introduction body conclusion in research

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Traditional Academic Essays In Three Parts

Part i: the introduction.

An introduction is usually the first paragraph of your academic essay. If you’re writing a long essay, you might need 2 or 3 paragraphs to introduce your topic to your reader. A good introduction does 2 things:

• Gets the reader’s attention. You can get a reader’s attention by telling a story, providing a statistic, pointing out something strange or interesting, providing and discussing an interesting quote, etc. Be interesting and find some original angle via which to engage others in your topic.
• Provides a specific and debatable thesis statement. The thesis statement is usually just one sentence long, but it might be longer—even a whole paragraph—if the essay you’re writing is long. A good thesis statement makes a debatable point, meaning a point someone might disagree with and argue against. It also serves as a roadmap for what you argue in your paper.

Part II: The Body Paragraphs

Body paragraphs help you prove your thesis and move you along a compelling trajectory from your introduction to your conclusion. If your thesis is a simple one, you might not need a lot of body paragraphs to prove it. If it’s more complicated, you’ll need more body paragraphs. An easy way to remember the parts of a body paragraph is to think of them as the MEAT of your essay:

Main Idea. The part of a topic sentence that states the main idea of the body paragraph. All of the sentences in the paragraph connect to it. Keep in mind that main ideas are…

• like labels. They appear in the first sentence of the paragraph and tell your reader what’s inside the paragraph.
• arguable. They’re not statements of fact; they’re debatable points that you prove with evidence.
• focused. Make a specific point in each paragraph and then prove that point.

Evidence. The parts of a paragraph that prove the main idea. You might include different types of evidence in different sentences. Keep in mind that different disciplines have different ideas about what counts as evidence and they adhere to different citation styles. Examples of evidence include…

• quotations and/or paraphrases from sources.
• facts , e.g. statistics or findings from studies you’ve conducted.
• narratives and/or descriptions , e.g. of your own experiences.

Analysis. The parts of a paragraph that explain the evidence. Make sure you tie the evidence you provide back to the paragraph’s main idea. In other words, discuss the evidence.

Transition. The part of a paragraph that helps you move fluidly from the last paragraph. Transitions appear in topic sentences along with main ideas, and they look both backward and forward in order to help you connect your ideas for your reader. Don’t end paragraphs with transitions; start with them.

Keep in mind that MEAT does not occur in that order. The “ T ransition” and the “ M ain Idea” often combine to form the first sentence—the topic sentence—and then paragraphs contain multiple sentences of evidence and analysis. For example, a paragraph might look like this: TM. E. E. A. E. E. A. A.

Part III: The Conclusion

A conclusion is the last paragraph of your essay, or, if you’re writing a really long essay, you might need 2 or 3 paragraphs to conclude. A conclusion typically does one of two things—or, of course, it can do both:

• Summarizes the argument. Some instructors expect you not to say anything new in your conclusion. They just want you to restate your main points. Especially if you’ve made a long and complicated argument, it’s useful to restate your main points for your reader by the time you’ve gotten to your conclusion. If you opt to do so, keep in mind that you should use different language than you used in your introduction and your body paragraphs. The introduction and conclusion shouldn’t be the same.
• For example, your argument might be significant to studies of a certain time period .
• Alternately, it might be significant to a certain geographical region .
• Alternately still, it might influence how your readers think about the future . You might even opt to speculate about the future and/or call your readers to action in your conclusion.

Handout by Dr. Liliana Naydan. Do not reproduce without permission.

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Purdue OWL - Introductions, Body Paragraphs, and Conclusions for an Expository/Persuasive Essay

Introductions, Body Paragraphs, and Conclusions for an Expository/Persuasive Essay

Introduction

The introduction is the broad beginning of the paper that answers three important questions:

• What is this?
• Why am I reading it?
• What do you want me to do?

You should answer these questions by doing the following:

• Set the context – provide general information about the main idea, explaining the situation so the reader can make sense of the topic and the claims you make and support
• State why the main idea is important – tell the reader why s/he should care and keep reading. Your goal is to create a compelling, clear, and convincing essay people will want to read and act upon
• State your thesis/claim – compose a sentence or two stating the position you will support with logos (sound reasoning: induction, deduction), pathos (balanced emotional appeal), and ethos (author credibility).

Thesis Checklist

Your thesis is more than a general statement about your main idea. It needs to establish a clear position you will support with balanced proofs (logos, pathos, ethos). Use the checklist below to help you create a thesis.

This section is adapted from Writing with a Thesis: A Rhetoric Reader by David Skwire and Sarah Skwire:

Make sure you avoid the following when creating your thesis:

• A thesis is not a title: Homes and schools (title) vs. Parents ought to participate more in the education of their children (good thesis).
• A thesis is not an announcement of the subject: My subject is the incompetence of the Supreme Court vs. The Supreme Court made a mistake when it ruled in favor of George W. Bush in the 2000 election.
• A thesis is not a statement of absolute fact: Jane Austen is the author of Pride and Prejudice.
• A thesis is not the whole essay: A thesis is your main idea/claim/refutation/problem-solution expressed in a single sentence or a combination of sentences.
• Please note that according to the MLA Handbook for Writers of Research Papers , Sixth Edition, "A thesis statement is a single sentence that formulates both your topic and your point of view" (Gibaldi 56). However, if your paper is more complex and requires a thesis statement, your thesis may require a combination of sentences .

Make sure you follow these guidelines when creating your thesis:

• A good thesis is unified: Detective stories are not a high form of literature, but people have always been fascinated by them, and many fine writers have experimented with them (floppy). vs. Detective stories appeal to the basic human desire for thrills (concise).
• A good thesis is specific: James Joyce’s Ulysses is very good. vs. James Joyce’s Ulysses helped create a new way for writers to deal with the unconscious.
• Try to be as specific as possible (without providing too much detail) when creating your thesis: James Joyce’s Ulysses helped create a new way for writers to deal with the unconscious. vs. James Joyce’s Ulysses helped create a new way for writers to deal with the unconscious by utilizing the findings of Freudian psychology and introducing the techniques of literary stream-of-consciousness.

Quick Checklist:

_____ The thesis/claim follows the guidelines outlined above

_____ The thesis/claim matches the requirements and goals of the assignment

_____ The thesis/claim is clear and easily recognizable

_____ The thesis/claim seems supportable by good reasoning/data, emotional appeal

Body Paragraphs

Summary: This resource outlines the generally accepted structure for introductions, body paragraphs, and conclusions in an academic argument paper. Keep in mind that this resource contains guidelines and not strict rules about organization. Your structure needs to be flexible enough to meet the requirements of your purpose and audience.

Body Paragraphs: Moving from General to Specific Information

Your paper should be organized in a manner that moves from general to specific information. Every time you begin a new subject, think of an inverted pyramid - the broadest range of information sits at the top, and as the paragraph or paper progresses, the author becomes more and more focused on the argument ending with specific, detailed evidence supporting a claim. Lastly, the author explains how and why the information she has just provided connects to and supports her thesis (a brief wrap up or warrant).

  The four elements of a good paragraph (TTEB)

A good paragraph should contain at least the following four elements: T ransition, T opic sentence, specific E vidence and analysis, and a B rief wrap-up sentence (also known as a warrant) – TTEB!

• A T ransition sentence leading in from a previous paragraph to assure smooth reading. This acts as a hand off from one idea to the next.
• A T opic sentence that tells the reader what you will be discussing in the paragraph.
• Specific E vidence and analysis that supports one of your claims and that provides a deeper level of detail than your topic sentence.
• A B rief wrap-up sentence that tells the reader how and why this information supports the paper’s thesis. The brief wrap-up is also known as the warrant. The warrant is important to your argument because it connects your reasoning and support to your thesis, and it shows that the information in the paragraph is related to your thesis and helps defend it.

Rebuttal Sections

In order to present a fair and convincing message, you may need to anticipate, research, and outline some of the common positions (arguments) that dispute your thesis. If the situation (purpose) calls for you to do this, you will present and then refute these other positions in the rebuttal section of your essay.

It is important to consider other positions because in most cases, your primary audience will be fence-sitters. Fence-sitters are people who have not decided which side of the argument to support.

People who are on your side of the argument will not need a lot of information to align with your position. People who are completely against your argument - perhaps for ethical or religious reasons - will probably never align with your position no matter how much information you provide. Therefore, the audience you should consider most important are those people who haven't decided which side of the argument they will support - the fence-sitters.

In many cases, these fence-sitters have not decided which side to align with because they see value in both positions. Therefore, to not consider opposing positions to your own in a fair manner may alienate fence-sitters when they see that you are not addressing their concerns or discussion opposing positions at all.

Organizing your rebuttal section

Following the TTEB method outlined in the Body Paragraph section, forecast all the information that will follow in the rebuttal section and then move point by point through the other positions addressing each one as you go. The outline below, adapted from Seyler's Understanding Argument , is an example of a rebuttal section from a thesis essay.

When you rebut or refute an opposing position, use the following three-part organization:

The opponent’s argument – Usually, you should not assume that your reader has read or remembered the argument you are refuting. Thus at the beginning of your paragraph, you need to state, accurately and fairly, the main points of the argument you will refute.

Your position – Next, make clear the nature of your disagreement with the argument or position you are refuting. Your position might assert, for example, that a writer has not proved his assertion because he has provided evidence that is outdated, or that the argument is filled with fallacies.

Your refutation – The specifics of your counterargument will depend upon the nature of your disagreement. If you challenge the writer’s evidence, then you must present the more recent evidence. If you challenge assumptions, then you must explain why they do not hold up. If your position is that the piece is filled with fallacies, then you must present and explain each fallacy.

Conclusions

Conclusions wrap up what you have been discussing in your paper. After moving from general to specific information in the introduction and body paragraphs, your conclusion should begin pulling back into more general information that restates the main points of your argument. Conclusions may also call for action or overview future possible research. The following outline may help you conclude your paper:

In a general way,

• restate your topic and why it is important,
• restate your thesis/claim,
• address opposing viewpoints and explain why readers should align with your position,
• call for action or overview future research possibilities.

Remember that once you accomplish these tasks, unless otherwise directed by your instructor, you are finished. Done. Complete. Don't try to bring in new points or end with a whiz bang(!) conclusion or try to solve world hunger in the final sentence of your conclusion. Simplicity is best for a clear, convincing message.

The preacher's maxim is one of the most effective formulas to follow for argument papers:

• Tell what you're going to tell them (introduction).
• Tell them (body).
• Tell them what you told them (conclusion).

 Copyright ©1995-2011 by The Writing Lab & The OWL at Purdue and Purdue University .

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Guide to Writing Introductions and Conclusions

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First and last impressions are important in any part of life, especially in writing. This is why the introduction and conclusion of any paper – whether it be a simple essay or a long research paper – are essential. Introductions and conclusions are just as important as the body of your paper. The introduction is what makes the reader want to continue reading your paper. The conclusion is what makes your paper stick in the reader’s mind.

Introductions

Your introductory paragraph should include:

1) Hook:  Description, illustration, narration or dialogue that pulls the reader into your paper topic. This should be interesting and specific.

2) Transition: Sentence that connects the hook with the thesis.

3) Thesis: Sentence (or two) that summarizes the overall main point of the paper. The thesis should answer the prompt question.

The examples below show are several ways to write a good introduction or opening to your paper. One example shows you how to paraphrase in your introduction. This will help you understand the idea of writing sequences using a hook, transition, and thesis statement.

» Thesis Statement Opening

This is the traditional style of opening a paper. This is a “mini-summary” of your paper.

For example:

» Opening with a Story (Anecdote)

A good way of catching your reader’s attention is by sharing a story that sets up your paper. Sharing a story gives a paper a more personal feel and helps make your reader comfortable.

This example was borrowed from Jack Gannon’s The Week the World Heard Gallaudet (1989):

Astrid Goodstein, a Gallaudet faculty member, entered the beauty salon for her regular appointment, proudly wearing her DPN button. (“I was married to that button that week!” she later confided.) When Sandy, her regular hairdresser, saw the button, he spoke and gestured, “Never! Never! Never!” Offended, Astrid turned around and headed for the door but stopped short of leaving. She decided to keep her appointment, confessing later that at that moment, her sense of principles had lost out to her vanity. Later she realized that her hairdresser had thought she was pushing for a deaf U.S. President. Hook: a specific example or story that interests the reader and introduces the topic.

Transition: connects the hook to the thesis statement

Thesis: summarizes the overall claim of the paper

» Specific Detail Opening

Giving specific details about your subject appeals to your reader’s curiosity and helps establish a visual picture of what your paper is about.

» Open with a Quotation

Another method of writing an introduction is to open with a quotation. This method makes your introduction more interactive and more appealing to your reader.

» Open with an Interesting Statistic

Statistics that grab the reader help to make an effective introduction.

» Question Openings

Possibly the easiest opening is one that presents one or more questions to be answered in the paper. This is effective because questions are usually what the reader has in mind when he or she sees your topic.

Source : *Writing an Introduction for a More Formal Essay. (2012). Retrieved April 25, 2012, from http://flightline.highline.edu/wswyt/Writing91/handouts/hook_trans_thesis.htm

Conclusions

The conclusion to any paper is the final impression that can be made. It is the last opportunity to get your point across to the reader and leave the reader feeling as if they learned something. Leaving a paper “dangling” without a proper conclusion can seriously devalue what was said in the body itself. Here are a few effective ways to conclude or close your paper. » Summary Closing Many times conclusions are simple re-statements of the thesis. Many times these conclusions are much like their introductions (see Thesis Statement Opening).

» Close with a Logical Conclusion

This is a good closing for argumentative or opinion papers that present two or more sides of an issue. The conclusion drawn as a result of the research is presented here in the final paragraphs.

» Real or Rhetorical Question Closings

This method of concluding a paper is one step short of giving a logical conclusion. Rather than handing the conclusion over, you can leave the reader with a question that causes him or her to draw his own conclusions.

» Close with a Speculation or Opinion This is a good style for instances when the writer was unable to come up with an answer or a clear decision about whatever it was he or she was researching. For example:

» Close with a Recommendation

A good conclusion is when the writer suggests that the reader do something in the way of support for a cause or a plea for them to take action.

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9.4 Writing Introductory and Concluding Paragraphs

Learning objectives.

• Recognize the importance of strong introductory and concluding paragraphs.
• Learn to engage the reader immediately with the introductory paragraph.
• Practice concluding your essays in a more memorable way.

Picture your introduction as a storefront window: You have a certain amount of space to attract your customers (readers) to your goods (subject) and bring them inside your store (discussion). Once you have enticed them with something intriguing, you then point them in a specific direction and try to make the sale (convince them to accept your thesis).

Your introduction is an invitation to your readers to consider what you have to say and then to follow your train of thought as you expand upon your thesis statement.

An introduction serves the following purposes:

• Establishes your voice and tone, or your attitude, toward the subject
• Introduces the general topic of the essay
• States the thesis that will be supported in the body paragraphs

First impressions are crucial and can leave lasting effects in your reader’s mind, which is why the introduction is so important to your essay. If your introductory paragraph is dull or disjointed, your reader probably will not have much interest in continuing with the essay.

Attracting Interest in Your Introductory Paragraph

Your introduction should begin with an engaging statement devised to provoke your readers’ interest. In the next few sentences, introduce them to your topic by stating general facts or ideas about the subject. As you move deeper into your introduction, you gradually narrow the focus, moving closer to your thesis. Moving smoothly and logically from your introductory remarks to your thesis statement can be achieved using a funnel technique , as illustrated in the diagram in Figure 9.1 “Funnel Technique” .

Figure 9.1 Funnel Technique

On a separate sheet of paper, jot down a few general remarks that you can make about the topic for which you formed a thesis in Section 9.1 “Developing a Strong, Clear Thesis Statement” .

Immediately capturing your readers’ interest increases the chances of having them read what you are about to discuss. You can garner curiosity for your essay in a number of ways. Try to get your readers personally involved by doing any of the following:

• Appealing to their emotions
• Using logic
• Beginning with a provocative question or opinion
• Opening with a startling statistic or surprising fact
• Raising a question or series of questions
• Presenting an explanation or rationalization for your essay
• Opening with a relevant quotation or incident
• Opening with a striking image
• Including a personal anecdote

Remember that your diction, or word choice, while always important, is most crucial in your introductory paragraph. Boring diction could extinguish any desire a person might have to read through your discussion. Choose words that create images or express action. For more information on diction, see Chapter 4 “Working with Words: Which Word Is Right?” .

In Chapter 8 “The Writing Process: How Do I Begin?” , you followed Mariah as she moved through the writing process. In this chapter, Mariah writes her introduction and conclusion for the same essay. Mariah incorporates some of the introductory elements into her introductory paragraph, which she previously outlined in Chapter 8 “The Writing Process: How Do I Begin?” . Her thesis statement is underlined.

If you have trouble coming up with a provocative statement for your opening, it is a good idea to use a relevant, attention-grabbing quote about your topic. Use a search engine to find statements made by historical or significant figures about your subject.

Writing at Work

In your job field, you may be required to write a speech for an event, such as an awards banquet or a dedication ceremony. The introduction of a speech is similar to an essay because you have a limited amount of space to attract your audience’s attention. Using the same techniques, such as a provocative quote or an interesting statistic, is an effective way to engage your listeners. Using the funnel approach also introduces your audience to your topic and then presents your main idea in a logical manner.

Reread each sentence in Mariah’s introductory paragraph. Indicate which techniques she used and comment on how each sentence is designed to attract her readers’ interest.

Writing a Conclusion

It is not unusual to want to rush when you approach your conclusion, and even experienced writers may fade. But what good writers remember is that it is vital to put just as much attention into the conclusion as in the rest of the essay. After all, a hasty ending can undermine an otherwise strong essay.

A conclusion that does not correspond to the rest of your essay, has loose ends, or is unorganized can unsettle your readers and raise doubts about the entire essay. However, if you have worked hard to write the introduction and body, your conclusion can often be the most logical part to compose.

