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AIBS Announces Winner of the IDEAL Leadership Award

The American Institute of Biological Sciences is pleased to announce Dr. Nyeema C. Harris as the 2023 winner of its Inspiring Inclusivity, Diversity, Equity, Acceptance, and Learning (IDEAL) Leadership Award. The IDEAL Award recognizes commendable leadership in advancing inclusion, diversity, equity, acceptance, accessibility, and learning in the biological sciences community. The award was presented by past awardee Dr. Steward T. A. Pickett on 30 November 2023 at AIBS’s Council of Member Societies and Organizations meeting, entitled “Expanding the Idea.”

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From PLOS Biology , Eggs of the mosquito Aedes aegypti survive desiccation by rewiring their polyamine and lipid metabolism

From PLOS Computational Biology , Rapid automated 3-D pose estimation of larval zebrafish using a physical model-trained neural network

From PLOS Pathogens , Divergence of TORC1-mediated stress response leads to novel acquired stress resistance in a pathogenic yeast

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Plos biology article types.

PLOS Biology considers works of exceptional significance, originality, and relevance in all areas of biological science, including both primary research, meta-analyses and Magazine articles . Our publication options are outlined below.

Research-based content

PLOS Biology publishes seven different types of research reports. All research articles are compatible with our easy, format-free submission process, and offer options for preprints, published peer review history, and publishing uncorrected proofs. Most, with rare exception, are also protected by our scooping policy , ensuring that your research will not be rejected for novelty within six-months of the publication of a complementary or confirmatory research publication. We evaluate all research based on the important questions it answers and its potential to impact an international scientific community as well as educators, policy makers, patient advocacy groups, and society more broadly.

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Preregistered research articles, methods & resources articles, meta-research articles, short reports, discovery reports, update articles, magazine articles.

Our magazine section features non-research articles that cover topical issues and are accessible to a broad audience while remaining scientifically rigorous. Magazine readers include scientists, scientific educators, students, physicians, patients, and the interested public. There is no publication fee for magazine articles.

Our magazine section is divided into Front Matter and In-depth Analysis subsections. Front Matter articles are short, focussed and provide opinion on topical issues, community resources or commentary on PLOS Biology articles. In-depth Analyses are long-form articles providing forward-looking analysis of a given topic, highlighting gaps in our current understanding or putting forward community recommendations or guidelines. We do not publish traditional review articles.

Most magazine articles are comissioned by the editors, but we do publish some unsolicted content. If you have a suggestion for any of our current magazine categories, we will consider the idea. Before finalizing the piece, we recommend that you discuss it with us by sending a presubmission enquiry via email to [email protected] . Before submitting, consult our General Guidelines for Magazine Submissions , as well as the guidelines for the specific article type you are interested in.

If you have any questions about submitting an idea or article for consideration in the magazine section, email our editorial team at  [email protected] .

General Guidelines for Magazine Submissions

Magazine content is intended for the broader biological community including students, scientists, and the educated general reader. It is therefore important that the writing style be concise, clear and accessible. Avoid specialist terms, abbreviations and jargon. Editors will make suggestions to make your piece more accessible, as well as cuts or additions that could strengthen the article. Our aim is to make the editorial process rigorous and consistent, but not intrusive or overbearing.

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As part of the PLOS Biology submission form you’ll be asked to provide a funding statement, which will be published with the article if the manuscript is accepted. Your funding statement should describe any funding that helped to support the work, as follows:

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Submit your manuscript through our submission system . When you start your submission, select the appropriate article type from the dropdown menu. Manuscripts can be submitted as DOC, DOCX, or PDF files.

Front Matter

Editorials are written in-house by members of the editorial staff or by members of the Editorial Board.

Perspectives

The Perspectives section provides experts with a forum to comment on topical or controversial issues of broad interest. They address controversial issues or those at the interface between science and policy or science and society; present a policy position aimed at influencing policy decisions; examine and make recommendations on scientific and publishing practices. These are meant to be short, opinionated, Op-ed type of pieces.

The ideal Perspective conveys a sense of urgency. Some things to think about would be:

  • Is this topic of immediate concern?
  • Is the topic relevant to a pressing regional or global issue
  • Does the piece offer a novel point of view on a scientific or policy issue, or on topical events?
  • Does is make specific, practical proposals to address the issue?

Controversial articles are welcomed, but the text should acknowledge that a position is in fact controversial and provide readers with enough background on the differing views.

Our Perspectives aim to engage a broad and diverse audience—it is therefore important to ensure that they are written in an accessible, persuasive, and stimulating style that appeals to both specialists and non-specialist readers. Perspectives are usually assessed in-house with our Editorial Board, but we reserve the right to peer-review them if needed. Commissioning does not guarantee publication. Editors work closely with authors to ensure that articles are written in an engaging, succinct, yet rigorous manner.

Guidelines for a Perspective

Example perspectives.

Lowe-Power T, Dyson L, Polter AM (2021) A generation of junior faculty is at risk from the impacts of COVID-19. PLoS Biol 19(5): e3001266. https://doi.org/10.1371/journal.pbio.3001266

Bourne PE (2021) Is “bioinformatics” dead? PLoS Biol 19(3): e3001165. https://doi.org/10.1371/journal.pbio.3001165

Primers provide concise and accessible context to a PLOS Biology research article of broad and current interest. Primers are commissioned and published alongside a research article that would benefit from additional context and/or explanation.

Above all, Primers should demystify an area of biology, avoid and/or explain technical jargon and provide critical and forward-thinking analysis about how the research article fits into the current state of the field and its future. A good Primer will briefly discuss (but not exhaustively review) what we know and what questions we have yet to answer for a particular field. It will then introduce the new findings and describe in roughly three paragraphs the advance represented in the related research article, highlighting its significance, not only for the discipline in question, but across disciplines. The Primer should then explain what the findings suggest in terms of next steps: what new avenues of investigation are opened, what new experiments can be tried, what new ideas can now be tested going forward? Ideally, Primers also offer insight into what big questions are likely to remain unanswered for many years (for whatever reasons).

We encourage the use of a figure to illustrate key concepts/mechanism/conclusions in an informative, easy-to-understand manner.

Primers are usually assessed by our Editorial Board, but we reserve the right to peer-review externally if needed. Commissioning does not guarantee publication. Editors work closely with authors to ensure that articles are written in an engaging, succinct, yet rigorous manner.

Guidelines for a Primer

Example primer.

Kazanova A, Rudd CE (2021) Programmed cell death 1 ligand (PD-L1) on T cells generates Treg suppression from memory. PLoS Biol 19(5): e3001272. https://doi.org/10.1371/journal.pbio.3001272

Guizetti J, Frischknecht F (2021) Apicomplexans: A conoid ring unites them all. PLoS Biol 19(3): e3001105. https://doi.org/10.1371/journal.pbio.3001105

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Community Pages provide individuals, networks and organizations with the opportunity to highlight resources, tools, or initiatives of benefit to the scientific community and beyond (including science education and public engagement in science). All resources or tools, and the outputs of initiatives must be open and accessible to all.

Contributors must resist the temptation of self-promotion and instead focus on conveying information to a diverse audience.Community Pages should be written in a succinct, accessible, semi-journalistic style that captures the interest of both specialists and non-specialist readers. We encourage the use of 1-2 figures to illustrate key concepts in an informative, easy to grasp manner; or the use of text boxes for background, self-contained information.

Community Pages are peer-reviewed and commissioning does not guarantee publication. Editors work closely with authors to ensure that articles are written in an engaging, succinct, yet rigorous manner.

Guidelines for a Community Page

Example Community Pages

Weissgerber TL (2021) Training early career researchers to use meta-research to improve science: A participant-guided “learn by doing” approach. PLoS Biol 19(2): e3001073. https://doi.org/10.1371/journal.pbio.3001073

McCullagh EA, Nowak K, Pogoriler A, Metcalf JL, Zaringhalam M, Zelikova TJ (2019) Request a woman scientist: A database for diversifying the public face of science. PLoS Biol 17 (4): e3000212. https://doi.org/10.1371/journal. pbio.3000212

Formal Comments

Formal Comments are intended to provide a formal outlet for the discussion and interpretation of research findings associated with specific articles published in PLOS Biology . They are designed to ensure that readers obtain a balanced view of a scientific or meta-scientific/policy question, especially in areas of debate/controversy. Formal Comments are peer-reviewed and indexed in PubMed.

Formal Comments must be coherent, concise, and well-argued, and are subject to the PLOS Biology criteria for publication . Editors will, as a matter of course, invite the authors of the original article to submit a response to the Formal Comment. Any revisions (of the Formal Comment or the response) will be shared with the authors of the associated comment.

Guidelines for Formal Comments

Example formal comment.

Rees WE, Wackernagel M (2013) The Shoe Fits, but the Footprint is Larger than Earth. PLoS Biol 11(11): e1001701. https://doi.org/10.1371/journal.pbio.1001701​ ​

Example Formal Comment - Response

Blomqvist L, Brook BW, Ellis EC, Kareiva PM, Nordhaus T, Shellenberger M (2013) The Ecological Footprint Remains a Misleading Metric of Global Sustainability. PLoS Biol 11(11): e1001702. https://doi.org/10.1371/journal.pbio.1001702

In-Depth Analysis

Essays are opinionated articles on a topic of interest to scientists and to a broader audience, including the general public. Unlike traditional review articles, which include a comprehensive account of a field, Essays take an imaginative approach to a provocative question, with an engaging but rigorous investigation of the problem. We encourage authors of Essays to select the most representative references to convey their points and avoid exhaustively covering the relevant literature.

