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Manuscript preparation


1. General information

Development is moving to a new submission system; for more information, see our FAQ page. To submit a manuscript, please use our online submission system.

Authors are required to read our journal policies before preparing their manuscripts, and all manuscripts should adhere to the journal’s terms of submission.

All pre-submission or general editorial queries should be directed to the Editorial Office.

1.1. New submissions – format free

In order to make manuscript submission as easy as possible for authors, Development has introduced format-free submission.

At first submission, authors may submit their manuscript in any format; however, we do still encourage authors to read the manuscript preparation guidelines below and to consider how easy a manuscript is to read by reviewers and editors (e.g. line spacing, line numbers).

All manuscripts must adhere to our guidelines regarding manuscript length.

1.2. Revised submissions

On Development, >95% of revised submissions are accepted for publication.

All revised manuscripts should adhere to the guidelines below on preparing text and tables, figures, movies and supplementary information.

Authors should complete and submit our submission checklist with their manuscripts. This form asks authors to confirm that they have followed best practice guidelines regarding experimental subjects, data reporting and statistics. The checklist is based on the NIH Principles and Guidelines for Reporting Preclinical Research and is intended to help ensure high standards for reporting and to aid reproducibility.

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2. Manuscript length

Research articles: the total length of the article should not exceed 7000 words, including the main text and figure legends, but not the title page, abstract, materials and methods section or reference list (reference citations in the text do count towards the word limit).

Research reports: the total length of the article should not exceed 3000 words, including the main text and figure legends, but not the title page, abstract, materials and methods section or reference list (reference citations in the text do count towards the word limit). The total number of display items (Figures and Tables) must not exceed four (4).

Note that final word limits will depend on the paper submitted and are at the discretion of the Editors.

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3. Preparing the text and tables

The information below relates to a standard Research Article. For all other article types, please refer to the style and layout guidelines provided on our article types page.


3.1 File formats

For manuscript text and tables, our preferred file format is Microsoft Word .docx (or .doc). We also accept Pages (rtf format) and LaTeX.

Please include tables as part of the manuscript file. Tables must be editable and not embedded as an image.

Authors working in LaTeX can download and use our template. Please upload a single PDF at first submission and include any component files, such as .st (style file), .cls (class file) and .bib (bibliography file) at revision submission. Please note that LaTeX files will be converted to Microsoft Word files during the production process and that authors will be required to check the conversion of symbols and special characters carefully at the proofing stage.

For mathematical equations, our preferred file format is MathType. We also accept Equation Editor (Microsoft Word) and LaTex.


3.2. Article sections

3.2.1.Title page

This section should include a title of 120 characters or less that clearly and concisely summarises your specific findings and avoids specialist abbreviations, a running title of 32 characters or less, the full names (including middle initials) and affiliations of all authors (including present addresses for authors who have moved), and the corresponding author’s email address. Please note any cases where authors contributed equally to the work. Please also include 3-6 key words for indexing purposes (select key words that will make your manuscript easily searchable).

3.2.2. Abstract

Provide a brief abstract of no more than 180 words. This should succinctly and clearly introduce the topic of the paper, summarise the main findings and highlight the significance of the data and main conclusions. The abstract is used by abstracting services without modification and is often read more frequently than the full paper and therefore needs to be comprehensible in its own right. Do not include subheadings or references, and avoid any non-standard abbreviations.

3.2.3. Summary statement

Provide a brief Summary Statement for use in emailed and online tables of content alerts. The text should be between 15 and 30 words, and should explain, without overstatement, why someone should read the article. Please do not simply repeat the title, and avoid unfamiliar terms and abbreviations, as the text should be comprehensible to non-experts. We reserve the right to edit the text.

3.2.4. Introduction

This section should succinctly provide the background information that is required to set the results into their proper biological context. It should not contain subheadings.

3.2.5. Results

This section should describe the results of the experiments performed and should be broken up by subheadings to organise the findings presented and walk the reader through the results. Reproducibility of results must be included – see our submission checklist for further information. Please ensure that the distinction between new results and published findings/established facts is clear.

