generic structure of a biography text is

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Kinds of Text, Biography Text: Definition, Purposes, Generic Structures, Language Features - mediainggris.com

Definition of biography text, the function of biography text (social function/purpose of biography text).

• To know a person’s story about his/her life outside of any accomplishments this person may be known for.
• To give many information easily and educate the readers.

Types Of Biography Text

1. autobiography, 2. biography, the structure of biography text, orientation (description), re-orientation (closing), biography text features / language features of biography text, use of simple past tense, temporal sequence and temporal conjunction, focus on specific person, use of action verbs., examples of biography text, short biography text about albert einsten, long biography text of jokowi, biography text of national heroes, biography text of cut nyak dhien, biography text of ir. soekarno, biography text exercise, handy 2022-05-15 tags: english , kinds of text.

How to Write a Biography

Biographies are big business. Whether in book form or Hollywood biopics, the lives of the famous and sometimes not-so-famous fascinate us.

While it’s true that most biographies are about people who are in the public eye, sometimes the subject is less well-known. Primarily, though, famous or not, the person who is written about has led an incredible life.

In this article, we will explain biography writing in detail for teachers and students so they can create their own.

While your students will most likely have a basic understanding of a biography, it’s worth taking a little time before they put pen to paper to tease out a crystal-clear definition of one.

What Is a Biography?

A biography is an account of someone’s life written by someone else . While there is a genre known as a fictional biography, for the most part, biographies are, by definition, nonfiction.

Generally speaking, biographies provide an account of the subject’s life from the earliest days of childhood to the present day or, if the subject is deceased, their death.

The job of a biography is more than just to outline the bare facts of a person’s life.

Rather than just listing the basic details of their upbringing, hobbies, education, work, relationships, and death, a well-written biography should also paint a picture of the subject’s personality and experience of life.

Full Biographies

Teaching unit.

Teach your students everything they need to know about writing an AUTOBIOGRAPHY and a BIOGRAPHY.

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Features of a Biography

Before students begin writing a biography, they’ll need to have a firm grasp of the main features of a Biography. An excellent way to determine how well they understand these essential elements is to ask them to compile a checklist like the one-blow

Their checklists should contain the items below at a minimum. Be sure to help them fill in any gaps before moving on to the writing process.

The purpose of a biography is to provide an account of someone’s life.

Biography structure.

ORIENTATION (BEGINNING) Open your biography with a strong hook to grab the reader’s attention

SEQUENCING: In most cases, biographies are written in chronological order unless you are a very competent writer consciously trying to break from this trend.

COVER: childhood, upbringing, education, influences, accomplishments, relationships, etc. – everything that helps the reader to understand the person.

CONCLUSION: Wrap your biography up with some details about what the subject is doing now if they are still alive. If they have passed away, make mention of what impact they have made and what their legacy is or will be.

BIOGRAPHY FEATURES

LANGUAGE Use descriptive and figurative language that will paint images inside your audience’s minds as they read. Use time connectives to link events.

PERSPECTIVE Biographies are written from the third person’s perspective.

DETAILS: Give specific details about people, places, events, times, dates, etc. Reflect on how events shaped the subject. You might want to include some relevant photographs with captions. A timeline may also be of use depending upon your subject and what you are trying to convey to your audience.

TENSE Written in the past tense (though ending may shift to the present/future tense)

THE PROCESS OF WRITING A BIOGRAPHY

Like any form of writing, you will find it simple if you have a plan and follow it through. These steps will ensure you cover the essential bases of writing a biography essay.

Firstly, select a subject that inspires you. Someone whose life story resonates with you and whose contribution to society intrigues you. The next step is to conduct thorough research. Engage in extensive reading, explore various sources, watch documentaries, and glean all available information to provide a comprehensive account of the person’s life.

Creating an outline is essential to organize your thoughts and information. The outline should include the person’s early life, education, career, achievements, and any other significant events or contributions. It serves as a map for the writing process, ensuring that all vital information is included.

Your biography should have an engaging introduction that captivates the reader’s attention and provides background information on the person you’re writing about. It should include a thesis statement summarising the biography’s main points.

Writing a biography in chronological order is crucial . You should begin with the person’s early life and move through their career and achievements. This approach clarifies how the person’s life unfolded and how they accomplished their goals.

A biography should be written in a narrative style , capturing the essence of the person’s life through vivid descriptions, anecdotes, and quotes. Avoid dry, factual writing and focus on creating a compelling narrative that engages the reader.

Adding personal insights and opinions can enhance the biography’s overall impact, providing a unique perspective on the person’s achievements, legacy, and impact on society.

Editing and proofreading are vital elements of the writing process. Thoroughly reviewing your biography ensures that the writing is clear, concise, and error-free. You can even request feedback from someone else to ensure that it is engaging and well-written.

Finally, including a bibliography at the end of your biography is essential. It gives credit to the sources that were used during research, such as books, articles, interviews, and websites.

Tips for Writing a Brilliant Biography

Biography writing tip #1: choose your subject wisely.

There are several points for students to reflect on when deciding on a subject for their biography. Let’s take a look at the most essential points to consider when deciding on the subject for a biography:

Interest: To produce a biography will require sustained writing from the student. That’s why students must choose their subject well. After all, a biography is an account of someone’s entire life to date. Students must ensure they choose a subject that will sustain their interest throughout the research, writing, and editing processes.

Merit: Closely related to the previous point, students must consider whether the subject merits the reader’s interest. Aside from pure labors of love, writing should be undertaken with the reader in mind. While producing a biography demands sustained writing from the author, it also demands sustained reading from the reader.

Therefore, students should ask themselves if their chosen subject has had a life worthy of the reader’s interest and the time they’d need to invest in reading their biography.

Information: Is there enough information available on the subject to fuel the writing of an entire biography? While it might be a tempting idea to write about a great-great-grandfather’s experience in the war. There would be enough interest there to sustain the author’s and the reader’s interest, but do you have enough access to information about their early childhood to do the subject justice in the form of a biography?

Biography Writing Tip #2: R esearch ! Research! Research!

While the chances are good that the student already knows quite a bit about the subject they’ve chosen. Chances are 100% that they’ll still need to undertake considerable research to write their biography.

As with many types of writing , research is an essential part of the planning process that shouldn’t be overlooked. If students wish to give as complete an account of their subject’s life as possible, they’ll need to put in the time at the research stage.

An effective way to approach the research process is to:

1. Compile a chronological timeline of the central facts, dates, and events of the subject’s life

2. Compile detailed descriptions of the following personal traits:

•      Physical looks
•      Character traits
•      Values and beliefs

3. Compile some research questions based on different topics to provide a focus for the research:

• Childhood : Where and when were they born? Who were their parents? Who were the other family members? What education did they receive?
• Obstacles: What challenges did they have to overcome? How did these challenges shape them as individuals?
• Legacy: What impact did this person have on the world and/or the people around them?
• Dialogue & Quotes: Dialogue and quotations by and about the subject are a great way to bring color and life to a biography. Students should keep an eagle eye out for the gems that hide amid their sources.

As the student gets deeper into their research, new questions will arise that can further fuel the research process and help to shape the direction the biography will ultimately go in.

Likewise, during the research, themes will often begin to suggest themselves. Exploring these themes is essential to bring depth to biography, but we’ll discuss this later in this article.

Research Skills:

Researching for biography writing is an excellent way for students to hone their research skills in general. Developing good research skills is essential for future academic success. Students will have opportunities to learn how to:

• Gather relevant information
• Evaluate different information sources
• Select suitable information
• Organize information into a text.

Students will have access to print and online information sources, and, in some cases, they may also have access to people who knew or know the subject (e.g. biography of a family member).

These days, much of the research will likely take place online. It’s crucial, therefore, to provide your students with guidance on how to use the internet safely and evaluate online sources for reliability. This is the era of ‘ fake news ’ and misinformation after all!

COMPLETE TEACHING UNIT ON INTERNET RESEARCH SKILLS USING GOOGLE SEARCH

Teach your students ESSENTIAL SKILLS OF THE INFORMATION ERA to become expert DIGITAL RESEARCHERS.

⭐How to correctly ask questions to search engines on all devices.

⭐ How to filter and refine your results to find exactly what you want every time.

⭐ Essential Research and critical thinking skills for students.

⭐ Plagiarism, Citing and acknowledging other people’s work.

⭐ How to query, synthesize and record your findings logically.

BIOGRAPHY WRITING Tip #3: Find Your Themes In Biography Writing

Though predominantly a nonfiction genre, the story still plays a significant role in good biography writing. The skills of characterization and plot structuring are transferable here. And, just like in fiction, exploring themes in a biographical work helps connect the personal to the universal. Of course, these shouldn’t be forced; this will make the work seem contrived, and the reader may lose faith in the truthfulness of the account. A biographer needs to gain and maintain the trust of the reader.

Fortunately, themes shouldn’t need to be forced. A life well-lived is full of meaning, and the themes the student writer is looking for will emerge effortlessly from the actions and events of the subject’s life. It’s just a case of learning how to spot them.

One way to identify the themes in a life is to look for recurring events or situations in a person’s life. These should be apparent from the research completed previously. The students should seek to identify these patterns that emerge in the subject’s life. For example, perhaps they’ve had to overcome various obstacles throughout different periods of their life. In that case, the theme of overcoming adversity is present and has been identified.

Usually, a biography has several themes running throughout, so be sure your students work to identify more than one theme in their subject’s life.

BIOGRAPHY WRITING Tip: #4 Put Something of Yourself into the Writing

While the defining feature of a biography is that it gives an account of a person’s life, students must understand that this is not all a biography does. Relating the facts and details of a subject’s life is not enough. The student biographer should not be afraid to share their thoughts and feelings with the reader throughout their account of their subject’s life.

The student can weave some of their personality into the fabric of the text by providing commentary and opinion as they relate the events of the person’s life and the wider social context at the time. Unlike the detached and objective approach we’d expect to find in a history textbook, in a biography, student-writers should communicate their enthusiasm for their subject in their writing.

This makes for a more intimate experience for the reader, as they get a sense of getting to know the author and the subject they are writing about.

Biography Examples For Students

• Year 5 Example
• Year 7 Example
• Year 9 Example

“The Rock ‘n’ Roll King: Elvis Presley”

Elvis Aaron Presley, born on January 8, 1935, was an amazing singer and actor known as the “King of Rock ‘n’ Roll.” Even though he’s been dead for nearly 50 years, I can’t help but be fascinated by his incredible life!

Elvis grew up in Tupelo, Mississippi, in a tiny house with his parents and twin brother. His family didn’t have much money, but they shared a love for music. Little did they know Elvis would become a music legend!

When he was only 11 years old, Elvis got his first guitar. He taught himself to play and loved singing gospel songs. As he got older, he started combining different music styles like country, blues, and gospel to create a whole new sound – that’s Rock ‘n’ Roll!

In 1954, at the age of 19, Elvis recorded his first song, “That’s All Right.” People couldn’t believe how unique and exciting his music was. His famous hip-swinging dance moves also made him a sensation!

Elvis didn’t just rock the music scene; he also starred in movies like “Love Me Tender” and “Jailhouse Rock.” But fame came with challenges. Despite facing ups and downs, Elvis kept spreading happiness through his music.

Tragically, Elvis passed away in 1977, but his music and charisma live on. Even today, people worldwide still enjoy his songs like “Hound Dog” and “Can’t Help Falling in Love.” Elvis Presley’s legacy as the King of Rock ‘n’ Roll will live forever.

Long Live the King: I wish I’d seen him.

Elvis Presley, the Rock ‘n’ Roll legend born on January 8, 1935, is a captivating figure that even a modern-day teen like me can’t help but admire. As I delve into his life, I wish I could have experienced the magic of his live performances.