The remit of Essays is very broad. They might:

  • take stock of progress in a field from a personal point of view
  • explore the implications of recent advances that promise to have broad-ranging consequences on a field
  • comment on a topical or controversial area of research
  • discuss key ideas or educational strategies to enhance understanding of fundamental biological questions
  • offer historical/philosophical reflections on contemporary biology
  • analyze scientific issues with policy implications

Our Essays aim to engage a broad and diverse audience—it is therefore important to ensure that they are written in an accessible, semi-journalistic style that captures the interest of both specialists and non-specialist readers. We encourage the use of figures to illustrate key concepts in an informative, easy to grasp manner; as well as the use of text boxes for background, self-contained information.

Essays are peer-reviewed and commissioning does not guarantee publication.

Guidelines for an Essay

Example essays.

Rees T, Bosch T, Douglas AE (2018) How the microbiome challenges our concept of self. PLoS Biol 16(2): e2005358. https://doi.org/10.1371/journal.pbio.2005358 Konig C, Weigelt P, Schrader J, Taylor A, Kattge J, Kreft H (2019) Biodiversity data integration—the significance of data resolution and domain. PLoS Biol 17(3): e3000183. https://doi.org/10.1371/journal.pbio.3000183

Unsolved Mysteries

Unsolved Mysteries discuss a topic of biological or medical importance that is poorly understood and in need of research attention—e.g., an unexplored or challenging question, an emerging opportunity, or a recent puzzling phenomenon. The articles are intended to stimulate the scientific community to think about future research possibilities outside their areas of expertise. The articles should be aimed at a very broad audience of biologists—an unsolved mystery in a neuroscience topic should be accessible to ecologists and biophysicists, for example.   The article should include a discussion of the basic science relevant to the topic, why it is biologically or medically important, what work has been done on the topic (if any), major challenges to understanding the question at hand, competing hypotheses, and what advances would be necessary to shed light on the problem. Ideally the structure of the article should reflect the mystery (e.g. subsections with questions as headings). The article should end with a discussion of possible means to a solution

Unsolved Mysteries are peer-reviewed and commissioning does not guarantee publication. Editors work closely with authors to ensure that articles are written in an engaging, succinct, yet rigorous manner.

Guidelines for an Unsolved Mystery

Example unsolved mysteries.

Margolis L, Sadovsky Y (2019) The biology of extracellular vesicles: The known unknowns. PLoS Biol 17(7): e3000363. https://doi.org/10.1371/journal.pbio.3000363

Vogels CBF, Ru¨ckert C, Cavany SM, Perkins TA, Ebel GD, Grubaugh ND (2019) Arbovirus coinfection and co-transmission: A neglected public health concern? PLoS Biol 17(1): e3000130. https://doi.org/10.1371/journal.pbio.3000130

Consensus View

Consensus View articles present a comprehensive analysis by an independent and usually multidisciplinary panel of experts who make specific recommendations on important scientific, publishing or policy issues.

Consensus Views are peer-reviewed and commissioning does not guarantee publication. Editors work closely with authors to ensure that articles are written in an engaging, succinct, yet rigorous manner.

Guidelines for a Consensus View

Example consensus view.

Brown TM, Brainard GC, Cajochen C, Czeisler CA, Hanifin JP, Lockley SW, et al. (2022) Recommendations for daytime, evening, and nighttime indoor light exposure to best support physiology, sleep, and wakefulness in healthy adults. PLoS Biol 20(3): e3001571. https://doi.org/10.1371/journal.pbio.3001571

Kent BA, Holman C, Amoako E, Antonietti A, Azam JM, Ballhausen H, et al. (2022) Recommendations for empowering early career researchers to improve research culture and practice. PLoS Biol 20(7): e3001680. https://doi.org/10.1371/journal.pbio.3001680

Vimercati G, Probert AF, Volery L, Bernardo-Madrid R, Bertolino S, Céspedes V, et al. (2022) The EICAT+ framework enables classification of positive impacts of alien taxa on native biodiversity. PLoS Biol 20(8): e3001729. https://doi.org/10.1371/journal.pbio.3001729

Retired Article Types

Book Review/Science in the Media. These short reviews critiqued books, films, plays, and other media that deal with some aspect of the biological sciences.  

  • Education. Although we no longer publish Education articles as a separate article type, we continue publishing them as part of the Education Series. The format selected (Essay, Perspective, or Community Page) depends on the aims of each article.
  • Historical and Philosophical Perspectives. The Historical and Philosophical Perspectives section provided professional historians and philosophers of science with a forum to reflect on topical issues in contemporary biology.
  • Obituaries.
  • Open Highlights. Written in-house by members of the editorial staff, Open Highlights used recent publication as keystones around which to nucleate a short synthesis of several related research articles from PLOS and the wider Open Access corpus.
  • Research Matters. Brief pieces by leading scientists explaining why the research carried out in their laboratories - and those of their collaborators and their colleagues - matters to lay audiences. 
  • Series. Series were recurrent themed articles on specific topics, including Education, Public Engagement with Science, Cool Tools, and Where Next?
  • Synopses. Selected PLOS Biology research articles are accompanied by a synopsis written for a general audience to provide non-experts with insight into the significance of the published work. They are commissioned only.

Post-Publication Notices

PLOS publishes Editorial Notes, Corrections, Expressions of Concern, and Retraction notices, as needed, to address issues that arise after a PLOS article has been published.

Biological Research

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Special series on Microbial Interactions

The nine articles of this special issue of  Biological Research  address biochemical and genetic determinants of microbial response and tolerance to stressors in different biological models and environmental contexts. Individual articles provide a broad exploration of our current knowledge of response to stressors, with a special emphasis on metal metabolism and toxic compounds.

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Special series on Antarctic Research

This special issue on Antarctic research in Biological Research comprises of recent studies, related to the discovery of several new enzymes and biotechnological applications that allow to expand the knowledge of Antarctic organisms and their potential applications.

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Cx43 hemichannels and panx1 channels contribute to ethanol-induced astrocyte dysfunction and damage

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Galectins in epithelial-mesenchymal transition: roles and mechanisms contributing to tissue repair, fibrosis and cancer metastasis

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Glutaminolysis regulates endometrial fibrosis in intrauterine adhesion via modulating mitochondrial function

Authors: Pei Chen, Chaoshuang Ye, Yunke Huang, Bingning Xu, Tianyu Wu, Yuanhang Dong, Yang Jin, Li Zhao, Changchang Hu, Jingxia Mao and Ruijin Wu

The long-chain flavodoxin FldX1 improves the biodegradation of 4-hydroxyphenylacetate and 3-hydroxyphenylacetate and counteracts the oxidative stress associated to aromatic catabolism in Paraburkholderia xenovorans

Authors: Laura Rodríguez-Castro, Roberto E. Durán, Valentina Méndez, Flavia Dorochesi, Daniela Zühlke, Katharina Riedel and Michael Seeger

MicroRNA-148b secreted by bovine oviductal extracellular vesicles enhance embryo quality through BPM/TGF-beta pathway

Authors: Karina Cañón-Beltrán, Yulia N Cajas, Vasileios Almpanis, Sandra Guisado Egido, Alfonso Gutierrez-Adan, Encina M González and Dimitrios Rizos

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Awareness and current knowledge of breast cancer

Authors: Muhammad Akram, Mehwish Iqbal, Muhammad Daniyal and Asmat Ullah Khan

Stress and defense responses in plant secondary metabolites production

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Fate of nitrogen in agriculture and environment: agronomic, eco-physiological and molecular approaches to improve nitrogen use efficiency

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Biological Research , formerly Archives of Experimental Medicine and Biology , was founded in 1964 and transferred to BioMed Central in 2014. An electronic archive of articles published between 1999 and 2013 can be found in the SciELO database.

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Manuel J Santos, Editor-in-Chief

Editor’s profile

Manuel J Santos, Editor-in-Chief

Dr Santos is an Associate Professor in the Faculty of Biological Sciences and Medicine at the Pontificia Catholic University of Chile.

Dr Santos received his MD from the University of Chile and his PhD in Cell and Molecular Biology from the Pontificia Catholic University of Chile. He majored in Medical Genetics at The John Hopkins University (USA) and The René Descartes University of Paris (France), and held a post doctorate position in Cell Biology and Genetics at the Rockefeller University (USA).

His research has focused on the biogenesis of cellular organelles, particularly peroxisomes. A pioneer in this field, his research lead him to discover a new type of human genetic disease, the peroxisomal biogenesis disorders, which include Zellweger Syndrome. More recently his research has centered on studying the role of peroxisomes in Alzheimer’s disease, and he also works in the field of bioethics.

Over the span of his career, Dr Santos has published more than 70 peer reviewed papers and been the President of the Society of Biology of Chile, the Genetics Society of Chile and the Bioethical Society of Chile.

About the Society

The Chilean Biology Society (Sociedad de Biología de Chile), previously the Biological Society of Santiago, was founded in late 1928 as a subsidiary of The Societé de Biologie of Paris, France. For several years the summaries of its communications were published in Comps Rendú of the Societé de Biologie du Paris. The Society is currently a member of the International Union of Biological Sciences (IUBS).

