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Formatting Research Paper Headings and Subheadings

heading in a research paper

Different style guides have different rules regarding the formatting of headings and subheadings in a paper, but what information you should actually put into your subheadings is a different question and often up to personal taste. Here we quickly summarize general guidelines, different approaches, and what not to do when choosing headings for a research paper.

Does it matter how I name my sections and subsections?

The main sections of a research paper have general headers and are often journal-specific, but some (e.g., the methods and discussion section) can really benefit from subsections with clear and informative headers. The things to keep in mind are thus the general style your paper is supposed to follow (e.g., APA, MLA), the specific guidelines the journal you want to submit to lists in their author instructions , and your personal style (e.g., how much information you want the reader to get from just reading your subsection headers). 

Table of Contents:

  • Style Guides: Rules on Headings and Subheadings
  • What Sections and Subsections Do You Need? 
  • How Should You Name Your Sections and Subsections?
  • Avoid These Common Mistakes

research paper headings

Style Guides: Research Paper Heading and Subheading Format

Headers identify the content within the different sections of your paper and should be as descriptive and concise as possible. That is why the main sections of research articles always have the same or very similar headers ( Introduction, Methods, Results, Discussion ), with no or only small differences between journals. However, you also need to divide the content of some of these sections (e.g., the method section) into smaller subsections (e.g., Participants, Experimental Design, and Statistical Analysis ), and make sure you follow specific journal formatting styles when doing so. 

If the journal you submit to follows APA style , for example, you are allowed to use up to five levels of headings, depending on the length of your paper, the complexity of your work, and your personal preference. To clearly indicate how each subsection fits into the rest of the text, every header level has a different format – but note that headers are usually not numbered because the different formatting already reflects the text hierarchy.

APA style headings example structure

Level 1 Centered, Bold, Title Case

Text begins as a new paragraph.

Level 2 Left-aligned, Bold, Title Case

Level 3 Left-aligned, Bold Italic, Title Case

Level 4     Indented, Bold, Title Case, Period . Text begins on the same                    

                                 line and continues as a regular paragraph.

Level 5     Indented, Bold Italic, Title Case, Period. Text begins on the                           

                                 same line and continues as a regular paragraph.

If you only need one section header (e.g. Methods ) and one level of subsection headers (e.g., Participants, Experimental Design, and Statistical Analysis ), use Level 1 and Level 2 headers. If you need three levels of headings, use Levels 1, 2, and 3 (and so on). Do not skip levels or combine them in a different way. 

If you write a paper in Chicago style or MLA style , then you don’t need to follow such exact rules for headings and subheadings. Your structure just has to be consistent with the general formatting guidelines of both styles (12-pts Times New Roman font, double-spaced text, 0.5-inch indentation for every new paragraph) and consistent throughout your paper. Make sure the different formatting levels indicate a hierarchy (e.g., boldface for level 1 and italics for level 2, or a larger font size for level 1 and smaller font size for level 2). The main specifics regarding Chicago and MLA headings and subheadings are that they should be written in title case (major words capitalized, most minor words lowercase) and not end in a period. Both styles allow you, however, to number your sections and subsections, for example with an Arabic number and a period, followed by a space and then the section name. 

MLA paper headings example structure

1. Introduction

2. Material and Methods

2.1 Subject Recruitment

2.2 Experimental Procedure

2.3 Statistical Analysis

3.1 Experiment 1

3.2 Experiment 2

4. Discussion

5. Conclusion

What research paper headings do you need?

Your paper obviously needs to contain the main sections ( Introduction, Methods, Results, Discussion, and maybe Conclusion ) and you need to make sure that you name them according to the target journal style (have a look at the author guidelines if you are unsure what the journal style is). The differences between journals are subtle, but some want you to combine the results and discussion sections, for example, while others don’t want you to have a separate conclusion section. You also need to check whether the target journal has specific rules on subsections (or no subsections) within these main sections. The introduction section should usually not be subdivided (but some journals do not mind), while the method section, for example, always needs to have clear subsections.

How to Name Your Sections and Subsections

The method section subheadings should be short and descriptive, but how you subdivide this section depends on the structure you choose to present your work – which can be chronological (e.g., Experiment 1, Experiment 2 ) or follow your main topics (e.g., Visual Experiment, Behavioral Experiment, Questionnaire ). Have a look at this article on how to write the methods for a research paper if you need input on what the best structure for your work is. The method subheadings should only be keywords that tell the reader what information is following, not summaries or conclusions. That means that “ Subject Recruitment ” is a good methods section subheading, but “ Subjects Were Screened Using Questionnaires ” is not.  

The subheadings for the result section should then follow the general structure of your method section, but here you can choose what information you want to put in every subheading. Some authors keep it simple and just subdivide their result section into experiments or measures like the method section, but others use the headings to summarize their findings so that the reader is prepared for the details that follow. You could, for example, simply name your subsections “ Anxiety Levels ” and “ Social Behavior ,” if those are the measures you studied and explained in the method section. 

Or, you could provide the reader with a glimpse into the results of the analyses you are going to describe, and instead name these subsections “ Anxiety-Like Behaviors in Mutant Mice ” and “ Normal Social Behaviors in Mutant Mice .” While keeping headings short and simple is always a good idea, such mini-summaries can make your result section much clearer and easier to follow. Just make sure that the target journal you want to submit to does not have a rule against that. 

Common Heading and Subheading Mistakes 

Subheadings are not sentences.

If your heading reads like a full sentence, then you can most probably omit the verb or generally rephrase to shorten it. That also means a heading should not contain punctuation except maybe colons or question marks – definitely don’t put a period at the end, except when you have reached heading level 4 in the APA formatting style (see above) and the rules say so.  