Growing up in Tupelo, Mississippi, Elvis faced challenges but found solace in music. At 11, he got his first guitar, a symbol of his journey into the world of sound. His fusion of gospel, country, and blues into Rock ‘n’ Roll became a cultural phenomenon.

The thought of being in the audience during his early performances, especially when he recorded “That’s All Right” at 19, sends shivers down my spine. Imagining the crowd’s uproar and feeling the revolutionary energy of that moment is a dream I wish I could have lived.

Elvis wasn’t just a musical prodigy; he was a dynamic performer. His dance moves, the embodiment of rebellion, and his roles in films like “Love Me Tender” and “Jailhouse Rock” made him a true icon.

After watching him on YouTube, I can’t help but feel a little sad that I’ll never witness the King’s live performances. The idea of swaying to “Hound Dog” or being enchanted by “Can’t Help Falling in Love” in person is a missed opportunity. Elvis may have left us in 1977, but he was the king of rock n’ roll. Long live the King!

Elvis Presley: A Teen’s Take on the Rock ‘n’ Roll Icon”

Elvis Presley, born January 8, 1935, was a revolutionary force in the music world, earning his title as the “King of Rock ‘n’ Roll.” Exploring his life, even as a 16-year-old today, I’m captivated by the impact he made.

Hailing from Tupelo, Mississippi, Elvis grew up in humble beginnings, surrounded by the love of his parents and twin brother. It’s inspiring to think that, despite financial challenges, this young man would redefine the music scene.

At 11, Elvis got his first guitar, sparking a self-taught journey into music. His early gospel influences evolved into a unique fusion of country, blues, and gospel, creating the electrifying genre of Rock ‘n’ Roll. In 1954, at only 19, he recorded “That’s All Right,” marking the birth of a musical legend.

Elvis wasn’t just a musical innovator; he was a cultural phenomenon. His rebellious dance moves and magnetic stage presence challenged the norms. He transitioned seamlessly into acting, starring in iconic films like “Love Me Tender” and “Jailhouse Rock.”

However, fame came at a cost, and Elvis faced personal struggles. Despite the challenges, his music continued to resonate. Even now, classics like “Hound Dog” and “Can’t Help Falling in Love” transcend generations.

Elvis Presley’s impact on music and culture is undeniable. He was known for his unique voice, charismatic persona, and electrifying performances. He sold over one billion records worldwide, making him one of the best-selling solo artists in history. He received numerous awards throughout his career, including three Grammy Awards and the Grammy Lifetime Achievement Award.

Elvis’s influence can still be seen in today’s music. Many contemporary artists, such as Bruno Mars, Lady Gaga, and Justin Timberlake, have cited Elvis as an inspiration. His music continues to be featured in movies, TV shows, and commercials.

Elvis left us in 1977, but his legacy lives on. I appreciate his breaking barriers and fearlessly embracing his artistic vision. Elvis Presley’s impact on music and culture is timeless, a testament to the enduring power of his artistry. His music has inspired generations and will continue to do so for many years to come.

Teaching Resources

Use our resources and tools to improve your student’s writing skills through proven teaching strategies.

BIOGRAPHY WRITING TEACHING IDEAS AND LESSONS

We have compiled a sequence of biography-related lessons or teaching ideas that you can follow as you please. They are straightforward enough for most students to follow without further instruction.

BIOGRAPHY LESSON IDEA # 1:

This session aims to give students a broader understanding of what makes a good biography.

Once your students have compiled a comprehensive checklist of the main features of a biography, allow them to use it to assess some biographies from your school library or on the internet using the feature checklist.

When students have assessed a selection of biographies, take some time as a class to discuss them. You can base the discussion around the following prompts:

• Which biographies covered all the criteria from their checklist?
• Which biographies didn’t?
• Which biography was the most readable in terms of structure?
• Which biography do you think was the least well-structured? How would you improve this?

Looking at how other writers have interpreted the form will help students internalize the necessary criteria before attempting to produce a biography. Once students have a clear understanding of the main features of the biography, they’re ready to begin work on writing a biography.

When the time does come to put pen to paper, be sure they’re armed with the following top tips to help ensure they’re as well prepared as possible.

BIOGRAPHY LESSON IDEA # 2:

This session aims to guide students through the process of selecting the perfect biography subject.

Instruct students to draw up a shortlist of three potential subjects for the biography they’ll write.

Using the three criteria mentioned in the writing guide (Interest, Merit, and Information), students award each potential subject a mark out of 5 for each of the criteria. In this manner, students can select the most suitable subject for their biography.

BIOGRAPHY LESSON IDEA # 3:

This session aims to get students into the researching phase, then prioritise and organise events chronologically.

Students begin by making a timeline of their subject’s life, starting with their birth and ending with their death or the present day. If the student has yet to make a final decision on the subject of their biography, a family member will often serve well for this exercise as a practice exercise.

Students should research and gather the key events of the person’s life, covering each period of their life from when they were a baby, through childhood and adolescence, right up to adulthood and old age. They should then organize these onto a timeline. Students can include photographs with captions if they have them.

They can present these to the class when they have finished their timelines.

BIOGRAPHY LESSON IDEA # 4:

Instruct students to look over their timeline, notes, and other research. Challenge them to identify three patterns that repeat throughout the subject’s life and sort all the related events and incidents into specific categories.

Students should then label each category with a single word. This is the thematic concept or the broad general underlying idea. After that, students should write a sentence or two expressing what the subject’s life ‘says’ about that concept.

This is known as the thematic statement . With the thematic concepts and thematic statements identified, the student now has some substantial ideas to explore that will help bring more profound meaning and wider resonance to their biography.

BIOGRAPHY LESSON IDEA # 5:

Instruct students to write a short objective account of an event in their own life. They can write about anyone from their past. It needn’t be more than a couple of paragraphs, but the writing should be strictly factual, focusing only on the objective details of what happened.

Once they have completed this, it’s time to rewrite the paragraph, but they should include some opinion and personal commentary this time.

The student here aims to inject some color and personality into their writing, to transform a detached, factual account into a warm, engaging story.

A COMPLETE UNIT ON TEACHING BIOGRAPHIES

Teach your students to write AMAZING BIOGRAPHIES & AUTOBIOGRAPHIES using proven RESEARCH SKILLS and WRITING STRATEGIES .

• Understand the purpose of both forms of biography.
• Explore the language and perspective of both.
• Prompts and Challenges to engage students in writing a biography.
• Dedicated lessons for both forms of biography.
• Biographical Projects can expand students’ understanding of reading and writing a biography.
• A COMPLETE 82-PAGE UNIT – NO PREPARATION REQUIRED.

FREE Biography Writing Graphic Organizer

Use this valuable tool in the research and writing phases to keep your students on track and engaged.

WRITING CHECKLIST & RUBRIC BUNDLE

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To Conclude

By this stage, your students should have an excellent technical overview of a biography’s essential elements.

They should be able to choose their subject in light of how interesting and worthy they are, as well as give consideration to the availability of information out there. They should be able to research effectively and identify emerging themes in their research notes. And finally, they should be able to bring some of their personality and uniqueness into their retelling of the life of another.

Remember that writing a biography is not only a great way to develop a student’s writing skills; it can be used in almost all curriculum areas. For example, to find out more about a historical figure in History, to investigate scientific contributions to Science, or to celebrate a hero from everyday life.

Biography is an excellent genre for students to develop their writing skills and to find inspiration in the lives of others in the world around them.

HOW TO WRITE A BIOGRAPHY TUTORIAL VIDEO

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Personal Narrative Writing Guide

Definition of Biography

A biography is the non- fiction , written history or account of a person’s life. Biographies are intended to give an objective portrayal of a person, written in the third person. Biographers collect information from the subject (if he/she is available), acquaintances of the subject, or in researching other sources such as reference material, experts, records, diaries, interviews, etc. Most biographers intend to present the life story of a person and establish the context of their story for the reader, whether in terms of history and/or the present day. In turn, the reader can be reasonably assured that the information presented about the biographical subject is as true and authentic as possible.

Biographies can be written about a person at any time, no matter if they are living or dead. However, there are limitations to biography as a literary device. Even if the subject is involved in the biographical process, the biographer is restricted in terms of access to the subject’s thoughts or feelings.

Biographical works typically include details of significant events that shape the life of the subject as well as information about their childhood, education, career, and relationships. Occasionally, a biography is made into another form of art such as a film or dramatic production. The musical production of “Hamilton” is an excellent example of a biographical work that has been turned into one of the most popular musical productions in Broadway history.

Common Examples of Biographical Subjects

Most people assume that the subject of a biography must be a person who is famous in some way. However, that’s not always the case. In general, biographical subjects tend to be interesting people who have pioneered something in their field of expertise or done something extraordinary for humanity. In addition, biographical subjects can be people who have experienced something unusual or heartbreaking, committed terrible acts, or who are especially gifted and/or talented.

As a literary device, biography is important because it allows readers to learn about someone’s story and history. This can be enlightening, inspiring, and meaningful in creating connections. Here are some common examples of biographical subjects:

• political leaders
• entrepreneurs
• historical figures
• serial killers
• notorious people
• political activists
• adventurers/explorers
• religious leaders
• military leaders
• cultural figures

Famous Examples of Biographical Works

The readership for biography tends to be those who enjoy learning about a certain person’s life or overall field related to the person. In addition, some readers enjoy the literary form of biography independent of the subject. Some biographical works become well-known due to either the person’s story or the way the work is written, gaining a readership of people who may not otherwise choose to read biography or are unfamiliar with its form.

Here are some famous examples of biographical works that are familiar to many readers outside of biography fans:

• Alexander Hamilton (Ron Chernow)
• Prairie Fires: The American Dreams of Laura Ingalls Wilder (Caroline Fraser)
• Steve Jobs (Walter Isaacson)
• Churchill: A Life (Martin Gilbert)
• The Professor and the Madman: A Tale of Murder, Insanity, and the Making of the Oxford English Dictionary (Simon Winchester)
• A Beautiful Mind (Sylvia Nasar)
• The Black Rose (Tananarive Due)
• John Adams (David McCullough)
• Into the Wild ( Jon Krakauer )
• John Brown (W.E.B. Du Bois)
• Frida: A Biography of Frida Kahlo (Hayden Herrera)
• The Immortal Life of Henrietta Lacks (Rebecca Skloot)
• Team of Rivals: The Political Genius of Abraham Lincoln (Doris Kearns Goodwin)
• Shirley Jackson : A Rather Haunted Life ( Ruth Franklin)
• the stranger in the Woods: The Extraordinary Story of the Last True Hermit (Michael Finkel)

Difference Between Biography, Autobiography, and Memoir

Biography, autobiography , and memoir are the three main forms used to tell the story of a person’s life. Though there are similarities between these forms, they have distinct differences in terms of the writing, style , and purpose.

A biography is an informational narrative and account of the life history of an individual person, written by someone who is not the subject of the biography. An autobiography is the story of an individual’s life, written by that individual. In general, an autobiography is presented chronologically with a focus on key events in the person’s life. Since the writer is the subject of an autobiography, it’s written in the first person and considered more subjective than objective, like a biography. In addition, autobiographies are often written late in the person’s life to present their life experiences, challenges, achievements, viewpoints, etc., across time.

Memoir refers to a written collection of a person’s significant memories, written by that person. Memoir doesn’t generally include biographical information or chronological events unless it’s relevant to the story being presented. The purpose of memoir is reflection and an intention to share a meaningful story as a means of creating an emotional connection with the reader. Memoirs are often presented in a narrative style that is both entertaining and thought-provoking.