The Chilean Biology Society promotes theoretical and experimental studies and research leading to advancement in and dissemination of the biological sciences for the benefit of the community. To accomplish this, the Society organizes periodic scientific meetings in which scientists communicate, comment and discuss research carried out in Chilean or foreign research laboratories. In addition, relations and cooperation with similar domestic and foreign institutions are stimulated, and communication by all appropriate means of biological research carried out in Chile. 

Members of the Society will receive a discount on Biological Research 's article-processing charge when they provide a discount code (which members can obtain by emailing the Society) during the submission process.  The discounted article-processing charge for Society members is £1150 in 2023.

The Society also publishes Revista Chilena de Historia Natural ( Chilean Journal of Natural History, founded in 1897).

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2022 Citation Impact 6.7 - 2-year Impact Factor 7.2 - 5-year Impact Factor 1.241 - SNIP (Source Normalized Impact per Paper) 1.294 - SJR (SCImago Journal Rank)

2023 Speed 25 days submission to first editorial decision for all manuscripts (Median) 155 days submission to accept (Median)

2023 Usage  489,080 downloads 731 Altmetric mentions 

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  • Research Articles & Papers

Korunes, KL; Myers, RB; Hardy, R; Noor, MAF

Drosophila pseudoobscura is a classic model system for the study of evolutionary genetics and genomics. Given this long-standing interest, many genome sequences have accumulated for D. pseudoobscura and closely related species D. persimilis, D. miranda, and D. lowei. To facilitate the exploration… read more about this publication  »

Zipple, MN; Roberts, EK; Alberts, SC; Beehner, JC

Bartoš et al. (2021; Mammal Review 51: 143–153; https://doi.org/10.1111/mam.12219) reviewed the mechanisms involved in the ‘Bruce effect’ – a phenomenon originally documented in inseminated female house mice Mus musculus, who block pregnancy following exposure to a novel (non-sire) male. They argue… read more about this publication  »

Byrne, M; Koop, D; Strbenac, D; Cisternas, P; Yang, JYH; Davidson, PL; Wray, G

The molecular mechanisms underlying development of the pentameral body of adult echinoderms are poorly understood but are important to solve with respect to evolution of a unique body plan that contrasts with the bilateral body plan of other deuterostomes. As Nodal and BMP2/4 signalling is involved… read more about this publication  »

Wang, Q; Xu, P; Sanchez, S; Duran, P; Andreazza, F; Isaacs, R; Dong, K

BackgroundInsects rely on their sense of smell to locate food and hosts, find mates and select sites for laying eggs. Use of volatile compounds, such as essential oils (EOs), to repel insect pests and disrupt their olfaction-driven behaviors has great practical significance in integrated pest… read more about this publication  »

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biological sciences research paper

Advice to a Young Mathematical Biologist

  • Perspectives
  • Open access
  • Published: 09 April 2024
  • Volume 86 , article number  52 , ( 2024 )

Cite this article

You have full access to this open access article

  • Paul A. Roberts   ORCID: orcid.org/0000-0001-5293-6431 1  

16 Altmetric

This paper offers advice to early-mid career researchers in Mathematical Biology from ten past and current Presidents of the Society for Mathematical Biology. The topics covered include deciding if a career in academia is right for you; finding and working with a mentor; building collaborations and working with those from other disciplines; formulating a research question; writing a paper; reviewing papers; networking; writing fellowship or grant proposals; applying for faculty positions; and preparing and giving lectures. While written with mathematical biologists in mind, it is hoped that this paper will be of use to early and mid career researchers across the mathematical, physical and life sciences, as they embark on careers in these disciplines.

Avoid common mistakes on your manuscript.

1 Introduction

Early-mid career researchers in Mathematical Biology face a particular set of challenges. As they develop in their career, a number of skills need to be learnt, most of which are not taught in a typical undergraduate degree. In this paper, ten leading mathematical biologists—all current or former Presidents of the Society for Mathematical Biology (SMB)—share their advice on a number of areas of particular interest to early and mid career researchers. While written with mathematical biologists in mind, much of the advice presented here is of relevance to any researcher working in the life, physical or mathematical sciences. It is hoped that this paper will prove a valuable resource to early and mid career researchers as they make the first steps in their academic journey, providing a helping hand from those who have trodden the road before them.

The idea for this paper occurred to me following the excellent past Presidents’ panel discussion, organised by Prof. Heiko Enderling, at the 2023 SMB conference, held at The Ohio State University in Columbus, Ohio. This was an inspiring session, with many useful insights shared by some of the greats in the field. It struck me that it would be good to capture the insights from some of these researchers in a permanent way, and that this would be of particular interest and benefit to early/mid career researchers.

All of the living past and current SMB Presidents were contacted, and to those who were able to contribute, a series of questions was posed, inviting their top tips and advice in a number of areas relevant to early/mid career researchers. These questions consisted of a subset of ten specific topics, together with two, more general questions, which were posed to all contributors. Responses were then compiled, ordered and edited to provide coherent guidance in each area.

The advice offered here is not intended to be exhaustive. Rather, it is hoped that this will be a starting point, bringing together guidance on a range of topics into a single place, leaving the reader to explore specific areas in greater depth as desired. As with any advice, it is left to the reader to follow or leave at their discretion.

The title of this article is a homage to Prof. Sir Peter Medawar’s book ‘Advice To A Young Scientist’ (Medawar, 1979 ) and to the later multi-author chapter ‘Advice to a Young Mathematician’ in The Princeton Companion to Mathematics (Atiyah et al., 2008 ); both of which are recommended. To the best of my knowledge, this is the first paper to offer guidance specifically to early/mid career mathematical biologists.

In what follows, we cover ten specific topics: ‘Deciding if a career in academia is right for you’, ‘Finding and working with a mentor’, ‘Building collaborations and working with those from other disciplines’, ‘Formulating a research question’, ‘Writing a paper’, ‘Reviewing papers’, ‘Networking’, ‘Writing fellowship or grant proposals’, ‘Applying for faculty positions’ and ‘Preparing and giving lectures’; together with two general topics: ‘What do you wish you had known when you were an early-mid career researcher?’ and ‘Some final words of advice’. These sections can be read in any order or in isolation, depending on the needs and interests of the reader.

2 Deciding if a Career in Academia is Right for You

Many of us may wonder if the academic path is the right one for us. This question might occur when deciding whether or not to pursue a doctorate, to apply for postdoctoral or faculty positions, or even whether to remain in academia, having obtained a permanent position. Whatever your stage, the following advice may be helpful to bear in mind.

Make a list of things that are important to you, what you want to accomplish in your professional life and what will make you happy going to work every day for the rest of your life.

If you are self-driven, have lots of questions, like to work and meet with people, and like to share your work in different venues (e.g. papers and presentations), you could consider a career in academia.

A career in academia is not easy.

You need to consider what kind of academic you would like to be: more research focused, or more teaching focused. Do you want to have a large or small group, or work at a large or small school?

You also need to consider that there is a lot more to an academic job than what you may have experienced during your undergrad/PhD/postdoc. Talk to PhD students, postdoctoral fellows and faculty to find out what they do from day to day, and get a sense of what the job entails. Learn what they like about their roles and what they wish was different.

Talk to many professionals outside academia about their experiences.

3 Finding and Working with a Mentor

The concept of a mentor is a familiar one, both historically and in popular culture: Plato had Socrates, Luke Skywalker had Obi-Wan Kenobi, Bertrand Russell had Alfred North Whitehead and Frodo had Gandalf. Though familiar, finding and developing such a relationship can be difficult. Here are some expert tips on how to navigate this area.

The mentor is probably the most important part of your academic career.

Finding a mentor

You can find a mentor in many different ways:

Get to know the faculty in your research institution;

Talk to people at conferences;

Participate in mentoring programs.

Have a one-on-one conversation about ideas and what the potential mentor looks for.

Identify what YOU need from a mentor. Make sure that you communicate your needs to a prospective mentor and evaluate if they can help you in your academic journey.

Be honest about your interests.

The best science is not necessarily done by the mentor that best serves your needs, though make sure the research approach of the mentor excites you.

Try to visit and meet members of the potential mentor’s group, talking with former/current students/collaborators/mentees. This is important, not least because it will enable you to check the potential mentor’s reputation. This will also help you to evaluate if their mentorship style is right for you.

Take into account the breadth of the institution, and especially the department, and the potential to interact with others outside the group.

Working with a mentor

Expect the relationship to develop and change over time.

Mentorship can be developed very naturally—through discussion at conferences and workshops, and then some emails in between.

Make time for the relationship to develop in social contexts in connection with or outside of research discussions (e.g. coffee/tea/beer etc. time).

Do not agree to work on a project if it does not align with your interests, but be open to suggestions of new projects or research questions/approaches.

4 Building Collaborations and Working with Those from Other Disciplines

Given the intrinsic interdisciplinarity of mathematical biology, the ability to build and grow fruitful collaborations is key to developing biologically faithful and impactful models. This is not something that is usually taught at the undergraduate level, but rather is learned on-the-job, with a degree of trial-and-error(/-improvement). While this is a rite-of-passage that all mathematical biologists must pass through—and, indeed, a lifelong learning process—here are few tips to smooth the way.