The Anatomy of a Strong Conclusion

Keep in mind that the ideas in your conclusion must conform to the rest of your essay. In order to tie these components together, restate your thesis at the beginning of your conclusion. This helps you assemble, in an orderly fashion, all the information you have explained in the body. Repeating your thesis reminds your readers of the major arguments you have been trying to prove and also indicates that your essay is drawing to a close. A strong conclusion also reviews your main points and emphasizes the importance of the topic.

The construction of the conclusion is similar to the introduction, in which you make general introductory statements and then present your thesis. The difference is that in the conclusion you first paraphrase , or state in different words, your thesis and then follow up with general concluding remarks. These sentences should progressively broaden the focus of your thesis and maneuver your readers out of the essay.

Many writers like to end their essays with a final emphatic statement. This strong closing statement will cause your readers to continue thinking about the implications of your essay; it will make your conclusion, and thus your essay, more memorable. Another powerful technique is to challenge your readers to make a change in either their thoughts or their actions. Challenging your readers to see the subject through new eyes is a powerful way to ease yourself and your readers out of the essay.

When closing your essay, do not expressly state that you are drawing to a close. Relying on statements such as in conclusion , it is clear that , as you can see , or in summation is unnecessary and can be considered trite.

It is wise to avoid doing any of the following in your conclusion:

• Introducing new material
• Contradicting your thesis
• Changing your thesis
• Using apologies or disclaimers

Introducing new material in your conclusion has an unsettling effect on your reader. When you raise new points, you make your reader want more information, which you could not possibly provide in the limited space of your final paragraph.

Contradicting or changing your thesis statement causes your readers to think that you do not actually have a conviction about your topic. After all, you have spent several paragraphs adhering to a singular point of view. When you change sides or open up your point of view in the conclusion, your reader becomes less inclined to believe your original argument.

By apologizing for your opinion or stating that you know it is tough to digest, you are in fact admitting that even you know what you have discussed is irrelevant or unconvincing. You do not want your readers to feel this way. Effective writers stand by their thesis statement and do not stray from it.

On a separate sheet of a paper, restate your thesis from Note 9.52 “Exercise 2” of this section and then make some general concluding remarks. Next, compose a final emphatic statement. Finally, incorporate what you have written into a strong conclusion paragraph for your essay.

Collaboration

Please share with a classmate and compare your answers

Mariah incorporates some of these pointers into her conclusion. She has paraphrased her thesis statement in the first sentence.

Make sure your essay is balanced by not having an excessively long or short introduction or conclusion. Check that they match each other in length as closely as possible, and try to mirror the formula you used in each. Parallelism strengthens the message of your essay.

On the job you will sometimes give oral presentations based on research you have conducted. A concluding statement to an oral report contains the same elements as a written conclusion. You should wrap up your presentation by restating the purpose of the presentation, reviewing its main points, and emphasizing the importance of the material you presented. A strong conclusion will leave a lasting impression on your audience.

Key Takeaways

• A strong opening captures your readers’ interest and introduces them to your topic before you present your thesis statement.
• An introduction should restate your thesis, review your main points, and emphasize the importance of the topic.
• The funnel technique to writing the introduction begins with generalities and gradually narrows your focus until you present your thesis.
• A good introduction engages people’s emotions or logic, questions or explains the subject, or provides a striking image or quotation.
• Carefully chosen diction in both the introduction and conclusion prevents any confusing or boring ideas.
• A conclusion that does not connect to the rest of the essay can diminish the effect of your paper.
• The conclusion should remain true to your thesis statement. It is best to avoid changing your tone or your main idea and avoid introducing any new material.
• Closing with a final emphatic statement provides closure for your readers and makes your essay more memorable.

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How to compose a research paper introduction, body and conclusion.

All research paper structures are the same which makes them pretty easy to create if you know what to put where. This article with give some insight into just that so that you can create great works of your own with ease. Keep in mind that research is a huge part of the process but after that is all done and you have notes on all of your research it is best to create an outline for your paper to make is easier to write. In order to do this though, you need to know what information goes where. Here is a guide for how to learn just how to organize your paper.

Three Parts of Any Research Paper

Introduction.

The introduction is where you introduce the topic of the work, so the reader knows what to expect to learn about in it. You want to hint at the ideas that will be going into it without giving too much away. It is a delicate balance between too much and too little information. You want to give them a taste of what is to come and entice them to read further so it should be interesting. It is the first thing they read, so it has to make them want to read further as well.

The body is where all of the real information goes. Usually, it is a five paragraph structure, so you have three body paragraphs. Each body paragraph should center around one single idea and relate well to the topic to prove something or show something about it. The topic is your main point, but these are your backup points to prove your main point. All of the information you gathered from each point goes in its respective body paragraph to give substance to your backup points

First you want to go back to your main point and explain once more why it is true or why it is what it is. This is your last chance to get them to understand you and your points. This should also be an ending. As the conclusion, it should conclude the paper nicely by wrapping up any loose ends and reiterating anything that needs to be said again.

As long as you know where everything goes and in what order, you should do just fine. Again, you do need the right information to make any of this work, and that takes some hard work as well but more or less they are pretty straight forward. Now that you know all of the steps you are ready to write one of your own!

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Introductions and Conclusions

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Introductions and conclusions play a special role in the academic essay, and they frequently demand much of your attention as a writer. A good introduction should identify your topic, provide essential context, and indicate your particular focus in the essay. It also needs to engage your readers’ interest. A strong conclusion will provide a sense of closure to the essay while again placing your concepts in a somewhat wider context. It will also, in some instances, add a stimulus to further thought. Since no two essays are the same, no single formula will automatically generate an introduction and conclusion for you. But the following guidelines will help you to construct a suitable beginning and end for your essay.

Some general advice about introductions

• Some students cannot begin writing the body of the essay until they feel they have the perfect introduction. Be aware of the dangers of sinking too much time into the introduction. Some of that time can be more usefully channeled into planning and writing.
• You may be the kind of writer who writes an introduction first in order to explore your own thinking on the topic. If so, remember that you may at a later stage need to compress your introduction.
• It can be fine to leave the writing of the introduction for a later stage in the essay-writing process. Some people write their introduction only after they have completed the rest of the essay. Others write the introduction first but rewrite it significantly in light of what they end up saying in the body of their paper.
• The introductions for most papers can be effectively written in one paragraph occupying half to three-quarters of the first page. Your introduction may be longer than that, and it may take more than one paragraph, but be sure you know why. The size of your introduction should bear some relationship to the length and complexity of your paper. A twenty page paper may call for a two-page introduction, but a five-page paper will not.
• Get to the point as soon as possible. Generally, you want to raise your topic in your very first sentences. A common error is to begin too broadly or too far off topic. Avoid sweeping generalizations.
• If your essay has a thesis, your thesis statement will typically appear at the end of your introduction, even though that is not a hard-and-fast rule. You may, for example, follow your thesis with a brief road map to your essay that sketches the basic structure of your argument. The longer the paper, the more useful a road map becomes.

How do I write an interesting, effective introduction?

Consider these strategies for capturing your readers’ attention and for fleshing out your introduction:

• Find a startling statistic that illustrates the seriousness of the problem you will address.
• Quote an expert (but be sure to introduce him or her first).
• Mention a common misperception that your thesis will argue against .
• Give some background information necessary for understanding the essay.
• Use a brief narrative or anecdote that exemplifies your reason for choosing the topic. In an assignment that encourages personal reflection, you may draw on your own experiences; in a research essay, the narrative may illustrate a common real-world scenario.
• In a science paper, explain key scientific concepts and refer to relevant literature. Lead up to your own contribution or intervention.
• In a more technical paper, define a term that is possibly unfamiliar to your audience but is central to understanding the essay.

In fleshing out your introduction, you will want to avoid some common pitfalls:

• Don’t provide dictionary definitions, especially of words your audience already knows.
• Don’t repeat the assignment specifications using the professor’s wording.
• Don’t give details and in-depth explanations that really belong in your body paragraphs. You can usually postpone background material to the body of the essay.

Some general advice about conclusions

• A conclusion is not merely a summary of your points or a re-statement of your thesis. If you wish to summarize—and often you must—do so in fresh language. Remind the reader of how the evidence you’ve presented has contributed to your thesis.
• The conclusion, like much of the rest of the paper, involves critical thinking. Reflect upon the significance of what you’ve written. Try to convey some closing thoughts about the larger implications of your argument.
• Broaden your focus a bit at the end of the essay. A good last sentence leaves your reader with something to think about, a concept in some way illuminated by what you’ve written in the paper.
• For most essays, one well-developed paragraph is sufficient for a conclusion. In some cases, a two-or-three paragraph conclusion may be appropriate. As with introductions, the length of the conclusion should reflect the length of the essay.

How do I write an interesting, effective conclusion?

The following strategies may help you move beyond merely summarizing the key points of your essay:

• If your essay deals with a contemporary problem, warn readers of the possible consequences of not attending to the problem.
• Recommend a specific course of action.
• Use an apt quotation or expert opinion to lend authority to the conclusion you have reached.
• Give a startling statistic, fact, or visual image to drive home the ultimate point of your paper.
• If your discipline encourages personal reflection, illustrate your concluding point with a relevant narrative drawn from your own life experiences.
• Return to an anecdote, example, or quotation that you introduced in your introduction, but add further insight that derives from the body of your essay.
• In a science or social science paper, mention worthwhile avenues for future research on your topic.

How does genre affect my introduction or conclusion?

Most of the advice in this handout pertains to argumentative or exploratory academic essays. Be aware, however, that different genres have their own special expectations about beginnings and endings. Some academic genres may not even require an introduction or conclusion. An annotated bibliography, for example, typically provides neither. A book review may begin with a summary of the book and conclude with an overall assessment of it. A policy briefing usually includes an introduction but may conclude with a series of recommendations. Check your assignment carefully for any directions about what to include in your introduction or conclusion.

Academic Writing

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• Basic Essay Structure

Writing an Essay

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Basic academic essays have three main parts:

• introduction

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Writing an Introduction

• Section  One  is a neutral sentence that will engage the reader’s interest in your essay.
• Section  Two  picks up the topic you are writing about by identifying the issues that you are going to explore.
• Section  Three  is an indication of how the question will be answered. Give a brief outline of how you will deal with each issue, and in which order.

An introduction   generally does three things. The first section is usually a  general comment  that shows the reader why the topic is important, gets their interest, and leads them into the topic. It isn’t actually part of your argument. The next section of the introduction is the  thesis statement . This is your response to the question; your final answer. It is probably the most important part of the introduction. Finally, the last section of an introduction tells the reader what they can expect in the essay body. This is where you  briefly outline your arguments .

Here is an example of the introduction to the question - Discuss how media can influence children. Use specific examples to support your view.

Writing Body Paragraphs

• The topic sentence  introduces  the topic of your paragraph.
• The sentences that follow the topic sentence will  develop and support the central idea  of your topic.
• The concluding sentence of your paragraph  restates the idea  expressed in the topic sentence.

The essay body itself is organized into paragraphs, according to your plan. Remember that each paragraph focuses on one idea, or aspect of your topic, and should contain at least 4-5 sentences so you can deal with that idea properly.

Each body paragraph has three sections. First is the  topic sentence . This lets the reader know what the paragraph is going to be about and  the main point it will make. It gives the paragraph’s point straight away. Next, come the  supporting sentences , which expand on the central idea, explaining it in more detail, exploring what it means, and of course giving the evidence and argument that back it up. This is where you use your research to support your argument. Then there is a  concluding sentence . This restates the idea in the topic sentence, to remind the reader of your main point. It also shows how that point helps answer the question.

Writing a Conclusion

• Re-read your introduction – this information will need to be restated in your conclusion emphasizing what you have proven and how you have proven it.
• Begin by  summarizing  your main arguments and restating your thesis ; e.g. "This essay has considered….."
• State your general conclusions,  explaining  why these are important.
• The final sentences should  draw together  the evidence you have presented in the body of the essay to restate your conclusion in an interesting way (use a transitional word to get you started e.g. Overall, Therefore).

The last section of an academic essay is the conclusion. The conclusion should reaffirm your answer to the question, and briefly summarize key arguments. It does not include any new points or new information.

A conclusion has three sections. First,  repeat the thesis statement . It won’t use the exact same words as in your introduction, but it will repeat the point: your overall answer to the question based on your arguments. Then set out your  general conclusions , and a short explanation of why they are important.  Finally,  draw together the question, the evidence in the essay body, and the conclusion. This way the reader knows that you have understood and answered the question. This part needs to be clear and concise.

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How To Write An Essay – Introduction, Body & Conclusion

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An essay is a structured piece of writing that presents an argument, tells a story, or explores a topic in depth. In academic writing , the term academic essay is frequently used. This denotes a carefully crafted piece of writing that adheres to certain standards and conventions, aiming to contribute to existing discourse or to provide a fresh perspective. With this article, we will help you understand the basics of how to write an essay, so you can receive good grades on your next work.

Inhaltsverzeichnis

• 1 How to write an essay in a nutshell
• 2 Definition: How to write an essay
• 3 Different types of how to write an essay
• 4 Step-by-step guide on how to write an essay
• 5 Structuring the paragraphs
• 6 Essay examples
• 7 Dos and don‘ts of how to write an essay

How to write an essay in a nutshell

Before you start on how to write an essay, you should read the essay question or topic carefully. Know what’s being asked of you. In the next step, you gather information and ideas about the topic. Use books, articles, or other reputable sources. Afterward, outline your main points and decide on a thesis (your main argument or stance) and supporting arguments.

An essay is typically made up of three parts :

Introduction

After you finish writing your essay, review your writing by paying attention to errors, clarity, and flow. Make sure your arguments are logical and well-presented. Check format, and citations (if any), and ensure it adheres to any guidelines given.

Definition: How to write an essay

How to write an essay refers to the systematic process of creating a structured written piece that presents and supports a specific idea or argument. This process typically involves selecting a topic, conducting research, planning and organizing one’s thoughts, drafting the content, and revising for clarity and coherence. The final product, an essay, is often a combination of an introduction that presents the main idea (thesis), body paragraphs that provide evidence or examples supporting the thesis, and a conclusion that summarizes and reinforces the main points.

Different types of how to write an essay

If you are eager to learn how to write an essay, keep these five types in mind:

• Narrative essay
• Descriptive essay
• Persuasive essay
• Compare-and-contrast essay
• Expository essay

Note: It is important to know what type of academic essay you have to write for your assignment. The type helps you to decide on a topic to write about as well as how to structure your essay outline.

Essay at university and high school

When you are given a typical five-paragraph expository essay , you would simply spend most of your time writing in high school. However, if you are at university, a college-level argumentative essay is bound to be a more complex piece of writing. It demands extensive independent research from varied sources, has stricter guidelines, and often requires deeper critical thinking compared to the more straightforward or surface-level student papers in high school. Depending on where you are in your academic journey, there is a vast difference when it comes to how to write an essay.

Step-by-step guide on how to write an essay

The process of how to write an essay can be broadly distilled into three main points or stages: Pre-writing and planning, drafting, and revising and editing.

For the planning, you should:

• Understand the essay question or prompt
• Conduct preliminary research to gather relevant sources
• Work on your essay outline

During the drafting, you:

• Craft a compelling introduction
• Develop the body of the essay
• Construct a conclusion

In the last step, you revise and edit your text. For this, you:

• Review for coherence, consistency, and logical flow
• Proofread for grammar, spelling, and punctuation errors
• Ensure the essay follows the required format or style guide (e.g., APA Style , MLA )
• Seek feedback from peers, tutors, or mentors and make necessary adjustments

Below you find the steps on how to write an entire essay.

Finishing touches

How to write an essay introduction is not difficult if you know what you should do. You have to lead into the topic and essay question, attract the reader’s attention, and give them a good idea of the focus of the essay. Use attention grabbers, also called hooks , like startling information, an anecdote, a dialogue, a strong statement, or a summary of the topic in general. Add a few more sentences to link the hook to your thesis statement, also called the topic sentence, that marks the end of the essay introduction .

From a child’s first taste of honey to the blooms in our gardens, honeybees touch our lives in unseen, myriad ways. These tiny workers, buzzing from flower to flower, play a crucial role in pollination, ensuring the reproduction of many of our favorite plants. However, the mysterious decline in honeybee populations poses a significant threat to our ecosystem. This essay will explore the significance of honeybees in our ecosystem, delve into the potential reasons behind their alarming decline, and propose solutions to address this growing crisis.

• Thesis statement
• Structure overview

Each of the main ideas in your outline will become one paragraph. Each of those paragraphs follows the same basic structure. First, you have to write down your main ideas. Then you add your supporting points as well as an elaboration (description, explanation, etc.) for each point. Lastly, round it up with a closing sentence. Make sure to use connections between sentences with the help of transition words , so the change in topic does not come abruptly.

Honeybees are not merely producers of honey; they are pivotal players in the world’s food chain. According to a report by the United Nations Food and Agriculture Organization (FAO), over 75% of the world’s food crops rely to some extent on animal pollination, with honeybees being among the most effective pollinators. This means that fruits such as apples, nuts like almonds, and even the coffee beans that make our morning brew, owe their existence in large part to the tireless work of these bees. The evidence underscores the gravity of the situation: a world with a declining bee population is one that risks significant disruption in its food supply chain. Such a decline doesn’t only spell trouble for the plants directly dependent on bees, but also for the animals and humans that consume those plants, creating a cascading effect on the larger ecosystem.

• Topic sentence

You have to summarize your main points as well as give a final perspective on the topic. Help your reader to draw a logical conclusion from what they just read. Repack your thesis statement in your conclusion so that the reader can remember the individual steps taken to come to this conclusion. Moreover, you should answer questions like: What are the implications of your topic sentence being true? What comes next? What questions remained unanswered?