Examples of Biography in Literature

An important subset of biography is literary biography. A literary biography applies biographical study and form to the lives of artists and writers. This poses some complications for writers of literary biographies in that they must balance the representation of the biographical subject, the artist or writer, as well as aspects of the subject’s literary works. This balance can be difficult to achieve in terms of judicious interpretation of biographical elements within an author’s literary work and consideration of the separate spheres of the artist and their art.

Literary biographies of artists and writers are among some of the most interesting biographical works. These biographies can also be very influential for readers, not only in terms of understanding the artist or writer’s personal story but the context of their work or literature as well. Here are some examples of well-known literary biographies:

Example 1:  Savage Beauty: The Life of Edna St. Vincent Millay  (Nancy Milford)

One of the first things Vincent explained to Norma was that there was a certain freedom of language in the Village that mustn’t shock her. It wasn’t vulgar. ‘So we sat darning socks on Waverly Place and practiced the use of profanity as we stitched. Needle in, . Needle out, piss. Needle in, . Needle out, c. Until we were easy with the words.’

This passage reflects the way in which Milford is able to characterize St. Vincent Millay as a person interacting with her sister. Even avid readers of a writer’s work are often unaware of the artist’s private and personal natures, separate from their literature and art. Milford reflects the balance required on the part of a literary biographer of telling the writer’s life story without undermining or interfering with the meaning and understanding of the literature produced by the writer. Though biographical information can provide some influence and context for a writer’s literary subjects, style, and choices , there is a distinction between the fictional world created by a writer and the writer’s “real” world. However, a literary biographer can illuminate the writer’s story so that the reader of both the biography and the biographical subject’s literature finds greater meaning and significance.

Example 2:  The Invisible Woman: The Story of Nelly Ternan and Charles Dickens  (Claire Tomalin)

The season of domestic goodwill and festivity must have posed a problem to all good Victorian family men with more than one family to take care of, particularly when there were two lots of children to receive the demonstrations of paternal love.

Tomalin’s literary biography of Charles Dickens reveals the writer’s extramarital relationship with a woman named Nelly Ternan. Tomalin presents the complications that resulted for Dickens from this relationship in terms of his personal and family life as well as his professional writing and literary work. Revealing information such as an extramarital relationship can influence the way a reader may feel about the subject as a person, and in the case of literary biography it can influence the way readers feel about the subject’s literature as well. Artists and writers who are beloved , such as Charles Dickens, are often idealized by their devoted readers and society itself. However, as Tomalin’s biography of Dickens indicates, artists and writers are complicated and as subject to human failings as anyone else.

Example 3:  Virginia Woolf  (Hermione Lee)

‘A self that goes on changing is a self that goes on living’: so too with the biography of that self. And just as lives don’t stay still, so life-writing can’t be fixed and finalised. Our ideas are shifting about what can be said, our knowledge of human character is changing. The biographer has to pioneer, going ‘ahead of the rest of us, like the miner’s canary, testing the atmosphere , detecting falsity, unreality, and the presence of obsolete conventions’. So, ‘There are some stories which have to be retold by each generation’. She is talking about the story of Shelley, but she could be talking about her own life-story.

In this passage, Lee is able to demonstrate what her biographical subject, Virginia Woolf, felt about biography and a person telling their own or another person’s story. Literary biographies of well-known writers can be especially difficult to navigate in that both the author and biographical subject are writers, but completely separate and different people. As referenced in this passage by Lee, Woolf was aware of the subtleties and fluidity present in a person’s life which can be difficult to judiciously and effectively relay to a reader on the part of a biographer. In addition, Woolf offers insight into the fact that biographers must make choices in terms of what information is presented to the reader and the context in which it is offered, making them a “miner’s canary” as to how history will view and remember the biographical subject.

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How to Write a Biography (Examples & Templates)

A biography is a written account of a person’s life that details their life in chronological order. Another person usually writes this detailed account, and it contains reports of their childhood, career, major life events, relationships, and social impact. It also details their relationships with their family, children, and life accomplishments.

The best way to find out more about a popular figure is through reading their biographies, so you need to make sure you get the correct information. Before writing a biography, you need to do a lot of research and interviews to represent a person’s life accurately.

Types of Biography

A biography is the story of someone’s life as written by another writer. Most biographies of popular figures are written years, or even decades, after their deaths. Authors write biographies of popular figures due to either a lack of information on the subject or personal interest.

A biography aims to share a person’s story or highlight a part of their life.

There are different types of biographies, depending on the story. Some biographies are written true to the story, while some are written as fictional works. Biographies can give you true understanding of a person on an internal as well as external level along with a lot of life lessons.

Autobiography

An autobiography is different from a biography because it is written by the subject of the story, themselves. The author writes in the first-person narrative, and it flows step-by-step like a story of their life. Autobiographies contain personal accounts of the subject’s life, along with their perspectives and opinions on events in their life.

How To Write a Biography

Pick a subject.

Picking a subject is the first step in writing a biography. You can pick an already famous person or a relatively unknown person with a great life story. If you already have a few in mind, you can start by asking yourself some questions such as;

• What has the subject accomplished that makes them a good subject?
• Have they had an impact on society?
• Is the subject a celebrity or a well-known personality?
• Will the biography appeal to a wide audience?

Get Permission

When you pick a subject, the next thing to do is to get permission from them or their family or rights owners. Although, with some historical figures, there may not be any need for permission. Getting permission from your subject makes it easier for you to get stories to put into your book. You can get the chance to obtain additional personal stories and anecdotes that will make your book more interesting by doing so as well.

Do The Research

Research is the most important part of a biography’s process as the entire content of the book is dependent on it. Irrespective of what you know about the subject, you need to carry out as much research as possible to get the story’s facts precisely.

Biography research comes from various sources, depending on the book’s subject. Firsthand reports from family, friends, or personal accounts from the subjects are primary sources. They are usually the most accurate and reliable, and they are crucial for a biography. Secondary sources come from other sources like magazines or documentaries.

Pick a Format

Biographies come in various formats, with each of them having their pros and cons. A typical biography will start at the beginning, usually with the birth and childhood of the subject. Yet, if the biography’s theme involves a different event in their life, the author may want to explore the flashback option or one with concurrent events from different times.

Usually, biographies have a theme or a general life lesson at the center. The author’s role is to tell the subject’s story leading up to the major event.

Which-ever format you choose should place the theme at the center, with the other events detailing the journey.

Create a Timeline Of The Story

Since a biography takes place in chronological order, there needs to be a timeline of the events in the right order. The timeline should contain the key events in the subject’s life, in the order the author plans on revealing them. A great way to declutter the story and keep it interesting is to use flashbacks . This way, the author can introduce past events and explain later events excluding the element of monotony.

Add In Your Thoughts

The good thing about biographies is that you don’t have to stick to the hard facts only. As the author, you can share your opinions and emotions in writing. The author has the freedom to do this by commenting on a significant action by the subject in a manner that describes why they feel the subject may have done what they did.

The author can also include commentary on events depicted in the biography – how it was influenced society or its impact on the lives around them. Recounting these events through a different perspective can make the biography more relatable and interesting to read.

FAQ’s

Why is a biography template important.

A biography template has an outline that makes the writing easier for the author. Biography templates usually contain a sample timeline, format, and questions that provide more information about the subject. With a great biography template, you can cut your writing time in half and spend less time coming up with an outline.

How are biographies better in comparison to autobiographies

Since a different person writes biographies, they tend to be more objective and somewhat accurate than autobiographies. An autobiography tells things from the author’s perspective, so their views and perspective cloud it. Thus, a biography will likely tell a more factual story.

These are the important steps you need to take to help you write a great biography. Now, to make things easier for you, we have a free customizable autobiography and biography template that you can use to start your first book. Get the template and start writing today

What are some of the most important elements to keep in consideration while writing a biography?

Any author looking to write a biography must consider the factors below. They aren’t the only important factors, but a biography isn’t complete without them. • Date and place of their birth • Academic background • Professional expertise • Death, if deceased • Facts and anecdotes about the person • Main accomplishments • Detailed accounts of their child and adult life

Biographies tell the untold stories of some incredibly relevant people in the world. But biographies are not always strictly accurate. So, every biographer needs to follow the necessary steps to provide a biography with all the requirements.

Related Documents

Module 3: Writing Process

Text: text structures.

A text structure is the framework of a text’s beginning, middle, and end. Different narrative and expository genres have different purposes and different audiences, and so they require different text structures. Beginnings and endings help link the text into a coherent whole.

BEGINNINGS: HOOKING YOUR READER

WHAT’S IN THE MIDDLE?

The organization of the middle of a piece of writing depends on the genre. Researchers have identified five basic organizational structures: sequence ,  description , cause and effect , compare and contrast , and problem and solution .

Sequence uses time, numerical, or spatial order as the organizing structure. Some narrative genres that use a chronological sequence structure are personal narrative genres (memoir, autobiographical incident, autobiography), imaginative story genres (fairytales, folktales, fantasy, science fiction), and realistic fiction genres. Narrative story structures include an initiating event, complicating actions that build to a high point, and a resolution. Many narratives also include the protagonist’s goals and obstacles that must be overcome to achieve those goals.

Description is used to describe the characteristic features and events of a specific subject (”My Cat”) or a general category (”Cats”). Descriptive reports may be arranged according to categories of related attributes, moving from general categories of features to specific attributes.

Cause and Effect structure is used to show causal relationships between events. Essays demonstrate cause and effect by giving reasons to support relationships, using the word “because.” Signal words for cause and effect structures also include if/then statements, “as a result,” and “therefore.”

Comparison and Contrast structure is used to explain how two or more objects, events, or positions in an argument are similar or different. Graphic organizers such as venn diagrams, compare/contrast organizers, and tables can be used to compare features across different categories. Words used to signal comparison and contrast organizational structures include “same,” “alike,” “in contrast,” “similarities,” “differences,” and “on the other hand.”

Problem and Solution requires writers to state a problem and come up with a solution. Although problem/solution structures are typically found in informational writing, realistic fiction also often uses a problem/solution structure.

ENDINGS: BEYOND “HAPPILY EVER AFTER”

Anyone who has watched a great movie for ninety minutes only to have it limp to the finish with weak ending knows that strong endings are just as critical to effective writing as strong beginnings. And anyone who has watched the director’s cut of a movie with all the alternate endings knows that even great directors have trouble coming up with satisfying endings for their movies. Just like directors, writers have to decide how to wrap up the action in their stories, resolving the conflict and tying up loose ends in a way that will leave their audience satisfied. 

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All About the Structure and Sample of A Biography And How To Write It

A biography is a non-fiction work about a specific person’s life. Usually written in the third person point of view, all examples of biography you will find are meant to give you an objective understanding and an authentic characterization. It is written following a person’s narration of it, their friends or family members’ accounts, or researching materials like recordings, journals, diaries, old interviews, experts, and the likes. It is not uncommon to find a sample of a biography of a deceased person. All kinds of biography samples can be found online or in your local library about both the heroes of old and modern heroes of today.

Most biography samples that you would find are about famous people. It’s not surprising that people would write and read about politicians, historical figures, artists, authors, inventors, all kinds of leaders, and even notorious people and criminal masterminds. It’s always interesting how an author of a biography example plots their narrative. Look for different examples of biography. You can note how the writers of it help you imagine what it was like to be there, at the exact moment that someone famous or good is doing something that is only known to us through history books. However, as you write your own, remember that, unlike fiction writing or other kinds, it is very rare to find a sample of a biography that would tell you how the subject is or was feeling. All biography samples that you would read would tell you the subject’s each step on their whole life journey but not how they were feeling during those times or not necessarily or accurately.

