The Anatomical Record

Cover image for Vol. 299 Issue 10

Edited By: Kurt H. Albertine, PhD

Online ISSN: 1932-8494

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Note to NIH Authors:

AAA JOURNALS SUPPORT AUTHORS VIA SUBMISSION TO PUBMED CENTRAL : AAA journal authors whose research is funded by NIH will not have to worry about submitting their accepted manuscripts to PubMed Central (PMC). AAA’s publisher, Wiley-Blackwell, will support authors by posting the accepted version of articles by NIH grant-holders to PubMed Central upon acceptance by the journal. The accepted version is the version that incorporates all amendments made during peer review, but prior to the publisher’s copy-editing and typesetting. This accepted version will be made publicly available on PMC 12 months after publication. The Anatomical Record is already freely available one year following publication on Wiley Online Library. The NIH public access mandate applies to all articles based on research that has been wholly or partially funded by the NIH and that are accepted for publication on or after April 7, 2008. In addition to depositing manuscripts on behalf of NIH-funded authors, Wiley has reached an agreement with the Howard Hughes Medical Institute (HHMI) to make post-peer reviewed articles publicly available six months after final publication. HHMI will cover the cost of this service. For other authors of primary research articles whose funding agency requires earlier public access, Wiley-Blackwell offers a $3,000 funded access option for public availability in PubMed Central and on the journal’s website immediately upon publication. Details are available at Author Resources - Funded Access.

The Anatomical Record is an official publication of the American Association of Anatomists. The Anatomical Record's purpose is to publish new discoveries in the morphological aspects of molecular, cellular, organ systems, whole-organism, and evolutionary biology. Areas of particular interest include: studies using an integrative approach that incorporates methods and technologies from different disciplines; hypothesis-driven studies; studies that use imaging approaches in contemporary biological, biomedical, and evolutionary biology research, particularly when framed in the context of experimental models or questions. The Anatomical Record is particularly interested in the morphological consequences of normal growth, disease, and/or interventions such as gene disruption, activation or over-expression on cell, tissue, organ, or whole-organism architecture. Submission of new findings made through the use of imaging modalities ranging from real-time signaling processes to gene or protein expression in individual cells, tissues or whole organisms to structural assessment of organ systems or whole-organism phenotype are also strongly encouraged.

In recognition and support of the growing importance of imaging, we are pleased to cover costs for use of color illustrations and images. High-quality, glossy paper is used for each journal issue, making color figures large and brilliant. The journal also does not charge for printed pages.


The Anatomical Record accepts four categories of publications:
1. Full-Length Papers
2. Reviews
3. Commentaries
4. Letters to the Editor

Full length papers present results or original studies dealing with morphological aspects of any vertebrate or invertebrate molecular pathway, cell type, organ system, whole-organism, or evolutionary biology. Inclusive topics are developmental, comparative, or evolutionary biology. With respect to developmental biology, our sister journal Developmental Dynamics focuses on biochemical and molecular mechanisms of vertebrate and invertebrate development. Thus, some overlap exists between the two journals. However, the area of coverage for developmental biological studies published in The Anatomical Record will be directed primarily to the organ, organ system, or whole-organism level, where descriptive studies of normal and abnormal development may become an important consideration in characterizing phenotypes.

Reviews of topics related to imaging or morphological aspects of any vertebrate or invertebrate molecular pathway, cell type, organ system, whole-organism, or topics in evolutionary biology are encouraged, either through invitation or original submission. Educational topics dealing with the anatomical disciplines will be referred to our sister journal Anatomical Sciences Education. The criteria for acceptance of all review papers are the quality of the writing, originality, and significance.

Commentaries are welcome from the readership. Commentaries should be focused to highlight a topic or controversy, and may include references and figures. These contributions will be peer-reviewed by the Editorial Board and/or Board of Reviewers.

Letters to the Editor also are welcome from the readership. Letters should be focused to highlight a topic or controversy, and may include references. These contributions will be peer-reviewed by the Editorial Board and/or Board of Reviewers.