3.2.6. Discussion

This section should explain the significance of the results and should place them into the broader context of the current literature. The Discussion may contain subheadings to highlight important areas that are expanded on in the text. Please note that, for Research Reports, the Results and Discussion sections should be combined, while they should be kept separate for Research Articles.

3.2.7. Materials and methods

This section should include sufficient detail to understand and to replicate the experiments performed, in conjunction with cited references. To facilitate detailed description of materials and methods (allowing the reader to fully understand and replicate the experimental protocols), this section does not count towards the word limit for article length. The materials and methods should be divided into sections, and should include subsections detailing reagents, animal models and statistical analysis. Please provide names for ALL equipment and reagent suppliers. Give Latin names for all experimental species. Reporting standards should follow those recommended in our journal policies and submission checklist. Where the Materials and Methods are particularly lengthy, more detailed experimental protocols or descriptions of computational analyses should be included as Supplementary Materials and Methods (though all methods must be mentioned in the main text). Materials such as primers, antibodies and strains may be provided as supplementary text or tables. Note that the editor or production team may request changes to the materials and methods to ensure compliance with these guidelines.

Authors who use AI tools in the writing of a manuscript, generation of code or production of images, or in the collection and analysis of data, must disclose this in the Materials and Methods section. They should describe which AI tool was used and how it was used (note that this section does not count towards the word limit for article length). Please note that authors are fully responsible for the content of their manuscript, including those parts produced by an AI tool.

3.2.8. Acknowledgements

This section should mention any individuals or groups that do not qualify for authorship according to our authorship guidelines, but have contributed to the research presented (e.g. in terms of reagents, time, expertise) or writing of the manuscript. Please also include details of support from core facilities (e.g. microscopy, sequencing, proteomics facilities); an example of how this can be done is included below.

“The authors thank the WEHI core facilities for their important contributions to this work, particularly The WEHI Advanced Genomics Facility, The Centre for Dynamic Imaging and the WEHI Bioservices Facility.”

3.2.9. Competing interests

Include a statement to identify any potential influences that readers may need to know about when thinking about the implications of the presented research. For more specific information regarding the affiliations and associations that must be disclosed, please see our journal policies page. Authors without financial or competing interests should explicitly assert this and include the statement ‘No competing interests declared’.

3.2.10. Funding

Details of all funding sources must be provided. It is the responsibility of the corresponding author to provide the relevant funding information from ALL authors. Please provide the official funding agency name as listed on the Crossref Open Funder Registry, i.e. 'National Institutes of Health', not 'NIH', and all relevant grant numbers. If your Funder is not listed in the Registry, please provide the name in full.

Where individuals need to be specified for certain sources of funding, please add initials after the relevant agency or grant number. Please use the following format: This work was supported by the National Institutes of Health [AA123456 to C.S., BB765432 to M.H.]; and the Alcohol & Education Research Council [hfygr667789]. Where no specific funding has been provided for the research, please state ‘This research received no specific grant from any funding agency in the public, commercial or not-for-profit sectors’.

3.2.11. Data availability

All publicly available datasets supporting your work should be included in the Data availability section. Details should include repository name, identifier such as accession number or doi and, where possible, include a hyperlink to the URL of the dataset. Datasets should be made publicly available at the time of publication. For more information on our data deposition requirements, see please see our Journal Policies.

Please note that Development endorses the Force 11 Data Citation Principles and recommends that references to datasets should also be included in the reference list with DOIs/accession numbers and hyperlinks, where available.

If all data are included only within the manuscript itself and/or its Supplementary information, please include a default statement as follows:
Data availability: All relevant data can be found within the article and its supplementary information.

3.2.12. Diversity and inclusion statement

Authors may provide a Diversity and inclusion statement in their paper. This can document diversity and inclusion practices relevant to both the scientific content of the paper, and authorship and attribution.

3.2.13. References

All references cited in the text, tables and figure legends should be included in a single reference list at the end of the article. We strongly encourage the citation of the primary literature over review articles wherever possible, and for this reason do not have a limit on the number of references that can be included. For specific information about reference formatting, please see the references section below.