There are many biography samples that you can find and read about. But in recent years, some biographies have been turned into another form of art, like a movie or a play. Some examples of biography that has been turned into a movie that you might recognize include:

• The Theory of Everything (Stephen Hawking)
• Wolf of Wall Street (Jordan Belfort)
• Ford vs Ferrari (Carroll Shelby and Ken Miles)
• Schindler’s List (Oskar Schindler)
• Bohemian Rhapsody (Rock band Queen)
• Molly’s Game (Molly Bloom)
• Godfellas (Henry Hill)
• Braveheart (William Wallace)
• Bonnie and Clyde (Bonnie Parker and Clyde Barrow)
• On the Basis of Sex (Ruth Bader Ginsberg)

Interesting enough, some biographies are even used as the basis for some musical plays. For example, the 2004 biography written by Ron Chernow, Alexander Hamilton, is a good sample of an amazingly transformed biography into a musical play (and even a movie) by Lin-Manuel Miranda.

Tips And Tricks on How To Write A Biography

1. a biography is not a resume.

Going into the task of writing a biography is serious business. This is why before even starting to write a biography, you should already have it stuck in your head that you are not writing a resume . This is a tendency for most writers because examples of biography typically include the same information as your resume. Only a sample of a biography would have to be more detailed. Nevertheless, do not make the mistake of writing a biography as if you were writing a detailed resume because it’s not. The whole idea of a biography is to narrate the life story of a person. This goes even further than their accomplishments and their educational background.

2. You Are Writing About a Real Person

Another thing to keep in mind is that a biography is about a real person. This narrative is not something that is made up or imagined. Any sample of a biography that you would find tells the story of a person that lived on the same earth you are now. Be careful not to alienate the person you are writing about. That is, do not be afraid to portray their human side and have them be relatable. As far as examples of biography go, the more readers find themselves relating to the person in the narrative, the better their chances of reading. You have to remember that your goal as a biography writer is to present the truth and authenticity of your subject. In all its bits and pieces, do not be afraid to put in the details about the birth, the childhood, the teenage years, all the way to adulthood, and maybe, even the death of your subject.

3. Find Your Era or Not

Examples of Biography that you would find typically start at the subject’s birth and end either at their death or the peak of their careers. And this is something that you can do. You can start on the day they were born and narrate how they spent their time when they were children, their experiences and experiments as teenagers and young adults, and their careers as adults. This is a sample of a biography that can go long enough to be a novel or a series of books. However, there is no harm in entertaining the idea of focusing and writing about only a specific period of a person’s life. You can find examples of biography that skip the childhood and teenage part of the person’s life and goes straight to their career as an adult. Some do the opposite and focus on the person’s early years to establish what honed their subject to the person they are known to be. Other examples of biography just focus on the few recent years of a person’s life and career , while certain writers tend to focus on the peak of the life and career of their subject. Regardless of what you decide to do, just be mindful not to lose sight of the reason and purpose of why you’re writing.

4. Research, Interview, Research

It might be a tad easier to write a biography about yourself or someone you know. This would save you a lot of time and trouble finding out details to include in your biography. However, most examples of biography are those of famous personalities’ lives and careers. So unless you are close friends with the celebrity or prominent figure you want to write about, you have to go through a lot of research. More or less, any sample of a biography that you would find and read contains the tiniest and most specific details about their subject. This was produced from the sweat and tears of hours, and even days, worth going through books, journals, newspapers, and any other records that the writer can find. Don’t be afraid to go above and beyond what you can find online or in your local libraries. Think creatively, try to find sources that are hidden in the cracks of the field of knowledge. Formulate a questionnaire and try to interview your subject, if possible, or find people connected to them that you can interview. Try to dive into public records, old school records, and any other reference material you can find. We’re telling you now that writing a biography can prove to be extremely tiresome and time-consuming. But we can also assure you that it is incredibly satisfying once you finish it.

5. Write, Edit, Write

You might be one of those writers who prefer to write as they gather information or maybe the ones that collect all the information they can find first and write after. Regardless of your methods, one thing is constant - your writing journey will not be a short one, especially if you are writing a whole life story. It is important to remember that you have to be accurate in your accounts and that authenticity plays a vital role in writing a biography. Your narrative has to be as detailed as you get it to be. It is also helpful to know early on that there is a point in the writing process of any sample of a biography you have read that you have to edit your writing. You have to rephrase, rewrite or completely revise the pieces that you have already written. It can be a good idea to outline how you want your narrative to go but don’t be afraid to explore the idea of reshaping or maybe even overhauling the biography you are writing about if you suddenly think of a better idea in the middle of your writing journey.

Share Your Written Biography Online

It might not be a bad idea to write a biography about a specific person in today’s modern world and post it online. You can help put out valuable and relevant information to your readers and maybe even make a name for yourself as a writer. Strikingly is a website builder that can help you create a home for your sample of a biography. You have two options:

• Showcase Your Biography Using A Simple Blog

You can use the Simple Blog feature of Strikingly to post chapters or parts of the biography you are writing. Posting chapters or parts of the biography adds uniqueness to entice your readers as not many people post examples of biography through blogspot websites . Here’s how you can use Simple Blog for your sample of a biography:

• Sign up or Log in with Strikingly.

Image taken from Strikingly Product

• Open your site editor and click “Add New Section”.
• Select the layout you want your Simple Blog to be.

• Click “Manage Blog Posts”.

• Click “Write a New Post” or “Edit”.

• Write your blog post.

• Let Your Biography be the Main Content of Your Website

If you are not very keen on the idea of posting your biography as blog posts, you can also just display your writing as plain text for your website. Here’s how Strikingly can help you write and present your sample of a biography on your website:

• Open your site editor and click “ Add New Section ”.

• Select “Plain Text”.

• Add a Title and write your own sample of a biography.

Writing your own sample of a biography is no easy task, but accomplishing it is a big feat you can be proud of. With Strikingly, you not only have an avenue to place yourself and your writing, but you also get to share the prowess of storytelling you have created with your own time, your efforts, and your magnificent mind. Sign up with Strikingly now and get started on adding to the collection of biography examples that the next generations would be reading about!

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Definition, Generic Structure, & Linguistics Features of Descriptive Text

• October 11, 2023
• Writing Corner

Descriptive texts are like word paintings. They use words to show us things, people, or places in our minds. When we read them, it is as if we are looking at a picture. In this article, we are going to learn what descriptive texts are, how they’re structured, and what makes them interesting to read. Let us start our journey into the world of descriptive texts!

Definition of Descriptive Text 

According to Anderson and Anderson (2003, p.26), the descriptive text describes a particular person, place, or thing. It means that descriptive text is designed especially about a person, a place, or things. They also stated descriptive text to tell about the subject by describing its features without including personal opinions.

In line with this statement, Gerot  &  Wignell (1994) and Knapp  &  Watkins  (2005) said that descriptive text is a type of text the writer or speaker uses to describe a particular thing, person, animal, place, and or event to the readers or hearers. The process of describing is done by ordering their characteristics clearly, starting from naming them, classifying them, and dealing with their attributes, behaviors, functions, and so on so that the readers or hearers can notice what the writer is writing about as if they could directly see it through their own eyes.

In addition, according to Pardiyono (2007, p.33), descriptive text gives descriptions of living or non-living things to the reader. Thus, this kind of text can tell an object that is still alive or not.

Generic Structure of Descriptive Text

Like other genres, descriptive text also has its structure or stages. Jenny Hammond (1992) divides generic descriptive text structures into two parts: Identification and Description.

• Identification: it is aimed at introducing and identifying a specific participant, such as a person, a  thing,  a  place,  an animal,  and or an event. 
• Description: it describes that  participant from  its  characteristics, appearances, personality, and habits or qualities (Butt, Fahey, Feez,  Spinks,  &  Yallop,  2000;  Derewianka, 1990;  Gerot  & Wignell,  1994; Knapp  &  Watkins, 2005)

Linguistics Features of Descriptive Text

Regarding  its  linguistics  features,  Derewianka  (1990), Emilia  and  Christie (2013), Gerot and Wignell (1994), Knapp and Watkins (2005) state that  descriptive text employs some linguistics features as summarized below:

• Focus on specific participants as the main character (My English teacher, Andini’s cat, My favorite place).
• Use present tense as dominant tenses.
• Use linking verbs or relational processes frequently (is, are, has, have, belongs to) to classify and describe the participant’s appearance or qualities and parts or functions.
• Use of descriptive adjectives (strong legs, white hair)

Example of Descriptive Text

My Mother Clean Kitchen

• Butt, D., Fahey, R., Feez, S. S. S., & Yallop, C. (2000). Using Functional Grammar: An Explorer’s Guide 2 nd Edition.  National Centre for English Language Teaching and Research: Macquarie University, Sydney .
• Derewianka,  B.  (1990). Exploring How Text Work.   Australia:  Primary English Teaching Association.
• Emilia, E., & Christie, F. (2013). Factual Genres in English: Learning to Write, Read, and Talk about Factual Information. Bandung: Rizqi Press.
• Gerot, L., & Wignell, P. (1994).  Making Sense of Functional Grammar: An Introductory Workbook . Queensland: Antipodean Educational Enterprises.
• Knapp, P., & Watkins, M. (2005).  Genre, Text, Grammar: Technologies for Teaching and Assessing Writing . Unsw Press.
• https://www.shutterstock.com/cs/image-photo/interior-modern-sunny-kitchen-scandinavianstyle-apartment-1464758027

Tenry Colle

Hi! My name is A. Tenry Lawangen Aspat Colle. I am a motivated and resourceful English educator. In addition, as the owner of @rymari.translation17 has shaped me to be a punctual and dependable translator of Indonesian to English and vice versa.

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4th grade reading & vocabulary

Course: 4th grade reading & vocabulary   >   unit 1.

• Summarizing nonfiction | Reading
• What is a main idea? | Reading
• What language shows cause and effect? | Reading

The structures of informational texts | Reading

• Superheroes: reading informational text; Superheroes around the World 4

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Video transcript

BRITISH Course

Space to find reference of english, explanation text; definition, generic structures, purposes, language features.

December 12, 2017 British Course Explanation of Text Genre 9

Assalamualaikum Warahmatullahi Wabarakatuh😊

Bismillahirrahmanirrahim

Hallo everybody BRITISH Course – I’m sure my friends have been familiar with the word “explanation”. Because in the world of education and even in daily life we usually meet explanation, maybe from our teachers, our parents, even our friends, right? For example, do you ever explain something to your friends? When your friend asked, how is the process of rain, or how is the process of human creation? etc. Then you explained to your friends about it. Those explanations are examples of Explanation Text.

Well, for my friends who want to learn about explanation text, my friends have come in the right place because in this occasion I will try to present explanations and examples of Explanation Text in detail and complete. I hope my explanation about Explanation Text below can be useful for all of the readers. Amen

Definition of Explanation Text

Explanation is a text which tells processes relating to forming of natural, social, scientific and cultural phenomena. Explanation text is to say ‘why’ and ‘how’ of the forming of the phenomena. It is often found in science, geography and history text books.

Generic Structure of Explanation Text

– General statement General statement; stating the phenomenon issues which are to be explained. – Sequenced of explanation Sequenced explanation; stating a series of steps which explain the phenomena.

Purpose of Explanation Text

– Explanation is a text which tells processes relating to forming of natural, social, scientific, and cultural phenomena. – To explain how or why something happens.

According to Mark Anderson and Kathy Anderson (1997: 82) says that the explanation text type is often used to tell how and why thing (phenomena) occur in nature.