Be consistent

Always check your numbering, for example for spaces and periods before and after numbers (e.g., 3.2. vs 3.2 ), because readability depends on such features. But also make sure that your headings are consistent in structure and content: Switching between short keyword headings (e.g., “ Experiment 2 ”) and summary headings (e.g., “ Mice Do not Recognize People ”) is confusing and never a good idea. Ideally, subheadings within a section all have a similar structure. If your first subsection is called “ Mice Do not Recognize People ,” then “ People Do not Recognize Mice” is a better subheader for the next subsection than “Do People Recognize Mice? ”, because consistency is more important in a research paper than creativity. 

Don’t overdo it

Not every paragraph or every argument needs a subheading. Only use subheadings within a bigger section if you have more than one point to make per heading level, and if subdividing the section really makes the structure clearer overall.

Before submitting your journal manuscript to academic publishers, be sure to get English editing services , including manuscript editing or paper editing from a trusted source. And receive instant proofreading with Wordvice AI, our AI online text editor , which provides unlimited editing while drafting your research work.

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What are headings?

Headings, sections, subsections, or levels of subordination are a style of dividing your research paper into major parts, then minor subsections. Most college papers do not need headings, especially if you are only producing two to five pages. However, if your professor requests you use headings or your are writing an especially long or detailed paper, then use headings to help readers navigate your text. Follow the APA style rules for creating the correct level of heading. Always start with a level one heading and drill down to the last subsection possible (five) in order as seen below. Instructions and examples for headings are available on p. 47- 49 of the new APA 7th Edition manual.

Levels of Headings

Additional headings resources.

  • APA Style: Headings This page of the APA Style Blog provides more details about styling paper section headings in APA style.
  • Heading Levels Template: Student Paper APA Style 7th Edition This example student paper clearly illustrates how to style section headings including the paper title and the Introduction section (which should not be labeled Introduction as APA assumes all papers begin with an introduction section).

Proper Title Case vs. lowercase paragraph heading

Proper title case is using both uppercase and lowercase letters in a title. It calls for the major words to be capitalized while any small conjunctions are made smaller, i.e., 

The Title of this Paper is Lengthy

Lowercase paragraph heading calls for the first word to be capitalized along with any proper nouns contained within the heading, i.e., 

        The title of this heading is much shorter and all lowercase except for the first word.

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heading in a research paper

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13.1 Formatting a Research Paper

Learning objectives.

  • Identify the major components of a research paper written using American Psychological Association (APA) style.
  • Apply general APA style and formatting conventions in a research paper.

In this chapter, you will learn how to use APA style , the documentation and formatting style followed by the American Psychological Association, as well as MLA style , from the Modern Language Association. There are a few major formatting styles used in academic texts, including AMA, Chicago, and Turabian:

  • AMA (American Medical Association) for medicine, health, and biological sciences
  • APA (American Psychological Association) for education, psychology, and the social sciences
  • Chicago—a common style used in everyday publications like magazines, newspapers, and books
  • MLA (Modern Language Association) for English, literature, arts, and humanities
  • Turabian—another common style designed for its universal application across all subjects and disciplines

While all the formatting and citation styles have their own use and applications, in this chapter we focus our attention on the two styles you are most likely to use in your academic studies: APA and MLA.

If you find that the rules of proper source documentation are difficult to keep straight, you are not alone. Writing a good research paper is, in and of itself, a major intellectual challenge. Having to follow detailed citation and formatting guidelines as well may seem like just one more task to add to an already-too-long list of requirements.

Following these guidelines, however, serves several important purposes. First, it signals to your readers that your paper should be taken seriously as a student’s contribution to a given academic or professional field; it is the literary equivalent of wearing a tailored suit to a job interview. Second, it shows that you respect other people’s work enough to give them proper credit for it. Finally, it helps your reader find additional materials if he or she wishes to learn more about your topic.

Furthermore, producing a letter-perfect APA-style paper need not be burdensome. Yes, it requires careful attention to detail. However, you can simplify the process if you keep these broad guidelines in mind:

  • Work ahead whenever you can. Chapter 11 “Writing from Research: What Will I Learn?” includes tips for keeping track of your sources early in the research process, which will save time later on.
  • Get it right the first time. Apply APA guidelines as you write, so you will not have much to correct during the editing stage. Again, putting in a little extra time early on can save time later.
  • Use the resources available to you. In addition to the guidelines provided in this chapter, you may wish to consult the APA website at http://www.apa.org or the Purdue University Online Writing lab at http://owl.english.purdue.edu , which regularly updates its online style guidelines.

General Formatting Guidelines

This chapter provides detailed guidelines for using the citation and formatting conventions developed by the American Psychological Association, or APA. Writers in disciplines as diverse as astrophysics, biology, psychology, and education follow APA style. The major components of a paper written in APA style are listed in the following box.

These are the major components of an APA-style paper:

Body, which includes the following:

  • Headings and, if necessary, subheadings to organize the content
  • In-text citations of research sources
  • References page

All these components must be saved in one document, not as separate documents.

The title page of your paper includes the following information:

  • Title of the paper
  • Author’s name
  • Name of the institution with which the author is affiliated
  • Header at the top of the page with the paper title (in capital letters) and the page number (If the title is lengthy, you may use a shortened form of it in the header.)

List the first three elements in the order given in the previous list, centered about one third of the way down from the top of the page. Use the headers and footers tool of your word-processing program to add the header, with the title text at the left and the page number in the upper-right corner. Your title page should look like the following example.

Beyond the Hype: Evaluating Low-Carb Diets cover page

The next page of your paper provides an abstract , or brief summary of your findings. An abstract does not need to be provided in every paper, but an abstract should be used in papers that include a hypothesis. A good abstract is concise—about one hundred fifty to two hundred fifty words—and is written in an objective, impersonal style. Your writing voice will not be as apparent here as in the body of your paper. When writing the abstract, take a just-the-facts approach, and summarize your research question and your findings in a few sentences.