3.2.14. Figure legends

Figure legends should be listed at the end of the manuscript. The first sentence of the legend should summarise the figure and be in bold. Each figure legend should stand alone and should contain enough information to ensure that the figure is understandable without having to refer to the main text. Figure panels should be labelled with uppercase letters (A, B, C, etc.), and each panel should be described in the legend. Any abbreviations not given in the main text should be defined. For further details on what should be included in figure legends, please refer to our submission checklist.


3.3. Preparing the text

3.3.1. General information

  • Prepare manuscripts in English (either US or UK spelling is acceptable but be consistent within the manuscript). Your writing should be comprehensible to editors and reviewers, and your writing style should be concise and accessible. If English is not your first language, please consider using a language editing service prior to submission.
  • Ensure that the language in your manuscript is original and does not contain previously published passages of text (including those from your own publications) – see our journal policies for more details. All accepted manuscripts are routinely screened using plagiarism-detection software.
  • Use 1.5 line spacing and continuous line numbering throughout the paper in order to facilitate online reviewing.
  • Do not embed figures in the text.
  • Cite each figure, table and movie in the text in numerical order. Figure or table parts should be labelled with uppercase letters (A, B, C, etc.). Use the following format for citations: Fig. 1A,B or Figs 1, 2 or Table 1 or Movie 1.
  • If necessary, display equations should be cited using the following format: Eqn 1.
  • For supplementary figures, tables and equations, cite as Fig. S1, Table S1, Eqn S1.
  • Define abbreviations at first mention.
  • For special characters not available on a standard keyboard (e.g. Greek characters, mathematical symbols), use the Symbol font or the ‘Insert Symbol’ function in Microsoft Word, where possible. For special characters that are not available via this route, please use MathType characters; do not use embedded images (e.g. GIF).

3.3.2. Units and nomenclature


3.3.3. References References in text

References in the text should be cited using the Harvard (name, date) referencing system.

Each reference cited in the text must be listed in the Reference list and vice versa: please check these carefully. Where references are cited only in supplementary information, please provide a separate supplementary reference list and do not include these in the main reference list.

Literature citations in text are as follows.

  • One author – (Jones, 1995) or (Jones, 1995; Smith, 1996).
  • Two authors – (Jones and Kane, 1994) or (Jones and Kane, 1994; Smith, 1996).
  • More than two authors – (Jones et al., 1995) or (Jones et al., 1995a,b; Smith et al., 1994, 1995).
  • Manuscripts accepted for publication but not yet published: include in Reference list and cite as (Jones et al., in press).
  • Manuscripts posted on preprint servers but not yet published: include in Reference list and cite as (Smith et al., 2016 preprint).
  • Citation of unpublished data: we strongly discourage the citation of unpublished data or data not shown. Where it is necessary, use the format (S.P. Jones, unpublished observations/data not shown); note that the editor or journal office may request that these data should be included prior to publication. Personal communications (the unpublished observations of scientists other than the authors) can only be cited with written permission from the scientist in question, and should be cited in the text using the format (full name, institution, personal communication). Unpublished work can not be included in the Reference list.
  • PhD theses: include in Reference list and cite as (Smith, 2016 ).
  • Datasets: we recommend that all publicly available datasets are fully referenced in the reference list with an accession number or unique identifier such as a DOI. Cite as (Jones and Jane, 1994).
  • Authors should avoid citing articles from journals that are suspected to be predatory in nature (see for an online resource designed to help researchers identify trusted journals).
  • Citation of retracted articles is strongly discouraged. If it is necessary to cite a retracted paper, the notice of retraction must also be cited and it must be obvious to the reader that the article has been retracted. Editors may question why a retracted publication has been cited. Reference List

References are listed in alphabetical order according to surname and initials of first author.

  • Use the following style:


    Rivera, A. R. V., Wyneken, J. and Blob, R. W. (2011). Forelimb kinematics and motor patterns of swimming loggerhead sea turtles (Caretta caretta): are motor patterns conserved in the evolution of new locomotor strategies? J. Exp. Biol. 214, 3314-3323.