The waning number of honeybees in our environment is not just a matter of ecological concern, but a looming crisis that touches every facet of our lives. As we’ve explored, these industrious insects are instrumental in the pollination of a vast majority of our food crops, a process vital to our global food supply chain. The evidence from reputable sources, such as the United Nations Food and Agriculture Organization, affirms the profound role honeybees play in sustaining our diets and of countless species. But beyond the tangible effects on food, the decline of honeybees serves as a potent reminder of the intricate, interconnected web of life and our role within it. If such a small creature can have such a vast impact on our world, it behooves us to take their decline as a clarion call. The broader implication is clear: preserving and nurturing our environment is not just an ethical duty; it’s a matter of survival, urging us to act with purpose.

• Return to the thesis
• Review of the key points
• Stating the broader implications

Come up with an intriguing title that arouses the reader’s interest. Furthermore, take your time to do the formatting of your paper. You also might want to put the paragraphs in a different order. Check the instructions again because you might have to include other information (name, date, etc.). Handing in a well-formatted academic essay makes a good impression on your instructor.

When it comes to how to write an essay, revision is the key to success. You have to analyze your writing to figure out if it makes logical sense and if there is a natural flow that makes it easy to read. Is every main idea supported by enough evidence, did you make clear how ideas are linked? Run a spelling and grammar checker to be on the safe side. Moreover, ask a friend to read your academic essay to give you feedback. Occasionally, you cannot see the mistakes when it comes to your writing. Having another opinion on your paper helps you with your revisions.

Structuring the paragraphs

Each paragraph should have an introductory, topic-based sentence as well as a concluding sentence that draws a link to the topic and critically summarizes your argument.

Follow with sentences that provide evidence or examples to back up the topic sentence. This can include data, quotations, anecdotes, or explanations. Delve deeper into the significance of the supporting details in relation to your main argument. Explain how the evidence supports the topic sentence and contributes to the overall thesis of the essay.

Furthermore, you should pay attention to coherence, consistency, flow, variety, and relevance.

• Stay consistent in tense, perspective, and style.
• Use transition sentences , a link between sentences, to guide the reader.
• Vary sentence structure and length to keep the reader engaged.
• Every paragraph should relate back to and support the essay’s overall thesis or argument.
• Avoid digressions or unnecessary details.

Essay examples

In the following, you will find samples of how to write an essay. Here, you can read several essay types , whether to help you get started or if you’re simply unsure how to distinguish them.

Dos and don‘ts of how to write an essay

Below, you will find a list of the dos and don’ts of how to write an essay.

• Signposting language
• Stay focused
• Write the body first
• Revise your writing
• Plain and clear writing style
• Procrastination
• Generalizations
• Use of personal pronouns
• Writing without an outline
• Contractions

How do you structure an essay?

The typical essay structure is easier to understand than the structure of a dissertation or thesis. There are many types of essays, but the structure remains mostly unchanged. You start with the introduction, then the body paragraphs, and finally, the conclusion.

How do you start writing an essay?

To start your essay, you first need an appropriate research paper topic . Ensure that your topic fits within the guidelines set by your institution, and it’s not too broad or narrow. Then, formulate your thesis statement and begin outlining a plan for your academic essay. Once you’re finished, you can start on how to write an essay.

What is a good essay introduction?

A good essay introduction will begin with an opening statement that grabs the reader’s attention and draws them in. Then, you give a bit of background information and lay out the structure for the reader. The thesis statement should be placed towards the end of the introduction, as it provides one to two sentences of a summary of your essay and the main idea.

What are the five steps of writing an essay?

The five steps on how to write an essay are the following.

• Planning: Understand the prompt and organize your ideas.
• Research: Gather relevant information and evidence.
• Drafting: Write the initial version of the essay.
• Revising: Refine content for clarity and coherence.
• Editing: Proofread for grammar, punctuation, and formatting errors.

What makes an essay good?

A good essay is clear, coherent, well-organized, presents strong arguments supported by relevant evidence, and is written with a consistent style and proper grammar. Furthermore, it starts with a bold statement and ends with an impactful conclusion.

I’m so happy with how my dissertation turned out! The order process was very...

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How to Write an Essay Introduction (with Examples)   

The introduction of an essay plays a critical role in engaging the reader and providing contextual information about the topic. It sets the stage for the rest of the essay, establishes the tone and style, and motivates the reader to continue reading. 

Table of Contents

What is an essay introduction , what to include in an essay introduction, how to create an essay structure , step-by-step process for writing an essay introduction , how to write an introduction paragraph , how to write a hook for your essay , how to include background information , how to write a thesis statement .

• Argumentative Essay Introduction Example: 
• Expository Essay Introduction Example 

Literary Analysis Essay Introduction Example

Check and revise – checklist for essay introduction , key takeaways , frequently asked questions .

An introduction is the opening section of an essay, paper, or other written work. It introduces the topic and provides background information, context, and an overview of what the reader can expect from the rest of the work. 1 The key is to be concise and to the point, providing enough information to engage the reader without delving into excessive detail. 

The essay introduction is crucial as it sets the tone for the entire piece and provides the reader with a roadmap of what to expect. Here are key elements to include in your essay introduction: 

• Hook : Start with an attention-grabbing statement or question to engage the reader. This could be a surprising fact, a relevant quote, or a compelling anecdote. 
• Background information : Provide context and background information to help the reader understand the topic. This can include historical information, definitions of key terms, or an overview of the current state of affairs related to your topic. 
• Thesis statement : Clearly state your main argument or position on the topic. Your thesis should be concise and specific, providing a clear direction for your essay. 

Before we get into how to write an essay introduction, we need to know how it is structured. The structure of an essay is crucial for organizing your thoughts and presenting them clearly and logically. It is divided as follows: 2  

• Introduction:  The introduction should grab the reader’s attention with a hook, provide context, and include a thesis statement that presents the main argument or purpose of the essay.  
• Body:  The body should consist of focused paragraphs that support your thesis statement using evidence and analysis. Each paragraph should concentrate on a single central idea or argument and provide evidence, examples, or analysis to back it up.  
• Conclusion:  The conclusion should summarize the main points and restate the thesis differently. End with a final statement that leaves a lasting impression on the reader. Avoid new information or arguments. 

Here’s a step-by-step guide on how to write an essay introduction: 

• Start with a Hook : Begin your introduction paragraph with an attention-grabbing statement, question, quote, or anecdote related to your topic. The hook should pique the reader’s interest and encourage them to continue reading. 
• Provide Background Information : This helps the reader understand the relevance and importance of the topic. 
• State Your Thesis Statement : The last sentence is the main argument or point of your essay. It should be clear, concise, and directly address the topic of your essay. 
• Preview the Main Points : This gives the reader an idea of what to expect and how you will support your thesis. 
• Keep it Concise and Clear : Avoid going into too much detail or including information not directly relevant to your topic. 
• Revise : Revise your introduction after you’ve written the rest of your essay to ensure it aligns with your final argument. 

Here’s an example of an essay introduction paragraph about the importance of education: 

Education is often viewed as a fundamental human right and a key social and economic development driver. As Nelson Mandela once famously said, “Education is the most powerful weapon which you can use to change the world.” It is the key to unlocking a wide range of opportunities and benefits for individuals, societies, and nations. In today’s constantly evolving world, education has become even more critical. It has expanded beyond traditional classroom learning to include digital and remote learning, making education more accessible and convenient. This essay will delve into the importance of education in empowering individuals to achieve their dreams, improving societies by promoting social justice and equality, and driving economic growth by developing a skilled workforce and promoting innovation. 

This introduction paragraph example includes a hook (the quote by Nelson Mandela), provides some background information on education, and states the thesis statement (the importance of education). 

This is one of the key steps in how to write an essay introduction. Crafting a compelling hook is vital because it sets the tone for your entire essay and determines whether your readers will stay interested. A good hook draws the reader in and sets the stage for the rest of your essay.  

• Avoid Dry Fact : Instead of simply stating a bland fact, try to make it engaging and relevant to your topic. For example, if you’re writing about the benefits of exercise, you could start with a startling statistic like, “Did you know that regular exercise can increase your lifespan by up to seven years?” 
• Avoid Using a Dictionary Definition : While definitions can be informative, they’re not always the most captivating way to start an essay. Instead, try to use a quote, anecdote, or provocative question to pique the reader’s interest. For instance, if you’re writing about freedom, you could begin with a quote from a famous freedom fighter or philosopher. 
• Do Not Just State a Fact That the Reader Already Knows : This ties back to the first point—your hook should surprise or intrigue the reader. For Here’s an introduction paragraph example, if you’re writing about climate change, you could start with a thought-provoking statement like, “Despite overwhelming evidence, many people still refuse to believe in the reality of climate change.” 

Including background information in the introduction section of your essay is important to provide context and establish the relevance of your topic. When writing the background information, you can follow these steps: 

• Start with a General Statement:  Begin with a general statement about the topic and gradually narrow it down to your specific focus. For example, when discussing the impact of social media, you can begin by making a broad statement about social media and its widespread use in today’s society, as follows: “Social media has become an integral part of modern life, with billions of users worldwide.” 
• Define Key Terms : Define any key terms or concepts that may be unfamiliar to your readers but are essential for understanding your argument. 
• Provide Relevant Statistics:  Use statistics or facts to highlight the significance of the issue you’re discussing. For instance, “According to a report by Statista, the number of social media users is expected to reach 4.41 billion by 2025.” 
• Discuss the Evolution:  Mention previous research or studies that have been conducted on the topic, especially those that are relevant to your argument. Mention key milestones or developments that have shaped its current impact. You can also outline some of the major effects of social media. For example, you can briefly describe how social media has evolved, including positives such as increased connectivity and issues like cyberbullying and privacy concerns. 
• Transition to Your Thesis:  Use the background information to lead into your thesis statement, which should clearly state the main argument or purpose of your essay. For example, “Given its pervasive influence, it is crucial to examine the impact of social media on mental health.” 

A thesis statement is a concise summary of the main point or claim of an essay, research paper, or other type of academic writing. It appears near the end of the introduction. Here’s how to write a thesis statement: 

• Identify the topic:  Start by identifying the topic of your essay. For example, if your essay is about the importance of exercise for overall health, your topic is “exercise.” 
• State your position:  Next, state your position or claim about the topic. This is the main argument or point you want to make. For example, if you believe that regular exercise is crucial for maintaining good health, your position could be: “Regular exercise is essential for maintaining good health.” 
• Support your position:  Provide a brief overview of the reasons or evidence that support your position. These will be the main points of your essay. For example, if you’re writing an essay about the importance of exercise, you could mention the physical health benefits, mental health benefits, and the role of exercise in disease prevention. 
• Make it specific:  Ensure your thesis statement clearly states what you will discuss in your essay. For example, instead of saying, “Exercise is good for you,” you could say, “Regular exercise, including cardiovascular and strength training, can improve overall health and reduce the risk of chronic diseases.” 

Examples of essay introduction 

Here are examples of essay introductions for different types of essays: 

Argumentative Essay Introduction Example:  

Topic: Should the voting age be lowered to 16? 

“The question of whether the voting age should be lowered to 16 has sparked nationwide debate. While some argue that 16-year-olds lack the requisite maturity and knowledge to make informed decisions, others argue that doing so would imbue young people with agency and give them a voice in shaping their future.” 

Expository Essay Introduction Example  

Topic: The benefits of regular exercise 

“In today’s fast-paced world, the importance of regular exercise cannot be overstated. From improving physical health to boosting mental well-being, the benefits of exercise are numerous and far-reaching. This essay will examine the various advantages of regular exercise and provide tips on incorporating it into your daily routine.” 

Text: “To Kill a Mockingbird” by Harper Lee 

“Harper Lee’s novel, ‘To Kill a Mockingbird,’ is a timeless classic that explores themes of racism, injustice, and morality in the American South. Through the eyes of young Scout Finch, the reader is taken on a journey that challenges societal norms and forces characters to confront their prejudices. This essay will analyze the novel’s use of symbolism, character development, and narrative structure to uncover its deeper meaning and relevance to contemporary society.” 

• Engaging and Relevant First Sentence : The opening sentence captures the reader’s attention and relates directly to the topic. 
• Background Information : Enough background information is introduced to provide context for the thesis statement. 
• Definition of Important Terms : Key terms or concepts that might be unfamiliar to the audience or are central to the argument are defined. 
• Clear Thesis Statement : The thesis statement presents the main point or argument of the essay. 
• Relevance to Main Body : Everything in the introduction directly relates to and sets up the discussion in the main body of the essay. 

Writing a strong introduction is crucial for setting the tone and context of your essay. Here are the key takeaways for how to write essay introduction: 3  

• Hook the Reader : Start with an engaging hook to grab the reader’s attention. This could be a compelling question, a surprising fact, a relevant quote, or an anecdote. 
• Provide Background : Give a brief overview of the topic, setting the context and stage for the discussion. 
• Thesis Statement : State your thesis, which is the main argument or point of your essay. It should be concise, clear, and specific. 
• Preview the Structure : Outline the main points or arguments to help the reader understand the organization of your essay. 
• Keep it Concise : Avoid including unnecessary details or information not directly related to your thesis. 
• Revise and Edit : Revise your introduction to ensure clarity, coherence, and relevance. Check for grammar and spelling errors. 
• Seek Feedback : Get feedback from peers or instructors to improve your introduction further. 

The purpose of an essay introduction is to give an overview of the topic, context, and main ideas of the essay. It is meant to engage the reader, establish the tone for the rest of the essay, and introduce the thesis statement or central argument.  

An essay introduction typically ranges from 5-10% of the total word count. For example, in a 1,000-word essay, the introduction would be roughly 50-100 words. However, the length can vary depending on the complexity of the topic and the overall length of the essay.

An essay introduction is critical in engaging the reader and providing contextual information about the topic. To ensure its effectiveness, consider incorporating these key elements: a compelling hook, background information, a clear thesis statement, an outline of the essay’s scope, a smooth transition to the body, and optional signposting sentences.  

The process of writing an essay introduction is not necessarily straightforward, but there are several strategies that can be employed to achieve this end. When experiencing difficulty initiating the process, consider the following techniques: begin with an anecdote, a quotation, an image, a question, or a startling fact to pique the reader’s interest. It may also be helpful to consider the five W’s of journalism: who, what, when, where, why, and how.   For instance, an anecdotal opening could be structured as follows: “As I ascended the stage, momentarily blinded by the intense lights, I could sense the weight of a hundred eyes upon me, anticipating my next move. The topic of discussion was climate change, a subject I was passionate about, and it was my first public speaking event. Little did I know , that pivotal moment would not only alter my perspective but also chart my life’s course.” 

Crafting a compelling thesis statement for your introduction paragraph is crucial to grab your reader’s attention. To achieve this, avoid using overused phrases such as “In this paper, I will write about” or “I will focus on” as they lack originality. Instead, strive to engage your reader by substantiating your stance or proposition with a “so what” clause. While writing your thesis statement, aim to be precise, succinct, and clear in conveying your main argument.  

To create an effective essay introduction, ensure it is clear, engaging, relevant, and contains a concise thesis statement. It should transition smoothly into the essay and be long enough to cover necessary points but not become overwhelming. Seek feedback from peers or instructors to assess its effectiveness. 

References  

• Cui, L. (2022). Unit 6 Essay Introduction.  Building Academic Writing Skills . 
• West, H., Malcolm, G., Keywood, S., & Hill, J. (2019). Writing a successful essay.  Journal of Geography in Higher Education ,  43 (4), 609-617. 
• Beavers, M. E., Thoune, D. L., & McBeth, M. (2023). Bibliographic Essay: Reading, Researching, Teaching, and Writing with Hooks: A Queer Literacy Sponsorship. College English, 85(3), 230-242. 

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How to Write a Body of a Research Paper

The main part of your research paper is called “the body.” To write this important part of your paper, include only relevant information, or information that gets to the point. Organize your ideas in a logical order—one that makes sense—and provide enough details—facts and examples—to support the points you want to make.

Logical Order

Transition words and phrases, adding evidence, phrases for supporting topic sentences.

• Transition Phrases for Comparisons
• Transition Phrases for Contrast
• Transition Phrases to Show a Process
• Phrases to Introduce Examples
• Transition Phrases for Presenting Evidence

How to Make Effective Transitions

Examples of effective transitions, drafting your conclusion, writing the body paragraphs.

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• The third and fourth paragraphs follow the same format as the second:
• Transition or topic sentence.
• Topic sentence (if not included in the first sentence).
• Supporting sentences including a discussion, quotations, or examples that support the topic sentence.
• Concluding sentence that transitions to the next paragraph.

The topic of each paragraph will be supported by the evidence you itemized in your outline. However, just as smooth transitions are required to connect your paragraphs, the sentences you write to present your evidence should possess transition words that connect ideas, focus attention on relevant information, and continue your discussion in a smooth and fluid manner.

You presented the main idea of your paper in the thesis statement. In the body, every single paragraph must support that main idea. If any paragraph in your paper does not, in some way, back up the main idea expressed in your thesis statement, it is not relevant, which means it doesn’t have a purpose and shouldn’t be there.

Each paragraph also has a main idea of its own. That main idea is stated in a topic sentence, either at the beginning or somewhere else in the paragraph. Just as every paragraph in your paper supports your thesis statement, every sentence in each paragraph supports the main idea of that paragraph by providing facts or examples that back up that main idea. If a sentence does not support the main idea of the paragraph, it is not relevant and should be left out.

A paper that makes claims or states ideas without backing them up with facts or clarifying them with examples won’t mean much to readers. Make sure you provide enough supporting details for all your ideas. And remember that a paragraph can’t contain just one sentence. A paragraph needs at least two or more sentences to be complete. If a paragraph has only one or two sentences, you probably haven’t provided enough support for your main idea. Or, if you have trouble finding the main idea, maybe you don’t have one. In that case, you can make the sentences part of another paragraph or leave them out.

Arrange the paragraphs in the body of your paper in an order that makes sense, so that each main idea follows logically from the previous one. Likewise, arrange the sentences in each paragraph in a logical order.