No paper having appeared previously will be accepted, regardless of the language in which the previous publication was written. Simultaneous publication elsewhere will not be considered. All submissions will be evaluated for plagiarism. If plagiarism is discovered, management of the manuscript will stop while the Editorial Office communicates with the authors.


Submission of Manuscripts: Each manuscript must be accompanied by a cover letter which includes the following elements:

  • The category of manuscript (full-length papers)
  • The names, e-mail addresses, telephone, and fax numbers of the authors
  • The title of the paper and a statement of its specific results and significance
  • A statement that the material has not been published and is not under current consideration elsewhere
  • The names, addresses, telephone numbers, and fields of interest of three to five persons outside your institution who are qualified to referee the paper. Similarly, identify those whom you would not want to referee the paper.

NOTE: submissions that do not include at least three (3) potential reviewers will be delayed in the review process while the editorial office communicates with the corresponding author to provide at least three (3) potential reviewers.


  • Any paper that is in press or being considered elsewhere and includes information that would be helpful in evaluating the work submitted to The Anatomical Record.
  • Written permission from any author whose work is cited as a personal communication, unpublished work or work in press, who is not an author of your manuscript. Please contact the Editorial Office if you have an element which requires permission; the Editorial Office will supply you with the proper permission form to use when soliciting written permission.

SUBMISSION PROCESS: Online submission of manuscripts is required at ScholarOne Manuscripts. Check for an existing account. If you are submitting for the first time, and you do not have an existing account, create a new account.

Submit manuscript and all figures as separate files. DO NOT MAIL ANY PAPER COPIES. At the end of a successful submission, a confirmation screen with manuscript number will appear and you will receive an e-mail confirming that the manuscript has been received by the journal. If this does not happen, please check your submission and/or contact tech support at

Notice of Wiley's Compliance with NIH Grants and Contracts Policy. The National Institutes of Health (NIH) has requested that its grantees submit copies of manuscripts upon their acceptance for publication to PubMed Central (PMC), a repository housed within the National Library of Medicine. On behalf of our authors who are also NIH grantees, Wiley will deposit in PMC at the same time that the article is published in our journal the peer-reviewed version of the author's manuscript. Wiley will stipulate that the manuscript may be available for public access in PMC 12 months after the date of publication. By assuming this responsibility, Wiley will ensure that authors are in compliance with the NIH request, as well as make certain that the appropriate version of the manuscript is deposited. When an NIH grant is mentioned in the Acknowledgements or any other section of a manuscript, Wiley will assume that the author wants the manuscript deposited in PMC, unless the author states otherwise. The author can communicate this via email, or a note in the manuscript. The version of the manuscript that Wiley sends to PMC will be the accepted version, i.e., the version that the journal's Editor-in-Chief sends to Wiley for publication. Wiley will notify the author when the manuscript has been sent to PMC. Wiley reserves the right to change or rescind this policy. For further information, please get in touch with your editorial contact at Wiley, or see the NIH Policy on Public Access, located at:


The manuscript should have uniform style according to The Anatomical Record, as detailed below. It should be written in English and be as concise as possible, without omitting relevant results. Literature surveys, overly detailed methods, or extensive bibliographies will not be published.

Supplemental material will be published electronically by The Anatomical Record. However, supplemental material is limited to video clips, 3-D files, sound files, and/or description of detailed methods. Not permitted as supplementary material is primary data (results), such as digital images, schemas, or data tables. Primary data are to be included as part of the manuscript files so that reviewers can evaluate the data. The editorial office will look at uploaded files and if digital image files, schema files, or data table files are uploaded as supplemental material, the editorial office will contact the corresponding author to correct the submission. Review will be delayed until the correction is made to the uploaded submission.

Text should be supplied in a format compatible with Microsoft Word for Windows. Mac users are asked to save their files with their appropriate file extension (i.e., .doc, .xls, .tif, .eps, etc.). For example, when saving a Word document on a Mac, please add a suffix of ".doc". ScholarOne Manuscripts (formerly known as Manuscript Central) does NOT accept .pdf files.