    Hochachka, P. W. and Somero, G. N. (2002). Biochemical Adaptation: Mechanism and Process in Physiological Evolution. Oxford, UK: Oxford University Press.

    Book chapter

    Feller, G. (2008). Enzyme function at low temperatures in psychrophiles. In Protein Adaptation in Extremophiles (ed. K. S. Siddiqui and T. Thomas), pp. 35-69. New York: Nova Science Publishers, Inc.

    Preprint server

    Baillie-Johnson, P., van den Brink, S. C., Balayo, T., Turner, D. A. and Martinez Arias, A. (2014). Generation of aggregates of mouse ES cells that show symmetry breaking, polarisation and emergent collective behaviour in vitro. bioRxiv doi:10.1101/005215.

    PhD thesis

    Jones, A. R. (2016). Title of thesis. PhD thesis, University of Washington, Seattle, WA.

    Dataset with persistent identifier

    Zheng, L.-Y., Guo, X.-S., He, B., Sun, L.-J., Peng, Y. and Dong, S.-S. (2011). Genome data from sweet and grain sorghum (Sorghum bicolor). GigaScience Database.

    Kingsolver, J. G., Hoekstra, H. E., Hoekstra, J. M., Berrigan, D., Vignieri, S. N., Hill, C. E., Hoang, A., Gibert, P. and Beerli, P. (2001). Data from: The strength of phenotypic selection in natural populations. Dryad Digital Repository.

  • If there are more than 10 authors, use 'et al.' after the 10th author.
  • Within a group of papers with the same first author, list single author papers first, then papers with two authors, then et al. papers. If more than one reference exists for each type, arrange in date order. Use a and b for papers published in the same year.
  • 'In press' citations must have been accepted for publication and the name of the journal or publisher included.


3.4. Preparing tables

Prepare tables in ‘cell’ format and include in the same file as the main text. Tables must be editable and not embedded as an image.

The title of the table should be a single sentence and should summarise the contents of the table. Details referring to one or more isolated item(s) in the table are best given in a table footnote. Units should be given in parentheses at the top of each column (do not repeat in the table).


3.5. Preparing display equations

Our preferred file format for equations is MathType. We also accept Equation Editor (Microsoft Word) or LaTex.

Please number all display equations, consecutively. They should take the form:


Units should be defined in the text rather than included in the equation.

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4. Preparing figures

4.1. General information

Figures should be numbered in a single series that reflects the order in which they are referred to in the text.

Figures should be prepared at the smallest size that will convey the essential scientific information; final figure size is at the discretion of the journal. For further information on how to arrange your figures to optimise viewing by reviewers and readers, download our figure layout guidelines.

At initial submission, you may submit a single PDF file containing all text and figures. Once an article has been accepted for publication, you are required to submit separate files for each figure (see below for file formats).

Figure legends should be included in the main text file and not in the figure file.

There are no charges for the use of colour in figures, although gratuitous use of colour in graphs and diagrams should be avoided and colour should only be used to improve scientific clarity.

For best practice and transparency, and to allow better assessment of the quality of the data and whether the data support the conclusions, we strongly recommend that you use graphs that allow the reader to see the true data spread (unless an N=1 is stated). See this article for further information. For bar graphs with error bars, please show individual data points. If data points are too numerous, please indicate variance using an acceptable alternative approach (e.g. box-and-whisker plots, super plots). If using box-and-whisker plots, please state what the whiskers represent (more information can be found here).

Development strongly encourages the use of colours that are suitable for readers with colour-vision impairment (a.k.a. colour blindness), particularly in the preparation of fluorescence microscopy images and in heatmaps. Authors should avoid the use of red/green for the display of 2-channel images; authors should consider an alternative colour combination (e.g. magenta/green). Guidance is available here and here. Free tools are available for you to check whether your images are compliant with BiO’s policy. These tools will change the image colouring to replicate common forms of colour blindness, showing which adjustments need to be made.