If you carefully organized your notes and made your outline, your ideas will fall into place naturally as you write your draft. The main ideas, which are building blocks of each section or each paragraph in your paper, come from the Roman-numeral headings in your outline. The supporting details under each of those main ideas come from the capital-letter headings. In a shorter paper, the capital-letter headings may become sentences that include supporting details, which come from the Arabic numerals in your outline. In a longer paper, the capital letter headings may become paragraphs of their own, which contain sentences with the supporting details, which come from the Arabic numerals in your outline.

In addition to keeping your ideas in logical order, transitions are another way to guide readers from one idea to another. Transition words and phrases are important when you are suggesting or pointing out similarities between ideas, themes, opinions, or a set of facts. As with any perfect phrase, transition words within paragraphs should not be used gratuitously. Their meaning must conform to what you are trying to point out, as shown in the examples below:

• “Accordingly” or “in accordance with” indicates agreement. For example :Thomas Edison’s experiments with electricity accordingly followed the theories of Benjamin Franklin, J. B. Priestly, and other pioneers of the previous century.
• “Analogous” or “analogously” contrasts different things or ideas that perform similar functions or make similar expressions. For example: A computer hard drive is analogous to a filing cabinet. Each stores important documents and data.
• “By comparison” or “comparatively”points out differences between things that otherwise are similar. For example: Roses require an alkaline soil. Azaleas, by comparison, prefer an acidic soil.
• “Corresponds to” or “correspondingly” indicates agreement or conformity. For example: The U.S. Constitution corresponds to England’s Magna Carta in so far as both established a framework for a parliamentary system.
• “Equals,”“equal to,” or “equally” indicates the same degree or quality. For example:Vitamin C is equally as important as minerals in a well-balanced diet.
• “Equivalent” or “equivalently” indicates two ideas or things of approximately the same importance, size, or volume. For example:The notions of individual liberty and the right to a fair and speedy trial hold equivalent importance in the American legal system.
• “Common” or “in common with” indicates similar traits or qualities. For example: Darwin did not argue that humans were descended from the apes. Instead, he maintained that they shared a common ancestor.
• “In the same way,”“in the same manner,”“in the same vein,” or “likewise,” connects comparable traits, ideas, patterns, or activities. For example: John Roebling’s suspension bridges in Brooklyn and Cincinnati were built in the same manner, with strong cables to support a metallic roadway. Example 2: Despite its delicate appearance, John Roebling’s Brooklyn Bridge was built as a suspension bridge supported by strong cables. Example 3: Cincinnati’s Suspension Bridge, which Roebling also designed, was likewise supported by cables.
• “Kindred” indicates that two ideas or things are related by quality or character. For example: Artists Vincent Van Gogh and Paul Gauguin are considered kindred spirits in the Impressionist Movement. “Like” or “as” are used to create a simile that builds reader understanding by comparing two dissimilar things. (Never use “like” as slang, as in: John Roebling was like a bridge designer.) For examples: Her eyes shone like the sun. Her eyes were as bright as the sun.
• “Parallel” describes events, things, or ideas that occurred at the same time or that follow similar logic or patterns of behavior. For example:The original Ocktoberfests were held to occur in parallel with the autumn harvest.
• “Obviously” emphasizes a point that should be clear from the discussion. For example: Obviously, raccoons and other wildlife will attempt to find food and shelter in suburban areas as their woodland habitats disappear.
• “Similar” and “similarly” are used to make like comparisons. For example: Horses and ponies have similar physical characteristics although, as working farm animals, each was bred to perform different functions.
• “There is little debate” or “there is consensus” can be used to point out agreement. For example:There is little debate that the polar ice caps are melting.The question is whether global warming results from natural or human-made causes.

Other phrases that can be used to make transitions or connect ideas within paragraphs include:

• Use “alternately” or “alternatively” to suggest a different option.
• Use “antithesis” to indicate a direct opposite.
• Use “contradict” to indicate disagreement.
• Use “on the contrary” or “conversely” to indicate that something is different from what it seems.
• Use “dissimilar” to point out differences between two things.
• Use “diverse” to discuss differences among many things or people.
• Use “distinct” or “distinctly” to point out unique qualities.
• Use “inversely” to indicate an opposite idea.
• Use “it is debatable,” “there is debate,” or “there is disagreement” to suggest that there is more than one opinion about a subject.
• Use “rather” or “rather than” to point out an exception.
• Use “unique” or “uniquely” to indicate qualities that can be found nowhere else.
• Use “unlike” to indicate dissimilarities.
• Use “various” to indicate more than one kind.

Writing Topic Sentences

Remember, a sentence should express a complete thought, one thought per sentence—no more, no less. The longer and more convoluted your sentences become, the more likely you are to muddle the meaning, become repetitive, and bog yourself down in issues of grammar and construction. In your first draft, it is generally a good idea to keep those sentences relatively short and to the point. That way your ideas will be clearly stated.You will be able to clearly see the content that you have put down—what is there and what is missing—and add or subtract material as it is needed. The sentences will probably seem choppy and even simplistic.The purpose of a first draft is to ensure that you have recorded all the content you will need to make a convincing argument. You will work on smoothing and perfecting the language in subsequent drafts.

Transitioning from your topic sentence to the evidence that supports it can be problematic. It requires a transition, much like the transitions needed to move from one paragraph to the next. Choose phrases that connect the evidence directly to your topic sentence.

• Consider this: (give an example or state evidence).
• If (identify one condition or event) then (identify the condition or event that will follow).
• It should go without saying that (point out an obvious condition).
• Note that (provide an example or observation).
• Take a look at (identify a condition; follow with an explanation of why you think it is important to the discussion).
• The authors had (identify their idea) in mind when they wrote “(use a quotation from their text that illustrates the idea).”
• The point is that (summarize the conclusion your reader should draw from your research).
• This becomes evident when (name the author) says that (paraphrase a quote from the author’s writing).
• We see this in the following example: (provide an example of your own).
• (The author’s name) offers the example of (summarize an example given by the author).

If an idea is controversial, you may need to add extra evidence to your paragraphs to persuade your reader. You may also find that a logical argument, one based solely on your evidence, is not persuasive enough and that you need to appeal to the reader’s emotions. Look for ways to incorporate your research without detracting from your argument.

Writing Transition Sentences

It is often difficult to write transitions that carry a reader clearly and logically on to the next paragraph (and the next topic) in an essay. Because you are moving from one topic to another, it is easy to simply stop one and start another. Great research papers, however, include good transitions that link the ideas in an interesting discussion so that readers can move smoothly and easily through your presentation. Close each of your paragraphs with an interesting transition sentence that introduces the topic coming up in the next paragraph.

Transition sentences should show a relationship between the two topics.Your transition will perform one of the following functions to introduce the new idea:

• Indicate that you will be expanding on information in a different way in the upcoming paragraph.
• Indicate that a comparison, contrast, or a cause-and-effect relationship between the topics will be discussed.
• Indicate that an example will be presented in the next paragraph.
• Indicate that a conclusion is coming up.

Transitions make a paper flow smoothly by showing readers how ideas and facts follow one another to point logically to a conclusion. They show relationships among the ideas, help the reader to understand, and, in a persuasive paper, lead the reader to the writer’s conclusion.

Each paragraph should end with a transition sentence to conclude the discussion of the topic in the paragraph and gently introduce the reader to the topic that will be raised in the next paragraph. However, transitions also occur within paragraphs—from sentence to sentence—to add evidence, provide examples, or introduce a quotation.

The type of paper you are writing and the kinds of topics you are introducing will determine what type of transitional phrase you should use. Some useful phrases for transitions appear below. They are grouped according to the function they normally play in a paper. Transitions, however, are not simply phrases that are dropped into sentences. They are constructed to highlight meaning. Choose transitions that are appropriate to your topic and what you want the reader to do. Edit them to be sure they fit properly within the sentence to enhance the reader’s understanding.

Transition Phrases for Comparisons:

• We also see
• In addition to
• Notice that
• Beside that,
• In comparison,
• Once again,
• Identically,
• For example,
• Comparatively, it can be seen that
• We see this when
• This corresponds to
• In other words,
• At the same time,
• By the same token,

Transition Phrases for Contrast:

• By contrast,
• On the contrary,
• Nevertheless,
• An exception to this would be …
• Alongside that,we find …
• On one hand … on the other hand …
• [New information] presents an opposite view …
• Conversely, it could be argued …
• Other than that,we find that …
• We get an entirely different impression from …
• One point of differentiation is …
• Further investigation shows …
• An exception can be found in the fact that …

Transition Phrases to Show a Process:

• At the top we have … Near the bottom we have …
• Here we have … There we have …
• Continuing on,
• We progress to …
• Close up … In the distance …
• With this in mind,
• Moving in sequence,
• Proceeding sequentially,
• Moving to the next step,
• First, Second,Third,…
• Examining the activities in sequence,
• Sequentially,
• As a result,
• The end result is …
• To illustrate …
• Subsequently,
• One consequence of …
• If … then …
• It follows that …
• This is chiefly due to …
• The next step …
• Later we find …

Phrases to Introduce Examples:

• For instance,
• Particularly,
• In particular,
• This includes,
• Specifically,
• To illustrate,
• One illustration is
• One example is
• This is illustrated by
• This can be seen when
• This is especially seen in
• This is chiefly seen when

Transition Phrases for Presenting Evidence:

• Another point worthy of consideration is
• At the center of the issue is the notion that
• Before moving on, it should be pointed out that
• Another important point is
• Another idea worth considering is
• Consequently,
• Especially,
• Even more important,
• Getting beyond the obvious,
• In spite of all this,
• It follows that
• It is clear that
• More importantly,
• Most importantly,

How to make effective transitions between sections of a research paper? There are two distinct issues in making strong transitions:

• Does the upcoming section actually belong where you have placed it?
• Have you adequately signaled the reader why you are taking this next step?

The first is the most important: Does the upcoming section actually belong in the next spot? The sections in your research paper need to add up to your big point (or thesis statement) in a sensible progression. One way of putting that is, “Does the architecture of your paper correspond to the argument you are making?” Getting this architecture right is the goal of “large-scale editing,” which focuses on the order of the sections, their relationship to each other, and ultimately their correspondence to your thesis argument.

It’s easy to craft graceful transitions when the sections are laid out in the right order. When they’re not, the transitions are bound to be rough. This difficulty, if you encounter it, is actually a valuable warning. It tells you that something is wrong and you need to change it. If the transitions are awkward and difficult to write, warning bells should ring. Something is wrong with the research paper’s overall structure.

After you’ve placed the sections in the right order, you still need to tell the reader when he is changing sections and briefly explain why. That’s an important part of line-by-line editing, which focuses on writing effective sentences and paragraphs.

Effective transition sentences and paragraphs often glance forward or backward, signaling that you are switching sections. Take this example from J. M. Roberts’s History of Europe . He is finishing a discussion of the Punic Wars between Rome and its great rival, Carthage. The last of these wars, he says, broke out in 149 B.C. and “ended with so complete a defeat for the Carthaginians that their city was destroyed . . . .” Now he turns to a new section on “Empire.” Here is the first sentence: “By then a Roman empire was in being in fact if not in name.”(J. M. Roberts, A History of Europe . London: Allen Lane, 1997, p. 48) Roberts signals the transition with just two words: “By then.” He is referring to the date (149 B.C.) given near the end of the previous section. Simple and smooth.

Michael Mandelbaum also accomplishes this transition between sections effortlessly, without bringing his narrative to a halt. In The Ideas That Conquered the World: Peace, Democracy, and Free Markets , one chapter shows how countries of the North Atlantic region invented the idea of peace and made it a reality among themselves. Here is his transition from one section of that chapter discussing “the idea of warlessness” to another section dealing with the history of that idea in Europe.

The widespread aversion to war within the countries of the Western core formed the foundation for common security, which in turn expressed the spirit of warlessness. To be sure, the rise of common security in Europe did not abolish war in other parts of the world and could not guarantee its permanent abolition even on the European continent. Neither, however, was it a flukish, transient product . . . . The European common security order did have historical precedents, and its principal features began to appear in other parts of the world. Precedents for Common Security The security arrangements in Europe at the dawn of the twenty-first century incorporated features of three different periods of the modern age: the nineteenth century, the interwar period, and the ColdWar. (Michael Mandelbaum, The Ideas That Conquered the World: Peace, Democracy, and Free Markets . New York: Public Affairs, 2002, p. 128)

It’s easier to make smooth transitions when neighboring sections deal with closely related subjects, as Mandelbaum’s do. Sometimes, however, you need to end one section with greater finality so you can switch to a different topic. The best way to do that is with a few summary comments at the end of the section. Your readers will understand you are drawing this topic to a close, and they won’t be blindsided by your shift to a new topic in the next section.

Here’s an example from economic historian Joel Mokyr’s book The Lever of Riches: Technological Creativity and Economic Progress . Mokyr is completing a section on social values in early industrial societies. The next section deals with a quite different aspect of technological progress: the role of property rights and institutions. So Mokyr needs to take the reader across a more abrupt change than Mandelbaum did. Mokyr does that in two ways. First, he summarizes his findings on social values, letting the reader know the section is ending. Then he says the impact of values is complicated, a point he illustrates in the final sentences, while the impact of property rights and institutions seems to be more straightforward. So he begins the new section with a nod to the old one, noting the contrast.

In commerce, war and politics, what was functional was often preferred [within Europe] to what was aesthetic or moral, and when it was not, natural selection saw to it that such pragmatism was never entirely absent in any society. . . . The contempt in which physical labor, commerce, and other economic activity were held did not disappear rapidly; much of European social history can be interpreted as a struggle between wealth and other values for a higher step in the hierarchy. The French concepts of bourgeois gentilhomme and nouveau riche still convey some contempt for people who joined the upper classes through economic success. Even in the nineteenth century, the accumulation of wealth was viewed as an admission ticket to social respectability to be abandoned as soon as a secure membership in the upper classes had been achieved. Institutions and Property Rights The institutional background of technological progress seems, on the surface, more straightforward. (Joel Mokyr, The Lever of Riches: Technological Creativity and Economic Progress . New York: Oxford University Press, 1990, p. 176)

Note the phrase, “on the surface.” Mokyr is hinting at his next point, that surface appearances are deceiving in this case. Good transitions between sections of your research paper depend on:

• Getting the sections in the right order
• Moving smoothly from one section to the next
• Signaling readers that they are taking the next step in your argument
• Explaining why this next step comes where it does

Every good paper ends with a strong concluding paragraph. To write a good conclusion, sum up the main points in your paper. To write an even better conclusion, include a sentence or two that helps the reader answer the question, “So what?” or “Why does all this matter?” If you choose to include one or more “So What?” sentences, remember that you still need to support any point you make with facts or examples. Remember, too, that this is not the place to introduce new ideas from “out of the blue.” Make sure that everything you write in your conclusion refers to what you’ve already written in the body of your paper.

Back to How To Write A Research Paper .

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• Open access
• Published: 27 June 2024

Effects of Staphylococcus aureus on stem cells and potential targeted treatment of inflammatory disorders

• Zi-xian Liu 1 , 2 , 3   na1 ,
• Guan-qiao Liu 1 , 2   na1 ,
• Ze-xin Lin 1 , 2   na1 ,
• Ying-qi Chen 1 , 2 ,
• Peng Chen 1 , 2 ,
• Yan-jun Hu 1 , 2 ,
• Bin Yu   ORCID: orcid.org/0000-0002-3109-2062 1 , 2 &
• Nan Jiang   ORCID: orcid.org/0000-0003-2416-1653 1 , 2  

Stem Cell Research & Therapy volume  15 , Article number:  187 ( 2024 ) Cite this article

116 Accesses

Metrics details

Due to the advanced studies on stem cells in developmental biology, the roles of stem cells in the body and their phenotypes in related diseases have not been covered clearly. Meanwhile, with the intensive research on the mechanisms of stem cells in regulating various diseases, stem cell therapy is increasingly being attention because of its effectiveness and safety. As one of the most widely used stem cell in stem cell therapies, hematopoietic stem cell transplantation shows huge advantage in treatment of leukemia and other blood-malignant diseases. Besides, due to the effect of anti-inflammatory and immunomodulatory, mesenchymal stem cells could be a potential therapeutic strategy for variety infectious diseases. In this review, we summarized the effects of Staphylococcus aureus ( S. aureus ) and its components on different types of adult stem cells and their downstream signaling pathways. Also, we reviewed the roles of different kinds of stem cells in various disease models caused by S. aureus , providing new insights for applying stem cell therapy to treat infectious diseases.

Introduction

As one of the most frequently detected bacteria in human infectious diseases [ 1 ], S. aureus is closely related to the occurrence of lung infection, bacteremia, infective endocarditis, osteomyelitis, and many other inflammatory disorders [ 2 ]. The preferred treatment for S. aureus -related infection is β-lactam antibiotics [ 3 ]. However, antibiotic resistance has developed rapidly in recent years. It has been reported that the death per year caused by antibiotic resistance has exceeded 10 million and will exceed that caused by cancer by 2050 [ 4 ]. Based on this emergency issue, exploring new strategies against such antibiotic resistance is crucial. Recently, many studies have reported that stem cells have powerful immune regulatory functions, playing an essential role in treating various infectious diseases [ 5 ].

Stem cells are a group of cells with self-renewal and self-differentiation functions [ 6 ], which are classified into four categories according to their sources: embryonic stem cells (ESCs), fetal and adult stem cells, and induced pluripotent stem cells (iPSCs) [ 7 ]. Since adult stem cells do not cause rejection or ethical controversy, relevant studies on them have been applied in the models of various infectious diseases [ 8 ]. Although many studies have confirmed that stem cells play positive roles in infectious diseases, they can also be influenced by the infectious environment. Understanding the mechanisms of how S. aureus and its components affect the functions of stem cells may help us exert their anti-infectious function more effectively and play more excellent value in the management of infectious diseases.

