with data.

Information that the results section of the abstract should ideally present

Examples of the results section of an abstract

CONCLUSIONS

This section should contain the most important take-home message of the study, expressed in a few precisely worded sentences. Usually, the finding highlighted here relates to the primary outcome measure; however, other important or unexpected findings should also be mentioned. It is also customary, but not essential, for the authors to express an opinion about the theoretical or practical implications of the findings, or the importance of their findings for the field. Thus, the conclusions may contain three elements:

• The primary take-home message
• The additional findings of importance
• The perspective

Despite its necessary brevity, this section has the most impact on the average reader because readers generally trust authors and take their assertions at face value. For this reason, the conclusions should also be scrupulously honest; and authors should not claim more than their data demonstrate. Hypothetical examples of the conclusions section of an abstract are presented in Table 7 .

Examples of the conclusions section of an abstract

MISCELLANEOUS OBSERVATIONS

Citation of references anywhere within an abstract is almost invariably inappropriate. Other examples of unnecessary content in an abstract are listed in Table 8 .

Examples of unnecessary content in a abstract

It goes without saying that whatever is present in the abstract must also be present in the text. Likewise, whatever errors should not be made in the text should not appear in the abstract (eg, mistaking association for causality).

As already mentioned, the abstract is the only part of the paper that the vast majority of readers see. Therefore, it is critically important for authors to ensure that their enthusiasm or bias does not deceive the reader; unjustified speculations could be even more harmful. Misleading readers could harm the cause of science and have an adverse impact on patient care.[ 12 ] A recent study,[ 13 ] for example, concluded that venlafaxine use during the second trimester of pregnancy may increase the risk of neonates born small for gestational age. However, nowhere in the abstract did the authors mention that these conclusions were based on just 5 cases and 12 controls out of the total sample of 126 cases and 806 controls. There were several other serious limitations that rendered the authors’ conclusions tentative, at best; yet, nowhere in the abstract were these other limitations expressed.

As a parting note: Most journals provide clear instructions to authors on the formatting and contents of different parts of the manuscript. These instructions often include details on what the sections of an abstract should contain. Authors should tailor their abstracts to the specific requirements of the journal to which they plan to submit their manuscript. It could also be an excellent idea to model the abstract of the paper, sentence for sentence, on the abstract of an important paper on a similar subject and with similar methodology, published in the same journal for which the manuscript is slated.

Source of Support: Nil

Conflict of Interest: None declared.

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• Published: 07 May 2024

Contextual and combinatorial structure in sperm whale vocalisations

• Pratyusha Sharma   ORCID: orcid.org/0000-0001-9053-4943 1 , 2 ,
• Shane Gero   ORCID: orcid.org/0000-0001-6854-044X 2 , 3 , 4 ,
• Roger Payne 2 ,
• David F. Gruber   ORCID: orcid.org/0000-0001-9041-2911 2 , 5 ,
• Daniela Rus   ORCID: orcid.org/0000-0001-5473-3566 1 , 2   na1 ,
• Antonio Torralba 1 , 2   na1 &
• Jacob Andreas   ORCID: orcid.org/0000-0002-3141-5845 1 , 2   na1  

Nature Communications volume  15 , Article number:  3617 ( 2024 ) Cite this article

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• Animal behaviour
• Marine biology
• Marine mammals

Sperm whales ( Physeter macrocephalus ) are highly social mammals that communicate using sequences of clicks called codas. While a subset of codas have been shown to encode information about caller identity, almost everything else about the sperm whale communication system, including its structure and information-carrying capacity, remains unknown. We show that codas exhibit contextual and combinatorial structure. First, we report previously undescribed features of codas that are sensitive to the conversational context in which they occur, and systematically controlled and imitated across whales. We call these rubato and ornamentation. Second, we show that codas form a combinatorial coding system in which rubato and ornamentation combine with two context-independent features we call rhythm and tempo to produce a large inventory of distinguishable codas. Sperm whale vocalisations are more expressive and structured than previously believed, and built from a repertoire comprising nearly an order of magnitude more distinguishable codas. These results show context-sensitive and combinatorial vocalisation can appear in organisms with divergent evolutionary lineage and vocal apparatus.

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Introduction.