Listen carefully to lectures on topics from other disciplines, and read review papers carefully to identify what questions motivate that discipline/topic. Ask yourself in what way you could contribute to answering such questions using your skill-set.

Learn a lot about the subject matter. Attend experiments when they are being done.

Follow your heart and make the effort to work with people who you find interesting and exciting.

Find someone who is open to theoretical approaches and who is a person with whom you get along really well.

It can take a while to build a good collaboration, so be patient, and invest in a few possible directions. Usually one or another will eventually pan out.

Trust that your collaborators know what they are talking about.

Ask a lot of stupid questions, balancing keeping expectations low with occasional moments of surprising brilliance.

Be clear about shared responsibilities.

Be willing to suppress your ego. Remember that what makes the work interesting is the experiments rather than the theory.

Learn the jargon of the biological discipline(s) relevant to your research.

Explain your ideas in plain English. Do not expect potential collaborators to know or be familiar with mathematical jargon or methods.

Explain what your methods could do to help test hypotheses or to analyse data, or to help with the design of experiments.

Try to get in a situation where you can help design the experiments to provide data needed for analysis.

Biological experiments usually cost a lot of money and take a lot of time. Do not expect that a collaborator will immediately agree to do your favourite experiment. (Sometimes, you have to make-do with data from the literature.)

Be willing to pay any students who may work on the theory and perhaps other costs associated with doing the experiments. Working on joint grants is one way to do this but that takes patience.

5 Formulating a Research Question

As any Douglas Adams fan will know, the key to making discoveries lies in asking the right questions. The following advice may be helpful in deciding upon a research topic and what question(s) to ask.

Find a problem that really interest you, about which you are passionate and want to know the answer, and do not care what others think.

Be driven by the research question, not by the methods you will use.

Find a topic that will potentially expand the field, not something that is just incremental.

There are many kinds of research questions: explaining a puzzling data set; testing a hypothesis for some mechanism; finding some optimal strategy; making a long-term prediction. Each case would imply a different strategy.

To find new interesting quantitative questions, read a number of recent review papers on the topic of your choice. Find sentences such as ‘The mechanism for this observed behaviour is poorly understood’, and look for key areas where a knowledge-gap is identified. Be sure that these questions are not just experimental ones. Be sure that some facts are known and/or some data is available on which to construct your model, for example.

Be open to approach by colleagues from other disciplines. Listen to their ideas and motivation, and assess whether your skills could be useful, or whether other colleagues have just the right tools to be helpful.

6 Writing a Paper

Most mathematical biologists begin by taking an undergraduate degree in mathematics, spending the bulk of their time working through a series of problem sheets. As such, when they come to do a doctorate and begin writing their first paper, it may be some years since they were required to write at any length. Further, the process of writing an academic article is unlike that of writing a secondary/high school essay. The following advice should be of help in providing a possible approach to writing papers, while also highlighting some common pitfalls.

Some general points

Do your literature review well: you do not want to submit a manuscript that is missing important references.

Spend time critically reviewing your results. Do they make sense? What are some questions that reviewers might have? Are any results difficult to understand?

Do not make the paper too long. Figure out what you want to say in a direct way.

A possible approach

Let us assume you have wrapped up an original piece of research and you are ready to write your first paper. The first step is to get your work organized in a logical, convincing fashion. You have probably already done this in preparing for your committee meetings, student presentations and poster sessions. A good MS PowerPoint presentation is a great place to start.

Next, consider the audience you want to reach. Defining your audience will dictate what journal to submit to and also what background information you need to include in the introduction.

Write an outline, using the standard format of a scientific publication.

Title: start with a working title; it may change later.

Abstract: write this last!

Introduction: make an outline, with your target audience in mind.

arrange your research in a logical fashion;

sketch your figures in some detail (and write cogent legends);

consider what tables you will need;

push some results to ‘Supplementary Material’ to stress the main points.

Discussion: make notes along the way, but write this part later.

Now that you are ready to start writing, keep the following Four Cs in mind.

Correct. Everything you write must be scientifically correct, to the best of your knowledge. Check each sentence and every equation. Make sure you have provided the correct parameter values for all your calculations.

Clear. Now that everything is correct, you must communicate your results clearly to your audience. You do not have to tell people what DNA means, but do not skip over important things that the reader needs to know. It is helpful here to get someone else’s point of view—on joint authored papers, it is the responsibility of all authors to make sure that what is written is clear. Some important points to note:

Often papers are not structured in a logical way, and read like a stream of consciousness. Look at the logical structure of your flow of ideas to make sure that your argument will make sense to your readers.

In this regard, basic grammar rules are important, especially coherent paragraphs with topical sentences. Do not let your paragraphs get too long; most long paragraphs can be broken into two or more separate ideas.

Watch how you use pronouns—they can be dangerous. You may know what your pronoun is referring to, but your reader may not. When a reader comes across a pronoun, he/she typically assumes that the pronoun refers to the last noun mentioned in the previous sentence. If the reader has to look further back, he/she will likely get lost. The simple fix is to repeat the noun, so it is absolutely clear what you are talking about.

Another mistake of non-English writers is overloading the subject of a sentence, using too many modifiers for a noun, or other nouns as modifiers of the main noun. It can be difficult for the reader to figure out what the noun of the sentence is, and which words are modifiers. The simple fix is to use prepositional phrases and dependent clauses to expand on a noun, rather than going beyond a few adjectives. For example, ‘the budding yeast cell cycle spindle assembly checkpoint’ should be ‘the spindle assembly checkpoint of the budding-yeast cell cycle’. Another good example of an ‘overloaded noun’ of a sentence is: ‘Initiation and progression of the cell cycle are considered to occur in response to the timely ordered transcriptional, post-transcriptional, and posttranslational regulation of the cell cycle (cyclin/cyclin dependent kinase [CDK]) machinery components ’. The italicised phrase is the object of the passive verb construction ‘are considered to occur in response to’. The object is ‘components’ and the preceding words all modify ‘components’. It would be clearer to write: ‘Progression through the cell cycle is thought to be based on the temporally ordered activation of cyclin/cyclin dependent kinases (CDKs), which are regulated by a complex molecular network of transcriptional, post-transcriptional, and post-translational controls’.

Concise. After you are sure your text is correct and clear, then go through it carefully to get rid of annoying repetitions that may have crept in. Pare things down to a minimum without destroying clarity. State your main points several times (in the Abstract, Results and Discussion); as for everything else, just say it once.

Compelling. Finally, polish up the writing. Use MS Word’s thesaurus to find exactly the right word to get your idea across. Make the paper easy/pleasant/attractive to read, so people will recommend it to others.

7 Reviewing Papers

Reviewing your first paper can feel like a daunting task, with a weight of responsibility to make an accurate and fair assessment. The following tips should prove useful both to first time reviewers, and to those with some experience under their belts.

Only accept reviews for manuscripts you are competent to assess.

Make sure you are familiar with other research in the field, so you know how novel the work is.

Do not take on another review if you already have one.

Negotiate with the editor a timeline that suits you and not just them.

Do not allow deadlines to make you do a superficial job.

Try to be fair and write the kind of review you would like to receive.

Read the introduction and discussion first, to get a feel for what the authors want you to get from the paper, then read the whole manuscript to see if the results match with this.

Do not question the motives but focus on the results.

Do not be sucked in by overhype.

Always ask for codes to be shared if they are not already.

8 Networking

Our scientific research is not conducted in isolation, but rather as part of a community. As such, developing relationships with fellow scientists and mathematicians is an important part of any mathematical biologist’s career. Indeed, the contacts we make now could be our future collaborators, reviewers or employers. We often use the word ‘networking’ to denote the practice of making and developing these relationships, particularly in the context of conferences. While most would agree that networking is important, many of us are unsure of how best to go about it. This problem is especially acute for early and mid career researchers, who may wish to speak with senior researchers, but are unsure of how to introduce themselves, or manage the conversation. Here is some advice on how to approach it.

Study the conference program before the meeting. Identify 4–6 people with whom you might be interested in meeting. These include people that are senior to you and also people that may be more junior. Email them ahead of time and schedule meetings during coffee or lunch breaks early in the conference.

Do your homework before approaching a specific scientist. If you have some knowledge of their research, then a simple introduction can be had through a compliment or question about a specific piece of work. All scientists love to discuss their research, so if you have a question or insight to share they almost always want to hear it.

Find an appropriate time to approach someone and be polite. A good time to introduce yourself might be at a reception or poster session; another meeting can always happen after the initial introduction.

Go to poster sessions, or better yet, present a poster. Poster sessions are a great networking opportunity.

Go to after program events (e.g. dinner, drinks and hikes). The best networking happens off campus.

Ask a mentor, or another scientist who knows the researcher you would like to meet to introduce you and help break the ice.

9 Writing Fellowship or Grant Proposals

Writing good fellowship and grant proposals is something of an art form in itself. As an early/mid career researcher writing your first proposal, it is easy to feel bewildered, not knowing quite where to start. It is hoped that the following guidance will set you in the right direction.

Know your audience. Grant proposals are diverse and depend on the specific call in regard to what is required, what the review procedures will be and who will be the reviewers; therefore, always read the specific call/request for proposals carefully, so that you know what is expected and what the deadlines are.

If appropriate, discuss your proposal with the specific program officer / agency’s program manager, if there is one, to be certain that what you are proposing fits the guidelines for support. They can often give good advice on what will be received well versus what will not be. Ask if the proposal will be reviewed by more than one group.