Effects of S. Aureus on stem cells

Mesenchymal stem cells (mscs).

As a common pluripotent stem cell, MSCs have recently received extensive attention in regenerative medicine, among which bone marrow-derived mesenchymal stem cells (BMSCs) are the most widely investigated [ 9 ]. In addition to bone marrow, MSCs can be isolated from different mature tissues, such as skeletal muscle, adipocytes, umbilical cord, amniotic fluid, peripheral blood, intima synovium, dental pulp, lung, and liver [ 10 , 11 ]. Such MSCs can differentiate into bone, chondrocytes, fat, muscle, neurons, islet cells, and liver cells under specific conditions [ 10 , 11 , 12 ]. Recent studies have found that S. aureus can affect migration and recruitment of lineage differentiation and activity of MSCs (Fig.  1 ; Table  1 ).

The mechanism of how S. aureus interacts with stem cells

As an essential therapeutic tool in regenerative medicine, MSCs have been proven to enhance proliferation during tissue injury, inflammation, and tumorigenesis. Then, they differentiate into different types of cells under different microenvironment stimulations and participate in tissue repair [ 13 , 14 ]. Recent studies in vitro have indicated that the number of MSCs at the site of infection caused by S. aureus significantly increased, possibly due to the local inflammatory specificity that promotes the migration and implantation of MSCs [ 15 , 16 ]. Yang et al. [ 17 ] indicated that the intestinal epithelial cells alone did not affect the migration of cord blood-derived MSCs. In contrast, the intestinal epithelial cells infected with S. aureus significantly enhanced the above biological process. It suggests that S. aureus may regulate the migration of cord blood-derived MSCs by influencing the secretion of migration-related chemokines. Further analysis revealed that S. aureus enhanced the expressions of tumor necrosis factor α (TNF-α) and C-C motif chemokine ligand 20 (CCL20) by activating the NF-κB signaling pathway in the intestinal epithelial cells. However, Ward et al. [ 18 ] found that the conditioned medium to produce S. aureus biofilm resulted in an increased caspase 3/7 activity in primary human bone marrow stromal cells (hBMSCs), leading to apoptosis and loss of viability. They also observed that the migration ability of cells exposed to soluble factors in the biofilm was significantly lower than that in the control group. However, the expressions of wound-healing promoting cytokines, such as stromal derived factor 1 (SDF-1) and vascular endothelial growth factor (VEGF), and antimicrobial peptides LL-37, were significantly increased. It demonstrates that the migration ability of MSCs differed among different stages of infection caused by S. aureus . During the acute phase, the inflammatory factors and chemokines produced by S. aureus may promote the migration of MSCs. However, after the establishment of bacterial biofilm, it can inhibit MSCs migration and reduce MSCs activity through the up-expression of wound-healing associated factors. These results imply that the formation of S. aureus biofilm impairs cells’ ability to recruit MSCs after bacterial stimulation, which provides new targets and therapeutic strategies for clinical diagnosis and treatment.

Osteogenic and adipogenic differentiations

Fractures, one of the most common orthopedic diseases, usually take over three months to heal. However, about 10-20% of patients still suffer from delayed union or non-union. One of the critical factors influencing bone healing is bone remodeling [ 19 , 20 ], a dynamic process between osteoblast (bone formation) and osteoclast (bone resorption) [ 21 ]. During this process, MSCs play an essential role owing to the differentiation abilities of osteogenesis, chondrogenesis, and adipogenesis [ 22 ]. In recent years, many studies in vitro have indicated that bacteria and their components significantly impact the differentiation of MCSs in osteogenesis and adipogenesis, among which S. aureus is one of the most frequently analyzed [ 18 , 23 , 24 , 25 , 26 ]. Ding et al. [ 23 ] found that, after infecting hBMSCs with different concentrations of S. aureus (0, 0.5, 1, 10, and 50 µg/ mL), the expressions of osteogenesis genes were inhibited after 14-days osteogenesis differentiation in a dose-dependent manner. They also found that the expression of miR-29b-3p in hBMSCs significantly increased. Moreover, when miR-29b-3p was inhibited, expressions of the osteogenesis genes in hBMSCs were noted to grow considerably, indicating that miR-29b-3p might be a negative regulator against hBMSCs osteogenic differentiation during infection [ 23 ]. As the rapid growth of bacteria in cell media leads to rapid nutrient depletion and acidification, Tomas et al. [ 24 ] used heat-inactivated S. aureus (HKSA) instead of alive S. aureus to conduct experiments. They observed that human adipose-tissue-derived mesenchymal stem cells (adMSCs) disposed of with HKSA significantly reduced the activity of alkaline phosphatase (ALP) as well as decreased adipogenic differentiation activity. Similarly, the soluble factors of biofilm produced by S. aureus in chronic infection played the same role. A previous study also showed that hBMSCs exposed to the conditioned medium of S. aureus biofilm significantly reduced the intracellular calcium deposits and oil droplets at 7, 14, and 21 days after osteogenic differentiation, with decreased expressions of osteogenesis and adipogenesis markers [ 18 ]. These results indicate that S. aureus and its biofilms can affect MSCs functions by inhibiting osteogenic and adipogenic differentiation abilities.

However, as virulence factors of S. aureus , α-hemolysin, lipoteichoic acid (LTA), and Staphylococcal enterotoxin C2 (Sect. 2) may have opposite functions on MSCs. Our previous study in vitro has shown that administrated mice with 40 µg/ml α-hemolysin significantly induced bone destruction directly by suppressing osteogenesis by stimulating the expression of caveolin-1 and activated lipid rafts accumulation in BMSCs [ 27 ]. However, LTA, as one of the primary components of the cell wall of S. aureus , was not found to affect the proliferation and differentiation of adMSCs in vitro [ 24 ]. Liu et al. [ 25 ] found that 10 µg/mL LTA could enhance the autophagy activity of MSCs in mice, manifested as increased expression of LC3-II protein and decreased expression of P62, thus promoting the osteogenic differentiation of MSCs. And osteogenic differentiation genes, such as ALP, collagen type I (ColI), and runt-related transcription factor-2 (Runx2), were upregulated. The difference in conclusions between the two studies may be due to differences in LTA concentrations. Besides, Wu et al. found that Sect. 2 also had the same bone-promoting effect on MSCs. The results showed that Sect. 2 had no significant impact on the proliferation of rat BMSCs at different concentrations from 1pg/ml to 500pg/ml. However, the formation of calcium nodules in BMSCs was significantly increased, and the expressions of osteogenic markers such as ALP, Runx2, OCN, and OPN were upregulated [ 26 ]. These results indicate that S. aureus and its related components have inconsistent effects on the osteogenic and adipogenic differentiation of MSCs, suggesting that the inhibition of MSCs by S. aureus is not caused by virulence factors alone. The changes in bone mass caused by these virulence factors may provide new targets and strategies for osteoporosis prevention and osteogenesis promotion in clinical.

Hematopoietic stem cells and progenitor cells (HSPCs)

In addition to maintaining hemostasis, such as oxygen and nutrient transport, and innate and adaptive immune responses, the cells in the blood generated by hematopoietic stem cells (HSCs) also can promote tissue regeneration and repair. HSCs can self-renewal and have a variety of differentiation potentials. Through asymmetric division, they can generate daughter cells that maintain the HSCs potential and a hematopoietic progenitor cell (HPCs). Although HPCs lose their self-renewal ability, they can further differentiate into various mature blood cells, such as red blood cells, macrophages, neutrophils, and T and B lymphocytes (T and B cells). Therefore, HSPCs are generally regarded as the cornerstone of the biological hematopoietic and immune systems [ 28 , 29 , 30 ].

As an essential part of the immune system, HSPCs can respond to the inflammatory environment in various ways. Recently studies have indicated that HSPCs are the first responder during infection, and various inflammatory factors released, such as TNF and interleukin-1 (IL-1), can affect the function of HSPCs [ 31 , 32 ]. S. aureus is one of the main bacteria which cause human skin and soft tissue infection [ 33 , 34 ]. According to the severity and duration of infection, S. aureus showed two different effects on HSPCs ultimately. When the infection is mild or at the early stage, it could induce various acute inflammatory changes in the microenvironment of the infection site, resulting in the rapid recruitment of HSPCs in the peripheral blood circulation to the infection situation, and enhancing the survival and proliferation and differentiation ability of local HSPCs, thus generating many immune cells (such as polymorphonuclear neutrophils, PMNs) to eliminate the inflammation caused by infection. However, when the infection is severe or enters a chronic stage, HSPCs would cause chronic inflammatory persistence and destroy the ecological niche, thus leading to the failure and dysfunction of HSPCs [ 31 , 35 , 36 , 37 , 38 , 39 ].

In addition, S. aureus can also affect the function of HSPCs by directly targeting it (Fig.  1 ; Table  1 ). Many previous studies in vivo have shown that the lipopeptides and LTA of S. aureus could directly interact with toll-like receptor 2 (TLR2) on HSPCs, thus activating the TLR2/ myeloid differentiation factor 88 (MyD88) downstream signaling pathway, leading to the enhancement of the proliferation and myeloid differentiation of HSPCs and increased the number of PMNs at the infection situation [ 40 , 41 ]. By constructing TLR2 or MyD88 deficient transgenic mice, Granick et al. [ 42 ] also found little difference in the number of HSPCs in TLR2 or MyD88 deficient mice and WT mice. However, the number of HSPCs and its daughter cells (such as promyelocytes and PMNs) at the wound site in TLR2 or MyD88 defective mice infected with S. aureus showed an apparent decreasing trend, indicating that S. aureus does affect the functions of HSPCs by directly activating TLR2/MyD88 signaling pathway. By injecting isolated HSPCs from WT mice into the S. aureus-infected WT mice and TLR2 or MyD88 deficient mice, they found that HSPCs had similar proliferation and differentiation abilities in different wound environments. In addition, HSPCs from WT, MyD88, or TLR2 deficient mice pretreated with prostaglandin E2 (PGE2) were injected into wounds of S. aureus -infected WT mice and results showed TLR2-dependent PGE2 production could regulate the proliferation and differentiation of HSPCs in mice. Their study indicated that S. aureus could induce the production of PGE2 by directly activating the TLR2/MyD88 signaling pathway of HSPCs, which in turn target on HSPCs themselves through PGE2, thus increasing the survival, proliferation, and differentiation ability of HPSCs. This result improved the internal molecular mechanism of S. aureus , stimulating the proliferation and differentiation of HSPCs. In addition, Maneu et al. [ 43 ] co-cultured purified mouse bone marrow HPCs with inactivated S. aureus and found that it could directly induce the differentiation of HPSCs into the myeloid system. These results highlight the importance of HSPCs in anti-infection in S. aureus infection. Modulating the TLR2 signaling pathway or focusing on the severity or stage of the infection may provide a new strategy by intervening with the function of HSPCs in clinical infection.

Dental pulp stem cells(DPSCs) and hair follicle stem cells

In addition to MSCs and HSPCs, S. aureus and its virulence factors can affect other stem cells in vitro differently (Fig.  1 ; Table  1 ). Shayegan et al. [ 44 ] analyzed the interaction between adult DPSCs and different concentrations of LTA. They found that LTA activated NF-κB signaling pathway through TLR2 in DPSCs, leading to the enhancement of proliferation and migration of DPSCs. In addition, as one of the most common pathogenic microorganisms in the human body, S. aureus can not only cause pulpitis, osteomyelitis, food poisoning, and other common inflammatory diseases but also be the leading cause of some inflammatory diseases, such as nail-free folliculitis (FD). As a rare scalp inflammatory disease, the pathogenesis of FD has been confirmed to be closely related to S. aureus infection and autoimmune dysfunction. Also, FD treatment primarily relies on antibiotics and immunomodulators. Studies have shown that FD can permanently destroy hair follicle stem cells, which indicates that S. aureus may adversely affect the function of hair follicle stem cells in the human body [ 45 , 46 ].

Effects of stem cells on S. Aureus

S. aureus infection is the primary microorganism that causes infections in the skin, soft tissue, respiratory system, bone and joint, and vascular systems [ 1 ]. Due to antibiotics abuse and the developed ability of antibiotic resistance, the prevalence of methicillin-resistant S. aureus (MRSA), the mortality rate of sepsis, and other diseases have been increasing in recent years [ 4 ]. Given this public health problem, it is crucial to develop new treatment strategies. As the most widely distributed adult stem cells in the human body, MSCs have been proven to help the host resist bacteria and reduce tissue damage and inflammation [ 5 ]. In this part, we summarize the role of mesenchymal stem cells in various diseases caused by S. aureus and their potential therapeutic targets (Fig.  2 ; Table  2 ), providing new possibilities for cell therapy in treating infections and inflammatory diseases.

The role of stem cells in alleviating infectious and inflammatory disease

Pneumonia in cystic fibrosis (CF)

CF is a fatal genetic disease that is caused by the mutation of the cystic fibrosis transmembrane conduction regulator (CFTR) gene, resulting in the defective activity of the chloride channel and increased viscosity of mucus, leading to severe lung infection and inflammation [ 47 , 48 , 49 ]. In recent years, with the development of CFTR enhancers and other drugs, new progress has been made in treating CF. However, these new drugs still find it challenging to improve the pulmonary bronchiectasis and inflammation caused by bacterial colonization, which leads to severe pulmonary edema and even death [ 50 ]. The latest research results in vivo found that treating human mesenchymal stem cells (hMSCs) significantly alleviates the CF lung infection caused by S. aureus [ 51 ]. Further studies in vitro showed that the bioactive molecules in the supernatant of hMSCs reduced the overall bacterial load, and the antimicrobial peptides (e.g., LL-37) produced by hMSCs enhanced the sensitivity of bacteria to antibiotics, thus improving the ability of antibiotics to kill bacteria. In addition, they found that hMSCs with impaired CFTR function may produce fewer LL-37 [ 51 ], suggesting that mutation of the CFTR gene in patients’ MSCs may also be one reason why S. aureus -associated pneumonia was persistent. In conclusion, exogenous hMSCs may provide us with a unique strategy for the treatment of CF patients.

Chronic skin trauma

Skin is the largest organ in the human body, regulating body fluid and balancing body temperature and antibacterial infection. It is the body’s first defense to resist harmful external stimuli. Therefore, the integrity of the skin plays a crucial role in the homeostasis of the internal environment [ 52 ]. The incidence of chronic skin trauma has been increasing recently. Delayed healing or even non-healing of chronic wounds leads to long-term skin defects, which further affect the functions of other tissues and organs in the human body and heavily burden public medical resources. A common feature of chronic skin wounds is that they are colonized by pathogenic bacteria such as S. aureus . The wounds infected by S. aureus generally show expansion and delayed healing [ 53 ]. At the same time, MSCs can be anti-inflammatory and bactericidal and promote tissue repair [ 54 , 55 , 56 ]. Therefore, the potential role of MSCs in chronic skin trauma attracts more and more attention.

Horses and humans all suffer chronic wounds due to S. aureus infection. The pathogenesis mechanism of delayed or non-healed is similar so that horses can be used as a physiologically related model of human wound healing [ 57 , 58 ]. Harman et al. [ 59 ] used an equine MSC suspension and supernatant from healthy horses to culture S. aureus . They evaluated the growth of S. aureus and the damage to the bacterial membrane. The results showed that both the suspension and supernatant could effectively inhibit the growth of S. aureus and depolarize the bacterial membrane, leading to S. aureus biofilm destruction. At the same time, they also identified four specific antimicrobial peptides (AMP [ 60 ], a class of alkaline polypeptides with extensive antibacterial effects produced in multicellular organisms) secreted by equine MSCs, demonstrating that equine MSCs participated in antibacterial effects through the secretion of AMP. Subsequently, Marx et al. [ 61 , 62 ] also used equine MSCs to study the therapeutic effect of MSCs on various chronic wound pathogens with biofilm-forming ability. They found that the MSCs could effectively inhibit the S. aureus biofilm formation in vitro by secreting cysteine protease. In addition, MSCs could also stimulate keratinocytes through the secretion of CCL2 to increase their AMP secretion capacity and antibacterial function, thus indirectly affecting the formation of S. aureus biofilm.

In addition to the horse model, mice and dogs commonly use physiologically related models for human wound healing. Johnson et al. [ 63 ] observed the effect of intravenous MSCs combined with antibiotics in treating chronic S. aureus wound infection in mouse and dog models and found that MSCs could effectively limit the severity of S. aureus infection in vivo and significantly enhance the efficiency of the antibiotics. Chow et al. [ 64 ] also studied the MSCs activity in the mouse biofilm infection model. They found that various cytokines secreted by MSCs could directly or indirectly target pathogenic microorganisms such as S. aureus , effectively inhibiting the microorganisms’ colonization in chronic wounds and thus improving wound healing.

Although MSCs have not yet been applied in clinical practice in the treatment of chronic wounds and the role of MSCs in delayed or non-healed human wounds caused by S. aureus infection is not clear, the antibacterial effect of MSCs in biological models such as horses, mice, and dogs provide a rich theoretical basis for the future application of MSCs in the treatment of chronic wounds.

Adipose stem cells and progenitor cells

Adipose tissue-derived stem cells (ascs).

ASCs, also known as adMSCs, are a particular type of MSCs that can be directly extracted from adipose tissue obtained through lipoplasty or liposuction without further expansion in a culture medium [ 65 ]. As one of the primary sources of mature adipocytes in adipose tissue, ASCs can self-renewal and have various differentiation potentials. Recently studies have shown that ASCs have the function of anti-inflammatory, cell microenvironment protection, and tissue regeneration [ 66 , 67 ]. Therefore, its potential target role in treating various inflammatory diseases is gradually attracting research attention.