Language Features of Explanation Text

In an explanation text, there are linguistic features as below : – Using simple present tense – Using abstract noun (no visible noun) – Using Passive voice – Using Action verbs – Containing explanation of the process ..

Examples of Explanation Text

Example of explanation text about natural phenomenon, how does rain happen.

Rain is the primary source of fresh water for most areas of the world, providing suitable conditions for diverse ecosystems, as well as water for hydroelectric power plants and crop irrigation.

The phenomenon of rain is actually a water circle. The concept of the water cycle involves the sun heating the Earth’s surface water and causing the surface water to evaporate. The water vapor rises into the Earth’s atmosphere. The water in the atmosphere cools and condenses into liquid droplets. The droplets grow until they are heavy and fall to the earth as precipitation which can be in the form of rain or snow.

However, not all rain reaches the surface. Some evaporates while falling through dry air. This is called virga, a phenomenon which is often seen in hot, dry desert regions.

Example of Explanation Text about Process of Making Chocolate

How chocolate is made.

Have we wondered how we get chocolate from? Well this time we will enter the amazing world of chocolate so we can understand exactly we are eating.

Chocolate starts a tree called cacao tree. This tree grows in equatorial regions, especially in place such as South America, Africa, and Indonesia. The cacao tree produces a fruit about the size of a small pine apple. In side the fruits are the tree’s seeds. They are also known as coco beans.

Next, the beans are fermented for about a week, dried in the sun. After that they are shipped to the chocolate maker. The chocolate maker starts processing by roasting the beans to bring out the flavour. Different beans from different places have different qualities and flavour. So they are often shorted and blended to produce a distinctive mix.

The next process is winnowing. The roasted beans are winnowed to remove the meat nib of the cacao bean from its shell. Then the nibs are blended. The blended nibs are grounded to make it liquid. The liquid is called chocolate liquor. It tastes bitter.

All seeds contain some amount of fat and cacao beans are not different. However, cacao beans are half fat, which is why they ground nibs from liquid. It is pure bitter chocolate.

Example of Explanation Text – How a Cancer is Formed

How a cancer is formed.

What is cancer? It is actually a group of more than one hundred separate diseases. Most of us are fear from cancer. It is reasonable because next to heart disease, cancer is the second leading cause of death.

Cancer cells come from normal cells because of mutations of DNA. Those mutations can occur spontaneously. The mutations may be also induced by other factors such as: nuclear and electromagnetic radiation, viruses, bacteria and fungi, parasites, heat, chemicals in the air, water and food, mechanical cell-level injury, free radicals, evolution and ageing of DNA, etc. All such factors can produce mutations that may start cancer.

Cancer cells are formed continuously in the organism. It is estimated that there are about 10,000 cancer cells at any given time in a healthy person. Why do some result in macroscopic-level cancers and some do not? First, not all damaged cells can multiply and many of them die quickly. Second, those which potentially divide and form cancer are effectively destroyed by the mechanisms available to the immune system. Therefore cancer develops if the immune system is not working properly or the amount of cells produced is too great for the immune system to eliminate.

Bagaimana Coklat Terbentuk

Sudahkah kita bertanya-tanya bagaimana kita mendapatkan cokelat? Nah kali ini kita akan memasuki dunia coklat yang menakjubkan sehingga kita bisa mengerti persis yang kita makan.

Coklat memulai pohon yang disebut pohon kakao. Pohon ini tumbuh di daerah khatulistiwa, terutama di tempat seperti Amerika Selatan, Afrika, dan Indonesia. Pohon kakao menghasilkan buah seukuran apel pinus kecil. Di sisi buah adalah biji pohon. Mereka juga dikenal sebagai coco beans.

Selanjutnya, biji difermentasi selama sekitar satu minggu, dikeringkan di bawah sinar matahari. Setelah itu mereka dikirim ke pembuat cokelat. Pembuat cokelat mulai memproses dengan memanggang biji coklat untuk mengeluarkan rasa. Biji yang berbeda dari tempat yang berbeda memiliki kualitas dan rasa yang berbeda. Jadi mereka sering disortir dan dicampur untuk menghasilkan campuran yang khas.

Proses selanjutnya adalah menampi. Biji coklat yang dipanggang diminyaki untuk mengeluarkan nib daging dari biji kakao dari cangkangnya. Kemudian nibs dicampur. Nibs yang dicampur digiling agar cair. Cairan itu disebut minuman cairan coklat. Rasanya pahit.

Semua biji mengandung sejumlah besar biji kakao dan lemak tidak berbeda. Namun, biji kakao setengah lemak, itulah sebabnya mereka menggiling nib dari cairan. Cokelat itu murni pahit.

Note on the Generic Structure of Explanation Sample

Every genre has its special purpose or social function. However if we see the generic structure point, we will get the understanding which both the explanation and procedure text have similar purposes. Both explain how to make or form something. However the procedure text will explain how to form or make something completely by instruction way. That is why most of procedure text is composed in command sentences. In the other hand, explanation text will show a knowledge about how thing is formed.

The above example of explanation text has the following generic structure :

General statement; it is a statement which says about chocolate and how it is formed

Sequenced explanation; it is a series of explanation on how chocolate is formed before we eat. First, the chocolate is coming from the cacao tree. Then it is fermented and ship to the chocolate producer. The cacao bean then are roasted and winnowed.

Example of Explanation Text – How A Fuel Light Works

How a fuel light works.

Many cars, motorcycles and other modern vehicles have fuel warning light devices. the warning light is usually red which switches on automatically when the level of fuel in the tank is very low. The warning light gives the driver information about the amount of petrol in the tank. When the light switches on red, it tells the driver that the petrol in the tank is almost empty. Therefore we have to put more fuel into the tank. However do you know how the fuel warning works?

Well this is the way the fuel warning light work and gives the driver information about the accurate amount of the petrol in the tank. When the level of the fuel falls, the float inside the tank moves downwards. When this condition happens, the arm also moves downwards and it make the lever touch an electrical contact. This switches on the fuel light in the car dashboard.

The red light which appears in the fuel panel of the dashboard tells the driver that he needs more petrol for his car. When he pours more petrol into the tank, this condition makes the fuel level rise and it pushes the float upwards. In return it disconnects to the electrical contact and makes the red light switch off.

Cara Kerja Cahaya Bahan Bakar

Banyak mobil, sepeda motor dan kendaraan modern lainnya memiliki lampu peringatan bahan bakar. Lampu peringatan biasanya merah yang menyala secara otomatis bila tingkat bahan bakar di dalam tangki sangat rendah. Lampu peringatan memberi informasi tentang jumlah bensin di tangki. Bila lampu menyala merah, berarti pengemudi nya bensin di tangki hampir kosong. Karena itu kita harus memasukkan lebih banyak bahan bakar ke dalam tangki. Namun tahukah anda bagaimana bahan bakar bekerja?

Nah begitulah cara kerja lampu peringatan bahan bakar dan memberi informasi pengemudi tentang jumlah bensin yang akurat di tangki. Bila tingkat bahan bakar turun, pelampung di dalam tangki bergerak ke bawah. Bila kondisi ini terjadi, lengan juga bergerak ke bawah dan itu membuat tuas menyentuh kontak listrik. Ini akan menyalakan lampu bahan bakar di dasbor mobil.

Lampu merah yang muncul di panel bahan bakar di dasbor memberi tahu pengemudi bahwa ia membutuhkan lebih banyak bensin untuk mobilnya. Saat menuangkan bensin lagi ke tangki, kondisi ini membuat tingkat bahan bakar naik dan mendorong float ke atas. Sebagai gantinya diputus ke kontak listrik dan mematikan lampu merah.

(Grammar – xx)

Related Articles : Report Text ; Definition, Generic Structures, Purposes, Language Features

Oke. I think that’s all my explanation about Explanation Text. I hope it will be useful for us. I hope you are never bored to visit this site. Don’t forget to visit this site if you need English reference in your study. See you next time..

Reference : Rudi Hartono, Genre of Texts, (Semarang: English Department Faculty of Language and Art Semarang State University, 2005). Text Genre, Grammar: Technologies for Teaching and Assessing Writing, Peter Knapp & Megan Watkins, New South Wales Press, Ltd : Australia Mark Andersons and Kathy Andersons, Text Type in English 1-2, Australia: MacMillanEducation, 2003.

Related posts:

• Explanation Text
• Descriptive Text (Penjelasan Dan Contoh)
• Narrative Text (Complete Explanation)
• Hortatory Exposition Text; Definition, Generic Structures, Purposes, Language Features

Why is the Hello written as ‘Hallo’ (lines 1) ?

I won’t to change my phone number because my phone is lost yesterday 240am

1)the purpose of the explanation text is to explain the process involved in the formation of a natural

2)the generic structure of the explanation text : a.general statement (this part is general introduction of the phenomeneon that we are going to explain) b.sequenced explanation (explains about some process involved of the phenomeneon) c.closing (presented in this part)

3)language features of explanation text: a.using simple present tense b.using passive voice c.focus on natural

Saya ingin bergabung

It is good writing to complete my Englush. Thanks

I want to learn English better

Ok. Good Widaya 🙂

52/1Durga Charan doctar lean road Kolkata 700014

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The recount Text Social function, Definition, Generic structures, Language features and examples

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Quantitative Biology > Populations and Evolution

Title: ecology, spatial structure, and selection pressure induce strong signatures in phylogenetic structure.

Abstract: Evolutionary dynamics are shaped by a variety of fundamental, generic drivers, including spatial structure, ecology, and selection pressure. These drivers impact the trajectory of evolution, and have been hypothesized to influence phylogenetic structure. Here, we set out to assess (1) if spatial structure, ecology, and selection pressure leave detectable signatures in phylogenetic structure, (2) the extent, in particular, to which ecology can be detected and discerned in the presence of spatial structure, and (3) the extent to which these phylogenetic signatures generalize across evolutionary systems. To this end, we analyze phylogenies generated by manipulating spatial structure, ecology, and selection pressure within three computational models of varied scope and sophistication. We find that selection pressure, spatial structure, and ecology have characteristic effects on phylogenetic metrics, although these effects are complex and not always intuitive. Signatures have some consistency across systems when using equivalent taxonomic unit definitions (e.g., individual, genotype, species). Further, we find that sufficiently strong ecology can be detected in the presence of spatial structure. We also find that, while low-resolution phylogenetic reconstructions can bias some phylogenetic metrics, high-resolution reconstructions recapitulate them faithfully. Although our results suggest potential for evolutionary inference of spatial structure, ecology, and selection pressure through phylogenetic analysis, further methods development is needed to distinguish these drivers' phylometric signatures from each other and to appropriately normalize phylogenetic metrics. With such work, phylogenetic analysis could provide a versatile toolkit to study large-scale evolving populations.

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Identification and expression analysis of the Xyloglucan transglycosylase/hydrolase ( XTH ) gene family under abiotic stress in oilseed ( Brassica napus L.)