Abbreviations and style of references are contained in the current edition of the CBE style manual (sixth edition, 1994, Council of Biology Editors, Inc., Suite 230 N. Michigan Ave., Chicago, IL 60601). Spelling reference is to the current edition of Webster's International Dictionary. In items of nomenclature, this journal adheres to the principles specified in Nomina Anatomica, Nomina Embryologica, Nomina Anatomica Veterinaria, and Nomina Anatomica Avium, where appropriate.

Manuscripts should be subdivided into the following sequence:

Title Page
Literature Cited
Figure Legends

Each subdivision should start on a new page.

Title page: The first page of the manuscript should include:

Title of paper
Full name of author(s)
Institutional affiliation and complete address
Telephone and facsimile numbers and e-mail address of the corresponding author
Running title not to exceed 45 letters and spaces
Individual and address to whom correspondence concerning manuscript should be sent
All grant information in the following format: Grant sponsor(s): ________; Grant number(s): _________.

Abstract: Submit an abstract of 250 words or less that will serve in lieu of a concluding summary. The abstract must be written in complete sentences. It should concisely state the significant findings without reference to the rest of the paper. Append three to eight key words at the end of the abstract for the purposes of citing your work by the secondary services.

Text: This is divided into an Introduction, Materials and Methods, Results, and Discussion sections.

Literature Cited:

Text references to literature should be arranged chronologically by author's name followed by year of publication:

. . . studies by Gheerbrandt et al., (2005) reveal
. . . studies by Geisler and Uhen (2005) shown
. . . a recent report (Buchholtz, 2007)
. . . (Uhen, 2007)

When references are made to more than one paper by the same author, published in the same year, they are to be designated in the text as (Kelley, 1970a,b) and in the literature list as follows:

Kelley RO. 1970a. An electron microscopic study of mesenchyme during development of interdigital spaces in man. Anat Rec 168:43-54.
Kelley RO. 1970b. Fine structure of apical, digital and interdigital cells during limb morphogenesis in man. In: Proceedings of the VIIth International Congress of Electron Microscopy. Vol. III: p 381-382.

Literature Cited is to be arranged alphabetically in the following style: Author's name (or names), year of publication, complete title, volume, and inclusive pages.

Bartsiokas A. 2002. Hominid cranial bone structure: a histological study of Omo 1 specimens from Ethiopia using different microscopic techniques. Anat Rec 267:52-59.
Bräuer G, Collard M, Stringer C. 2004. On the reliability of recent tests of the Out of Africa hypothesis for modern human origins. Anat Rec 279:701-707.
Broadfield DC, Holloway RL, Mowbray K, Silvers A, Yuan MS, Márquez M. 2001. Endocast of Sambungmacan 3 (Sm 3): a new Homo erectus from Indonesia. Anat Rec 262:369-379.
Bruner E, Manzi G. 2005. CT-based description and phyletic evaluation of the archaic human calvarium from Ceprano, Ital. Anat Rec 285:643-657.
Bush EC, Simons EL, Allman JM. 2004. High-resolution computed tomography study of the cranium of a fossil anthropoid primate, Paraphithecus grangeri: new insights into the evolutionary history of primate sensory systems. Anat Rec 281:1083-1087.
Martin RD, MacLarnon AM, Phillips JL, Dobyns WB. 2006. The Flores hominid: new species or microcephalic dwarf? Anat Rec 288:1123-1145.

Book Chapter:
Depew MJ, Tucker AS, Sharpe PT. 2002. Craniofacial development. In: Rossant J, Tam PPL, editors. Mouse Development: Patterning, Morphogenesis and Organogenesis. San Diego: Academic Press. p 421-498.

Sternberger LA. 1986. Immunocytochemistry. 3rd ed. New York: John Wiley & Sons.