Chronic skin wounds

A recent case report on chronic venous leg ulcers showed that local autologous ASCs-enriched, high-density lipoaspirate (HDL), and timolol could promote the healing of ulcerative wounds [ 68 ]. In addition, Moradi et al. [ 69 ] also found that ASCs usage could significantly boost the healing of MRSA-infected wounds in type 2 diabetic rats model. Their findings suggest that ASCs may have a therapeutic effect on the delayed recovery of chronic skin wounds caused by S. aureus infection. Ruiz et al. [ 70 ] evaluated the phagocytic ability of human ASCs by flow cytometry, fluorescent latex beads, and transmission electron microscopy. The results showed that human ASCs had a strong phagocytic ability against common pathogenic microorganisms such as S. aureus in skin wounds. Wood et al. [ 71 ] also used scanning electron microscopy (SEM) and other techniques to evaluate human ASCs’ interaction with S. aureus. The results also showed that human ASCs had a strong phagocytosis effect on S. aureus . In addition, the growth and proliferation ability of S. aureus could be significantly inhibited even by using a conditioned medium obtained from ASCs without being infected with S. aureus , which was consistent with the results by Ruiz et al. The phagocytic ability of ASCs on pathogenic microorganisms such as S. aureus may be the intrinsic mechanism of ASCs in treating chronic skin wounds, providing a theoretical basis for future research on stem cell therapy in chronic skin wounds.

Bone and joint infection

Clinical treatment of bone and joint infection is limited. Currently, antibiotics and local debridement surgery are common strategies, but the treatment effects often fail to meet expectations. Therefore, cell therapy based on MSCs for bone infection has recently attracted wide attention. In the latest study in vitro, Yagi et al. [ 72 ] found that the conditioned medium of ASCs significantly inhibited the growth of S. aureus in joint synovial fluid. Previous studies have reported that the primary mechanism of antibacterial effect in MSCs is through cationic antimicrobial peptide LL-37 [ 73 ], which was further confirmed in this study. Pretreatment of ASC with 1, 25-dihydroxyvitamin D (1,25(OH)2D3) increased the expression of LL-37 and enhanced its antibacterial activity, and was reversed by vitamin D receptor antagonists [ 72 ], suggesting that the vitamin D signaling pathway plays a critical regulatory role.

Bone infection, which is most infected by S. aureus [ 74 , 75 ], is a common cause of bone defects [ 76 ]. After infection, it mainly activates the NF-κB signaling pathway, reducing the proliferation of infected osteoblasts and their ability to form calcium nodules [ 77 , 78 , 79 ]. Meanwhile, activated B cells can secrete RANKL under infection, increasing the activity of osteoclasts [ 80 ]. All the above results in extensive bone loss after infection. To improve the prognosis of bone infection, stem cell therapy gradually comes into researchers’ vision in recent years. Wagner et al. [ 81 ] found that adequate debridement followed by topical application of ASCs in a mouse model of osteomyelitis resulted in increased osteoblast proliferation and reduced osteoclast numbers via the RANKL/OPG axis; In addition, the expression level of B-cell activating factor (BAFF) treated by ASCs in bone was significantly decreased, while the galectin-9 (GAL9) was increased. Immunofluorescence and flow cytometry showed that B cell populations decreased significantly. These results suggest that ASCs can regulate the innate immune system, improve the bone microenvironment and reduce the damage to bone regeneration after osteomyelitis. However, Seebach et al. [ 82 ] proposed the opposite view. Their study pointed out that BMSCs implantation on the femur of osteomyelitis rats significantly increased osteomyelitis score and aggravated infectious bone defects. In addition, they found that BMSCs exposed to S. aureus significantly increased the expression of pro-inflammatory factors such as IL-6, IL-1β, TNF-α, monocyte chemotactic protein-1 (MCP-1), and anti-inflammatory, immune mediators prostaglandin E synthase 3 (PTGES3) and TNF-stimulated gene 6 protein (TSG-6). However, the gene expression of the cathelicidin antimicrobial peptide (CAMP/LL-37), which was previously discussed as contributing to the cell-mediated bacterial defense mechanism, was not significantly enhanced. In conclusion, the application of MSC in infectious diseases still needs further research, but MSCs still have their unique therapeutic advantages in aseptic osteomyelitis. Meanwhile, MSCs conditional medium has been proven to have antibacterial activity, which could provide a new therapeutic strategy in bone infection.

Acute lung injury (ALI)

ALI is a clinical syndrome characterized by the acute onset of clinical symptoms such as tachypnea and hypoxemia. The mortality rate remained around 40% in the past 20 years [ 83 , 84 ]. ALI can be caused by various causes, including S. aureus infection [ 85 ]. Qian et al. [ 86 ] studied the effect of ASCs on mice models of acute lung injury. They found that administrating ASCs through the airway significantly reduced the severity of lung inflammation and bacterial load of mice caused by S. aureus infection. It was also revealed that the antimicrobial activity of ASCs was mainly achieved through the secretion of TLR2-MyD88-JAK2/Stat3-dependent regenerating islet-derived IIIγ (RegIIIγ). Their findings revealed the therapeutic role of ASCs in acute lung injury caused by S. aureus infection and the internal mechanism of ASCs’ resistance to infection, indicating that ASCs played a bactericidal effect on S. aureus through the paracrine circuit. It suggests that ASCs may be a potential therapeutic target for S. aureus -caused ALI and provides a new strategy to treat ALI in the future.

S. aureus , the most common pathogen in bovine mastitis [ 87 ], can invade mammary epithelial cells, form abscesses and promote the formation of biofilms [ 88 ]. The effective rate of commonly used antibiotics (such as pirlimycin) in treating S. aureus -caused bovine mastitis is only 10-30% [ 89 ]. Therefore, sacrificing infected dairy cows is the most used strategy. But this strategy brings tremendous financial losses and seriously affects public health through unstable milk quality [ 90 ]. In the latest study, Peralta et al. [ 91 ] found that conditioned medium (CM) of MSC from fetal bovine bone marrow (BM-MSC) and adipose tissue (AT-MSC) significantly inhibited the growth of S. aureus isolated from clinical bovine mastitis cases in vitro. Among them, the primary AMP were β-defensin 4 A (bBD-4 A) and NK-lysine 1 (NK1). Still, only bBD-4 A was upregulated in BM-MSC, indicating differences in the antimicrobial efficacy of MSCs from different tissue sources against S. aureus . Torres et al. [ 92 ] verified the role of MSCs in treating S. aureus -caused mastitis in vivo. They found that the total bacterial load in milk from mastitis cows was significantly reduced after being treated with adipose tissue-derived stem cells (ASCs). Taken together, these results suggest a potential basis for the development of MSC-based therapies for mastitis. However, we should further explore the mechanism while optimizing its therapeutic effect.

Adipocyte progenitor cells (APCs)

In addition to ASCs, APCs are also one of the sources of mature adipocytes in biological adipose tissue [ 66 ]. The relationship between obesity and immune capacity has always attracted the attention of many researchers. Previous studies in vivo showed that the risk of bacterial infection in obese individuals was significantly higher than in normal individuals [ 93 , 94 ]. By studying the infectious resistance in diet-induced obese mice infected with S. aureus , Zhang et al. [ 95 ] found that the proliferation and differentiation of APCs were abnormal in adipose tissue, which caused the depletion of APCs and accumulation of mature adipose cells during obesity. Moreover, ASCs in the adipose tissue in normal individuals can secrete AMP to resist S. aureus infection. Then they observed changes in AMP expression in adipocyte maturation and found that mature adipocytes could not produce AMP. They also demonstrated that mature adipocytes could indirectly inhibit ASCs through secreting transforming growth factor-β (TGFβ) by treating APCs with TGFβ-receptor inhibitors or peroxisome proliferator-activated receptor-γ agonists. These findings further suggest that the decreased resistance of obese individuals to S. aureus infection may be caused by the reduced ability of APCs to survive, multiply, and differentiate in adipose tissue. However, some studies found that the formation of dermal fat was conducive to enhancing the anti-infection ability of the body [ 96 , 97 ], which needed further investigation.

With the continued understanding of the relationship between adipose-derived stem cells/ progenitor cells (ASPCs) and S. aureus , more and more studies have shown that ASPCs play an essential role in various inflammatory diseases caused by S. aureus (Fig.  2 ; Table  2 ). Although there are no clinical reports on using of ASPCs as the treatment of S. aureus -caused infectious diseases, the potential target role of ASPCs in the treatment of infectious diseases suggests that stem cell therapy may be a new therapeutic approach for delayed wound healing and soft tissue inflammation caused by S. aureus infection in the future.

Other stem cells

Human umbilical cord-derived mesenchymal stem cells (hucmscs).

In addition to ASPCs, umbilical cord-derived stem cells, also known as hucMSCs, are a particular type of MSCs. hucMSCs can be isolated from the human umbilical cord with low immunogenicity, which makes hucMSCs a promising candidate for stem cell therapy [ 98 ].

MRSA is one of the main pathogenic microorganisms of hospital infection [ 99 ], and severe pneumonia caused by MRSA often induced cytokine storm, eventually leading to multiple organ dysfunction syndromes (MODS) and even death [ 1 , 100 ]. Linezolid is the preferred antibiotic in hospital-acquired pneumonia [ 101 , 102 ]. Still, due to the imbalance between the pathogen and the host immune system during pneumonia, the therapeutic effect of antibiotics alone could be better [ 103 ]. MSCs have become one of the essential therapeutic methods in regenerative medicine in recent years due to their powerful immunomodulatory properties and their ability to ameliorate the storm of inflammatory factors through paracrine [ 104 , 105 , 106 ]. KONG et al. [ 107 ] found that in a rabbit model of severe pneumonia, the combination treatment of hucMSCs and linezolid significantly alleviated clinical symptoms such as cough, shortness of breath, decreased food intake, and less mucosal congestion and erosions under bronchoscopy. They also found that immune cell infiltration and inflammatory exudation were more limited, and the plasma levels of IL-8, IL-6, C-reactive protein (CRP), and TNF-α significantly decreased. Based on these findings, Mccarthy et al. [ 108 ] proposed that MSCs could be delivered directly to the lungs of infected people by atomized cell suspension. After atomization, the antibacterial capacity and the contents of factors such as hepcidin, lipid carrier protein-2, and IL-8 were not affected, but it was worth noting that the LL-37 level was significantly reduced. This may be related to the gas-liquid interface generated by the atomizer, which may affect protein stability [ 109 ]. In conclusion, hucMSCs and linezolid administration improve the survival rate of severe pneumonia and reduce lung impairment (Fig.  2 ; Table  2 ). In addition, Nebulization technologies provide us with new clinical insights. These findings highly indicated the clinical potential of MSCs in treating severe pneumonia.

By studying human DPSCs, Gronthos et al. [ 110 ] found a kind of cells that have similar immunophenotype with BMSCs and can form mineralized nodules. The cells, with spindle morphology, self-renewal ability, and multiple differentiation potential, isolated from pulp tissue, are called DPSCs. During infection and injury, DPSCs enhance their proliferation and differentiation ability and stimulate the migration of pulp progenitor cells to the site, where they generate a protective layer to protect and repair dentin [ 111 , 112 , 113 ]. Therefore, their potential therapeutic role in dental caries and other oral diseases has attracted increasing attention from researchers.

The reaction of dental pulp to dental caries is a complex process to prevent dental caries’ lesions and protect the dental pulp from bacterial invasion [ 114 ]. DPSCs differentiate into new odontogenic cells to accomplish repair, regeneration, and tertiary dentin formation. Since the elimination of local infection occurs before repair and regeneration, most people believe that DPSCs have both abilities to regenerate and antibacterial potential [ 115 ]. Lundy et al. [ 116 ] confirmed that, unlike previous studies, the classical AMP (LL-37, β-defensin 2, β-defensin 3, and lipocalin) had lower gene expression in DPSCs. However, they found that hepatocyte growth factor (HGF) in the culture medium of DPSCs can destroy the bacterial membrane of S. aureus in vitro , thereby exerting its bactericidal activity (Fig.  2 ; Table  2 ). In conclusion, with the deepening of our understanding of the antibacterial ability of DPSCs, more support will be provided for dental pulp repair and regeneration in the future.

Conclusions and future perspectives

This paper reviewed the relationship between S. aureus and stem cells. We found that S. aureus increased the migration, proliferation, and osteogenic differentiation of MSCs and increased the proliferation and differentiation of other stem cells, such as HSPCs and DPSCs, in the case of acute infection. However, when the infection progressed to the chronic stage, S. aureus could inhibit the average physiological ability of stem cells such as MSCs, HSPCs, and DPSCs, even leading to permanent destruction of stem cells (such as FD). It showed that the effect of S. aureus on stem cells was closely related to the toxicity and infection severity of bacteria strain. In addition, we also found that bone marrow MSCs, ASPCs, DPSCs, umbilical cord stem cells, and other stem cells had specific therapeutic effects on various S. aureus -caused infection. These stem cells could directly or indirectly target S. aureus and biofilm and then killed S. aureus and relived the local inflammation.

However, stem cells still face many challenges in the treatment of infectious diseases associated with S. aureus : (1) Complexity of Mechanisms: The intricacies of stem cell therapy in combating infections surpass those of conventional antibiotic therapy. Stem cells exhibit a diverse range of mechanisms including direct antimicrobial activity, immune response modulation, and tissue repair following infection-induced damage [ 117 , 118 ]. The multifaceted nature of these mechanisms presents obstacles for both research and practical implementation in clinical settings. (2) Evaluation of efficacy: The evaluation of the efficacy of stem cell therapy is difficult. For S. aureus -related infections, traditional assessment metrics focus on pathogen clearance and reduction of infection markers. However, stem cell therapy may affect multiple biological processes simultaneously, including immunomodulation and tissue repair, so more comprehensive methods of efficacy assessment need to be developed [ 92 , 119 ]. (3) Safety concerns: The safety implications of utilizing stem cell therapy in the context of infectious diseases are of particular concern. For instance, while stem cells may facilitate tissue regeneration, they may also hinder the immune system’s ability to combat pathogens, potentially resulting in the prolonged presence or reemergence of the infection [ 120 , 121 ]. (4) Antibiotic resistance: Staphylococcus aureus, particularly methicillin-resistant strains, have demonstrated significant resistance to various antibiotics [ 122 ]. The potential incorporation of stem cell therapy in addressing this issue may necessitate a concurrent use of antibiotics; however, the optimal strategy for this combination to prevent the emergence of resistance remains a subject of inquiry [ 123 , 124 , 125 ]. (5) Individual differences: Variability among patients, such as differences in immune status, pathogen characteristics, and the location and severity of infection, can impact the effectiveness of stem cell therapy [ 126 , 127 ]. Consequently, personalized treatment regimens are necessary, leading to heightened complexity in clinical implementation. (6) Ethical and regulatory: The ethical and regulatory issues associated with stem cell therapy apply equally to the use of stem cells in the treatment of S. aureus -related infections. In particular, research using embryonic stem cells may face more stringent ethical scrutiny and legal restrictions.

In summary, S. aureus and its related components affect the functional state of various stem cells, while the stem cells secrete polypeptides, chemokines and cytokines (AMP, HGF, etc.) that inhibit the activity of S. aureus . Therefore, further studies on the interaction between S. aureus and stem cells may provide new ideas for the treatment of infectious diseases in the future, but their clinical efficacy and safety need to be further investigated and confirmed.

Data availability

Not applicable.

Abbreviations

• Staphylococcus aureus

Methicillin-resistant S. aureus

Induced pluripotent stem cells

Embryonic stem cells

• Mesenchymal stem cells

Bone marrow-derived mesenchymal stem cells

Tumor necrosis factor α

C-C motif chemokine ligand 20

Stromal derived factor 1

Vascular endothelial growth factor

Adipose-tissue-derived mesenchymal stem cells

Alkaline phosphatase

Lipoteichoic acid

Staphylococcal enterotoxin C2

Runt-related transcription factor-2

Collagen type I

Hematopoietic stem cells and progenitor cells

Hematopoietic stem cells

Hematopoietic progenitor cell

Human mesenchymal stem cells

Adipose tissue-derived stem cells

Extracellular vesicles

Interleukin-1

Toll-like receptor 2

Myeloid differentiation factor 88

Prostaglandin E2

Dental pulp stem cells

Cystic fibrosis

Cystic fibrosis transmembrane conduction regulator

High-density lipoaspirate

Scanning electron microscopy

B-cell activating factor

Monocyte chemotactic protein-1

TNF-stimulated gene 6 protein

Prostaglandin E synthase 3

Regenerating islet-derived IIIγ

β-defensin 4 A

NK-lysine 1

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Transforming growth factor-β

Adipose-derived stem cells/ progenitor cells

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Acknowledgements

This research was funded by the National Natural Science Foundation of China, grant numbers: 82172197, 82272517, 82201732, the President Foundation of Nanfang Hospital, Southern Medical University, grant number: 2021L001.

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Zi-xian Liu, Guan-qiao Liu and Ze-xin Lin contributed equally to this work.

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Division of Orthopaedics & Traumatology, Department of Orthopaedics, Southern Medical University Nanfang Hospital, Guangzhou, 510515, China

Zi-xian Liu, Guan-qiao Liu, Ze-xin Lin, Ying-qi Chen, Peng Chen, Yan-jun Hu, Bin Yu & Nan Jiang

Guangdong Provincial Key Laboratory of Bone and Cartilage Regenerative Medicine, Southern Medical University Nanfang Hospital, Guangzhou, 510515, China

Department of Orthopedics, Lanzhou University Second Hospital, Lanzhou, 730000, China

Zi-xian Liu

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Conceptualization, N.J. and B.Y.; writing—original draft preparation, Z.L.1. and G.L.; Validation, Y.C. and Z.L.2.; Writing—Review and Editing, P.C. and Y.H.; Supervision, N.J. and B.Y.; Z.L.1., G.L., Z.L.2 contributed equally to this work. All authors have read and agreed to the published version of the manuscript. (Z.L.1 refer to Zi-xian Liu, Z.L.2 refer to Ze-xin Lin)

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Liu, Zx., Liu, Gq., Lin, Zx. et al. Effects of Staphylococcus aureus on stem cells and potential targeted treatment of inflammatory disorders. Stem Cell Res Ther 15 , 187 (2024). https://doi.org/10.1186/s13287-024-03781-6

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The writing process of preparation, writing, and revisions applies to every essay or paper, but the time and effort spent on each stage depends on the type of essay .