The social complexity hypothesis 1 , 2 posits that animals in complex societies—featuring coordination, distributed decision-making, social recognition, and social learning of cultural strategies 3 , 4 , 5 , 6 —require complex communication systems to mediate these behaviours and interactions 1 , 7 . In humans, communication plays a particularly large role in complex social behaviours like strategising and teaching 8 , 9 , 10 . These behaviours require the ability to generate and understand a vast space of possible messages. For example, the sentence ‘Let’s meet next to the statue of Claude Shannon on the fifth floor at 3 pm’ picks out a specific action at a specific place and time from an enormous space of possible activities. This ability to generate complex messages, in turn, is supported by the fact that all known human languages exhibit contextual and combinatorial structure. (Throughout this paper, we use ‘context’ to denote conversational context—e.g., neighbouring codas—rather than behavioural context—e.g., diving—as is standard in the study of natural and formal languages. 11 ) To enable efficient communication, the meaning of most human utterances is underspecified and derived in part from the conversation that precedes them 12 . To enable many distinct meanings to be communicated, humans access a large inventory of basic sounds (phonemes) by combining phonetic features like place of articulation, manner of articulation, then sequence phonemes to produce an unbounded set of distinct utterances 13 , 14 , 15 , 16 , 17 . Contextuality and combinatoriality, especially below the sequence level, have few analogues in communication systems outside of human language 18 , 19 , 20 , 21 , 22 . Understanding when and how aspects of human-like communication arise in nature offers one path toward understanding the basis of intelligence in other life forms.

Cetaceans are an important group for the study of evolution and development of sophisticated communication systems 23 . Among cetaceans, long-term observational studies of sperm whales (Physeter macrocephalus) have described both a culturally defined, multi-level, matrilineal society 24 and a socially transmitted communication system 25 , 26 , 27 . Sperm whales are known for complex social and foraging behaviour, as well as group decision-making 28 . They communicate using codes 29 : stereotyped sequences of 3–40 broadband clicks (see glossary in Table  1 for definitions). Codas are exchanged between whales when socialising or between long, deep, foraging dives 24 . To date, scientists have described the sperm whale communication system in terms of a finite repertoire of coda types, each defined by a characteristic sequence of inter-click intervals (ICIs) as seen in Fig.  1 A. Coda-type repertoires can be defined manually or using automated clustering procedures, and have been used to delineate cultural boundaries among socially segregated but sympatric ‘clans’ of whales 25 whose members differ in their behaviour 26 , 30 , 31 . Past research has identified around 150 discrete coda types globally, with 21 in the Caribbean. But there is an apparent contradiction between the social and behavioural complexity evinced by sperm whales and the comparative simplicity of a communication system with a fixed set of messages. This contradiction naturally raises the question of whether any additional, previously undescribed structure is present in sperm whale vocalisations.

Sperm whales communicate by producing sequences of clicks. A Shows a two-minute exchange between two whales (with clicks visualized in blue and orange respectively) from the Dominica Sperm Whale Dataset. B Projects these clicks over a time–time plot, in which the horizontal axis plots the time in the exchange at which a click occurs, and the vertical axis represents the time of the click from the first click in the coda. The vertical axis serves as a microscope over the horizontal axis, revealing the internal structure of each coda. C Shows a time–time visualisation for the entire two-minute exchange (with lines connecting matching clicks between adjacent codas), revealing complex, context-dependent variation in coda structure.

We first demonstrate that some coda structure is contextual: When analysing codas exchanged between whales, we observe fine-grained modulation of inter-click intervals relative to preceding codas, as well as modification of standard coda types via the addition of an extra click. We term these contextual features rubato and ornamentation. Next, we show that the coda repertoire has combinatorial structure: in addition to rubato and ornamentation, codas’ rhythms and tempos can independently be discretised into a small number of categories or types. We show that all four features are sensed and acted upon by participants in the vocal exchanges, and thus constitute deliberate components of the whale communication system rather than unconscious variation. Rhythm, tempo, rubato and ornamentation can be freely combined, together enabling whales to systematically synthesize an enormous repertoire of distinguishable codas. In a dataset of 8719 codas from the sperm whales of the Eastern Caribbean clan, this ‘sperm whale phonetic alphabet’ makes it possible to systematically explain observed variability in coda structure. While the communicative function of many codas remains an open question, our results show that the sperm whale communication system is, in principle, capable of representing a large space of possible meanings, using similar mechanisms to those employed by human sound production and representation systems (e.g., speech, text, Morse code, and musical notation).

Results and discussion

The dataset.