• Jingdong Chen 1 , 2 ,
• Heping Wan 2 ,
• Huixia Zhao 2 ,
• Xigang Dai 2 ,
• Wanjin Wu 2 ,
• Jin Liu 1 ,
• Jinsong Xu 1 ,
• Rui Yang 1 ,
• Benbo Xu 1 ,
• Changli Zeng 2 &
• Xuekun Zhang 1  

BMC Plant Biology volume  24 , Article number:  400 ( 2024 ) Cite this article

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XTH genes are key genes that regulate the hydrolysis and recombination of XG components and plays role in the structure and composition of plant cell walls. Therefore, clarifying the changes that occur in XTHs during plant defense against abiotic stresses is informative for the study of the plant stress regulatory mechanism mediated by plant cell wall signals. XTH proteins in Arabidopsis thaliana was selected as the seed sequences in combination with its protein structural domains, 80 members of the BnXTH gene family were jointly identified from the whole genome of the Brassica napus ZS11, and analyzed for their encoded protein physicochemical properties, phylogenetic relationships, covariance relationships, and interoperating miRNAs. Based on the transcriptome data, the expression patterns of BnXTHs were analyzed in response to different abiotic stress treatments. The relative expression levels of some BnXTH genes under Al, alkali, salt, and drought treatments after 0, 6, 12 and 24 h were analyzed by using qRT-PCR to explore their roles in abiotic stress tolerance in B. napus . BnXTHs showed different expression patterns in response to different abiotic stress signals, indicating that the response mechanisms of oilseed rape against different abiotic stresses are also different. This paper provides a theoretical basis for clarifying the function and molecular genetic mechanism of the BnXTH gene family in abiotic stress tolerance in rapeseed.

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Introduction

In complex and changeable natural environments, crops may be exposed to abiotic stresses such as temperature, drought, salt, heavy metals, etc., potentially compromising their yield and quality [ 1 ]. As the first barrier, the cell wall plays an important role in the defense of plants against external environmental stresses [ 2 ].

The cell wall has a complex structure consisting of pectin embedded microfibrils and non-cellulosic neutral polysaccharides, crosslinked with structural proteins and, depending on the tissue and organ, with lignin [ 3 ]. Plant cell walls are divided into the intercellular layer, primary and secondary walls [ 4 ]. The main component of the intercellular layer is pectin, which is located between two neighboring cells and is able to adhere to them and buffer intercellular extrusion [ 5 ]; the primary wall is mainly composed of cellulose, hemicellulose, and structural proteins, which has a large plasticity, which allows the cell to maintain a certain shape, but also can be extended with the growth of the cell [ 6 ]; the secondary wall, which is mainly composed of cellulose and often contains lignin, located between the plasma membrane and the primary wall, is usually thicker and harder, giving the cell wall great mechanical strength [ 7 ].

Xyloglucan (XG) is the main chain component of the polysaccharide hemicellulose that accounts for the largest proportion in the primary cell wall of dicotyledonous plants [ 8 ]. The glucan backbone of XG is composed of β-1,4-D-Glup, and the O-6 position is connected to α-D-Xylp. These Xyls can be further modified by other glycosyl groups [ 9 ]. The structures of XG are diverse, showing variability among plants, organs, tissues and even within the same plant [ 10 ]. The basic function of XG is to bind to the outside of cellulose fibrils to form a load-bearing network of the primary cell wall, which determines the tension of the cell wall. The modification and reconstruction of XG can adjust the elasticity and strength of the cell wall skeleton [ 11 ].

The recombination of XG requires the participation of Xyloglucan transglycosylase/hydrolase (XTH) of the GH16 family. XTH has dual activities of Xyloglucan endotransglycosylase (XET) and Xyloglucan hydrolase (XEH), allowing it to independently complete the cleavage and recombination process of XG, and can carry out the catalytic reaction of extending the XG chain without the direct involvement of ribose [ 12 ]. XTH genes play roles in plant growth and development, especially in response to abiotic stress. The cell wall modification of xth19 could affect freezing tolerance after cold and sub-zero acclimation. Compared with the Col-0 wild type, the cell wall structure and composition of the xth19 mutant of Arabidopsis changed, which resulted in a lower freezing tolerance after cold and sub-zero acclimation [ 13 ]. Compared to the control, the expressions of TaXTHs of wheat were significantly altered under drought stress; the drought tolerance of transgenic plants of Arabidopsis overexpressing TaXTH12.5a was improved, with the germination rate, root length, hypocotyl length, and the number of green leaves during the germination stage and nutrient growth stage significantly higher than that of the control lines [ 14 ]. Under salt stress, there were 11 differentially expressed PtrXTHs in the roots of poplar, nine differentially expressed PtrXTHs in the stems, and 7 differentially expressed PtrXTHs in the leaves [ 15 ]; the water retention capacity of the leaves in transgenic plants of tobacco overexpressing PeXTH of Populus were improved, and the net photosynthetic rate had a significant enhancement, which enhanced the salt tolerance of tobacco, indicating that the XTHs might play roles for plants in the response to salt stress [ 16 ]. It was found that many XTHs of soybean responded to ethylene and flooding treatments, The Arabidopsis thaliana AtXTH31 gene is overexpressed in soybean, and transgenic soybean plants with AtXTH31 overexpressing under flooding stress tolerance showed higher germination rate, and longer roots/ hypocotyls compared to the control lines at the seedling and nutrient growth stages, it was confirmed that XTHs played roles in regulating the response of soybean to abiotic stresses [ 17 ]. Du et al. found that aluminum (Al) induces ZmXTHs to be up-regulated in the maize root system, especially in the root tips, and this induction was Al 3+ specific; ZmXTHs overexpressing plants of Arabidopsis grew more healthily with lower Al content in their roots and root cell walls under Al stress compared to wild-type, suggesting that ZmXTHs could be endowed with aluminum tolerance on transgenic Arabidopsis plants by reducing the accumulation of aluminum in roots and cell walls [ 18 ]. XTHs also played important roles in the response of heavy metal stress in plants: most XTHs in ramie ( Boehmeria nivea ) were responsive to cadmium stress, while heterologous expression of BnXTH3, BnXTH6 and BnXTH15 significantly enhances the cadmium tolerance of transgenic yeast cells [ 19 ]; Xuan et al. identified 44 possible MtXTHs in the Medicago truncatula genome, 28 of which were responsive to HgCl 2 , and copper/mercury stresses would significantly induce the expression of the MtXTH3 both in the roots and shoots. In addition, the expression of MtXTH3 showed an increasing trend with the elevation of Cu and Hg concentrations [ 20 ].

While the impact of XTHs on plant cell wall composition and abiotic stress resilience has been thoroughly investigated in Arabidopsis and various crops, the expression profiles of these genes under abiotic stress conditions in Brassica napus remain comparatively underexplored. In this study, we undertook a comprehensive genome-wide identification of the BnXTH gene family members in rapeseed; additionally, we conducted an exhaustive analysis encompassing protein characteristics, gene structure, phylogenetic relationships, and collinearity the expression patterns of BnXTHs in two varieties of rapeseed were compared under alkaline (Na 2 CO 3 ) stress using RNA-seq analysis. At the same time, by qPCR, the relative expressions of BnXTHs under aluminum (AlCl 2 ), alkali (Na 2 CO 3 ), salt (NaCl) and drought (PEG 6000) stress at four time points (0, 6 h, 12 h and 24 h) were examined, which provided evidence for the response of BnXTHs to abiotic stress. The findings of our study establish a foundational framework for subsequent investigations into the functional roles of these genes in the abiotic stress response of rapeseed. Furthermore, they offer a theoretical underpinning for the selection and breeding of abiotic stress-tolerant rapeseed germplasms.

Identification and chromosomal localization of the BnXTH gene family

Chromosomal location of the BnXTHs in the B. napus genome

Joint identification was conducted using BLASTP and HMMER, and the NCBI-CDD tool was used to remove the sequences with incomplete structural domains. Finally, 80 BnXTH proteins were obtained, which were named BnXTH1 – BnXTH80 based on the order of the location of their encoding genes on the chromosomes. Among these 80 genes, 79 were mapped to 18 identified chromosomes (Fig.  1 ), and one was mapped to pseudochromosome Scaffold0027. Of these, 10 BnXTHs were localized on ChrA03 and ChrC07, nine on ChrC09, seven on ChrA01, six on ChrA08 and ChrC09, five on ChrA06 and ChrC01, three on ChrA07, ChrC02, ChrA05, and ChrA09, two were localized on ChrA10 and ChrC05, one was localized on ChrA02, ChrA04, and ChrC04, while no possible BnXTHs were detected on ChrC06. Number of BnXTHs’ exons ranged from one to eight, and the information of their positions in the chromosomes was provided on Table S1 .

Physicochemical properties and subcellular localization prediction of BnXTH proteins

By predictive analysis, the number of amino acids (AAs) of the BnXTH proteins ranged from 212 to 473, the relative molecular mass (MW) ranged from 24.42 to 55.10, and the theoretical isoelectric point (PI) ranged from 4.77 to 9.96 (Table S1 ). Subcellular localization prediction results showed that 54 XTH proteins may exist in the cytoplasm, 21 in the cell wall, and 5 in the nucleus.

Phylogenetic analysis of BnXTH proteins

In order to explore the phylogenetic relationship between XTH proteins, 80 XTH proteins of B. napus and 33 XTH proteins of Arabidopsis thaliana were combined to form a phylogenetic tree. Based on the genetic relationship between them and the position of the branch where the AtXTH proteins were located, XTH proteins were divided into four groups: Early diverging group, Group I/II, Group IIIA and Group IIIB (Fig.  2 A). According to the phylogenetic tree, it could be seen that the number of proteins contained within each group varies, among which the most proteins were clustered into Group I/II, with 22 AtXTHs and 49 BnXTHs; in Group IIIB, there were 5 AtXTHs and 16 BnXTHs; in the Early diverging group, there were 4 AtXTHs and 8 BnXTHs; Group IIIA had the fewest proteins, with 2 AtXTHs and 7 BnXTHs (Table  1 ).

To further confirm the phylogenetic relationships of XTHs within Brassicaceae plants, a total of 52 XTH proteins from Brassica rapa and 47 XTH proteins from Brassica oleracea were identified by using BLASTP and HMMER methodologies for the construction of a phylogenetic tree (Table S2 , Fig.  2 B). The analysis revealed that these proteins could also be classified into four groups, with the groupings of BnXTHs remaining unchanged, thereby validating the reliability of the phylogenetic tree. Notably, the Group I/II contained the highest number of proteins, with 35 BrXTHs and 33 BoXTHs, followed by Group IIIB, which comprised 8 BrXTHs and 8 BoXTHs. Group IIIA included 4 BrXTHs and 4 BoXTHs, and the Early diverging group consisted of 4 BrXTHs and 4 BoXTHs (Table  1 ).

Phylogenetic tree of XTH proteins. ( A ) Phylogenetic tree of XTH proteins in B. napus and (A) Thaliana . ( B ) Phylogenetic tree of XTH proteins in (B) napus , B. rapa , and B. oleracea

Gene structures, conserved motifs and promoter cis-acting elements analysis of BnXTHs

Analysis of motifs in conserved domains is a powerful tool for understanding the function, structure, and evolution of proteins. Conserved motif analysis of BnXTH proteins shows that Motif2, Motif 3, Motif 5 and Motif 6 were common to all BnXTH proteins; Motif7 was merely absent from BnXTH75; Motif4 was absent from BnXTH33 and BnXTH75 of Group I/II; Motif1 was absent in BnXTH7 and BnXTH24 of the Early diverging group; most Group IIIB and a small number of Early diverging group proteins did not contain Motif10; Motif8 existed in some Group I/II and BnXTH35 proteins; Motif9 was present in most of the Group IIIB proteins. (Fig.  3 A). The protein structural characteristics of BnXTHs were identified, and all proteins were found to contain a Glyco_hydro_16 and a XET_C domain (Fig.  3 B). Analysis of promoter cis-acting elements showed that BnXTHs may have a response mechanism in response to stress and plant hormone signals (Fig.  3 C). Among them, there were as many as 910 light responsiveness elements, far more than other cis-acting elements. In addition, BnXTH41 and BnXTH45 contained the most cis-acting elements, amounting to 41, indicating that they may play roles in the growth and development of rapeseed. The study of the mature mRNA structure of BnXTHs revealed that they contained 1–8 CDS (Coding sequence) regions (Fig.  3 D), and some BnXTHs also contained 1–2 UTR (Untranslational region) regions.