In Chapter 12 “Writing a Research Paper” , you read a paper written by a student named Jorge, who researched the effectiveness of low-carbohydrate diets. Read Jorge’s abstract. Note how it sums up the major ideas in his paper without going into excessive detail.

Beyond the Hype: Abstract

Write an abstract summarizing your paper. Briefly introduce the topic, state your findings, and sum up what conclusions you can draw from your research. Use the word count feature of your word-processing program to make sure your abstract does not exceed one hundred fifty words.

Depending on your field of study, you may sometimes write research papers that present extensive primary research, such as your own experiment or survey. In your abstract, summarize your research question and your findings, and briefly indicate how your study relates to prior research in the field.

Margins, Pagination, and Headings

APA style requirements also address specific formatting concerns, such as margins, pagination, and heading styles, within the body of the paper. Review the following APA guidelines.

Use these general guidelines to format the paper:

  • Set the top, bottom, and side margins of your paper at 1 inch.
  • Use double-spaced text throughout your paper.
  • Use a standard font, such as Times New Roman or Arial, in a legible size (10- to 12-point).
  • Use continuous pagination throughout the paper, including the title page and the references section. Page numbers appear flush right within your header.
  • Section headings and subsection headings within the body of your paper use different types of formatting depending on the level of information you are presenting. Additional details from Jorge’s paper are provided.

Cover Page

Begin formatting the final draft of your paper according to APA guidelines. You may work with an existing document or set up a new document if you choose. Include the following:

  • Your title page
  • The abstract you created in Note 13.8 “Exercise 1”
  • Correct headers and page numbers for your title page and abstract

APA style uses section headings to organize information, making it easy for the reader to follow the writer’s train of thought and to know immediately what major topics are covered. Depending on the length and complexity of the paper, its major sections may also be divided into subsections, sub-subsections, and so on. These smaller sections, in turn, use different heading styles to indicate different levels of information. In essence, you are using headings to create a hierarchy of information.

The following heading styles used in APA formatting are listed in order of greatest to least importance:

  • Section headings use centered, boldface type. Headings use title case, with important words in the heading capitalized.
  • Subsection headings use left-aligned, boldface type. Headings use title case.
  • The third level uses left-aligned, indented, boldface type. Headings use a capital letter only for the first word, and they end in a period.
  • The fourth level follows the same style used for the previous level, but the headings are boldfaced and italicized.
  • The fifth level follows the same style used for the previous level, but the headings are italicized and not boldfaced.

Visually, the hierarchy of information is organized as indicated in Table 13.1 “Section Headings” .

Table 13.1 Section Headings

A college research paper may not use all the heading levels shown in Table 13.1 “Section Headings” , but you are likely to encounter them in academic journal articles that use APA style. For a brief paper, you may find that level 1 headings suffice. Longer or more complex papers may need level 2 headings or other lower-level headings to organize information clearly. Use your outline to craft your major section headings and determine whether any subtopics are substantial enough to require additional levels of headings.

Working with the document you developed in Note 13.11 “Exercise 2” , begin setting up the heading structure of the final draft of your research paper according to APA guidelines. Include your title and at least two to three major section headings, and follow the formatting guidelines provided above. If your major sections should be broken into subsections, add those headings as well. Use your outline to help you.

Because Jorge used only level 1 headings, his Exercise 3 would look like the following:

Citation Guidelines

In-text citations.

Throughout the body of your paper, include a citation whenever you quote or paraphrase material from your research sources. As you learned in Chapter 11 “Writing from Research: What Will I Learn?” , the purpose of citations is twofold: to give credit to others for their ideas and to allow your reader to follow up and learn more about the topic if desired. Your in-text citations provide basic information about your source; each source you cite will have a longer entry in the references section that provides more detailed information.

In-text citations must provide the name of the author or authors and the year the source was published. (When a given source does not list an individual author, you may provide the source title or the name of the organization that published the material instead.) When directly quoting a source, it is also required that you include the page number where the quote appears in your citation.

This information may be included within the sentence or in a parenthetical reference at the end of the sentence, as in these examples.

Epstein (2010) points out that “junk food cannot be considered addictive in the same way that we think of psychoactive drugs as addictive” (p. 137).

Here, the writer names the source author when introducing the quote and provides the publication date in parentheses after the author’s name. The page number appears in parentheses after the closing quotation marks and before the period that ends the sentence.

Addiction researchers caution that “junk food cannot be considered addictive in the same way that we think of psychoactive drugs as addictive” (Epstein, 2010, p. 137).

Here, the writer provides a parenthetical citation at the end of the sentence that includes the author’s name, the year of publication, and the page number separated by commas. Again, the parenthetical citation is placed after the closing quotation marks and before the period at the end of the sentence.

As noted in the book Junk Food, Junk Science (Epstein, 2010, p. 137), “junk food cannot be considered addictive in the same way that we think of psychoactive drugs as addictive.”

Here, the writer chose to mention the source title in the sentence (an optional piece of information to include) and followed the title with a parenthetical citation. Note that the parenthetical citation is placed before the comma that signals the end of the introductory phrase.

David Epstein’s book Junk Food, Junk Science (2010) pointed out that “junk food cannot be considered addictive in the same way that we think of psychoactive drugs as addictive” (p. 137).

Another variation is to introduce the author and the source title in your sentence and include the publication date and page number in parentheses within the sentence or at the end of the sentence. As long as you have included the essential information, you can choose the option that works best for that particular sentence and source.