For this study, we used the annotated coda dataset from The Dominica Sperm Whale Project (DSWP), the current largest sperm whale data repository. The recordings of the Eastern Caribbean 1 (EC-1) clan were used in the analysis, comprising 8719 codas making up 21 previously defined coda types. This dataset contains manually annotated coda clicks and extracted inter-click intervals in data recorded from various platforms and various recording systems between 2005 and 2018, including animal-worn, acoustic, biologging tags (DTags) deployed between 2014 and 2018. The EC-1 clan comprises 400 individuals. 42 tags were deployed on 25 different individuals in 11 different social units. Three of these are less-studied units, for which precise size estimates are not available. We conservatively estimate that at least 60 distinct whales are recorded in the DSWP dataset.

Exchange plots: visualizing multi-whale calls

Codas, considered to be the basic units of sperm whale communication, consist of click groups generally less than two seconds in duration. Previous work has defined coda types and characterised coda repertoires by examining single codas outside the context in which they were produced. However, codas are not produced in isolation, but in interactive exchanges between two or more chorusing whales. Individual whales within a chorus tend to produce sequences of codas with a periodicity of approximately 4 seconds (see Supplementary Discussion Section  1) . Across a chorus, interacting whales produce codas both alternately (i.e., turn-taking) or near-simultaneously (i.e., overlapping). Therefore, sperm whale vocalisations demonstrate complexity on two different timescales: a fine-grained time scale that determines the makeup of each individual coda, and a longer time scale that determines the overall structure of the interactive exchange across codas within a chorus.

We depict these vocalisations using a new visualisation we call an exchange plot (Fig.  1 B, C). Both axes of this plot measure time: the horizontal axis shows the time elapsed since the beginning of a conversation, and the vertical axis shows the time since the onset of each coda. Exchange plots reveal the structure of codas in their interactive context, making it possible to observe both fine-grained differences between adjacent codas made by interacting whales, and long-range trends over the course of an exchange.

Contextuality

Visualising whale vocalisations with an exchange plot, as in Fig.  1 B, C, makes it apparent that characterising sperm whale interactions as sequences of fixed coda types overlooks a great deal of information. First, coda duration varies smoothly over the course of an exchange; variation in the duration of a whale’s codas is systematically imitated by interlocutors, even when coda-internal click spacing differs (Fig.  1 C). Second, some ‘extra’ clicks in Fig.  1 C appear at the end of codas that otherwise match their neighbours. We hypothesised that these smooth variations and extra clicks constitute perceptible and controllable features of codas independent of their discrete type, pointing toward a more complex sperm whale communication system with a greater information-carrying capacity than previously reported.

Rubato: codas exhibit fine-grained duration variation in addition to discrete tempo

A coda’s duration is the sum of its inter-click intervals. While durations tend to cluster around a finite set of values (Fig.  2 A), there remains substantial continuous variation within these clusters. Past work has described these differences as unexplained ‘within-type variation’ of categorical coda types 25 , 32 . However, the structured nature of this variation—which, as shown in Fig.  1 C, is temporally correlated and imitated across whales—has never been previously documented. We demonstrate that variation in coda duration is not random: individual whales modulate coda durations smoothly over the course of multi-coda exchanges without necessarily imitating click counts or inter-click interval spacing. An example is depicted in Fig.  1 C: one whale produces a sequence of codas smoothly varying in duration, while its interlocutor closely matches these changes in overall coda duration but not the number of clicks (this refines the finding in ref. 33 that overlapping codas were more likely to be of the same coda type than expected by chance: sometimes codas share a duration but not a discrete type). By analogy to the corresponding musical phenomenon, we call this behaviour rubato.

Sperm whale codas were previously hypothesized to comprise 21 independent coda types. We show that this coda repertoire is built from two context-independent features (rhythm and tempo) and two context-sensitive features (rubato and ornamentation). A Tempo: (Left) The overall duration of a coda is the sum of its inter-click intervals. (Centre) Coda durations are distributed around a finite set of modes, which we call (tempo types). (Right) Snippets from exchange plots showing codas of different tempo types. B Rhythm: (Left) Normalising the vector of ICIs by the total duration returns a duration-independent coda representation, which we call rhythm. (Centre) Codas cluster around 18 rhythm types. (Right) Examples of normalised codas showing different rhythm types. C Rubato: (Left) Sperm whales slowly modulate coda duration across consecutive codas, a phenomenon we call rubato. (Centre) Rubato is gradual: adjacent codas have durations more similar to each other than codas of the same type from elsewhere in an exchange. (Right) Whale choruses with imitation of rubato represented in exchange plots. D Ornamentation: (Left) Some codas feature `extra clicks' (ornaments) not present in neighbouring codas that otherwise share the same ICIs. (Centre) A density plot showing the distribution of the ratio between final ICIs in ornamented codas versus unornamented codas. Ornamented codas have a significantly different ICI distribution compared to regular codas. (Right) Examples of ornaments in the DSWP dataset. E Thirty minutes of multi-whale choruses: Exchanges feature imitation of coda duration across whales, gradually accumulated changes in call structure, and rich contextual variability.