For example, if you’ve been assigned a five-paragraph expository essay for a high school class, you’ll probably spend the most time on the writing stage; for a college-level argumentative essay , on the other hand, you’ll need to spend more time researching your topic and developing an original argument before you start writing.

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Before you start writing, you should make sure you have a clear idea of what you want to say and how you’re going to say it. There are a few key steps you can follow to make sure you’re prepared:

• Understand your assignment: What is the goal of this essay? What is the length and deadline of the assignment? Is there anything you need to clarify with your teacher or professor?
• Define a topic: If you’re allowed to choose your own topic , try to pick something that you already know a bit about and that will hold your interest.
• Do your research: Read  primary and secondary sources and take notes to help you work out your position and angle on the topic. You’ll use these as evidence for your points.
• Come up with a thesis:  The thesis is the central point or argument that you want to make. A clear thesis is essential for a focused essay—you should keep referring back to it as you write.
• Create an outline: Map out the rough structure of your essay in an outline . This makes it easier to start writing and keeps you on track as you go.

Once you’ve got a clear idea of what you want to discuss, in what order, and what evidence you’ll use, you’re ready to start writing.

The introduction sets the tone for your essay. It should grab the reader’s interest and inform them of what to expect. The introduction generally comprises 10–20% of the text.

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The first sentence of the introduction should pique your reader’s interest and curiosity. This sentence is sometimes called the hook. It might be an intriguing question, a surprising fact, or a bold statement emphasizing the relevance of the topic.

Let’s say we’re writing an essay about the development of Braille (the raised-dot reading and writing system used by visually impaired people). Our hook can make a strong statement about the topic:

The invention of Braille was a major turning point in the history of disability.

2. Provide background on your topic

Next, it’s important to give context that will help your reader understand your argument. This might involve providing background information, giving an overview of important academic work or debates on the topic, and explaining difficult terms. Don’t provide too much detail in the introduction—you can elaborate in the body of your essay.

3. Present the thesis statement

Next, you should formulate your thesis statement— the central argument you’re going to make. The thesis statement provides focus and signals your position on the topic. It is usually one or two sentences long. The thesis statement for our essay on Braille could look like this:

As the first writing system designed for blind people’s needs, Braille was a groundbreaking new accessibility tool. It not only provided practical benefits, but also helped change the cultural status of blindness.

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In longer essays, you can end the introduction by briefly describing what will be covered in each part of the essay. This guides the reader through your structure and gives a preview of how your argument will develop.

The invention of Braille marked a major turning point in the history of disability. The writing system of raised dots used by blind and visually impaired people was developed by Louis Braille in nineteenth-century France. In a society that did not value disabled people in general, blindness was particularly stigmatized, and lack of access to reading and writing was a significant barrier to social participation. The idea of tactile reading was not entirely new, but existing methods based on sighted systems were difficult to learn and use. As the first writing system designed for blind people’s needs, Braille was a groundbreaking new accessibility tool. It not only provided practical benefits, but also helped change the cultural status of blindness. This essay begins by discussing the situation of blind people in nineteenth-century Europe. It then describes the invention of Braille and the gradual process of its acceptance within blind education. Subsequently, it explores the wide-ranging effects of this invention on blind people’s social and cultural lives.

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Lack of access to reading and writing put blind people at a serious disadvantage in nineteenth-century society. Text was one of the primary methods through which people engaged with culture, communicated with others, and accessed information; without a well-developed reading system that did not rely on sight, blind people were excluded from social participation (Weygand, 2009). While disabled people in general suffered from discrimination, blindness was widely viewed as the worst disability, and it was commonly believed that blind people were incapable of pursuing a profession or improving themselves through culture (Weygand, 2009). This demonstrates the importance of reading and writing to social status at the time: without access to text, it was considered impossible to fully participate in society. Blind people were excluded from the sighted world, but also entirely dependent on sighted people for information and education.

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Braille paved the way for dramatic cultural changes in the way blind people were treated and the opportunities available to them. Louis Braille’s innovation was to reimagine existing reading systems from a blind perspective, and the success of this invention required sighted teachers to adapt to their students’ reality instead of the other way around. In this sense, Braille helped drive broader social changes in the status of blindness. New accessibility tools provide practical advantages to those who need them, but they can also change the perspectives and attitudes of those who do not.

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Genome-wide association analysis identify candidate genes for feed efficiency and growth traits in Wenchang chickens

• Keqi Cai 1 , 2 ,
• Ranran Liu 2 ,
• Limin Wei 3 ,
• Xiuping Wang 4 ,
• Huanxian Cui 2 ,
• Jie Wen 2 ,
• Yuxiao Chang 1 &
• Guiping Zhao 2 , 3  

BMC Genomics volume  25 , Article number:  645 ( 2024 ) Cite this article

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Metrics details

Wenchang chickens are one of the most popular local chicken breeds in the Chinese chicken industry. However, the low feed efficiency is the main shortcoming of this breed. Therefore, there is a need to find a more precise breeding method to improve the feed efficiency of Wenchang chickens. In this study, we explored important candidate genes and variants for feed efficiency and growth traits through genome-wide association study (GWAS) analysis.

Estimates of genomic heritability for growth and feed efficiency traits, including residual feed intake (RFI) of 0.05, average daily food intake (ADFI) of 0.21, average daily weight gain (ADG) of 0.24, body weight (BW) at 87, 95, 104, 113 days of age (BW87, BW95, BW104 and BW113) ranged from 0.30 to 0.44. Important candidate genes related to feed efficiency and growth traits were identified, such as PLCE1, LAP3, MED28, QDPR, LDB2 and SEL1L3 genes.

The results identified important candidate genes for feed efficiency and growth traits in Wenchang chickens and provide a theoretical basis for the development of new molecular breeding technology.

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Introduction

Poultry production is an important entreprise worldwide. Chicken is considered a healthy white meat source that is lower in fat, calories and cholesterol than other red meat sources [ 1 ]. In recent years, the promotion of white meat consumption has gradually become a trend [ 2 ]. As an important part of the white meat market, chickens represent an efficient and inexpensive source of animal protein [ 3 ]. Since the 1980s, with the continuous development of modern broiler production, China has become the second largest country in the world in terms of chicken production and consumption [ 4 ]. The main breeds of chickens are native breeds and broilers [ 5 ]. Wenchang chicken (Fig.  1 ) is one of the most popular chicken breeds in the local chicken industry, originating from Hainan Island in the South China Sea and has been raised for 400 years. It is famous for its excellent meat quality and is one of the four famous dishes from Hainan [ 6 ]. It’s an economic mainstay of animal husbandry in Hainan Province, with an annual output of nearly 100 million chickens and a total output value of 1.78 billion dollars in 2020 [ 7 ].

Picture for Wenchang chicken. ( a ), roosters ( b ), hens

However, the primary shortcoming of Wenchang chickens is their low feed efficiency [ 8 ]. Therefore, there is an urgent need to find a more reliable breeding method to improve the feed efficiency of Wenchang chickens. Feed represents more than 70% of the total cost of poultry production, and improving feed efficiency has consistently been the goal of any chicken breeding strategy [ 9 ]. Feed efficiency is contingent upon the relation between the feed intake (FI) and the growth (or bodyweight gain) of an animal and is quantified by several indexes, such as RFI and feed conversion rate (FCR) [ 10 , 11 , 12 ]. Feed efficiency is influenced by several factors, including the breed and its sex, age, diet, and management [ 13 , 14 ].The RFI is an important index measuring feed efficiency, and it is defined as the difference between the actual and expected feed intake [ 15 ].In 1963, the concept of RFI was first proposed in beef cattle research [ 16 ], and it was first applied to chickens by Luiting in 1991 [ 17 ], covering the calculation methods of RFI, heritability calculation, and the phenotypic and genetic correlations with relevant traits. Since RFI reflects the variation in feed efficiency, it appears to be independent of growth traits [ 15 ]. Studies have shown that RFI is moderately heritable in poultry [ 12 , 18 ]. The study by Bai et al. demonstrates that selecting for low RFI can improve poultry feed efficiency without compromising growth performance [ 19 ]. Growth traits are key in poultry breeding, and the properties of growth traits need to be considered in breeding for feed efficiency. In recent years, GWAS has been applied in poultry to explore the association between host genetics and economic traits [ 20 ]. Earlier studies identified candidate genes including NSUN3 , EPHA6 , and AGK for broilers RFI, while LAP3 was identified as a candidate gene for broiler body weight [ 21 , 22 , 23 , 24 , 25 , 26 ]. By deeply studying the functions and regulatory mechanisms of these genes, our aim is to provide more efficient and sustainable solutions for the chicken breeding technologies [ 4 ].

In the breeding process of this study, the focus will be on maintaining the excellent meat quality while moderately improving the feed efficiency and growth rate. The objectives of this study were (1) to determine the inheritance pattern of feed efficiency and growth traits to provide a basis for formulating a better breeding method for Wenchang chickens, and (2) to find key variants affecting feed efficiency and growth traits of Wenchang chickens and also to elucidate the molecular mechanism of feed efficiency and growth traits.

Materials and methods

Experimental birds.

The study uses 1,547 chickens hatched and raised by Hainan Tanniu Wenchang Chicken Co., Ltd. These birds are from a commercial line of Wenchang chicken selected for 18 generations. All birds were raised in three-tier battery cages (one bird per cage) under the same management and nutritional conditions. The diet was formulated based on the National Research Council (NRC) requirements [ 27 ] and the Feeding Standards of Chickens established by the Ministry of Agriculture, Beijing, China [ 28 ]. The chickens were raised from 87 to 113 days to collect data on individual phenotypes.

The phenotypes measured in this study included body weight and feed intake of chickens.

For body weight measurements, an electronic scale with a precision of 0.1 g was utilized. The trough was emptied of any remaining feed 12 h prior to weighing, and body weights were recorded at 87, 95, 104, and 113 days of age.

Feed intake in this experiment was recorded for 26 days. Fresh feed was added at a fixed time every day and recorded. In addition to the above measured phenotypes, the phenotypes calculated from the measured phenotypes included FI, ADFI, ADG, Metabolic weight in the middle of the test (MWT), and RFI.

RFI, g/d: The RFI was obtained from the multiple regression equation of average daily feed intake and metabolic weight in the middle of the test and average daily weight gain [ 18 ]. The equation was as follows:

where ADFI represents the mean average daily feed intake, µ is the intercept, sex is a fixed effect, MWT and ADG are as defined above, β1 and β2 represent the partial regression coefficient.

Genotyping, imputation, and quality control (QC)

At 112 days of age, 2 mL of blood was collected from the wing vein, placed in an anticoagulant tube containing EDTA-K2, mixed, and stored at -20℃ until later analysis. Genomic DNA was extracted from blood samples with the phenol-chloroform method. Genotyping was conducted with a customized chicken 55 K SNP array (Beijing Compass Biotechnology Co., Ltd., Beijing, China) [ 29 ]. QC of the generated genotype data was achieved using PLINK (V2.0) ( https://www.cog-genomics.org/plink/2.0/ ) software. The specific process setting the individual genotype detection rate at ≥ 90%, single SNP detection rate at ≥ 90%, minimum allele frequency (MAF) of 95%, and retention of SNPs on autosomes 1–28. A total of 45,278 SNPs in 1,479 chickens (762 males and 717 females) passed the QC. Whole genome resequencing of 247 individuals from the 1,479 Wenchang chickens, including 25 males and 222 females was carried out. The sequencing generated 150 bp paired end reads on an Illumina NovaSeq 6000 platform with the average depth of approximately 10×, at the Shenzhen BGI Co., Ltd. The QC standards were as follows: setting the detection rate of individual genotype at ≥ 90%, the detection rate of single SNP site at ≥ 90%, a MAF of 95%, and Hardy-Weinberg equilibrium (HWE) at P  < 0.000001. A total of 12,590,784 autosomal SNPs in 247 individuals were passed the QC.

Beagle 5.2 software was used to impute the 55 K chip data to the whole-genome sequence (WGS) level [ 30 ]. Before imputation, inconsistencies between the target panel and the reference panel were checked using conform-gt software ( http://faculty.washington.edu/browning/conform-gt.html ). Then, the 55 K SNP chip data were populated to the resequencing level using Beagle 5.2 software. The QC condition: HWE at P  < 1.00e-6, setting the detection rate of individual genotype at ≥ 90%, the detection rate of single SNP site at ≥ 90%, and a MAF of ≥ 0.05. A total of 12,184,765 autosomal SNPs from 1,479 samples passed QC.

Estimation of heritability and genetic correlations

Phenotypic correlation coefficients were calculated using ggpairs within the GGally R package, and then genetic parameter estimation was performed using restricted maximum likelihood (REML) of GCTA v1.93.2 beta software [ 31 , 32 ]. The statistical model used was:

where, \(\text{y}\) is a vector of observations, \(\text{b}\) is a vector of fixed effects (i.e., sex), \({\upalpha }\) is the random vector representing the genomic effects, \(e\) is the vector of random residual effects, \(\text{X}\) and \(\text{Z}\) are incidence matrices. The distribution of the random animal effect \(\alpha\) is \(\alpha \ \sim N(0,\,{\rm{G}}\sigma _a^2)\) with \(\text{G}\) is being the genomic relationship matrix, and \({\sigma }_{a}^{2}\) being the additive genetic variance.

Genome-wide association study

A GWAS analysis was carried out between all the genotyped SNPs and feed efficiency and growth traits using a mixed linear model (MLM). The MLM for feed efficiency and growth traits was performed using sex (female or male) as a fixed effect and the top three principal components (PCs) as covariates. All association tests were performed using the MLM option in GCTA based on the following model [ 33 ]:

where y is a vector of observations, X and Z are incidence matrices for the vectors for parameters b and µ, b is a vector of fixed effects including the sex and three eigenvectors from principal component analysis (PCA), µ is the vector of the additive genetic effect of the candidate SNP to be tested for association, and e is the vector of the residual effect.

The Bonferroni correction method was used in this study to determine the significance thresholds, and the formula for performing the Bonferroni corrected multiple tests was as follows:

Where \(P\) is the corrected significance threshold, \(\alpha\) represents the significance threshold for a single test, and N represents the number of multiple hypothesis tests, i.e., the number of SNPs analysed by GWAS. We calculated the number of genome-wide independent markers using the PLINK (V1.9) command -indep-pairwise, with a window size of 25 SNPs, a step of five SNPs, and an r 2 threshold of 0.2. Manhattan and quantile‒quantile (Q-Q) plots were derived from the GWAS results using the qqman ( https://cran.r-project.org/web/packages/qqman/ ) and Cairo ( http://www.rforge.net/Cairo/ ) packages within R software ( http://www.r-project.org/ ). LD blocks of target regions were performed using Haploview v4.2 software [ 34 ]. For additive and dominance effects of important SNPs on traits, the calculation process in this study was done in ASReml v4.1 software ( https://asreml.kb.vsni.co.uk/knowledge-base/asreml_documentation ). The SNP positions were updated according to the newest release from Ensembl ( https://asia.ensembl.org/index.html ). Identification of the closest genes to genome-wide significant and suggestive variants was obtained using the ChIPpeakAnno package ( https://www.bioconductor.org/packages/devel/bioc/vignettes/ChIPpeakAnno/inst/doc/pipeline.html ). Gene function enrichment analysis was performed using bioinformatics ( https://www.bioinformatics.com.cn ).

Differential expression analysis

In this study, the RFI or BW of 127 broiler chickens was ranked from low to high. Using individuals with the highest ( n  = 15) and lowest RFI ( n  = 15) phenotypes, and individuals with the highest ( n  = 15) and lowest ( n  = 15) BW. The Wenchang chickens used in this study were a mix of males and females, so we did not differentiate between genders in the 30 chickens selected. Gene expression of important candidate genes in the different groups can demonstrate the reliability of the GWAS results. Figures were generated using GraphPad Prism 8 [ 4 ].

Description of phenotypic traits

Descriptive statistics of the feed efficiency traits of Wenchang chickens are shown in Table  1 ; Fig.  2 . According to the body weights at 87, 95, 104 and 113 days, Wenchang chickens exhibited slow growth, with an ADFI of 82.52 g/d and an ADG of 8.66 g/d. The RFI ranged from − 39.14 to 36.25 g/d.

Phenotypic data and correlation analysis of Wenchang chicken

Genetic parameters’ estimates

Comparison of density markers before and after the imputation in Fig.  3 revealed a significant increase in loci after imputation. The estimated genetic parameters of residual feed intake and body weight are shown in Table  2 . The heritability of the RFI, BW, ADFI and ADG ranged from moderate and low level (0.05–0.44). RFI had the highest genetic association with ADFI and ADG, at 0.92 and 0.86, respectively ( P  < 0.001). Regarding phenotypic correlation, the genetic correlation between RFI and ADFI was 0.63 and decreased with body weight to a minimum of 0.20. Based on the result, the heritability of RFI in Wenchang chicken is low, while the heritability of body weight is not significantly different from that of other breeds.