Citing a book with a single author is usually a straightforward task. Of course, your research may require that you cite many other types of sources, such as books or articles with more than one author or sources with no individual author listed. You may also need to cite sources available in both print and online and nonprint sources, such as websites and personal interviews. Chapter 13 “APA and MLA Documentation and Formatting” , Section 13.2 “Citing and Referencing Techniques” and Section 13.3 “Creating a References Section” provide extensive guidelines for citing a variety of source types.

Writing at Work

APA is just one of several different styles with its own guidelines for documentation, formatting, and language usage. Depending on your field of interest, you may be exposed to additional styles, such as the following:

  • MLA style. Determined by the Modern Languages Association and used for papers in literature, languages, and other disciplines in the humanities.
  • Chicago style. Outlined in the Chicago Manual of Style and sometimes used for papers in the humanities and the sciences; many professional organizations use this style for publications as well.
  • Associated Press (AP) style. Used by professional journalists.

References List

The brief citations included in the body of your paper correspond to the more detailed citations provided at the end of the paper in the references section. In-text citations provide basic information—the author’s name, the publication date, and the page number if necessary—while the references section provides more extensive bibliographical information. Again, this information allows your reader to follow up on the sources you cited and do additional reading about the topic if desired.

The specific format of entries in the list of references varies slightly for different source types, but the entries generally include the following information:

  • The name(s) of the author(s) or institution that wrote the source
  • The year of publication and, where applicable, the exact date of publication
  • The full title of the source
  • For books, the city of publication
  • For articles or essays, the name of the periodical or book in which the article or essay appears
  • For magazine and journal articles, the volume number, issue number, and pages where the article appears
  • For sources on the web, the URL where the source is located

The references page is double spaced and lists entries in alphabetical order by the author’s last name. If an entry continues for more than one line, the second line and each subsequent line are indented five spaces. Review the following example. ( Chapter 13 “APA and MLA Documentation and Formatting” , Section 13.3 “Creating a References Section” provides extensive guidelines for formatting reference entries for different types of sources.)

References Section

In APA style, book and article titles are formatted in sentence case, not title case. Sentence case means that only the first word is capitalized, along with any proper nouns.

Key Takeaways

  • Following proper citation and formatting guidelines helps writers ensure that their work will be taken seriously, give proper credit to other authors for their work, and provide valuable information to readers.
  • Working ahead and taking care to cite sources correctly the first time are ways writers can save time during the editing stage of writing a research paper.
  • APA papers usually include an abstract that concisely summarizes the paper.
  • APA papers use a specific headings structure to provide a clear hierarchy of information.
  • In APA papers, in-text citations usually include the name(s) of the author(s) and the year of publication.
  • In-text citations correspond to entries in the references section, which provide detailed bibliographical information about a source.

Writing for Success Copyright © 2015 by University of Minnesota is licensed under a Creative Commons Attribution-NonCommercial-ShareAlike 4.0 International License , except where otherwise noted.

APA Style 7th Edition: Citing Your Sources

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What are headings?

Levels of headings.

  • Academic Integrity and Plagiarism
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Headings are used to effectively organize ideas within a study or manuscript.  It can also highlight important items, themes or topics within sections.  By creating concise headings, the reader can anticipate key points and track the development of your argument.  The heading levels establish the hierarchy of each section and are designated by their formatting.

Adapted from American Psychological Association. (2009). Format for Five Levels of Heading in APA Journals. Publication manual of the American psychological association (6th ed., p. 62) Washington, D.C.: American Psychological Association.

  • If you need to use subsections in any given section, use a least two, otherwise omit their use.
  • Do not label headings with numbers or letters
  • Use of title case : Use of both upper and lower case letters, all major words are capitalized
  • Paragraph headings are immediately followed by text for that subsection, rather than starting on a new line.  The heading sits at the start of the first paragraph for that section.
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Practical Ideas on How to Properly Head a Paper

College provides numerous obstacles for students to overcome in different ways. The changes are very apparent, from the freedom to attend classes at your convenience to handling bulky assignments and study requirements.

The way you handle your research papers is also different from what you may be familiar with in high school. There are basic rules to follow in heading a college paper, and once you get those rules down, understanding how to head a paper becomes easy.

Of course, that is easier said than done. So, in this article, we provide you with simple and practical tips on how to properly head a paper.

What is the header for a paper?

A header is a line of text that appears at the top of a page. It usually consists of the author’s last name, the title, and the page number. Essay headers are often needed for different academic texts.

The page header appears at the top margin of every page of your research paper. It normally consists of the page number for student papers only. For professional papers, it consists of the page number and running head.

Headers for any page must be appropriately formatted. They should also be of similar fonts, sizes, and styles placed in the same position on every page. This is often at the top of the page, on the first line.

Why is research paper heading important?

Headers are crucial because they identify the information within the different sections of your paper. They should be as descriptive and concise as possible, with their main sections permitted to have very similar headers to describe each content accurately.

Secondly, headers make your writing look polished and organized. Moreover, a professor examining several student essays would find it easier to sort if the essay pages get mixed up.

Lastly, when writing a proper heading for paper, you may need to divide the content of some sections into smaller subsections. Doing this gives your research content clarity and makes the whole reading process less demanding. However, it is important to follow specific journal formatting styles.

How to write a header for a paper?

In learning how to write heading for your papers, some guidelines will help you along the way. Some tips for writing a good header for your paper include:

  • Consider Paper Style

Often, the MLA style is utilised for most papers, including homework assignments. When using this format on your research paper, don’t skip lines between the initial heading. Your heading on each page should contain your last name and the page number in a right-justified format.

Your heading structure will differ from other paper formats you might use in college, such as American Psychological Association (APA) and Chicago Style.

  • Understand Your Margins

After format consideration, your heading will often be placed in the upper left-hand corner of the page. Margin size and spacing differ in formatting styles and understanding your margins will enable you to accurately place your headings for any style.