To show that rubato is not random variation, we first evaluated whether changes in duration are smooth by measuring whether codas are more similar to their neighbours than other codas of the same type. To do so, we computed the tempo drift between two codas from the same speaker, defined as the difference in coda durations (Fig.  2 C). We computed the average absolute drift between (1) adjacent coda pairs and (2) random coda pairs of the same discrete coda type (according to its rhythm and tempo; see Section 5 for a discussion of rhythm and tempo and Supplementary Discussion Section  4 for additional details). We found that drift was significantly smaller between adjacent codas (0.05s on average) than would be expected under a null hypothesis that drift depends only on a coda’s discrete type (which would give a drift of 0.08s on average; test: permutation test (one-sided), p  = 0.0001, n  = 2593; see Supplementary Discussion Section  4.1 for details). Thus, within-type variation is context-dependent.

Second, we evaluated whether sequences of codas reflect longer-term trends. To do so, we collected coda triples of the same discrete coda type and measured the correlation between tempo drift across adjacent pairs. We found a significant positive correlation, compared to a null hypothesis that drift between adjacent pairs is uncorrelated (test: Spearman’s rank-order correlation (two-sided), r (2586) = 0.57, p  < 0.0001, 95% CI = [0.54, 0.60], n  = 2588). Thus, rubato is sustained across sequences of multiple codas.

Finally, we evaluated whether rubato is perceived and controlled by measuring whales’ ability to match their interlocutors’ coda durations when chorusing. We measured the average absolute difference in duration between (1) pairs of overlapping codas from different whales, and (2) pairs of non-overlapping codas of the same discrete coda type. Durations are significantly more closely matched for overlapping codas (0.099s on average) than would be expected under a null hypothesis that chorusing whales match only discrete coda type (which would give a drift of 0.129s on average) (test: permutation test (one-sided), p  = 0.0001, n  = 908; see Supplementary Discussion Section  4) .

Ornamentation: some clicks do not belong to standard tempo types

Figure  2 D depicts an exchange consisting of one six-click coda, followed by a long sequence of five-click codas. The first five clicks of the initial 6-click coda closely match those of neighbouring codas: if the sixth click were removed, we would identify the first coda as having nearly the same inter-click intervals as its neighbours (and assign it to the same discrete coda type). While not previously described, ‘extra clicks’ of this kind occur in (4%) of the codas in the exchanges of Eastern Caribbean whales. Additional examples appear in Fig.  2 D and Supplementary Discussion Section  5 . We hypothesised that ‘extra’ clicks play a different role from the other clicks in the codas in which they appear: they do not determine discrete coda type. Instead, like rubato, they constitute an independent feature of the sperm whale vocalisation system. We call these extra clicks ornaments.

We define an ornament as the final click in a coda containing one more click than the nearest preceding or following coda within a window of ten seconds, with this interval being based on the average response time (Supplementary Discussion Section  1) . To show that these ornaments play a distinct role from other clicks, we first computed the mean squared distance between each standardised, ornamented coda and the cluster centre of its assigned rhythm type. We then removed ornaments and computed mean squared distance between the standardised coda and the centres of rhythm clusters for adjacent codas produced by the same whale. The second quantity (0.0034 s 2 , reflecting a hypothesis that ornamented codas match their neighbours) is significantly smaller than the first (0.0053 s 2 , reflecting a null hypothesis that ornamented codas resemble other codas of the same type) (test: permutation test (one-sided), p  = 0.002, n  = 178). In other words, ornamented codas are less like other codas with a matching number of clicks, and more like neighbouring non-ornamented codas, if ornaments are modelled as a separate factor. To further verify that ornaments are distinct from other clicks, we compared ICIs (inter-click intervals) in ornamented vs. non-ornamented codas with the same number of clicks. We specifically compared the difference between the final two ICIs, normalised by the penultimate ICI, to reduce variance arising from rubato. This measurement exhibits a significant difference in distribution in ornamented vs non-ornamented codas test: Kolmogorov–Smirnov test (two-sample), D (150, 3688) = 0.51, p  < 0.0001, 95% CI = [0.42, 0.62], n  = (150, 3688), Fig.  2 D.