Follow faithfully any guidelines that are given by the funding body, e.g. if you are asked to write the proposal in 12pt Arial font.

Ask a successful grant writer to share some of their previous grants—the structure and level of detail as well as visual support for a proposal varies greatly and needs to be tailored to the specific call.

Collaborate with someone who has been successful in obtaining support in the past from the agency.

Try to plan ahead so that you have time to share a draft of your proposal with your peers or mentor for feedback.

Make sure you have an exciting and innovative idea in the first place! Remember that the person(s) reviewing your fellowship application / grant proposal will probably have many others as well, so it is important to ‘grab the reviewer’s attention’ from the outset. Aim to write a factual but stimulating first paragraph which will make the reviewer want to read on and find out about the exciting project you are proposing.

Ensure also that your idea is appropriate, carefully stating the goals of the proposed work somewhere near the start of the proposal.

Write passionately from the heart and be ‘achievably ambitious’.

Justify any claims you make and give as good an argument as you can that what you are about to do can be achieved.

Do not try to cram every possible thing you can think of into the proposal; rather, be focused and have a good timeline with appropriate milestones.

Most grants are scored badly because the reviewer could not understand what you really wanted to do. Far fewer fail because of a flawed idea, so make a big effort in articulating your ideas as clearly as possible; visual support can really help e.g. cartoons, schematics and graphs.

Emphasize why you are the appropriate person to do the work.

Almost nobody is successful with the first iteration of a grant, so it is good to submit to a call on the first round and then resubmit on subsequent rounds, integrating reviewer feedback.

For more on this topic, see ‘Notes on Writing and Getting Grants’ by Lou Gross: lgross.utk.edu/grantwriting.txt .

10 Applying for Faculty Positions

Many early/mid career researchers may be relatively inexperienced in writing job applications, or be unsure of how best to present themselves to potential employers. The following advice is given with faculty applications in mind, though many of the tips are also relevant to applications for postdoctoral positions.

Do not apply for a job you do not want—you might get it.

Publish your work when it is ripe, even if it is not perfect.

Collaborate, but be sure to establish your own identity.

Think about who you are: a fox or a hedgehog? This reference comes from a 1953 book by the philosopher Isaiah Berlin, in which he quotes the Ancient Greek poet Archilochus as saying that ‘ the fox knows many things, but the hedgehog knows one big thing ’ (Berlin 1953 ). In the context of mathematical biology, think about whether you see your research as centring around one topic, or as touching on many topics, perhaps with a more abstract common theme. Both are valid ways to work, but it is good to think about who you are, to avoid getting pushed or pulled in directions that might not fit.

Make sure your CV is up-to-date and is written well.

Do not try to exaggerate anything, e.g. do not list lots of unpublished papers.

Cover letter

Read the job advertisement carefully and write a relevant, engaging cover letter, outlining your background, your current research interests, your future research plans and your teaching philosophy / teaching experience.

Be explicit as to why you are appropriate for the position. Spend some time finding out about the department and the university in general, and aim to include in your cover letter how you feel you could fit in and connect with the teaching and research that is going on in the department and also potential collaborations elsewhere in the university (e.g. departments of biology / life sciences and medicine).

Be enthusiastic.

Statement of research interests

Summarize in one paragraph the main results from your prior work.

Lay out a research plan, possibly with several different components. Think of this as a research plan for the initial 5–10 years of your career. Where do you want to be, what ‘big’ questions do you want to work on and how do the smaller ones fit into this?

Statement of teaching interests

Summarize what your teaching experience has been.

Give a bit of your teaching philosophy and provide examples of how you have applied it (e.g. projects you developed/used in a course you taught, or implementation of computer-based examples).

State your teaching objectives over the next 5 years—what courses and seminars you might like to teach/develop, what texts you might be interested in developing. Tie this in to particular courses the university provides.

Make sure you have good referees who will provide strongly supportive but not hyperbolic references.

Make it as easy as possible for your referees to write a letter for you—give them all the material you are sending out, explaining how to address letters and providing the links to the adverts for positions you are applying to.

Make sure the referees know which jobs are the ones of most interest to you.

Perhaps ask your referees to contact (email or phone) anyone they know at your top choice positions to alert them to your application.

For more on this topic, see ‘Applying for a job, haggling for a job, and keeping a job’ by Lou Gross: lgross.utk.edu/gettingjobs.postdocs.mbi06.txt .

11 Preparing and Giving Lectures

The average early/mid career researcher will have attended hundreds of lectures during their undergraduate studies; some of them better than others. While many PhD students will get experience of leading or assisting with tutorials and problems classes, opportunities for lecturing experience arise less frequently. The following guidance should be of help to postdoctoral researchers and new faculty preparing to give their first lectures.

Find the lecturing style that you are most comfortable with e.g. ‘chalk and talk’, slides, iPad/Tablet etc., and practise at it.

Do not practise too much—talks can sound really canned with too much practice. Put another way, too much practice can stand in the way of ‘presence’ during a talk, thinking a little on your feet and taking a few chances.

Prepare your notes in advance and try to connect with external material e.g. books, research articles, online videos etc.

Think about your main point during your pre-lecture preparation.

Your lecture has to fit your audience. Do not attempt to give the same lecture to biologists and to mathematicians.

Optimise your slides: a maximum of 20 words per slide, brief bullet points, self-contained and easy to follow.

Do not include something on a slide if you do not want to talk about it.

Go to the lecture theatre before you start the course and work out where everything is so that you can begin the first lecture without any glitches or delays.

Try to be enthusiastic and passionate in your delivery and to ENJOY giving the lecture.

Never forget that it is about the material, not about you.

Consider introducing your talk with interesting scientific questions, and returning to those at the end to show that you ‘solved them’. Merely reproducing a behaviour with a model is not very interesting unless you can show new insights or novel predictions.

Aim to engage the students rather than just lecture for one hour e.g. stop regularly and ask questions, ask the students to suggest ways to complete a piece of algebra or offer the answer to a problem.

Provide plenty of motivation and background for the audience to understand the main ideas. Be sure to emphasize the significance and goals.

Give plenty of worked examples in the class which underpin any piece of theory you deliver.

Be sure to EXPLAIN everything. Your audience will appreciate that.

Make the lecture interesting. Use some colour, make fonts nice and large, consider some humour if possible, once you gain confidence.

Make a deliberate mistake now and again—this can encourage the students to engage and when they get the correct answer it gives them confidence. It also shows them that you are not infallible!

Never go over time.

12 What Do You Wish You Had Known When You were an Early-mid Career Researcher?

In addition to asking our seasoned professionals for guidance on specific questions, their advice to early and mid career researchers was also sought at a more general level, as recounted in this section and that which follows. First, in this section, we explore the hard-earned knowledge that our experts wish they had possessed when they were early/mid career researchers.

Seeking advice

Do not be shy about getting advice, particularly on grant proposals.

Understand how the system at your institution works, who to go to for advice/assistance and how to work around arcane rules that constrain your ability to advance your research and teaching.

Career planning

Think a few years ahead but do not let long-term planning stand in the way.

Early in your career, it is common not to know what you really want and that is OK, since you have not experienced enough yet.

‘When I started as a graduate student, I had a very specific plans about what I wanted to study: quantum chemistry. Like most mathematical biologists, I never intended to be one! I stumbled onto the field through my professors and mentors. So keep your eyes open, see what catches your interest, see where new research areas are opening up and where you can make a contribution. Be flexible, find your place in the world and have fun!’

Think strategically about what you will gain from a specific position and how it might lead you to new opportunities in the future.

‘Failure’ and rejections

Be ready to accept rejections and how to move on effectively from these, such as re-applying for grants to either the same agency which initially rejected it or to try someplace else.

Do not take failure personally; academia is a constant source of failure, whether it is papers, grants or even your science. Failure is the only way we can learn; of course it still stings, but know that this is a universal pain we all feel as scientists, and it is also temporary, as it will drive resubmissions, rewriting, reframing and ultimately success.

Lack of a job offer, or interview, may just be due to various political factors in a given department/unit that have nothing to do with your excellence. Therefore, do not let such ‘rejections’ affect your morale and work.

Do not skip your postdoc; exploit every second of it. It is a rare time in your scientific career that you will never have again—both scientific freedom and no financial concerns.

Have fun! Most research ideas come outside the laboratory; on a walk, while exercising, or while having dinner with friends. A lot of great ideas start out on a napkin.

If you are not excited about a problem, the work is not going to be worthwhile.

You can work on anything you want to, independent of your field, as long as you are willing to learn the new area.

Keep doing good work, even when the job-market looks bleak. Eventually this will pay off.

Take the time to learn new skills.

Do the hard work yourself.

‘I wish I had known’:

LaTeX —‘I wrote my PhD thesis using troff’ ( wikipedia.org/wiki/Troff ).

More numerical analysis.

The Sobolev Embedding Theorem (just kidding!).

Sharing your work

Put real effort into making your science as accessible as possible—the more people who understand it, the better it will be cited and shared.

Grab any opportunity you can to present your work, even if you find it difficult. It will help you understand your own work better and expose the community to what you are doing and critically provide valuable feedback.

Open science is a golden opportunity to share your work before it is published, embrace it. Share your papers on preprint servers (e.g. bioRxiv and arXiv), and your code and data on public repositories (e.g. GitHub).