  • Use Standard Fonts

After considering your paper style and understanding your margins, use a 12-point font and a standard font style. The Times New Roman font is often used, but Arial, Modern, Lucina, and Palermo can also be used for headers because they are not script-style fonts.

  • Writing Names

It would be best to always place your first and last names on the first line, then double space each line of the heading. All lines in the heading are typically justified at the left margin.

Next, put your professor’s name on the nearest line and use their first and last name preceded by Professor. For example, “Professor Eric Thomas” goes on this line. Afterward, place the name of your course on the next line.

The date usually enters the final line, and your first paragraph begins after double spacing.

In learning how to head papers in college or even in high school, you should take note of the guidelines above. Remember the paper format you’re asked to write in, and you don’t have to type a header for each page of your essay. Writing tools like Microsoft Word make it easy to write a header that appears in the correct format at the top of each page.

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How the brain is flexible enough for a complex world (without being thrown into chaos)

Many neurons exhibit “mixed selectivity,” meaning they can integrate multiple inputs and participate in multiple computations. Mechanisms such as oscillations and neuromodulators recruit their participation and tune them to focus on the relevant information.

Every day our brains strive to optimize a trade-off: With lots of things happening around us even as we also harbor many internal drives and memories, somehow our thoughts must be flexible yet focused enough to guide everything we have to do. In a new paper in Neuron, a team of neuroscientists describes how the brain achieves the cognitive capacity to incorporate all the information that’s relevant without becoming overwhelmed by what’s not.

The authors argue that the flexibility arises from a key property observed in many neurons: “mixed selectivity.” While many neuroscientists used to think each cell had just one dedicated function, more recent evidence has shown that many neurons can instead participate in a variety of computational ensembles, each working in parallel. In other words, when a rabbit considers nibbling on some lettuce in a garden, a single neuron might be involved in not only assessing how hungry it feels but also whether it can hear a hawk overhead or smell a coyote in the trees and how far away the lettuce is.

The brain does not multitask, said paper co-author Earl K. Miller , Picower Professor in The Picower Institute for Learning and Memory at MIT and a pioneer of the mixed selectivity idea, but many cells do have the capacity to be roped into multiple computational efforts (essentially “thoughts”). In the new paper the authors describe specific mechanisms the brain employs to recruit neurons into different computations and to ensure that those neurons represent the right number of dimensions of a complex task.

“These neurons wear multiple hats,” Miller said. “With mixed selectivity you can have a representational space that’s as complex as it needs to be and no more complex. That’s what flexible cognition is all about.”

Co-author Kay Tye , Professor at The Salk Institute and the University of California at San Diego, said mixed selectivity among neurons particularly in the medial prefrontal cortex is key to enabling many mental abilities.

"The mPFC is like a hum of whispers that represents so much information through highly flexible and dynamic ensembles," Tye said. “Mixed selectivity is the property that endows us with our flexibility, cognitive capacity, and ability to be creative.  It is the secret to maximizing computational power which is essentially the underpinnings of intelligence."

Origins of an idea

The idea of mixed selectivity germinated in 2000 when Miller and colleague John Duncan defended a surprising result from a study of cognition in Miller’s lab. As animals sorted images into categories, about 30 percent of the neurons in the prefrontal cortex of the brain seemed to be involved. Skeptics who believed that every neuron had a dedicated function scoffed that the brain would devote so many cells to just one task. Miller and Duncan’s answer was that perhaps cells had the flexibility to be involved in many computations. The ability to serve on one cerebral task force, as it were, did not preclude them from being able to serve many others.

But what benefit does mixed selectivity convey? In 2013 Miller teamed up with two co-authors of the new paper, Mattia Rigotti of IBM Research and Stefano Fusi of Columbia University, to show how mixed selectivity endows the brain with powerful computational flexibility. Essentially, an ensemble of neurons with mixed selectivity can accommodate many more dimensions of information about a task than a population of neurons with invariant functions.

“Since our original work, we've made progress understanding the theory of mixed selectivity through the lens of classical machine learning ideas,” Rigotti said. “On the other hand, questions dear to experimentalists about the mechanisms implementing it at a cellular level had been comparatively under-explored. This collaboration and this new paper set out to fill that gap.”

In the new paper the authors imagine a mouse who is considering whether to eat a berry. It might smell delicious (that’s one dimension). It might be poisonous (that’s another). Yet another dimension or two of the problem could come in the form of a social cue. If the mouse smells the berry scent on a fellow mouse’s breath, then the berry is probably OK to eat (depending on the apparent health of the fellow mouse). A neural ensemble with mixed selectivity would be able to integrate all that.

Recruiting neurons

While mixed selectivity has the backing of copious evidence—it has been observed across the cortex and in other brain areas such as the hippocampus and amygdala—there are still open questions. For instance, how are neurons recruited to tasks and how do neurons that are so “open-minded” remain tuned only to what really matters to the mission?

In the new study, the researchers who also include Marcus Benna of UC San Diego and Felix Taschbach of The Salk Institute, define the forms of mixed selectivity that researchers have observed, and argue that when oscillations (also known as “brain waves”) and neuromodulators (chemicals such as serotonin or dopamine that influence neural function) recruit neurons into computational ensembles, they also help them “gate” what’s important for that purpose.

To be sure, some neurons are dedicated to a specific input, but the authors note they are an exception rather than the rule. The authors say these cells have “pure selectivity.” They only care if the rabbit sees lettuce. Some neurons exhibit “linear mixed selectivity,” which means their response predictably depends on multiple inputs adding up (the rabbit sees lettuce and feels hungry). The neurons that add the most dimensional flexibility are the “nonlinear mixed selectivity” ones that can account for multiple independent variables without necessarily summing them. Instead they might weigh a whole set of independent conditions (e.g. there’s lettuce, I’m hungry, I hear no hawks, I smell no coyotes, but the lettuce is far and I see a pretty sturdy fence).