Finally, ornaments are not distributed randomly but appear in distinctive positions in longer exchanges. Within a single whale’s call sequences (defined as a sequence of codas separated by no more than eight seconds), a significantly greater proportion of ornamented codas appear at the beginning of call sequences than unornamented codas (test: Fisher’s exact test (two-sided), odds ratio: 2.00, p  = 0.0006). A significantly greater fraction of ornamented codas also appear at the end of call sequences compared to unornamented codas (test: Fisher’s exact test (two-sided), odds ratio: 1.71, p  = 0.008). Moreover, ornaments are predictive of changes in multi-whale interactions. We define a ‘change in chorusing behaviour’ as one of three events: a following whale begins chorusing with a leading whale, pauses chorusing, or ceases vocalizing for the remainder of the exchange. Compared to unornamented codas, ornamented codas from the leading whale are disproportionately succeeded by a change in chorusing behaviour from the following whale (test: Fisher’s exact test (two-sided), odds ratio: 1.56, p  = 0.009). Thus ornamentation, like rubato, appears to be perceived in multi-whale interactions.

Combinatoriality

The existence of ornamentation and rubato features demonstrates that codas can carry more information, and have more complex internal structure than their discrete type alone would indicate. Instead, they result from a combinatorial coding system in which discrete type, ornamentation and rubato combine to realise individual codas. Based on these findings, we hypothesized that categorical coda types might themselves arise from a combinatorial process, with a simpler set of features explaining the prototypical ICI vector for each coda type.

During rubato, consecutive calls from a single-whale gradually vary coda duration while preserving the relative relationship of ICIs (Supplementary Discussion Section  2) , suggesting that whales are capable of maintaining this relationship independent of its duration. Moreover, existing studies have noted that codas may also be discretely clustered according to standardised ICI 25 , 32 , 34 , a process that assigns codas with very different durations to the same cluster. Finally, some instances of chorusing involve whales producing codas with different ICIs (or different numbers of clicks) but matched durations. Together, these observations suggest that past inventories of discrete coda types (e.g., ref. 35 ) might specifically be interpretable in terms of two features: a normalised ICI category (which we term a coda’s rhythm) and a discrete duration category (independent of rubato, and which we term tempo). To validate this hypothesis, we measured (1) whether coda rhythms and tempos cluster around a discrete set of values, and (2) whether rhythm and tempo features are independently combinable (both with each other and with ornamentation and rubato features).

As shown in Fig.  2 A, codas with the same duration may have different internal click spacing (and even different numbers of clicks) but still span the same amount of time from the first click to the last click. Performing kernel density estimation (KDE) on scalar coda durations from the DSWP dataset reveals five distinct modes in the distribution of durations (Fig.  2 A), indicating that the number of realised coda durations is much smaller than the total number of identified coda types (Supplementary Discussion Section  3) .

Across codas, the relative relationships between ICIs are often repeated even independent of tempo. For example, in Fig.  2 A, note the existence of two five-click codas, one long and one short, but both characterised by the uniform spacing of the constituent ICIs. Past work has shown that these rhythms are reused; our analysis uses the 18 rhythm clusters proposed by ref. 35 (detailed breakdowns are given in Fig.  2 B and in Supplementary Discussion Section  2) .

Finally, to evaluate the combinability of these features, we computed the frequency with which each rhythm and tempo feature co-occurred in the DWSP dataset, as well as the frequency with which each combination appeared with ornamentation or rubato. Results are shown in Fig.  3 . Each rhythm type appears with at least one tempo types and each tempo type appears with at least three rhythm types. Moreover, (22%) of these combinations can appear with or without rubato and ornamentation.

Analogous to visualizations of the human phonetic repertoire, we propose a phonetic alphabet for sperm whales. Tempo types are plotted on the vertical axis, rhythm types are plotted on the horizontal axis, and the colour of each cell represents the number of occurrences of that rhythm/tempo combination in the DSWP dataset. Pie charts in each cell provide further information about the prevalence of rubato and ornamentation within each feature combination: the left pie shows the ratio of the number of codas that appear with rubato to those without, while the right pie shows the fraction of all ornaments that appear with that feature combination. While not all feature combinations are realised (as observed in human languages), sperm whale codas have a rich combinatorial structure with both discrete and continuous parameters and at least 143 combinations frequently realised (Supplementary Discussion Section  6) .