Footnotes: Number footnotes to the text consecutively. The corresponding reference numbers must be clearly indicated in the text. Additional references to the identical footnote must be numbered with the next consecutive number, for example:

1 Material used for this experiment was . . .
2 provided by . . .
3 See footnote 2, page . . .

Type table footnotes directly beneath the table and number them 1, 2, 3 etc. They must not be numbered in sequence with text footnotes.

Tables: All tables must be cited in the text and have titles. Table titles should be complete but brief. Information other than that defining the data should be presented as footnotes. Since tabular matter is expensive to reproduce, it should be simple and uncomplicated with as few vertical and horizontal rules as possible.

Figure legends: All figures must be cited in the text and must have legends. Number figures, including charts and graphs, consecutively throughout the text. Give text references to figures only in terms of the figure number. Whenever possible, integrate figures into the text. Group figures to fit a single page with their appropriate legend. References to relevant text passages can often reduce the length of legends and avoid redundancy.

Abbreviations: Spell out all nonstandard abbreviations the first time used. Abbreviations for all figures should be listed alphabetically and placed before the first figure in which they are mentioned, e.g.,




Cortical Plate


Primary somatosensory cortex



Digital Illustrations: The Anatomical Record is known for its free-of-charge publication of high-quality figures. To maintain this level of quality, it is necessary to require images to be submitted and processed with this goal in mind. Therefore, when preparing digital art, please submit figures in separte .tif or .eps file formats, with the following attributes:

  • CMYK format for color figures (color figures are FREE to all authors)
  • 300 DPI/PPI for picture-only (without text of any kind) figure files
  • 600 DPI/PPI for figures containing pictures and lie elements (i.e., text labels, thin lines, arrows, etc.)
  • 1200 DPI/PPI for black and white images such as line drawings, graphs, or charts

Scaling, cropping, and rotating should be performed in the originating application. To ensure that your figures will not be too large to upload, be sure to adjust the height and width to approximately 2500 pixels. If the figure(s) does not meet the specifications shown above (including not exceeding the maximum size of 2500 pixels), please use graphics software (e.g., Adobe Photoshop or Illustrator) to modify the figure(s).

ATTENTION AUTHORS: Please verify that figure files meet the printer's specifications for format and resolution, at the time that you submit the original version of your manuscript. Note that .tif (or .eps) file formats for figures (black and white, color, and grayscale) is recommended. We recommend creating your graphics, with all fonts and scale bars included, using Photoshop, Illustrator, or Freehand and then uploading the figure files into ScholarOne Manuscripts (formerly known as Manuscript Central). File formats that are NOT acceptable are JPG/JPEG, GIF, ONG, PCX, PNG, XBM, Word, and Excel. For further guidance on preparing digital figure files, authors are encouraged to visit . Figure files that do not conform to the required format and resolution will delay review of your manuscript, because the editorial office will return the files to you for correction before sending your manuscript to reviewers.

Cover Illustrations: Authors are encouraged to submit CMYK color figures for consideration as cover illustrations. These figures must be submitted with the manuscript, preferably sized to 21 x 26 cm, with 600 DPI/PPI resolution.

Care and Use of Experimental Animals: The Anatomical Record and the American Association of Anatomists require that all studies involving experimental animals be conducted in a humane manner and in accordance with all local, state and federal guidelines for the care and utilization of laboratory animals. Husbandry of the animals must meet the NIH Guidelines for the Care and Use of Laboratory Animals. Each manuscript must include details of the a) food and water regimen, b) light cycles, c) appropriate tranquilizers, analgesics, anesthetics, and care performed in association with all surgical procedures, d) manner by which the animals were euthanized, including drugs and their dosages, and e) written assurance that an Institutional Animal Care and Use Committee (or equivalent) approved the protocol.

Protection of Human Subjects: The Anatomical Record and the American Association of Anatomists also require that all studies involving human subject adhere to the principles of the Declaration of Helsinki or research in Human Beings. Each manuscript must include details of the a) number of subjects, b) age, c) gender, d) women, children and other minorities, e) inclusion and exclusion criteria, and f) written assurance that an Institutional Review Board (or equivalent) approved the protocol.