Collaboration and networking

Work with people you like, in labs that are happy and have a good community ethic. Do not try to work with people simply because of their prestige.

Use administrative roles to build collaborations.

Networking with others (in your unit and at conferences) is very important. Consider sending your e-publications to the top researchers with a short email. Many are busy, so may not answer, but some will.

Do not be shy at a conference. Schedule meetings ahead of time to make sure you are not alone during coffee breaks.

Maintain contact with those you have met who might help your career advance in the future.

Organising your time

‘I did not realize how much time I would spend in service-related activities. I sit on many committees. My service takes about one full day per week.’

‘I did not realize how much grant writing I would be doing. I had to learn how to write grants for many different reviewing bodies. This can take time, but can also be helpful in that you then understand how to talk about your research with many different audiences.’

Teaching and mentorship activities can occupy much of your time. Make sure that you structure your week so that you have research blocks that are long enough for you to remember what you are doing, and get some work done to review and advance your projects.

Get home in time for dinner with your family.

13 Some Final Words of Advice

In this last section, we offer some final words of advice, not covered by the previous sections.

Community, collaboration and care

Collaborate broadly and build your network of collaborators in ways that stretch your research to fields that might be far from your formal education.

Team science is truly a gift for mathematical biology. It is being embraced across many different disciplines and is a golden opportunity to work across fields with creative teams, where the team is far more powerful scientifically than any of the individuals. If you can work with a team, jump at the chance.

Develop a community around you, but do not feel that you need to collaborate with everyone. Deliberately keep some experts in your field at ‘arm’s-length’ as you will need people to review your file at tenure and promotion, for grants, and your manuscripts for publication.

Care about your community—take time to contribute, to nurture and enrich your community as it will not continue without it.

Make time for self-care; something outside of science even if it is with scientists. It is important to recharge your creative and non-creative batteries and that cannot happen if you use them all the time.

Most scientists are good people even if they may ask difficult questions and appear intimidating—they are a scientist just like you and care about similar things.

Always be honest, even if it means admitting mistakes, being truthful will always pay dividends in the end.

Work on what you want, not on what other people think you should.

Enjoy yourself, have fun, work on problems that you are really interested in and passionate about.

Aim high. Always ask ‘could my work be better?’ Do not settle for the first result and hurry to publish—do your due diligence and make that sure every piece of work has the highest impact it can.

Mathematical biology is a subfield of biology. Talk to biologists as often as you can. Let their questions guide your research.

Do not be afraid of data. Indeed, look at the data! You may find something that you did not expect that is more interesting than what you did expect.

Understand what it means to calibrate and validate a mathematical model. Not every curve that fits data makes a model plausible and it does not guarantee predictive power (if that is what you are aiming for).

Do not be a one-trick pony. It will help your career if you become the go-to person in the world on a particular topic, but do not constrain yourself to this area. Look for side-projects that may be well outside this area of focus.

Be willing to take risks and try out new/alternative things. It is only by failing that we discover what does not work and this helps put us on another track that perhaps will work. Do not be afraid to ‘fail’. The following quote from John Backus (who invented FORTRAN) illustrates this point:

‘ I, myself, have had many failures and I’ve learned that if you are not failing a lot, you are probably not being as creative as you could be—you aren’t stretching your imagination. You need the willingness to fail all the time. You have to generate many ideas and then you have to work very hard only to discover that they don’t work. And you keep doing that over and over until you find one that does work. ’ — mathshistory.st-andrews.ac.uk/Biographies/Backus/quotations/

Communication

Learning to communicate in writing and orally is just as important as doing advanced research. Your funding and the respect you achieve will depend on your ability to explain your work and convince others that it is significant.

Work to build your vocabulary to be able to communicate with experts in fields quite different from your own.

Get some formal training from science communication experts to assist you in being able to discuss your work with non-scientists and journalists. Do not be bashful about tooting your own horn.

For more on careers in academia, see ‘Careers in Academia: How to Enhance your Chances for Success’ by Lou Gross: lgross.utk.edu/eeb504Spring2021.html .

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Acknowledgements

This paper would not have been possible were it not for the sagacious advice, generously offered by the following current and former SMB Presidents: Prof. Frederick R. Adler (University of Utah); Prof. Alexander R. A. Anderson (H. Lee Moffitt Cancer Center & Research Institute); Prof. Mark A. J. Chaplain (University of St Andrews); Prof. Leah Edelstein-Keshet (University of British Columbia); Prof. Heiko Enderling (The University of Texas MD Anderson Cancer Center); Prof. Leon Glass (McGill University); Prof. Louis J. Gross (University of Tennessee); Prof. Jane M. Heffernan (York University); Prof. Simon A. Levin (Princeton University); and Prof. John J. Tyson (Virginia Polytechnic Institute and State University). PAR acknowledges financial support from the University of Birmingham Dynamic Investment Fund.

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Roberts, P.A. Advice to a Young Mathematical Biologist. Bull Math Biol 86 , 52 (2024). https://doi.org/10.1007/s11538-024-01269-1

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Received : 02 February 2024

Accepted : 12 February 2024

Published : 09 April 2024

DOI : https://doi.org/10.1007/s11538-024-01269-1

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The Principles of Biomedical Scientific Writing: Citation

Zahra bahadoran.

1 Nutrition and Endocrine Research Center, Research Institute for Endocrine Sciences, Shahid Beheshti University of Medical Sciences, Tehran, Iran

Parvin Mirmiran

2 Department of Clinical Nutrition and Human Dietetics, Faculty of Nutrition Sciences and Food Technology, National Nutrition and Food Technology Research Institute, Shahid Beheshti University of Medical Sciences, Tehran, Iran

Khosrow Kashfi

3 Department of Molecular, Cellular and Biomedical Sciences, Sophie Davis School of Biomedical Education, City University of New York School of Medicine, New York, United States

Asghar Ghasemi

4 Endocrine Physiology Research Center, Research Institute for Endocrine Sciences, Shahid Beheshti University of Medical Sciences, Tehran, Iran

Citation, the act of properly referring to others’ ideas, thoughts, or concepts, is a common and critical practice in scientific writing. Citations are used to give credit to own work, to support an argument, to acknowledge others’ work, to distinguish other authors’ ideas from one’s work, and to direct readers to sources of information. A good citation adds to the scientific prestige of the paper and makes it more valuable to the reader. The citation has three basic elements: quoting from others, an in-text reference to the source, and bibliographic details of the source. Beyond technical skills, the citation needs an in-depth knowledge of the field and should follow basic rules, including the selection of relevant and valid sources, stating information/facts from others’ work, and referring to others’ work accurately and ethically. Several systems and styles are used to cite scientific sources; however, the most commonly used systems in medical sciences are ‘author-date’ systems (e.g., Harvard system) and numerical systems (e.g., Vancouver system). Here, we discuss how to make an accurate, complete, and ethical citation, and provide simple and practical guides to organize references in a scientific medical paper.

According to Merriam Webster dictionary, the word citation is defined as “an act of quoting,” where quote means “to speak or write from another usually with credit acknowledgment.” In scientific communication, the citation is commonly considered a technical practice to refer to the source in the text to represent that the information is derived from an external source ( 1 , 2 ). The citation is more than just referencing; it provides more value than acknowledging the source of literature being reviewed and making a reference list ( 2 ). Citation helps the authors put their work in connection with previous ones, to tell the story artfully, to acknowledge others’ work, and to contextualize study findings ( 2 - 4 ). A well-referenced paper supports the novelty and value of the work and improves its visibility ( 5 ).

Regardless of its importance, the citation may be the least noticed aspect of a scientific manuscript ( 6 ). Citation errors, including misquotation and errors in the bibliography, are common in the medical literature. The prevalence of misquotations in different journals ranges from 10% to 20% ( 7 ), and about 50% to 70% of references of published papers contain at least one error ( 8 , 9 ). Inaccurate quotations are displeasing for the cited author, misleading for the reader, and initiate circulation of false facts ( 7 , 10 ). The consequences of bibliography errors include difficulty in reference retrieval, limitation for the reader to read more widely, failure to credit the cited author(s), and inaccuracies in citation indexes.

Following our previous guides on how to write Introduction ( 11 ), Material and Methods ( 12 ), Results ( 13 ), Discussion ( 14 ), Title ( 15 ), and Abstract and keyword ( 16 ) of a hypothesis-testing paper, here, we provide a practical guide on importance and function of the citation of a scientific medical paper. We discuss how to select sources for the citation, quote information from others’ work, refer to the sources, and make an accurate reference list for a scientific biomedical paper.

2. Functions of Citation

The citation is used to give credit to an author’s work, acknowledge other’s work, distinguish an author’s ideas from others, direct readers to original sources of information, and avoid plagiarism ( 2 - 4 , 17 ). Citation help readers to understand the work, to justify the conclusions, to judge the novelty and scope of the manuscripts ( 18 ), and critically evaluate what contribution the study makes ( 4 , 19 ). Using proper citation is the only way to use the work of others and not commit plagiarism ( 20 ). To avoid plagiarism, authors need to accurately refer to the most relevant publications and cite facts and conclusions ( 1 , 20 ). Functions of the citation in different sections of an original paper are shown in Box 1; note that the result section does not have any citation.