Like the International Phonetic Alphabet for human languages, this ‘Sperm Whale Phonetic Alphabet’ (Fig.  3 ) shows how a small set of axes of variation (place of articulation, manner of articulation, and voicedness in humans; rhythm, tempo, ornamentation, and rubato in sperm whales) give rise to the diverse set of observed phonemes (in humans) or codas (in sperm whales). As in human languages, not all theoretically realisable feature combinations are attested in the DSWP dataset, and some combinations are more frequent than others. As in human languages, most coda variation is discrete: though ICIs can vary continuously in principle, only specific patterns (associated with specific rhythms and tempos) are realised in practice. Supplementary Discussion Section  2 shows the full set of codas in the dataset, organised by rhythm, tempo, and the presence of rubato and ornamentation for each combination of rhythm and tempo. Notably, these factors of sub-coda variation exist alongside another combinatorial process—the sequential ordering of codas shown in Fig.  1 E—in which codas of different types are combined in sequence to give rise to an even larger family of distinct vocalisations, reminiscent of the bi-level combinatorial structure of speech production in humans (see Supplementary Discussion Section  7 ).

Figure  3 also demonstrates that these vocalisations have a significantly greater information capacity than was previously known. Prior work identified 21 discrete coda types, and the system could be understood to have an information rate of at most 5 bits/coda. However, our analysis suggests that with 18 rhythms, 5 tempos, optional ornamentation, and three variations (increasing, decreasing or constant duration) in rubato, the information rate could be up to twice as large (details in Supplementary Discussion Section  6 ). The role of rubato within this coding system remains an important open question: it might be discrete (with some simpler inventory of contours explaining the patterns in Figs.  1 C and  2 C, as in the songs of birds 36 , 37 , 38 , 39 , 40 , 41 , and humpback whales (Megaptera novaeangliae) 42 , 43 ). Or it might convey continuous-valued information, analogous to the orientation and duration features of the waggle dance in bees (Apis sp.) 44 .

Limitations

Our study investigates the basic structural elements, and not the semantics, of the sperm whale communication system. As in several foundational papers on call structure in animal communication systems 42 , 45 , 46 , it provides no characterisation of call semantics and features no playback experiments. We believe our work provides a foundation for future research on the semantics of whale calls. However, this future research additionally requires interactive playback experiments with whales in the wild to ground hypotheses about the semantics and functional role of sperm whale vocalizations. In the absence of playback experiments establishing a causal relationship between the features and the meaning they communicate is challenging. It is necessary to have a deep understanding of the structure of the communication system enabling the creation of specific and unconfounded test stimuli prior to undertaking whale-in-the-loop experiments with potentially long-term effects on the population 47 , 48 , which in this case is extremely vulnerable 49 .

Concluding remarks

Our results demonstrate that sperm whale vocalisations form a complex combinatorial communication system: the seemingly arbitrary inventory of coda types can be explained by combinations of rhythm, tempo, rubato, and ornamentation features. Sizable combinatorial vocalisation systems are exceedingly rare in nature; however, their use by sperm whales shows that they are not uniquely human, and can arise from dramatically different physiological, ecological, and social pressures.

These findings also offer steps towards understanding how sperm whales transmit meaning. In some organisms with combinatorial codes, such as honey bees (Apis sp.) , the constituent features of the code transparently encode semantics (e.g., direction and distance to food sources). Further research on sperm whale vocalisations may investigate if rhythm, tempo, ornamentation, and rubato function similarly, directly encoding whales’ communicative intents. Alternatively, one of the key differentiators between human communication and all known animal communication systems is duality of patterning: a base set of individually meaningless elements that are sequenced to generate a very large space of meanings. The existence of a combinatorial coding system-at either the level of sounds, sound sequences, or both-is a prerequisite for duality of patterning. Our findings open up the possibility that sperm whale communication might provide our first example of that phenomenon in another species.

Data collection and coda annotation

Data from The Dominica Sperm Whale Project were collected under scientific research permits from the Fisheries Division of the Government of Dominica. The field protocols for approaching, photographing, tagging, and recording sperm whales were approved by either the University Committee on Laboratory Animals of Dalhousie University, Canada; the Animal Welfare and Ethics Committee of the University of St Andrews, Scotland; or Aarhus University, Denmark; and sometimes several or all of these across years.

Social units of female and immature sperm whales were located and followed in an area that covered approximately 2000 squared kilometres along the entire western coast of the Island of Dominica (N15.30 W61.40) between 2005 and 2018.