Gene structure analysis of XTH family in B.napus . ( A ) Conserved motifs of BnXTH family proteins. ( B ) Pfam structure of BnXTH family proteins. ( C ) Promoter cis-acting element of BnXTHs . ( D ) The mRNA structure encoded by the BnXTHs

Collinearity analysis of XTH gene family

In order to investigate the collinearity of XTH genes, BnXTH80 located on the pseudochromosome was removed and the collinearity maps of rapeseed ( B. napus ), B. rapa and B. oleracea were drawn (Fig.  4 ). There were 92 pairs of collinear genes between the XTH genes of B. napus and B. rapa , and 88 pairs of collinear genes between the XTH genes of B. napus and B. oleracea (Table S3 ). Collinearity analysis of BnXTHs within the B. napus genome showed that there were 83 pairs of collinear genes among BnXTHs (Fig.  5 and Table S4 ). To understand the genetic relationship of BnXTH collinear gene pairs, KaKs values were calculated to understand their selection pressure relationships (Table S5 ). The results showed that the Ka/Ks values of these 83 collinear gene pairs were all less than 1, indicating that they were all affected by Purification selection.

Collinearity of XTH genes in B. napus , B. rapa , and B. oleracea

Collinearity of BnXTH s. The circles in the figure from inside to outside represent the unknown base N ratio, gene density, GC ratio, GC skew, and chromosome length of the B. napus genome

Prediction of targeting relationship between miRNA and BnXTHs

As shown in Fig.  6 , a total of 32 miRNAs of B. napus targeted 29 BnXTH genes through cleavage, and 5 miRNAs of B. napus target 3 BnXTH genes through translation. No miRNA was found to have both cleavage and translation inhibition effects on the same BnXTH gene. The results suggest that multiple miRNAs were involved in the post-transcriptional regulation of BnXTHs by targeting them through cleavage and translation.

Sankey diagram for the relationships of miRNA targeting to BnXTHs transcripts. The 3 columns represent miRNA, BnXTHs and inhibition effect

Analysis of transcriptome expression patterns of BnXTHs under abiotic stresses

In order to explore the gene expression patterns of BnXTHs under different abiotic stresses, we obtained the transcriptome TPM data of BnXTHs in B. napus ZS11 under CK, salt, drought, freezing, cold, heat and osmotic stress, and expression heat maps were plotted after log 10 (TPM + 1) treatment (Fig.  7 ). Some BnXTHs were not expressed or had low expression in leaves and roots under different stresses; however, BnXTH26 , BnXTH63 and BnXTH37 showed high expression in leaves and roots under various stresses; BnXTH13 and BnXTH52 had high expression in roots under various stresses and high expression in leaves under some of the stresses; the expression levels of BnXTH15 , BnXTH66 , BnXTH25 , and BnXTH53 were higher in leaves under various stresses; the expression levels of BnXTH17 , BnXTH68 , BnXTH22 , BnXTH58 , and BnXTH73 showed higher expression in roots under various stresses.

Analysis of expression patterns of BnXTHs under abiotic stress (CK, salt, drought, freezing, cold, heat, and osmotic) treatments. L: leaves; R: Roots

Detection of BnXTHs expression under various abiotic stresses by qPCR technique

The cell wall is the first barrier to protect plant cells from damage, and the XTH genes plays an important role in plants defense against abiotic stress. Therefore, to investigate the effect of BnXTHs on rapeseed response to abiotic stress, we performed qPCR on leaves of oilseed rape seedling ZS11 treated with 0.5 mmol·L -1 AlCl 3 , 0.2% (w/v) Na 2 CO 3 , 1.2% (w/v) NaCl, and 20% (w/v) PEG 6000 after 0, 6 h, 12 h and 24 h to analyze the relative expression of 9 different BnXTHs (Fig.  8 ). The results showed that they responded significantly to different abiotic stresses.

Under AlCl 3 treatment, the expression of BnXTH7 , BnXTH17 , and BnXTH6 did not change significantly at all time points; BnXTH72 and BnXTH37 increased with the extension of the treatment time; BnXTH24 , BnXTH9 , BnXTH20 , and BnXTH61 increased with the extension of the treatment time, reaching the maximum at 12 h, and then decreased at 24 h again.

Under Na 2 CO 3 treatment, the expression of BnXTH7 , BnXTH17 , BnXTH6 , BnXTH20 , and BnXTH37 increased with the prolongation of the treatment time; the expression of BnXTH24 , BnXTH19, and BnXTH61 increased and then decreased; and the expression of BnXTH72 showed a decreasing tendency first, but it was insignificant, and then increased significantly at 24 h. The expression of BnXTH72 increased significantly at 24 h, but it was not significant.

Under NaCl treatment, the expression of BnXTH7 and BnXTH6 did not change significantly; the expression of BnXTH72 , BnXTH37 , and BnXTH61 increased with the prolongation of time; and the expression of BnXTH24 , BnXTH17 , BnXTH9 , and BnXTH20 increased and then decreased.

The relative expression of BnXTH7 , BnXTH24 , BnXTH17 , BnXTH6 , BnXTH9 , BnXTH20 , BnXTH72 , BnXTH37 , and BnXTH61 in leaves under 0.5 mmol·L -1 AlCl 3 , 0.2% (w/v) Na 2 CO 3 , 1.2% (w/v) NaCl, and 20% (w/v) PEG 6000 after 0, 6 h, 12 h and 24 h. Data represent the mean ± standard error for threebiological experiments. Student’s t-test was used to determine differences. *: significant differences between treatments at p  ≤ 0.05. **: significant differences between treatments at p  ≤ 0.01

Under PEG 6000-induced drought stress, the expression of BnXTH6 , BnXTH72 , and BnXTH37 did not change significantly; the expression of BnXTH7 increased with the prolongation of time; the expression of BnXTH17 , BnXTH9 , BnXTH20 , and BnXTH61 increased and then decreased; the expression of BnXTH24 at 6 h and then decreased at 12 h. The expression of BnXTH24 was elevated at 6 h, and then increased again at 24 h. The expression of BnXTH24 increased at 6 h and then decreased at 12 h, and then increased again after 24 h.

Gene family analysis is conducive to mining key functional genes in crop genomes and provides a genetic research basis for the development of high-yield and high-quality germplasms. In this study, the Arabidopsis XTH protein sequences were used as the seed sequence, combined with the Pfam domain search results, and finally identified 80 members of the B. napus XTH gene family, distributed on 18 definite chromosomes and 1 pseudochromosome. In this study, 52 BrXTH proteins and 47 BoXTH proteins were identified, the numbers of which differ from previous studies (53 XTHs of B. rapa and 38 XTHs of B. oleracea ) [ 21 ]. This discrepancy may be due to differences in the sources of genome databases and the selection of thresholds. Compared with the 52 BrXTH proteins and 47 BoXTH proteins, B. napus , as an allotetraploid, was identified with 80 XTH genes, a number smaller than the sum of the former two, which also suggested that recombination or mutation at the gene or chromosome level may have occurred in the B. napus genome during the evolutionary process. Phylogenetic analysis was performed based on the XTH protein sequences of (A) thaliana and (B) napus , and the distribution of XTH proteins in the phylogenetic tree was statistically analyzed. It was found that BnXTH proteins were more uniformly distributed among the four groups of the phylogenetic tree, suggesting that gene duplication events might have occurred in the genome of B. napus at the whole-genome level in these four groups (Fig.  2 A). In the construction of phylogenetic trees for three species within the Brassicaceae family, it was observed that the distribution of XTHs within the four groups was broadly consistent (refer to Fig.  2 B; Table  1 ). According to the U-triangle model, B. napus (AACC: 2n = 38), an allopolyploid species, originates from the cross between B. rapa (AA: 2n = 20) and B. oleracea (CC: 2n = 18) [ 22 ]. This analogous distribution pattern of XTHs across the species underscores their genetic conservation. Furthermore, the total count of BnXTHs being lesser than the combined total of BrXTHs and BoXTHs highlights the evolutionary distinctiveness of BnXTHs.

Further analysis of the gene structures and conserved domains showed that evolutionary conservation may exist among species including (A) thaliana [ 23 ], Osmanthus fragrans [ 24 ], Schima superba [ 25 ], (B) rapa and B. oleracea [ 21 ]. Motif 2, Motif 3, Motif 5, and Motif 6 were identified as ubiquitous across all BnXTH proteins. This uniform presence suggests that these four motifs likely constitute characteristic sequences of BnXTHs, playing pivotal roles in the structural and functional attributes of these proteins. Analysis of promoter cis-acting elements showed that the function of BnXTHs may be related to photosynthesis due to the large number of light responsiveness elements. By modifying the cell wall, XTHs influence leaf thickness, which can affect the internal leaf architecture and, consequently, the efficiency of light capture and the distribution of light within the leaf [ 16 ]. XTHs are also involved in the development and functioning of stomata by modulating the flexibility and integrity of the cell walls surrounding guard cells [ 26 ]. While the direct relationship between XTH activity and photosynthesis requires further empirical study, it is evident that XTHs, through their role in cell wall modification, indirectly contribute to optimizing the conditions necessary for efficient photosynthesis.

The Ka/Ks analysis is a method used to measure the selective pressure on genes, commonly applied in the study of gene or gene family evolution. In this context, Ka represents the rate of nonsynonymous substitutions, which lead to changes in amino acids, while Ks denotes the rate of synonymous substitutions that do not result in amino acid changes [ 27 ]. Ka/Ks analysis serves as a crucial tool for understanding the dynamics of gene evolution and revealing the key mechanisms behind the adaptive evolution of organisms, suitable for discussion in scientific literature. By collinearity analysis between genomes, we found that there were 92 pairs of XTH collinear genes between B. napus and B. rapa , 88 pairs of XTH collinear genes between B. napus and B. oleracea (Fig.  4 ), and 83 XTH covariance pairs within the B. napus genome (Fig.  5 ), and their Ka/Ks values were all less than 1 (Table S5 ). The number of collinear gene pairs of XTHs among species was greater than the number of BnXTHs we identified, indicating that XTHs are evolutionarily correlated and have both inter- and intra-species evolutionary conservation.

miRNAs (microRNA) are small non-coding RNAs that regulate gene expression at the post-transcriptional level by inhibiting the translation of messenger RNAs (mRNAs) or by promoting mRNA degradation [ 28 ]. miRNAs are approximately 20–24 nucleotides (nt) in size and are encoded by plants, animals and some viruses [ 29 ]. In plants, miRNAs inhibit gene expression mainly by mediating target RNA cleavage or translational repression [ 30 ], and they control target genes at the post-transcriptional or translational level by controlling the level of protein synthesis, and have an impact on plant growth, development, and response to environmental stresses [ 31 ]. For example, bna-miR159, bna-miR6029, and bna-miR827 negatively regulate target genes related to nitrogen metabolism pathway in B. napus , thereby affecting nitrogen signaling within the B. napus plant and consequently its pod thickness [ 32 ]. bna- miR319 overexpressing transgenic lines of B. napus exhibited abnormal development of serrated leaves and stem tip meristematic tissues [ 33 ]. Fu et al. analyzed miRNA-mRNA expression of Cd-stressed B. napus seedlings and found that Cd treatment significantly affected the expression of 22 miRNAs belonging to 11 families in roots and 29 miRNAs belonging to 14 families in shoots, and identified 8 miRNA-mRNA interaction pairs in roots and 8 miRNA-mRNA interaction pairs in shoots. 8 miRNA-mRNA interaction pairs in roots and 8 miRNA-mRNA interaction pairs in shoots were identified [ 34 ]. In this study, a total of 37 miRNAs of B. napus were targeted to 29 BnXTH genes through cleavage and translational repression (Fig.  6 ), and it was hypothesized that miRNAs form a miRNA-target gene regulatory network with BnXTHs , which are involved in the process of reorganization and hydrolysis of XG in B. napus cell walls. Interestingly, BnXTHs have similar cis-acting elements (Fig.  3 ), but could have relatively different expression patterns (Fig.  7 ). For example, BnXTH2 , and BnXTH41 all targetted bna-miR167 and had relatively consistent expression patterns. They were not expressed in leaves under various abiotic stresses but have a certain amount of expression in roots. However, compared with the expression patterns of other BnXTHs , they were different.