So what brings neurons into the fold to focus on the salient factors, however many there are? One mechanism is oscillations, which are produced in the brain when many neurons all maintain their electrical activity at the same rhythm. This coordinated activity enables information sharing, essentially tuning them together like a bunch of cars all playing the same radio station (maybe the broadcast is about a hawk circling overhead). Another mechanism the authors highlight is neuromodulators. These are chemicals that upon reaching receptors within cells can influence their activity as well. A burst of acetylcholine, for instance, might similarly attune neurons with the right receptors to certain activity or information (like maybe that feeling of hunger).

“These two mechanisms likely work together to dynamically form functional networks,” the authors write.

Understanding mixed selectivity, they continue, is critical to understanding cognition.

“Mixed selectivity is ubiquitous,” they conclude. “It is present across species and across functions from high-level cognition to ‘automatic’ sensorimotor processes such as object recognition. The widespread presence of mixed selectivity underscores its fundamental role in providing the brain with the scalable processing power needed for complex thought and action.”

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16.1: Formatting a Research Paper

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Learning Objectives

  • Identify the major components of a research paper written using American Psychological Association (APA) style.
  • Apply general APA style and formatting conventions in a research paper.

In this chapter, you will learn how to use APA style, the documentation and formatting style followed by the American Psychological Association, as well as MLA style, from the Modern Language Association. There are a few major formatting styles used in academic texts, including AMA, Chicago, and Turabian:

  • AMA (American Medical Association) for medicine, health, and biological sciences
  • APA (American Psychological Association) for education, psychology, and the social sciences
  • Chicago—a common style used in everyday publications like magazines, newspapers, and books
  • MLA (Modern Language Association) for English, literature, arts, and humanities
  • Turabian—another common style designed for its universal application across all subjects and disciplines

While all the formatting and citation styles have their own use and applications, in this chapter we focus our attention on the two styles you are most likely to use in your academic studies: APA and MLA.

If you find that the rules of proper source documentation are difficult to keep straight, you are not alone. Writing a good research paper is, in and of itself, a major intellectual challenge. Having to follow detailed citation and formatting guidelines as well may seem like just one more task to add to an already-too-long list of requirements.

Following these guidelines, however, serves several important purposes. First, it signals to your readers that your paper should be taken seriously as a student’s contribution to a given academic or professional field; it is the literary equivalent of wearing a tailored suit to a job interview. Second, it shows that you respect other people’s work enough to give them proper credit for it. Finally, it helps your reader find additional materials if he or she wishes to learn more about your topic.

Furthermore, producing a letter-perfect APA-style paper need not be burdensome. Yes, it requires careful attention to detail. However, you can simplify the process if you keep these broad guidelines in mind:

  • Work ahead whenever you can. Chapter 11 includes tips for keeping track of your sources early in the research process, which will save time later on.
  • Get it right the first time. Apply APA guidelines as you write, so you will not have much to correct during the editing stage. Again, putting in a little extra time early on can save time later.
  • Use the resources available to you. In addition to the guidelines provided in this chapter, you may wish to consult the APA website at http://www.apa.org or the Purdue University Online Writing lab at http://owl.english.purdue.edu , which regularly updates its online style guidelines.

General Formatting Guidelines

This chapter provides detailed guidelines for using the citation and formatting conventions developed by the American Psychological Association, or APA. Writers in disciplines as diverse as astrophysics, biology, psychology, and education follow APA style. The major components of a paper written in APA style are listed in the following box.

These are the major components of an APA-style paper:

  • Headings and, if necessary, subheadings to organize the content
  • In-text citations of research sources
  • References page

All these components must be saved in one document, not as separate documents.

The title page of your paper includes the following information:

  • Title of the paper
  • Author’s name
  • Name of the institution with which the author is affiliated
  • Header at the top of the page with the paper title (in capital letters) and the page number (If the title is lengthy, you may use a shortened form of it in the header.)

List the first three elements in the order given in the previous list, centered about one third of the way down from the top of the page. Use the headers and footers tool of your word-processing program to add the header, with the title text at the left and the page number in the upper-right corner. Your title page should look like the following example.

Beyond the Hype: Evaluating Low-Carb Diets cover page

The next page of your paper provides an abstract, or brief summary of your findings. An abstract does not need to be provided in every paper, but an abstract should be used in papers that include a hypothesis. A good abstract is concise—about one hundred to one hundred fifty words—and is written in an objective, impersonal style. Your writing voice will not be as apparent here as in the body of your paper. When writing the abstract, take a just-the-facts approach, and summarize your research question and your findings in a few sentences.

In Chapter 12, you read a paper written by a student named Jorge, who researched the effectiveness of low-carbohydrate diets. Read Jorge’s abstract. Note how it sums up the major ideas in his paper without going into excessive detail.

Beyond the Hype: Abstract

Exercise \(\PageIndex{1}\)

Write an abstract summarizing your paper. Briefly introduce the topic, state your findings, and sum up what conclusions you can draw from your research. Use the word count feature of your word-processing program to make sure your abstract does not exceed one hundred fifty words.

Depending on your field of study, you may sometimes write research papers that present extensive primary research, such as your own experiment or survey. In your abstract, summarize your research question and your findings, and briefly indicate how your study relates to prior research in the field.

Margins, Pagination, and Headings

APA style requirements also address specific formatting concerns, such as margins, pagination, and heading styles, within the body of the paper. Review the following APA guidelines.