4.2. Preparing photographic images

4.2.1. General information

Photographic images (also known as bitmap images) are made up of pixels (e.g. light, fluorescence and electron microscopy, gels, and traditional photography)

  • The maximum figure size, including lettering and labels, is 180 mm x 210 mm.
  • Images should be saved at a resolution of 300 pixels per inch. Any image quality option should be set to maximum.
  • For figures that contain both photographs and line art or text, 600 pixels per inch is preferred.
  • For micrographs, use a scale bar to show the magnification and give the length of this in the figure legend.
  • Colour: supply images in RGB (not CMYK) mode, as this maximizes colour quality and is how the figures will be displayed online; do NOT use Spot, Pantone or Hex colours and do NOT assign a colour profile.
  • Text labelling: use 12 pt bold uppercase letters (A, B, C, etc.) to distinguish figure panels; other labelling should be 8 pt Arial font (sentence case) (headings should be bold); for gene sequences, use Courier font to ensure that each letter is the same width; use Symbol font for Greek characters.

4.2.2. File formats

Accepted file formats are: EPS/PDF/TIFF/JPEG

  • ARIAL or HELVETICA must be the font choice used throughout figure preparation.

PowerPoint images: we do NOT accept PowerPoint files. Instead, please save as PDF using the instructions below.

  • Go to ‘print’ and then choose 'Save as PDF' in the print dialogue box
  • You can download free software which will enable you to print EPS/PDF files to disk:


4.3. Preparing graphs and diagrams (line art)

4.3.1. General information

  • The maximum figure size, including lettering and labels, is 180 mm × 210 mm.
  • Line thicknesses and symbols should be of sufficient size to ensure clarity if the figure is reduced in size.
  • For graphs, our preferred symbols are filled and open circles, triangles, squares, or diamonds; where possible, the same symbol should be used for the same entity in different figures.
  • Colour: supply line art in RGB (not CMYK) mode, as this maximizes colour quality and is how the figures will be displayed online; do NOT use Spot, Pantone or Hex colours and do NOT assign a colour profile.
  • Text labelling: use 12 pt bold uppercase letters (A, B, C, etc.) to distinguish figure panels; other labelling should be 8 pt Arial font (sentence case) (headings should be bold); for gene sequences, use Courier font to ensure that each letter is the same width; use Symbol font for Greek characters.

4.3.2. File formats

Authors should submit their source figures in an editable format (vector graphic) that retains font, line and shape information. This format ensures that we can edit where necessary and produce high-quality print and online PDFs.

We accept the following file formats for graphs/line art: EPS, PDF, and WMF.

  • Applications such as Adobe Illustrator, Canvas, DeltaGraph, Corel Draw, Freehand, MatLab and SigmaPlot provide these formats.
  • Please ensure that you 'export' or 'save' with (text/font) information included
  • Save text/font information as ‘text’ not ‘curves’ or ‘outlines’.
  • If combining images, always 'embed' images; do NOT simply 'link' them. In Adobe Illustrator, copying and pasting or dragging an image directly from Adobe Photoshop will embed the image. Alternatively, if you use the 'Place' command, uncheck 'Link' in the dialogue box. For other software applications, please refer to the documentation (often there will be a 'link', 'proxy', 'OLE' or 'OPI' option, which must NOT be used with EPS files).
  • Note that submission of JPEG or TIFF format for graphs/line art may delay production of your article.


4.4. Image manipulation

Any alterations made to figures using computer software must be consistent with our image manipulation policy. The images presented in the manuscript must remain representative of the original data, and the corresponding author will be asked to confirm this at submission. Please read our requirements for preparing your figures to avoid a potential delay in the publication process or rejection on the basis of non-compliance with these guidelines. This guide also includes recommendations for improving figure layout to help reviewers and readers appreciate your data.

All accepted manuscripts are routinely screened by our production department for any indication of image manipulation. If evidence of inappropriate manipulation is detected, the journal's Editors might ask for the original data to be supplied and, if necessary, may revoke the acceptance of the article.


4.5. Figure permissions

It is the responsibility of the author to obtain permission to use figures from another publication in any article submitted to Development and to ensure that any such use is credited to the source. Any fees associated with use of the figure are the responsibility of the author. Written permission from the author and/or publisher of the original material, as appropriate, should be provided at the time of submission, otherwise publication may be delayed. If a figure has been modified from a previously published figure, please check with the copyright owners to see whether permission is required and include a complete citation/reference for the original article.