Codas were recorded using one of several recording setups: In 2005, we used a Fostex VF-160 multitrack recorder (44.1 kHz sampling rate) and a custom-built towed hydrophone (Benthos AQ-4 elements, frequency response: 0.1–30 kHz) with a filter box with high-pass filters up to 1 kHz resulting in a recording chain with a flat frequency response across a minimum of 2–20 kHz. No recordings were made during the short 2006 season. In the 2007, 2009, 2011, 2016, and 2017 seasons, we used a Zoom H4 portable field recorder (48 kHz sampling rate) and a Cetacean Research Technology C55 hydrophone (frequency response: 0.02–44 kHz) and no filters. During the 2008, 2010, 2012, 2015, and 2018 seasons, we used a custom-built towed hydrophone (Benthos AQ-4 elements, frequency response: 0.1–30 kHz) with a filter box with high-pass filters up to 1 kHz resulting in a recording chain with a flat frequency response across a minimum of 2–20 kHz. This was connected to a computer-based recording system as a part of the International Fund for Animal Welfare’s (IFAW) LOGGER software package (48 kHz sampling rate) or PAMGUARD (minimum 48 kHz sampling rate) 50 .

In addition, recordings were also made through the deployment of animal-borne sound and movement tags (DTag generation 3, Johnson and Tyack 2003). Tagging was undertaken between 2014 and 2018 on an 11-meter rigid-hulled inflatable boat (RHIB). Tags were deployed from a 9-meter, hand-held, carbon fiber pole, and were attached to the whales using four suction cups. DTags record two-channel audio at 120 kHz with a 16-bit resolution, providing a flat (±2 dB) frequency response between 0.4 and 45 kHz. Pressure and acceleration were sampled at a rate of 500 Hz with a 16-bit resolution, and were decimated to 25 Hz for analysis. DTag analysis was conducted using custom scripts in Matlab 2015b (The Mathworks, Inc., MA, USA). The variation in the frequency responses and sampling rates of the recording systems used did not affect our ability to record clean signals for both the coda and echolocation clicks produced by sperm whales, and as a result, the temporal patterning of clicks used in this analysis.

Whales, including the tagged whales, were identified through photographs of the trailing edge of their tails 51 . Identifications were used to ensure that only recordings from one of the two sympatric clans (EC-1, the Eastern Caribbean Clan) were included in the analysis to control for any differences in repertoires between vocal clans 27 .

To define the temporal structure of the codas recorded, absolute inter-click intervals were measured as in ref. 49 , using either custom-written Matlab tools and Rainbow Click Software (all years before 2014) or CodaSorter a custom-written tool (K. Beedholm, Marine Bioacoustics Lab, Aarhus University) in LabView (National Instruments, TX, USA). CodaSorter allows users to playback audio at various speeds and manually mark detected clicks as belonging to a specific coda. Estimates for each click for the angle of arrival, channel delay, centroid frequency, and inter-pulse interval (IPI, the time between the onset of the first pulse and the onset of the next pulse in the multi-pulse structure of sperm whales clicks 52 ) allowed for determining if the codas were produced by tagged whales or non-focal animals; and to ensure that, on days in which multiple tags were deployed, codas recorded by different tags were not double-counted. Photo-identification supported this process by identifying which whales were present and associated with the tagged whales at each surfacing.

There are two components to the dataset: Dataset 1, a large set of all 8719 codas that are annotated with information on their inter-click intervals; and Dataset 2, a smaller set of 3948 codas, which were recorded from animal-borne DTags, which remain in temporal order and are additionally annotated with information of the absolute time in the day of the first click of each of the codas and their associated speaker identities across the bouts. Experiments that do not require contextual information (those discussing the context-independent features of rhythm and tempo) use Dataset 1, whereas those requiring information about the relative ordering of the codas and their speaker IDs (those discussing the context-sensitive features of rubato and ornamentation), use Dataset 2. In both cases, rare, long codas were excluded from analysis (greater than 10 clicks, less than 5% of all codas recorded).

Additional discussion of statistical tests

Comparisons of coda durations (either with adjacent codas, when studying rubato, or with overlapping codas, when studying all features in the context of chorusing behaviour) use permutation tests to avoid making distributional assumptions about durations of codas and their absolute differences, some of which have non-normal distributions. All permutation tests are computed over 10,000 random resamplings of the data without replacement. Evaluation of Rubato additionally uses Spearman rank-correlation tests to measure longer-range trends across coda triplets (again based on initial observations that these trends appeared to be non-linear). Comparisons of coda-internal structure (e.g., durations of penultimate ICIs in ornamented codas) use Kolmogorov–Smirnov tests, as we are interested only in distributional differences rather than orderings of mean durations. Finally, measurements of changes in vocalization behaviour following Rubato use Fisher’s exact test to compare proportions of these changes in different vocal contexts.