Comparison of density markers before and after the imputation

Genome-wide association study of feed efficiency and growth traits

The GWAS results showed significant SNPs on GGA 2, 6 and 26 were associated with RFI (expansion coefficient λ of 0.987) (Fig.  4 a-b). One of significant SNPs 6_21123592 on GGA6 was located on the intron of the PLCE1 gene, with this SNP explaining 2.46% of the genetic variation. Additionally, significant SNPs 2_45795056 and 26_2851843 were located in the lncRNA introns of the ENSGALG00000052614 and ENSGALG00000001264 genes, respectively (Table  3 ). The GWAS results showed significant SNPs on GGA 2, 4, 6, 19, and 28 were associated with ADFI (expansion coefficient λ of 0.994) (Fig.  4 c-d). Specifically, the significant SNPs 4_75971941, 4_85488222, 19_2763444, and 28_388715 were found on the genes LAP3, IMMT, GATSL2 , and FBN3 , respectively (Table  3 ). The SNP 6_21123592 for RFI in the CT genotype exhibited significantly higher values than those in the CC genotype (Table  4 ). Another SNP 2_45795056 showed a significant additive effect on ADFI but a nonsignificant effect on the two feed efficiency traits assessed in this study. The estimated additive effect of candidate SNP 6_21123592 on RFI and ADFI were − 0.81 and 2.67 respectively, while the dominance effect was − 2.99 and 0.93 respectively. Significant dominance effects were observed on both RFI and ADFI, while the additive effect was not significant (Table  5 ).

Manhattan and Q-Q plots of GWAS for feed efficiency traits. The horizontal red lines indicate the thresholds for genome-wide significance ( P = 3.47E-08), and the horizontal blue lines indicate the thresholds for suggestive significance ( P = 6.93E-07)

The GWAS analysis results for growth traits at different ages are presented in Fig.  5 ; Table  6 . Significant QTLs affecting BW, ADG, and ADFI were observed in the 73.15–76.55 Mb interval of GGA4. Specifically, BW87 (Fig.  5 a, Table S1 ) had 661 SNPs in the significant interval, BW95 had 933 SNPs (Fig.  5 c, Table S2 ), BW104 had 1018 SNPs (Fig.  5 e, Table S3 ), BW113 had 1150 SNPs (Fig.  5 g, Table S4 ), and ADFI and ADG had 3 and 7 SNPs (Table  3 ; Fig.  4 c and e). The most prominent SNP among these traits was 4_75971941, located in the intron of the LAP3 gene. There were 13 important candidate genes in the colocalization interval of different day-ages, including LAP3, KCNIP4, NCAPG, LDB2, FAM184B, SEL1L3, ZCCHC4, PPARGC1A, PACRGL, SLIT2, LCORL, MED28 and QDPR (Table  6 ). Through gene function enrichment analysis (Figure S1 ), we identified significant enrichment primarily in functions such as RNA biosynthetic process, regulation of RNA metabolic process, and regulation of cellular macromolecule biosynthetic process.

Manhattan and Q-Q plots of GWAS for growth traits. The horizontal red lines indicate the thresholds for genome-wide significance ( P = 3.47E-08), and the horizontal blue lines indicate the thresholds for suggestive significance ( P = 6.93E-07)

The four LD blocks were detected within the common location of growth traits on GGA4:75971.28-75974.42 kb (Fig.  6 ), each containing 3–10 SNPs. The most significant SNP 4_75971941 was located on block1, which is located in the intron of the LAP3 gene. Genotype analysis showed that the body weight of the CC genotype at different ages was significantly higher than that of the CT genotype. Specifically, the weight of the CC genotype for BW87 was 54.89 g heavier than that of the CT genotype, and the weight of the CC genotype for BW113 was 82.80 g heavier than that of the CT genotype (Fig.  6 b-g). The estimated additive effects of candidate SNP 4_75971941 on body weight at different ages were 37.97, 46.67, 53.38 and 60.53, while the dominance effects were − 16.92, -16.19, -17.98 and − 22.26. SNP 4_75971941 had significant additive effects on BW and ADG ranging from 37.97 to 60.53. The additive effects increased with age, while the locus had significant dominance effects on BW and nonsignificant dominance effects on ADG and ADFI (Table  7 ).

Association results of the candidate SNPs on GGA4. ( a ), LD analysis of the 25 significant SNPs on GGA4. ( b , c , d , e , f , g ), The phenotypic differences of individuals with different genotypes at rs80610898 on GGA4. *, P < 0.05; **, P < 0.01; ***, P < 0.001; ns, no significance

The candidate sites for body weight were determined to be located in distinct LD blocks. In this study, haplotype association analysis was conducted using the glm model with sex as a fixed effect (Table  8 ). The analysis indicated that block1 had a highly significant effect ( P  < 0.01) on BW at various ages, while block2 exhibited a significant effect ( P  < 0.05) on BW across different age groups. The results revealed that block2, block3, and block4 all had a significant impact on ADFI ( P  < 0.05), while none of the four LD blocks showed a significant impact on RFI ( P  > 0.05).

We conducted gene differential expression analysis on all genes mapped to the feed efficiency and growth traits, respectively. Significant differences in phenotypes were observed between the high and low RFI groups (Fig.  7 a) and between the high and low BW groups (Fig.  7 c). Between the high and low RFI groups, the PLCE1 and IMMT genes showed significantly ( P  < 0.01) higher expression levels in the high RFI group than in the low RFI group (Fig.  7 b). Similarly, expression level of LAP3, MED28 , and QDPR genes was significantly higher in the high BW group than that in the low BW group ( P  < 0.01), while expression of LDB2 and SEL1L3 genes was significantly ( P  < 0.05) higher in the low BW group than the high BW group (Fig.  7 d).

The expressions of candidate genes for feed efficiency and growth traits. ( a ), Analysis of RFI Differences between high and low RFI groups. ( b ), Differential expression of related candidate genes between high and low RFI groups. ( c ), Analysis of BW Differences between high and low BW groups. ( d ), Differential expression of related candidate genes between high and low weight groups. Data were expressed as the mean ± SEM ( n = 15), *** P < 0.001, ** P < 0.01, * P < 0.05

Phenotypic and estimation of the genetic parameters of Wenchang chickens

Body weight traits (BW87-BW113) of Wenchang chickens were selected for the GWAS analysis because Wenchang chickens are listing age within that age range. Through different trait measurements at this stage, our study revealed that the BW87 was 1,409.48 g, that of BW113 was 1,894.36 g, the ADG was 18.66 g/d, and the relative growth rate was 25.60%. These findings are consistent with previous studies on yellow-feathered chickens in China [ 35 , 36 ]. The low heritability of RFI may be caused by the lack of systematic breeding efforts and the chickens are in the latter stage of growth, similar to the results of previous studies [ 37 , 38 ]. Heritability of BW, ADFI and ADG revealed in the current study ranged from medium to low (0.21–0.44), which was similar to that of other breeds [ 39 , 40 , 41 ]. The phenotypic and genetic correlations between RFI and ADFI were 0.92 and 0.63, similar to the research of Shirali et al [ 42 ]. To optimize breeding outcomes, it is recommended to integrate additional metrics, such as ADFI, and implement a multi-trait selection approach throughout the breeding process, ultimately enhancing breeding results in Wenchang chickens.

To investigate the genetic architecture of feed efficiency and growth traits in Wenchang chickens, a mixed linear model was used for GWAS analysis of related traits. Through GWAS analysis, we identified significant SNP 6_21123592 on GGA6 was located within the intron of the PLCE1 gene associated with RFI. Previous studies have shown that PLCE1 gene is highly expressed in the nervous system and belongs to the phosphoinositide-specific phospholipase C family. The production of second messenger molecules such as diacylglycerol is regulated by activated phosphatidylinositol-specific phospholipase C enzymes, which mediate small GTPases of the Ras superfamily through the activity of its Ras guanine exchange factor. As the effector of heterotrimer and small G protein, PLCE1 is involved in regulating cell growth, T-cell activation, actin organization and cell survival. Mapping the PLCE1 gene function were mostly related to nervous system activity, which regulates the function of the brain in different ways, and the brain was key to regulating diet behavior and body energy homeostasis [ 43 , 44 , 45 , 46 ].

The significant QTL affecting BW, ADG, and ADFI was located in the 73.15–76.55 Mb interval of GGA4.There were 13 important candidate genes in the colocalization interval related to BW, including LAP3, KCNIP4, NCAPG, LDB2, FAM184B, SEL1L3, ZCCHC4, PPARGC1A, PACRGL, SLIT2, LCORL, MED28 and QDPR . Among these candidates LAP3, MED28, QDPR, LDB2 and SEL1L3 demonstrated differential expression between high and low groups. Comparison with the Animal QTL database (Chicken QTL Database at (animalgenome.org)) reveals a total of 305 QTLs related to BW and 221 QTLs associated with ADG within 73.15–76.55 Mb interval of GGA4. LAP3 has been shown to catalyze the hydrolysis of the amino-terminal leucine residues of protein or peptide substrates, with diverse functions in mammals, invertebrates, microbes, and plants [ 47 ]. The primary function of LAP3 lies in protein maturation and degradation, processes crucial for metabolism, development, adaptation, and repair [ 48 ]. LAP3 gene variation may underlie variations in growth rates among species and significant genetic polymorphism of traits of interest in breeding, potentially leading to applications in animal breeding. Furthermore, studies have been conducted on its SNP and its association with growth traits in mammals, such as bovine [ 49 ]. Another study found that the LAP3 gene may have a potential function affecting muscle development in sheep [ 50 ]. Prenatal development stages are directly related to the growth and development of individual skeletal muscle, which determines the number of muscle fibers and postnatal muscle mass and further exerts long-term effects on the postnatal growth of animals [ 51 , 52 , 53 ]. Related research that tracked LAP3 mutations in Hu sheep populations reportedly linked, the mutations with body weight at different growth stages [ 54 ]. In poultry LAP3 gene was found to be associated with chicken growth traits [ 22 ].

Related study linked the MED28 gene with live weight in sheep [ 55 ].We found that the MED28 gene was related to muscle development in pig [ 56 ]. QDPR for an enzyme that regulates tetrahydrobiopterin (BH4), a cofactor for enzymes involved in neurotransmitter synthesis and blood pressure regulation. Therefore, QDPR gene are also important genes in the regulation of growth [ 57 ]. Previous studies have shown that the LDB2 gene located at GGA4 is important for chicken growth traits, and a 31-bp indel was significantly correlated with multiple growth and carcass traits in the F2 population and affected the expression of the LDB2 gene in muscle tissue [ 26 , 58 , 59 ]. It was also identified as an important candidate gene for rapid growth in chickens and had the strongest association with late body weight in Jinghai yellow chicken hens [ 22 , 60 , 61 ]. According to relevant studies, the KCNIP4 gene is located on GGA4, and the candidate gene belongs to the potassium channel interaction protein family and has a wide range of physiological functions, including heart rate regulation, insulin secretion, neurotransmitter release, and smooth muscle contraction. It was considered to be an important candidate gene for growth traits of chickens. In addition, it was reported in different breeds and different growth stages, which also verified that the KCNIP4 and FAM184B genes can affect the growth and development of chickens [ 58 , 62 , 63 ]. Studies have shown that the NCAPG gene is an important candidate gene for mammalian body size growth traits in growth trait association analysis of horses, sheep, and domestic donkeys and is involved in chromosome condensation and methylation [ 64 , 65 , 66 , 67 ]. The FAM184B gene has been found in previous studies to be associated with cattle carcass weight [ 68 ].

Transcriptomic data enable the quantification of DNA or RNA abundance and expression levels [ 69 ]. Differential analysis between different groups is conducted by measuring the expression levels of gene RNA, thereby identifying distinct patterns and variations in gene expression among different groups. The results of related studies also confirm gene expression data of important candidate genes in different groups can demonstrate the reliability of the GWAS results [ 70 , 71 ]. We showed that expression of PLCE1 in the high and low RFI groups of broilers, validatied it a key candidate gene for RFI. We found that the candidate gene LAP3 was related to the BW, ADFI and ADG traits, as demonstrated by the significant difference in the expression of LAP3 in high and low body recombination in broilers, and relevant research reports also indicated that this might be an important candidate gene affecting growth traits. This study also found that the candidate genes LDB2 , KCNIP4 , FAM184B , and NCAPG were located on GGA4 and were related to growth traits, which provides an important reference value for subsequent research on the growth traits of Wenchang chickens.

The validity of the SNPs and candidate genes obtained in this study is worth further extensive verification. The significant SNPs and candidate genes identified in this study can be incorporated into the chip markers in future research. By utilizing genomic selection breeding techniques, these markers can be used to breed and improve the target traits of WenChang chickens, ultimately enhancing breeding efficiency.

Conclusions

In this study, we identified that the significant SNP 6_21123592 was located in candidate gene PLCE1 for feed efficiency traits of Wenchang chickens, and the significant SNP 4_75971941 was located in candidate gene LAP3. Other candidate genes including MED28, QDPR, LDB2 , and SEL1L3 were identified for growth traits of Wenchang chickens. This provides a good theoretical basis for developing methods of Wenchang chickens breeding, and by further studying the functions and regulatory mechanisms of these genes, we could provide more efficient solutions for breeding of these chickens.

Data availability

The raw sequence data reported in this paper have been deposited in the Genome Sequence Archive (Genomics, Proteomics & Bioinformatics 2021) in National Genomics Data Center (Nucleic Acids Res 2022), China National Center for Bioinformation / Beijing Institute of Genomics, Chinese Academy of Sciences that are publicly accessible at https://bigd.big.ac.cn/gsa/browse/CRA016976.

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Acknowledgements

We thanks the Institute of Animal Science of CAAS, Chinese Academy of Agricultural Sciences for the support of this experiment.

This research was supported by the STI 2030-Major Project (2023ZD04072), the Wenchang Chicken superiority characteristic industrial cluster project (WCSCICP20211106), the Project of Sanya Yazhou Bay Science and Technology City (SKJC-2022-PTDX-002), the Special Project for Southern Propagation of the Chinese Academy of Agricultural Sciences (YYLH04), the National chicken industry technology system project (CARS-41), the Study of the Key Genetic Resources [JBGS (2021) 107].

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Keqi Cai & Yuxiao Chang

State Key Laboratory of Animal Nutrition, Key Laboratory of Animal (Poultry) Genetics Breeding and Reproduction, Ministry of Agriculture, Institute of Animal Sciences, Chinese Academy of Agricultural Sciences, Beijing, 100193, P.R. China

Keqi Cai, Ranran Liu, Huanxian Cui, Na Luo, Jie Wen & Guiping Zhao

The Sanya Research Institute, Hainan Academy of Agricultural Sciences, Sanya, 572025, P.R. China

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Cai, K., Liu, R., Wei, L. et al. Genome-wide association analysis identify candidate genes for feed efficiency and growth traits in Wenchang chickens. BMC Genomics 25 , 645 (2024). https://doi.org/10.1186/s12864-024-10559-w

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Design and research of a strain elastic element with a double-layer cross floating beam for strain gauge wireless rotating dynamometers.

1. Introduction

2. design and research of strain elastic element with double-layer cross floating beam, 2.1. structural design of strain gauge wireless rotating dynamometer, 2.2. structural design of the strain elastic element, 2.3. strain and deformation segmented rigid body model of double-layer cross floating beam, 2.3.1. strain and deformation analysis of double-layer cross floating beam under the action of f x, 2.3.2. strain and deformation analysis of double-layer cross floating beam under the action of f z, 2.4. comparison of fea solution and model solution for strain and deformation of double-layer cross floating beam, 2.5. size optimization of double-layer cross floating beam, 2.6. overall analysis of the strain elastic element, 3. strain gauge arrangement and testing, 3.1. strain gauge arrangement, 3.2. static calibration testing, 3.3. free modal testing, 3.4. cutting test, 4. conclusions.

• This paper designed a compact strain elastic element with a double-layer cross floating beam. The maximum outer diameter is only 75 mm and the maximum thickness is 25 mm. The design of the double-layer cross floating beam allows the strain elastic element to improve the sensitivity and reduce the cross-sensitivity error while ensuring the overall stiffness;
• Based on the proposed strain elastic element, a strain gauge wireless rotating dynamometer with compact size is designed. The overall diameter of the dynamometer is 170 mm and the axial size is 286 mm. In order to facilitate integration, a ring-shaped data acquisition and wireless transmission module PCB is designed;
• The static model of the double-layer cross floating beam on the strain elastic element was established by the segmented rigid body method. The rationality of the model was verified by comparison with the finite element results. According to the obtained static model, the structural parameters of the double-layer cross floating beam were optimized using the sequential quadratic programming algorithm to maximize the sensitivity of the floating beam;
• The strain elastic element was analyzed using finite element software, and the strain of the structure under simulation conditions was obtained, which provided a reference for subsequent calibration tests and circuit design;
• Through static calibration tests, the sensitivities of the strain elastic element in the four directions of F X , F Y , F Z , and M Z are determined to be 3.3 mV/N, 2.7 mV/N, 1.6 mV/N, and 104.1 mV/Nm, respectively, and the maximum cross-sensitivity error does not exceed 1%. Through modal tests in the free state, the natural frequency of the strain elastic element is determined to be 2954 Hz. The results of cutting tests show that the strain elastic element can obtain the change in cutting force and the tool-passing frequency. The results show that the designed strain elastic element can be applied to the strain gauge wireless rotating dynamometer to measure four-component cutting forces under medium- and low-speed conditions.

Author Contributions

Data availability statement, conflicts of interest.

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Wang, Q.; Wu, W.; Zhao, Y.; Cheng, Y.; Liu, L.; Yan, K. Design and Research of a Strain Elastic Element with a Double-Layer Cross Floating Beam for Strain Gauge Wireless Rotating Dynamometers. Micromachines 2024 , 15 , 857. https://doi.org/10.3390/mi15070857

Wang Q, Wu W, Zhao Y, Cheng Y, Liu L, Yan K. Design and Research of a Strain Elastic Element with a Double-Layer Cross Floating Beam for Strain Gauge Wireless Rotating Dynamometers. Micromachines . 2024; 15(7):857. https://doi.org/10.3390/mi15070857

Wang, Qinan, Wenge Wu, Yongjuan Zhao, Yunping Cheng, Lijuan Liu, and Kaiqiang Yan. 2024. "Design and Research of a Strain Elastic Element with a Double-Layer Cross Floating Beam for Strain Gauge Wireless Rotating Dynamometers" Micromachines 15, no. 7: 857. https://doi.org/10.3390/mi15070857

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