Use these general guidelines to format the paper:

  • Set the top, bottom, and side margins of your paper at 1 inch.
  • Use double-spaced text throughout your paper.
  • Use a standard font, such as Times New Roman or Arial, in a legible size (10- to 12-point).
  • Use continuous pagination throughout the paper, including the title page and the references section. Page numbers appear flush right within your header.
  • Section headings and subsection headings within the body of your paper use different types of formatting depending on the level of information you are presenting. Additional details from Jorge’s paper are provided.

Cover Page

Exercise \(\PageIndex{2}\)

Begin formatting the final draft of your paper according to APA guidelines. You may work with an existing document or set up a new document if you choose. Include the following:

  • Your title page
  • The abstract you created in Exercise 1
  • Correct headers and page numbers for your title page and abstract

APA style uses section headings to organize information, making it easy for the reader to follow the writer’s train of thought and to know immediately what major topics are covered. Depending on the length and complexity of the paper, its major sections may also be divided into subsections, sub-subsections, and so on. These smaller sections, in turn, use different heading styles to indicate different levels of information. In essence, you are using headings to create a hierarchy of information.

The following heading styles used in APA formatting are listed in order of greatest to least importance:

  • Section headings use centered, boldface type. Headings use title case, with important words in the heading capitalized.
  • Subsection headings use left-aligned, boldface type. Headings use title case.
  • The third level uses left-aligned, indented, boldface type. Headings use a capital letter only for the first word, and they end in a period.
  • The fourth level follows the same style used for the previous level, but the headings are boldfaced and italicized.
  • The fifth level follows the same style used for the previous level, but the headings are italicized and not boldfaced.

Visually, the hierarchy of information is organized as indicated in Table 13.1 “Section Headings”.

A college research paper may not use all the heading levels shown in Table 13.1 “Section Headings”, but you are likely to encounter them in academic journal articles that use APA style. For a brief paper, you may find that level 1 headings suffice. Longer or more complex papers may need level 2 headings or other lower-level headings to organize information clearly. Use your outline to craft your major section headings and determine whether any subtopics are substantial enough to require additional levels of headings.

Exercise \(\PageIndex{3}\)

Working with the document you developed in Note 13.11 “Exercise 2”, begin setting up the heading structure of the final draft of your research paper according to APA guidelines. Include your title and at least two to three major section headings, and follow the formatting guidelines provided above. If your major sections should be broken into subsections, add those headings as well. Use your outline to help you.

Because Jorge used only level 1 headings, his Exercise 3 would look like the following:

Citation Guidelines

In-text citations.

Throughout the body of your paper, include a citation whenever you quote or paraphrase material from your research sources. As you learned in Chapter 11, the purpose of citations is twofold: to give credit to others for their ideas and to allow your reader to follow up and learn more about the topic if desired. Your in-text citations provide basic information about your source; each source you cite will have a longer entry in the references section that provides more detailed information.

In-text citations must provide the name of the author or authors and the year the source was published. (When a given source does not list an individual author, you may provide the source title or the name of the organization that published the material instead.) When directly quoting a source, it is also required that you include the page number where the quote appears in your citation.

This information may be included within the sentence or in a parenthetical reference at the end of the sentence, as in these examples.

Epstein (2010) points out that “junk food cannot be considered addictive in the same way that we think of psychoactive drugs as addictive” (p. 137).

Here, the writer names the source author when introducing the quote and provides the publication date in parentheses after the author’s name. The page number appears in parentheses after the closing quotation marks and before the period that ends the sentence.

Addiction researchers caution that “junk food cannot be considered addictive in the same way that we think of psychoactive drugs as addictive” (Epstein, 2010, p. 137).

Here, the writer provides a parenthetical citation at the end of the sentence that includes the author’s name, the year of publication, and the page number separated by commas. Again, the parenthetical citation is placed after the closing quotation marks and before the period at the end of the sentence.

As noted in the book Junk Food, Junk Science (Epstein, 2010, p. 137), “junk food cannot be considered addictive in the same way that we think of psychoactive drugs as addictive.”

Here, the writer chose to mention the source title in the sentence (an optional piece of information to include) and followed the title with a parenthetical citation. Note that the parenthetical citation is placed before the comma that signals the end of the introductory phrase.

David Epstein’s book Junk Food, Junk Science (2010) pointed out that “junk food cannot be considered addictive in the same way that we think of psychoactive drugs as addictive” (p. 137).

Another variation is to introduce the author and the source title in your sentence and include the publication date and page number in parentheses within the sentence or at the end of the sentence. As long as you have included the essential information, you can choose the option that works best for that particular sentence and source.

Citing a book with a single author is usually a straightforward task. Of course, your research may require that you cite many other types of sources, such as books or articles with more than one author or sources with no individual author listed. You may also need to cite sources available in both print and online and nonprint sources, such as websites and personal interviews. Section 13.2 and Section 13.3 provide extensive guidelines for citing a variety of source types.

writing at work

APA is just one of several different styles with its own guidelines for documentation, formatting, and language usage. Depending on your field of interest, you may be exposed to additional styles, such as the following:

  • MLA style. Determined by the Modern Languages Association and used for papers in literature, languages, and other disciplines in the humanities.
  • Chicago style. Outlined in the Chicago Manual of Style and sometimes used for papers in the humanities and the sciences; many professional organizations use this style for publications as well.
  • Associated Press (AP) style. Used by professional journalists.

References List

The brief citations included in the body of your paper correspond to the more detailed citations provided at the end of the paper in the references section. In-text citations provide basic information—the author’s name, the publication date, and the page number if necessary—while the references section provides more extensive bibliographical information. Again, this information allows your reader to follow up on the sources you cited and do additional reading about the topic if desired.