AAA Guidelines for Research Involving Human Subjects and Vertebrate Animals: The authors signify by submission of their manuscript that research involving use of human subjects—including research for educational studies— has been conducted according to the principles of the Declaration of Helsinki and Title 45, U.S. Code of Federal Regulations, Part 46, Protection of Human Subjects, Revised November 13, 2001, unless regulated by more restrictive state or local laws. Research involving vertebrate animals must adhere to AAA’s Guiding Principles in the Care and Use of Animals. For investigations involving human subjects or vertebrate animals, a statement of protocol approval from an Institutional Review Board (IRB) or Institutional Animal Care and Use Committee (IACUC) or its equivalent, respectively, must be included in the Methods section of the paper. Editors/Associate Editors are expected to refuse papers in which evidence of adherence to these principles is not stated explicitly.

  • Do not hyphenate words at the end of the lines.
  • Corrections to the manuscript should be typed or printed legibly in ink.
  • Do not begin sentences with abbreviations.
  • Spell out the word Figure in the text except when it appears in parentheses: Figure 2, (Figs. 4-6).
  • Always spell out numbers when they stand as the first word in a sentence, abbreviations cannot follow such numbers. Numbers indicating time, weight and measurements are to be in Arabic numerals when followed by abbreviations (e.g., 2mm; 1sec; 3ml). In general, write out the numbers one to ten in the text. All higher numbers should be given as numerals.
  • The metric system should be used for all measurements, weights, etc.
  • The author will be asked to sign a Copyright Transfer Agreement transferring rights to the publisher, who reserves copyright.
  • It is the current policy to underwrite all normal black-and-white tabular and illustration costs.

Proofs: A single set of page proofs will be sent via E-mail to the corresponding author. All corrections should be marked clearly directly on page proofs.

Reprints: Reprints may be purchased at It is important to order initially a sufficient quantity of reprints, since the price is substantially higher is they are ordered after the paper has been published.

If your paper is accepted, the author identified as the formal corresponding author for the paper will receive an email prompting them to login into Author Services; where via the Wiley Author Licensing Service (WALS) they will be able to complete the license agreement on behalf of all authors on the paper.

For authors signing the non-standard CTA

If the OnlineOpen option is not selected the corresponding author will be presented with The Anatomical Record non-standard CTA to sign. The terms and conditions of the non-standard CTA can be previewed below:

Terms and Conditions. Please do not complete this PDF until you are prompted to login into Author Services as described above.

Note to Contributors on Deposit of Accepted Version

Funder arrangements

Certain funders, including the NIH, members of the Research Councils UK (RCUK) and Wellcome Trust require deposit of the Accepted Version in a repository after an embargo period. Details of funding arrangements are set out at the following website: Please contact the Journal production editor if you have additional funding requirements.


Wiley has arrangements with certain academic institutions to permit the deposit of the Accepted Version in the institutional repository after an embargo period. Details of such arrangements are set out at the following website:

For authors choosing OnlineOpen

If the OnlineOpen option is selected the corresponding author will have a choice of the following Creative Commons License Open Access Agreements (OAA):

Creative Commons Attribution License OAA

Creative Commons Attribution Non-Commercial License OAA

Creative Commons Attribution Non-Commercial -NoDerivs License OAA

To preview the terms and conditions of these open access agreements please visit the Copyright FAQs hosted on Wiley Author Services and visit

If you select the OnlineOpen option and your research is funded by The Wellcome Trust and members of the Research Councils UK (RCUK) you will be given the opportunity to publish your article under a CC-BY license supporting you in complying with Wellcome Trust and Research Councils UK requirements. For more information on this policy and the Journal’s compliant self-archiving policy please visit:

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Editorial office:

Kurt H. Albertine, Ph.D.
Univeristy of Utah School of Medicine
Department of Pediatrics
PO Box 581289
Salt Lake City, Utah 84158
Tel: 801-213-3213
Fax: 801-585-7395
Tel: 011-801-213-3213
Fax: 011-801-585-7395

AR Production