3. Components of the Citation

The citation has three components: (1) quotation, i.e., providing either a summary, a paraphrase or a direct quotation from others’ works, (2) in-text references, i.e., brief addressing to the source, and (3) bibliographic details, i.e., name of the authors, source of publication, date of publication.

3.1. Quotation

Quotations from other sources can be provided by direct quoting, paraphrasing, or providing a summary ( 3 ). In direct quoting, others statements (more than six consecutive words) are exactly copied, put in the quotation marks, and immediately followed by an in-text citation to the original source; every word and punctuation mark should be exactly the same as the original version ( 17 , 20 , 22 ). Direct quoting is used when original words express an idea distinctively or more concisely than your summary, as well as to present well-known statements or to provide historical context for a particular theory or construct ( 8 , 23 ). Direct quoting is not used for reporting findings of a published work, variable names, and operational definitions. Box 2 provides some practical tips for the appropriate use of direct quotations.

In paraphrasing, authors express others’ writing in their own words ( 23 ), followed by a reference to the original source. In cases of paraphrasing another’s work or idea, authors should check their statement to be accurate and fair ( 17 , 20 ). Use of synonyms and rephrasing are typical ways to change the original statements ( 9 ); however, if the paraphrase is too close to the source text in wording, syntax, and structure, it may be considered ‘patch-writing’, which is “the act of making small changes and substitutions to the copied source material” ( 24 ). Note that paraphrasing and summarizing are complex and critical academic skills and depend on one’s knowledge of the content ( 25 ).

In providing a summary, a brief statement of the main points of a work (paper, book or chapter) is presented ( 23 ) using a neutral, affiliating, or distancing approach; in affiliating and distancing approaches, not only existing knowledge is summarized but also authors present their stance/viewpoint ( 2 ). The authors need to present their approaches accurately and persuasively by using appropriate verbs ( 2 ). For a neutral summary, use of verbs ‘comment’, ‘explain’, ‘indicate’, ‘note’, ‘describe’, ‘observe’, remark’, ‘state’, and ‘find’ is recommended ( 2 ). These referring verbs can be used either in the present or the past tense; using the present tense indicates that the source is recent and still valid, whereas the past indicates that the source is older and may be out of date ( 3 ). To find further appropriate verbs and vocabularies to cite the literature being reviewed, readers can refer to other works ( 2 , 26 ).

3.2. In-Text References

In-text references (brief address to the source in text) are presented by three major systems: ‘citation-sequence’, ‘citation-name’, and ‘name-year’. In the ‘citation-sequence’ system, numbers are used to refer to the reference list ( 27 , 28 ), which is numbered sequentially according to the appearance in the text. In the ‘citation-name’ system, the numbering system is used to refer to the reference list, which is numbered in alphabetical order by authors’ names; it means numbers are used in the text regardless of the order in which they appear ( 28 ). The ‘name-year’ system consists of the surname of the author and the year of publication, and the reference list is alphabetically ordered first by author and then by year ( 28 ). If ‘name-year’ system is used, in case of referring to two references with the same first author, chronological hierarchy order is followed (e.g., Annesley, 2010; Annesley, 2011), and in case of the same first author and same publication year, the references need to be differentiated by alphabetical letters after the year of publication (e.g., Annesley, 2010a; Annesley, 2010b) ( 19 ). If the name of author/editor cannot be identified, use the title of the work and the year of publication instead; if the date is not identified, use the phrase ‘no date’ after author’s name and where both author and date are unknown, use the title followed by ‘no date’ ( 23 ).

Citation management software programs (e.g., EndNote, Reference Manager, RefWorks, ProCite, and Refbase) easily connect in-text references to the reference list. These programs can format in-text references and bibliographic details in a different style (discussed in section 4) and can change them from one style to another.

The general rule to refer to a reference in the text is to place the reference immediately after the idea or fact introduced. In other words, an in-text referring might appear in the middle of a sentence and not always at the end ( 6 , 19 ). Unless a sentence ends with a fact (in which case the citation follows), the authors should not pool all the references at the end of a sentence ( 29 ). In case of citing multiple facts in a sentence, it should be clarified which reference is corresponding to which fact ( 19 ). Wherever more than one reference is used to support a fact, the authors should refer to the references in chronological order (the oldest reference is listed as the first in-text reference) ( 19 ). For direct quoting or citing a specific idea or piece of information, the page number of the quote should be included in the in-text reference ( 23 , 30 ). For an in-text citation that refers to secondary sources (second-hand references), name the original source and then use the term ‘cited by’ followed by the reference for the work in which it is quoted (e.g., Schweer, cited by Harrison, 1992, p. 774) ( 31 ).

3.3. Bibliographic Details (Reference List)

Every in-text reference should have a corresponding entry in the reference list ( 28 ); the exceptions are ‘personal communications’ ( 28 ), and ‘unpublished data’ ( 6 ) that are referenced within the text, but do not appear in the reference list. According to the British Standards Institution, a reference is “a set of data describing a document, sufficiently precise and detailed to identify it and enable it to be located” ( 32 ). The essential elements and order of the most common forms of references, including journal articles, books, reports, and websites, are given in supplementary Box 1.

The bibliographic details provided in the reference list should be accurate and complete to ensure that readers will be able to locate the material as easily as possible ( 32 ). The accuracy of the reference list increases the credibility of the author, the journal, and the research itself ( 33 ). To increase the accuracy of the bibliographic details, ICMJE (International Committee of Medical Journal Editors) recommends that “References should be verified using either an electronic bibliographic source, such as PubMed, or print copies from original sources” ( 34 ). Errors in the reference list usually arise by copying bibliographic details from previous papers; thus, authors should not copy from reference lists or databases, and the only reliable source is the original paper published in the journal ( 8 ). In case of using second-hand information, bibliographic detail of a source, where the information is found, should be included in the reference list.

4. Referencing Systems and Styles

Broadly, three types of referencing systems are employed in the academic world ( 3 ). These are, (i) consecutive-numbering system (well-known as Vancouver), (ii) author name-publication year system (well-known as Harvard) ( 19 ); (iii) footnote/endnote system, where sources are listed at the bottom of the page or at the end of the paper and the numbers in superscript run consecutively throughout the paper ( 3 ). The first two systems are commonly used in medicine, whereas the third system is most often used in the humanities. Other less common reference styles that have been established and used within different fields of science and disciplines include Modern Language Association (MLA) ( 35 ), the American Psychological Association (APA) ( 36 ), the Chicago Manual of Style (CMS), and the American Medical Association (AMA) Style ( 29 ).

The first format for bibliographic references in medical science was adopted officially by the Vancouver Group and the National Library of Medicine (NLM), in 1979 ( 37 ). The Vancouver Group was a small group of editors of medical journals who met informally in 1978 in Vancouver, British Columbia, to establish guidelines for the format of manuscripts submitted to their journals ( 37 ); the group expanded into the ICMJE and developed ‘the Uniform Requirements for Manuscripts Submitted to Biomedical Journals,’ which is updated regularly ( 38 ) ( http://www.icmje.org ). The reference style is famed as the Vancouver style because of its origin, and it has become a broadly accepted bibliographic format ( 8 ). According to the Vancouver style (the author-number system), references are numbered consecutively in the order in which they are first mentioned in the text; references in text, tables, and legends should be identified by Arabic numerals in parentheses ( 38 ). The Vancouver style is used by PubMed and MEDLINE.

The Harvard system, so-called as the ‘parenthetical author-date method’, is another popular referencing system ( 31 ). The origin of the system is obscure; however, the first evidence of the system goes back to 1881, when Edward Laurens Mark, professor of anatomy and director of Harvard’s zoological laboratory, published a landmark cytological paper and used parenthetic author-year citation ( 39 ). As Chernin narrated ( 39 ), from an editorial note in the British Medical Journal in 1945, the expression ‘Harvard system’ was not introduced by the Harvard University, but an English visitor to the library of Harvard University was impressed by the system of bibliographical reference and named it as the ‘Harvard system’ upon their return to England. The Royal Society defined the Harvard referencing system in 1965 as “a system in which names and dates are given in the body of the text and the references alphabetically at the end of the paper” ( 31 ).

Although most biomedical journals have adopted the Vancouver style, some still prefer the Harvard system, because they like to know just what author(s) is/are being cited as they read the text ( 40 ). However, the Harvard system is criticized due to potential difficulties that it may create for the readers. For example, if they are interested in an item in the reference list, they need to look it up within the main text. This system may also disrupt the text when a large number of references need to be cited within a paragraph ( 40 ).

Although medical journals specify Vancouver or Harvard systems, some journals have made minor modifications ( 41 ). For example, they have modified the referencing systems, both in-text references, and bibliographies, they have changed punctuation marks, used bold, and italics enhancements, alphabetical or sequential ordering of references, or have made combinations of variations that create a unique reference style that is as large in number as the number of journals currently published ( 41 , 42 ). Authors should carefully follow the format used by the target journal that is usually given in the Information/Instructions for Authors.

5. Other Considerations

5.1. dealing with scientific sources.

The most important challenge of the citation process in a scientific work is ‘which sources must be selected for citation’ and ‘How do the authors deal with the literature being reviewed to cite others accurately and ethically?’ Existence of a large number of publications on the topic makes selection difficult; however, authors should artfully select them to cover all citation purposes and add to the manuscript scientific prestige ( 43 ). Among available sources, the most relevant, valid, methodologically sound, and those with a landmark contribution to the topic should be selected ( 9 , 17 ). If there are a significant number of prior studies on the topic, the most comprehensive and the most recent works should be selected because they presumably discuss and reference the older studies ( 44 ).