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5. Preparing movies

Our preferred file format for movies is .mp4, but we also accept .mov. Movies should be prepared at the smallest file-size that will convey the essential scientific information. We have a limit of 500 MB for all movie files. If your movies exceed this limit, please contact the Editorial Office for advice before submission.

Please include the titles and captions of all movies in your supplementary information PDF (see section on preparing supplementary information). Please keep captions as short as possible and ensure that they explain what is being shown in the movie and any necessary details of how the movie was made.

Movies should be numbered in a single series that reflects the order in which they are cited in the text, e.g. see Movie 1. Movie 2, etc. Please do not use alphabetical labelling, e.g. Movies 1A-C should be relabelled as Movies 1-3.

All movies will play in place in the full-text online version of the article and a link to each movie will be included in the supplementary information PDF.

When preparing your movies, please note the recommendations below:

  • Use a resolution of no greater than 1280x720 (720p), as most readers will view on a desktop or mobile device
  • Use a well-characterized video compression codec such as H.264 and use multi-pass encoding if available
  • Do not exceed a bitrate of 2500mbps for 720p H.264-encoded video
  • Keep duration to the minimum required to illustrate your point
  • Do not include an audio track unless it is essential
  • If including audio, use a well-characterized audio compression codec such as AAC
  • Do not exceed a bitrate of 128kbps, sample rate of 44.1kHz, or channel count of 2 for encoded audio

Please note that we reserve the right to make movies or other data forms available on an Open Access basis via The Company of Biologists’ website, YouTube and other online channels. Where we do, the movies and other data forms may on occasion be made available under the terms of the Creative Commons Attribution 4.0 International (CC-BY) Licence (the terms of which are set out at These terms permit the copying and/or adaptation of the movie and the distribution of the movie or any such adaptation by any means and in any medium or format to any other person, including for commercial purposes, provided that you are credited as the original author. There would be no additional cost to you, the author.

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6. Supplementary information

Data that are essential for interpretation of the results of the main paper should be included in the main paper. Supplementary information provides access to supporting data that do not appear in the printed article or PDF but that accompany the final version of a paper online.

These data are peer reviewed and subject to the same criteria as the data that are to be published in the paper itself. During peer review, editors and reviewers are asked to assess whether supplementary information is appropriate and essential for supporting the findings of a paper.

All supplementary data will be strictly limited to a total of 50 MB per article (excluding movie files and cover art submissions).

Supplementary material may include movies, figures and tables. Descriptions of computational/mathematical analyses and further details of experimental protocols already described in the Materials and Methods section of the main article may be included as supplementary Materials and Methods; the main article must however contain sufficient information in the Materials and Methods to allow the reader to understand all experiments performed. Tables comprising datasets or listing materials such as oligonucleotide primers, antibodies or strains may be provided as supplementary material. We do not accept additional Results or Discussion text as supplementary material. Very large files or those requiring specialist software are not suitable as supplementary information. For large datasets, e.g. imaging data, please see our guidelines on data deposition.

With the exception of movies (see section on preparing the movies) and large tables, all supplementary information, including movie titles and captions, should be collated into a single PDF file. If your table is very large, or you wish readers to be able to export and/or manipulate the data, we would prefer you to submit it as a Microsoft Excel file.

Use a separate numbering system from that used in the main article and use the format Fig. S1, Fig. S2, Table S1 etc. If a supplementary figure relates to a particular figure in the text, please cite it as close to this figure as possible. For the convenience of readers, please place each figure next to the corresponding legend in the supplementary information PDF. Please include a legend for each figure and a title for each table.

Please note that supplementary data files are not copyedited by Development and therefore authors must ensure that all files are checked carefully before submission and that the style of terms and figures conforms to that of the article. Modification of supplementary information after publication will require a formal correction.

Refer to each piece of supplementary information at least once within the text of the main article (the article that is published in the print issue of the journal).

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