Reporting summary

Further information on research design is available in the  Nature Portfolio Reporting Summary linked to this article.

Data availability

All data generated and in this study has been deposited on GitHub at https://github.com/pratyushasharma/sw-combinatoriality/tree/main/data 53 .

Code availability

Custom scripts used for this study are available at https://github.com/pratyushasharma/sw-combinatoriality 53 .

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Acknowledgements

This analysis was funded by Project CETI via grants from Dalio Philanthropies and Ocean X; Sea Grape Foundation; Virgin Unite, Rosamund Zander/Hansjorg Wyss, Chris Anderson/Jacqueline Novogratz through The Audacious Project: a collaborative funding initiative housed at TED to P.S., S.G., R.P., D.F.G., D.R., A.T. and J.A. Further funding was provided by the J.H. and E.V. Wade Fund at MIT. Fieldwork for The Dominica Sperm Whale Project was supported by through a FNU fellowship for the Danish Council for Independent Research supplemented by a Sapere Aude Research Talent Award (1325-00047A), a Carlsberg Foundation expedition grant (CF14-0789), two Explorer Grants from the National Geographic Society (WW-218R-17 and NGS-64863R-19), a grant from Focused on Nature, and supplementary grants from the Arizona Center for Nature Conservation, Quarters For Conservation, the Dansk Akustisks Selskab, Oticon Foundation, and the Dansk Tennis Fond all to S.G. Further funding was provided by a Discovery and Equipment grants from the Natural Sciences and Engineering Research Council of Canada (NSERC) to Hal Whitehead of Dalhousie University and a FNU large frame grant and a Villum Foundation Grant (13273) to Peter Madsen of Aarhus University. We thank the Chief Fisheries Officers and the Dominica Fisheries Division officers for research permits and their collaboration in data collection; all the crews of R/V Balaena and The DSWP team for data collection, curation, and annotation; as well as Dive Dominica, Al Dive, and W.E.T. Dominica for logistical support while in Dominica. A big thank you to Alex Boersma for providing us with sperm whale caricatures used in Fig.  1 and to Roger Levy and the CETI team for helpful discussions and valuable feedback on the paper.

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These authors contributed equally: Daniela Rus, Antonio Torralba, Jacob Andreas.

Authors and Affiliations

Computer Science and Artificial Intelligence Laboratory, Massachusetts Institute of Technology, Cambridge, MA, USA

Pratyusha Sharma, Daniela Rus, Antonio Torralba & Jacob Andreas

Project CETI, New York, NY, USA

Pratyusha Sharma, Shane Gero, Roger Payne, David F. Gruber, Daniela Rus, Antonio Torralba & Jacob Andreas

Department of Biology, Carleton University, 1125 Colonel By Drive, Ottawa, ON, K1S 5B6, Canada

The Dominica Sperm Whale Project, Roseau, Dominica

Baruch College and The Graduate Center, City University of New York, New York, NY, USA

David F. Gruber

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P.S., J.A., A.T., D.R., S.G., R.P. conceptualized the study, P.S., J.A., A.T. developed the methods, S.G. and The DSWP team collected the data, S.G. and The DSWP team annotated the original dataset, P.S., S.G. and The DSWP team curated the data, P.S. conducted the analyses and authored the code provided, P.S., J.A., A.T., S.G. verified the analytical results, S.G., D.F.G., D.R., A.T., J.A. funded the study, P.S. wrote the manuscript, all authors contributed review and editing to the manuscript and gave final approval for publication.

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Correspondence to Daniela Rus , Antonio Torralba or Jacob Andreas .

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Sharma, P., Gero, S., Payne, R. et al. Contextual and combinatorial structure in sperm whale vocalisations. Nat Commun 15 , 3617 (2024). https://doi.org/10.1038/s41467-024-47221-8

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DOI : https://doi.org/10.1038/s41467-024-47221-8

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Transition from covalent to noncovalent bonding between tetrel atoms.

The strength and nature of the bonding between tetrel (T) atoms in R2T··TR2 is examined by quantum calculations. T atoms cover the range of Group 14 atoms from C to Pb, and substituents R include Cl, F, and NH2. Systems vary from electrically neutral to both positive and negative overall charged radicals. There is a steady weakening progression in T-T bond strength as the tetrel atom grows larger, transitioning smoothly from a strong covalent to a much weaker noncovalent bond for the larger T atoms. The latter have some of the characteristics of a ditetrel bond, but there are also significant deviations from a classic bond of this type. The T2Cl4- anions are more strongly bonded than the corresponding cations, which are in turn stronger than the neutrals.

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