The most valid and available sources for citing are published peer-reviewed original journal articles; primary sources (research articles written by those who conducted the research) are preferred ( 21 ). Secondary sources (review papers) can be used when primary sources are unavailable, or a summary for elaborating research problem is more effective; however, they should not be taken as definitive word or fact on the topic ( 9 ). Citation of review articles rather than the original papers should be limited ( 45 , 46 ) since it fails to provide credit or acknowledge the effort of the authors of original research papers. In addition, it may lead to misinterpretation or oversimplification of original research findings ( 45 ). The use of high-quality systematic review is acceptable; otherwise, it should be acknowledged in the text as a review paper to prevent misleading the casual reader about the originality of the work ( 9 ).

Less valid sources (i.e., theses, conference proceeding papers, unpublished data, abstracts, and personal communications) are not recommended ( 19 ) unless they contain essential information not available from public sources ( 47 ). These less valid references can only be used for supporting the results of preliminary studies or citing parallel results in another study population ( 17 ). In case of referring to ‘unpublished sources’ or ‘personal communications,’ the written permission of the author is required to ensure the accuracy of the data and prior approval from the authors ( 6 , 17 ).

Sources that may not be found in public domain, e.g., submitted but unaccepted journal articles, meeting abstracts, and posters should not be used ( 6 , 17 ). Standard textbooks are not cited except for describing a theoretical or methodologic principle or a statistical procedure. As stated by ICMJE, authors are responsible for checking that the references cited not be retracted articles ( 34 ).

5.2. Statement Needs to be Supported by a Reference

As a general rule, findings/statements of other’s work need to be supported by references ( 9 ). Statements like ‘the literature suggest that…’ or ‘there is general agreement that…’ should be followed by addressing one or more references ( 9 ); it is, however, not appropriate and usually essential to support a statement with more than 3 or 4 references ( 9 ). In contrast, common knowledge in a field, defined as facts, dates, events, or information that are expected to be known by someone studying or working in a particular field (e.g., long-established facts or theories), or facts that can be found publicly (e.g., date of the second world war) and are likely to be known by many people (e.g., capital cities of the countries) do not generally have to be referenced ( 23 ).

5.3. Accuracy and Ethics of the Citation

Accurate citation is a crucial issue, enabling readers to follow the flow of ideas and statements in a scientific field and ensure the integrity of the science being communicated ( 19 , 48 ). Citing the sources without retrieving and reading their full-texts and understanding their entirety ( 19 ), giving multiple similar references to support a single statement, or using a single source to support multiple statements are among examples of inaccurate citations ( 48 ). Since an abstract is a brief summary of the work, its content may not accurately present details reported in the text, and therefore, it is a poor practice to cite references after skimming results of the abstract rather considering the whole text ( 9 ).

To avoid inaccurate quoting, the authors should review the entire original article to check the facts. They should be careful in case of paraphrasing or summarizing in order to make sure that the intent or meaning of the original author is not altered ( 49 ). To reduce the risk of misinterpretation of information, the use of secondary sources should be avoided ( 49 ). According to NLM, “The medical literature is full of references that have been cited from other references, serving only to perpetuate erroneous information”, thus, they emphasizes that the authors should never reference documents that they have not read ( 27 ).

Spurious citation, biased citation, and over self-citation are also common problems of citation (Box 3). Self-citation, defined as citing one’s own work in a scientific paper, is a common practice and is an essential part of scientific communication, which represents the continuous and cumulative nature of the research process ( 50 ). When a researcher works on a specific topic for years, 25% self-citation is not uncommon ( 46 ). However, either irrelevant self-citation or over self-citation are considered unethical practices, which affect the precision of the paper ( 50 , 51 ). The spurious citation occurs where sources are not needed but are included anyway, e.g., over-citation or redundant citation (i.e., where the extra sources do not add any value beyond the first source), or citing an obscure, historical reference to give an impression of erudition ( 4 ).

6. Common Errors and Problems of Citation

Citation errors reflect badly on the authors and the publishing journal and may reflect underlying flaws in other areas of the published research ( 33 ). Citation content errors (e.g., inaccurate quoting from others) and both major and minor errors in referencing (both in-text references and bibliographic details) can occur during the citation process ( 4 , 7 , 33 ). Inaccurate quotations or misquotations are perpetual citation errors that lead to circulate a false ‘accepted fact’, which are very hard to correct ( 10 ). A list of common citation errors is provided in Box 3.

Major errors in references, which are responsible for up to 21% of citation errors in the medical field ( 53 ), prevent the source being retrievable ( 4 ). With minor errors (i.e., punctuation and spelling mistakes in bibliography, i.e., name of authors, title, journal, volume, year, and page numbers), references can still be found ( 4 , 9 ). Potential pitfalls of citation-management software programs may also cause some citation errors ( 54 ). Several duplicate copies of a reference in the software may be inserted due to importing the same reference on a number of different occasions (maybe with different patterns); this may lead to the appearance of duplication in the reference list ( 6 , 48 ). In the case of anonymous papers (prepared by a committee or a group of authors), some mistakes may occur ( 54 ). Another common mistake relates to the journal-title; the journal-title may be imported in the abbreviated form, while both full title and conventional abbreviation need to be entered into the journal section of the reference manager ( 54 ). To sum up, the author is responsible for final checking the accuracy of the bibliographic details ( 19 ) and should correct reference manager databases before the reference is exported to the final bibliography.

7. Conclusion

Making good and accurate citations adds to the manuscript’s scientific prestige and signifies that authors have an in-depth knowledge of the literature and writing skills. An original research paper usually has 25 - 40 references and the authors should be note that citing too few or too many references may reflect poor intellectual attitude and work validity. The most important, most elegant and the most recent sources should be selected for citation.

Authors' Contribution: Study concept and design: Zahra Bahadoran and Asghar Ghasemi; drafting of the manuscript: Zahra Bahadoran, Parvin Mirmiran, and Asghar Ghasemi; critical revision of the manuscript for important intellectual content: Khosrow Kashfi and Parvin Mirmiran.

Conflict of Interests: The authors have no conflict of interest.

Funding/Support: This study was supported by the Research Institute for Endocrine Sciences, Shahid Beheshti University of Medical Sciences.

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Proceedings of The Nutrition Society publishes review papers presented by invited speakers who are recognised international experts in their field, and abstracts of Original Communications presented at the scientific conferences of The Nutrition Society.

Public Health Nutrition focuses on the promotion of good health through nutrition and the primary prevention of nutrition related illness in the population. This international journal is essential reading for all specialists involved with nutritional epidemiological research, the role of nutrition in public health, and the identification of practical solutions to public health problems.

Nutrition Research Reviews

Nutrition Research Reviews offers a comprehensive overview of nutritional science today. By distilling the latest research and linking it to established practice, the journal consistently delivers the widest range of in-depth articles in the field of nutritional science.

Journal of Nutritional Science is an internationally peer-reviewed open access journal which welcomes high-quality research articles in all aspects of nutrition. The underlying aim of all work should be, as far as possible, to develop nutritional concepts.

Publications by the Royal Society of Biology , Biochemical Society and Portland Press

emerging

Emerging Topics in Life Sciences , is a new hybrid open access reviews journal jointly owned with the Biochemical Society and published by Portland Press. Available in 2017, it will highlight new and growing fields across the life sciences.

The journal will feature reviews written for an interdisciplinary audience ranging from students and early career researchers to established scientists, each issue will focus on a new or growing field of life science research, guest edited by an expert in the field.

Publications by the Royal Society of Biology

JBE 50.1 March 16

Journal of Biological Education is an international journal that covers policy and curriculum developments, bringing you the latest results of research into the teaching, learning and assessment of biology.

It is owned by the Royal Society of Biology and published by Routledge.

The archive of the journal and the current issue can be accessed via the Taylor and Francis website.

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A new study report may help NASA define and align biological and physical sciences research to uniquely advance scientific knowledge, meet human and robotic exploration mission needs, and provide benefits for all over the next decade.

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The agency announced Tuesday it received the National Academies of Sciences, Engineering, and Medicine’s latest decadal survey on biological and physical sciences research.

“NASA looks forward to reviewing the recommendations in detail and appreciates the steering committee and panels for their dedicated work to deliver this report,” said Nicola Fox, associate administrator, Science Mission Directorate, at NASA Headquarters in Washington. “The survey will inform a compelling new chapter in biological and physical sciences research in space and provide recommendations on ways to support the technologies and infrastructure that will be needed to carry out the science.”

The report, “Thriving in Space – Ensuring the Future of Biological and Physical Sciences Research: A Decadal Survey for 2023-2032,” includes recommendations to implement a comprehensive strategy and vision for transformative science at the frontiers of biological and physical sciences research in space.

“The continued use of space to study biological and physical phenomena enables researchers to pursue innovations and discoveries not possible on Earth,” said Lisa Carnell, director, Biological and Physical Sciences Division, at the agency’s headquarters. “We look forward to using the survey to guide our next decade of transformative science as we maintain U.S. science leadership in space.”

To learn more about NASA’s science missions, visit:

Karen Fox / Melanie White Lyons Headquarters, Washington 202-358-1600 [email protected] / [email protected]

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