The specific format of entries in the list of references varies slightly for different source types, but the entries generally include the following information:

  • The name(s) of the author(s) or institution that wrote the source
  • The year of publication and, where applicable, the exact date of publication
  • The full title of the source
  • For books, the city of publication
  • For articles or essays, the name of the periodical or book in which the article or essay appears
  • For magazine and journal articles, the volume number, issue number, and pages where the article appears
  • For sources on the web, the URL where the source is located

The references page is double spaced and lists entries in alphabetical order by the author’s last name. If an entry continues for more than one line, the second line and each subsequent line are indented five spaces. Review the following example. (Section 13.3 provides extensive guidelines for formatting reference entries for different types of sources.)

b561934bebfadaf7ee8c8da990644aac.jpg

In APA style, book and article titles are formatted in sentence case, not title case. Sentence case means that only the first word is capitalized, along with any proper nouns.

Key Takeaways

  • Following proper citation and formatting guidelines helps writers ensure that their work will be taken seriously, give proper credit to other authors for their work, and provide valuable information to readers.
  • Working ahead and taking care to cite sources correctly the first time are ways writers can save time during the editing stage of writing a research paper.
  • APA papers usually include an abstract that concisely summarizes the paper.
  • APA papers use a specific headings structure to provide a clear hierarchy of information.
  • In APA papers, in-text citations usually include the name(s) of the author(s) and the year of publication.
  • In-text citations correspond to entries in the references section, which provide detailed bibliographical information about a source.

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  • Published: 06 May 2024

Electrophysiological signatures of veridical head direction in humans

  • Benjamin J. Griffiths   ORCID: orcid.org/0000-0001-8600-4480 1 , 2 ,
  • Thomas Schreiner   ORCID: orcid.org/0000-0003-2030-5998 1 ,
  • Julia K. Schaefer   ORCID: orcid.org/0000-0002-9246-267X 1 ,
  • Christian Vollmar 3 ,
  • Elisabeth Kaufmann   ORCID: orcid.org/0000-0002-7582-2215 3 ,
  • Stefanie Quach   ORCID: orcid.org/0000-0003-2914-3649 4 ,
  • Jan Remi 3 ,
  • Soheyl Noachtar 3 &
  • Tobias Staudigl   ORCID: orcid.org/0000-0003-2885-1280 1  

Nature Human Behaviour ( 2024 ) Cite this article

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  • Neural circuits

Information about heading direction is critical for navigation as it provides the means to orient ourselves in space. However, given that veridical head-direction signals require physical rotation of the head and most human neuroimaging experiments depend upon fixing the head in position, little is known about how the human brain is tuned to such heading signals. Here we adress this by asking 52 healthy participants undergoing simultaneous electroencephalography and motion tracking recordings (split into two experiments) and 10 patients undergoing simultaneous intracranial electroencephalography and motion tracking recordings to complete a series of orientation tasks in which they made physical head rotations to target positions. We then used a series of forward encoding models and linear mixed-effects models to isolate electrophysiological activity that was specifically tuned to heading direction. We identified a robust posterior central signature that predicts changes in veridical head orientation after regressing out confounds including sensory input and muscular activity. Both source localization and intracranial analysis implicated the medial temporal lobe as the origin of this effect. Subsequent analyses disentangled head-direction signatures from signals relating to head rotation and those reflecting location-specific effects. Lastly, when directly comparing head direction and eye-gaze-related tuning, we found that the brain maintains both codes while actively navigating, with stronger tuning to head direction in the medial temporal lobe. Together, these results reveal a taxonomy of population-level head-direction signals within the human brain that is reminiscent of those reported in the single units of rodents.

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Data availability.

Data acquired from the healthy participants are available at Ludwig-Maximilians-Universität via https://data.ub.uni-muenchen.de/439/ (ref. 87 ). Due to privacy laws, data acquired from the patients are not openly available, though (subject to privacy laws) can be provided by contacting the corresponding author. Source data are provided with this paper.

Code availability

Openly available at GitHub via https://github.com/benjaminGriffiths/human-hd (ref. 88 ).

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NITRIC https://www.nitrc.org/projects/wfu_pickatlas/

GitHub https://github.com/circstat/circstat-matlab

Staudigl, T. Data for: Griffiths et al.: Electrophysiological signatures of veridical head direction in humans (Nature Human Behaviour, 2024). (Open Data LMU, 2024). https://doi.org/10.5282/ubm/data.439

GitHub https://github.com/benjaminGriffiths/human-hd

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Acknowledgements

This work was supported by the European Research Council ( https://erc.europa.eu/ , starting grant 802681 awarded to T. Schreiner) and the Leverhulme Trust ( https://www.leverhulme.ac.uk/ , early career fellowship ECF-2021-628 awarded to B.J.G.). We thank all participants and in particular all patients who volunteered to participate in this study. We thank the staff and physicians at the Epilepsy Center, Department of Neurology, Ludwig-Maximilians-Universität, Munich, for assistance. We thank A. Chowdhury for valuable input. The funders had no role in study design, data collection and analysis, decision to publish or preparation of the manuscript.

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Contributions

B.J.G.: conceptualization, methodology, software, validation and formal analysis; investigation; data curation; writing—original draft; writing—review and editing; and visualization. T. Schreiner: conceptualization, methodology, investigation, data curation and writing—review and editing. J.K.S.: investigation, data curation and writing—review and editing. C.V.: resources and writing—review and editing. E.K.: resources and writing—review and editing. S.Q.: resources and writing—review and editing. J.R.: resources and writing—review and editing. S.N.: resources and writing—review and editing. T. Staudigl: conceptualization, methodology and investigation; writing—original draft; and writing—review and editing, supervision, project administration and funding acquisition.

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Griffiths, B.J., Schreiner, T., Schaefer, J.K. et al. Electrophysiological signatures of veridical head direction in humans. Nat Hum Behav (2024). https://doi.org/10.1038/s41562-024-01872-1

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