Various abiotic stresses may affect the integrity of plant cell walls, but plants are able to repair adverse changes in the cell wall, including changing its composition, structure, and mechanical properties to maintain growth [ 35 ]. There has been some research on how the cell wall responds to stress [ 36 ]. However, due to the constraints of imaging and biomechanical equipment and the potential crosstalk between cell wall signaling and stress signaling, it is difficult to draw general conclusions about the regulatory mechanisms of the cell wall in response to various abiotic stresses [ 2 ]. XTH genes are key genes that regulate the hydrolysis and recombination of XG components and plays an important role in the structure and composition of plant cell walls [ 25 ]. Therefore, clarifying the changes that occur in XTHs during plant defense against abiotic stresses is informative for the study of the plant stress regulatory mechanism mediated by plant cell wall signals. In this study, qPCR technology was used to detect the relative expression BnXTH7 , BnXTH24 , BnXTH17 , BnXTH6 , BnXTH9 , BnXTH20 , BnXTH72 , BnXTH37 , and BnXTH61 in leaves under 4 abiotic stresses treatment after 4 time points: 0, 6 h, 12 h, and 24 h (Fig.  8 ). It was found that in response to abiotic stresses, the expression of most of the BnXTHs gradually increased or first increased and then decreased with the growth of treatment time, indicating that oilseed rape needs large amounts of recombinant XGs in response to abiotic stress in order to repair the damage that may be caused by the stress to the cell wall. And with the time went, the plant slowly adapted to the stress, and some of the BnXTHs may have received the relevant signals, thus the expression fell back.

Materials and methods

Identification and chromosome mapping of bnxth family members.

The sequence files of ZS11 (a variety of B.napus ), cabbage ( B. rapa ), and kale ( B. oleracea ) were obtained from the BRAD database (BRAD: http://brassicadb.cn/#/ ) [ 37 ]. 33 AtXTH protein sequences from the A. thaliana genome database (TAIR: https://www.arabidopsis.org/ ) were downloaded as seed sequences, and searched for possible BnXTH proteins in the whole protein sequences of B.napus (e Value < 1e-10) by BLASTp [ 38 ]. The PF06955 and PF00722 conserved structural domain files were downloaded from the Pfam database to search the possible BnXTH proteins by HMMER software ( http://www.hmmer.org/ ) [ 39 , 40 ], combining the BLAST results to obtain the BnXTH hypothetical proteins. The putative protein sequence was uploaded to the NCBI-CDD website ( https://www.ncbi.nlm.nih.gov/cdd/ ) for further confirmation, and 80 members of the BnXTH gene family were finally identified. The BnXTHs were named according to their chromosomal position sequence, and were named as BnXTH1  ~  BnXTH80 . The Protparam function in the ExPASy website was used to predict the physical and chemical properties of BnXTH proteins [ 41 ], and the subcellular localization prediction results were implemented by the Plant-mPLoc website [ 42 ]. Chromosomal location information of BnXTHs was retrieved using downloaded annotation files and visualized by TBTools [ 43 ].

Phylogenetic analysis

The XTH protein sequences of A. thaliana and B.napus were imported into MEGA 11 software, and the parameters were adjusted to neighbor joining (NJ) and 1000 boots-trap repetitions to obtain a phylogenetic tree, and the phylogenetic tree was embellished in the online website iTOL ( https://itol.embl.de/ ) [ 44 , 45 ].

Prediction of gene structure, protein conserved motifs and promoter cis-acting elements

The CDS/UTR region and gene structure information of BnXTH gene family members were obtained from the NCBI-CDD website and Pfam database. The MEME 5.5.1 website was used to obtain the conserved motif of the BnXTH proteins [ 46 ], and the prediction of the cis-acting element 2000 bp upstream of the BnXTHs promoter region was predicted through the PlantCARE website [ 47 ]. All data information visualization was completed by TBTools.

Collinearity analysis of XTH genes

MCScanX software was used to analyze the collinear relationship between XTHs within the genomes of B. napus, B. rapa , and B. oleracea , as well as within the genomes of B. napus [ 48 ]. Collinearity information visualization was performed using TBTools. Ka/Ks value calculation was implemented using KaKs_calculator 3.0 software [ 49 ].

The CDS sequences of BnXTHs were uploaded to the psRNATarget website ( https://www.zhaolab.org/psRNATarget/analysis?function=2/ ) [ 50 ], and combined with the miRNA mature sequences information of B. napus collected by the website, the targeting relationships between miRNA and BnXTHs were analyzed. The data were visualized using the Bioinformatics website ( https://www.bioinformatics.com.cn/plot_basic_alluvial_plot_017 ).

Analysis of the expression pattern of BnXTHs under different abiotic stress

To investigate the gene expression patterns of BnXTHs under abiotic stress treatments, the BnXTH gene IDs were uploaded to the BNIR database [ 51 ] to obtain the Transcript per Kilobase per Million mapped reads (TPM) of B. napus ZS11 in control (CK), salt (200 mmol·L -1 NaCl treatment), drought (exposure to airflow for 1 h), freezing (recovering to 25 °C after 3 h of stress at -4 °C), cold (4 °C), heat (recovering to 25 °C after 3 h of stress at 38 °C), and osmosis (300 mmol-L-1 mannitol) after 24 h, and the data were visualized using TBTools software after calculating log 10 (TPM + 1).

Materials and treatment

B. napus ZS11 seeds were germinated on wet gauze (soaked with water) in a plant growth chamber at 20 to 22 °C and 65% humidity under a long-day condition (16-h-light/ 8-h-dark cycle) [ 52 ]. The one-week-old seedlings were then transferred into a previously described hydroponic system [ 53 ] under the same culture conditions for nearly 20 days until the fourth leaves had extended (plant samples used for transcriptome sequencing were also harvested for root system). For stress treatment research, leaf samples from 4-week-old plants of ZS11 were collected after 0, 6, 12 and 24 h of 0.5 mmol·L -1 AlCl 3 , 0.2% (w/v) Na 2 CO 3 , 1.2% (w/v) NaCl, and 20% (w/v) PEG 6000 treatment. Seedlings without any stress treatment were used as the control. Each treatment includes three biological replications. Leaves were harvested immediately frozen in liquid nitrogen and stored at -80 °C for RNA extraction.

Relative expression of BnXTHs under different abiotic stresses

Total RNA of ZS11 under control and abiotic treatments was extracted using the RNA simple Total RNA kit (Tiangen Biotechnology Co., Ltd., Beijing, China) according to the manufacturer’s protocol. cDNA was synthesized with 1 µg RNA from each sample with HiScript® II Q Select RT SuperMix with gDNA wiper (Vazyme, Nanjing, China). Gene-specific primers used for quantitative real-time PCR (qRT-PCR) listed in Table  2 . qRT-PCR was run on the AriaMx real-time PCR system (Agilent Technologies). The following cycling parameters were used: initial denaturation at 95 °C for 5 min; 40 amplification cycles consisting of denaturation at 95 °C for 10s, annealing and extension at 60 °C for 30 s; The melting curve was then tested at 65–95 °C. The internal standard was the B. napus actin gene ( BnaA01g27090D ). To investigate the expression patterns of BnXTHs under various environmental conditions, 9 BnXTHs were randomly selected from the 4 groups for qRT-PCR experiments. Three biotic replicates were performed for each sample, and each replicate contained three technical replicates. Relative expression levels were calculated according to the 2 −ΔΔCt method [ 54 ]. Data represent the mean ± standard error for threebiological experiments. Student’s t-test was used to determine differences. *: significant differences between treatments at p  ≤ 0.05. **: significant differences between treatments at p  ≤ 0.01.

Conclusions

B. napus genome was identified to contain 80 XTH genes distributed on 18 definite chromosomes and one pseudochromosome. Their phylogenetic relationships, protein physicochemical properties, subcellular localization, gene structures, promoter cis-acting elements, covariance relationships, and reciprocal miRNAs were predicted and analyzed, and their transcriptional expression patterns as well as differences in expression in response to abiotic stress treatments were investigated. The results showed that the expression patterns of BnXTHs under abiotic stress treatments were varied, suggesting that cell wall signaling in B. napus in response to various abiotic stresses changes depending on the type of stress. The analysis of the BnXTH gene family and the study of the response pattern in abiotic stresses provide a good theoretical basis for further research on this family of genes in resistance breeding of B. napus .

Data availability

All the data generated or analyzed during this study are included in this published article and its supplementary information files.

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Acknowledgements

We would like to thank the Hubei Province “515” Action (Cooperative Extension) “Rapeseed New Technology Demonstration and Science and Technology Service Rapeseed Industry Chain Project” and Ministry of Agriculture and Rural Affairs Plantation Management Department “Rapeseed Industry Policy Research and Main Pests and Diseases Prevention and Control Strategies " (15214011), National major biological breeding project of China (2022ZD04010). Thanks for their financial support.

This study was supported by the grants from the Hubei Province “515” Action (Cooperative Extension) “Rapeseed New Technology Demonstration and Science and Technology Service Rapeseed Industry Chain Project”, Ministry of Agriculture and Rural Affairs Plantation Management Department “Rapeseed Industry Policy Research and Main Pests and Diseases Prevention and Control Strategies " (15214011), National major biological breeding project of China (2022ZD04010) and Jianghan University scientific research project funding scheme (2022XKZX17).

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Hubei Engineering Research Center for Protection and Utilization of Special Biological Resources in the Hanjiang River Basin, College of Life Science, Jianghan University, Wuhan, 430056, Hubei, China

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Contributions

X.Z. and C.Z. designed and managed the project. J.C. and H.W. completed experiments and paper writing. H.Z., X.D., W.W., J.L., J.X. and R.Y. performed experiments, material sampling. Laboratory data measurements. X.Z., C.Z. and B.X. revised the manuscript. All authors read and approved the final manuscript.

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Correspondence to Changli Zeng or Xuekun Zhang .

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12870_2024_5121_MOESM1_ESM.xlsx

Supplementary Material 1: Table S1: Physicochemical properties and subcellular localization prediction of BnXTH proteins.

Supplementary Material 2: Table S2: Table S2. XTH IDs of B. rapa and B. oleracea .

Supplementary material 3: table s3: collinearity of xth genes in b. napus , b. rapa , and b. oleracea ., supplementary material 4: table s5: collinearity of xth genes in b. napus ., supplementary material 5: table s4: collinearity of xth genes in b. napus ., rights and permissions.

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Chen, J., Wan, H., Zhao, H. et al. Identification and expression analysis of the Xyloglucan transglycosylase/hydrolase ( XTH ) gene family under abiotic stress in oilseed ( Brassica napus L.). BMC Plant Biol 24 , 400 (2024). https://doi.org/10.1186/s12870-024-05121-5

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Comprehensive identification of maize ZmE2F transcription factors and the positive role of ZmE2F6 in response to drought stress

• Yang Cao 1   na1 ,
• Kexin Wang 1   na1 ,
• Fengzhong Lu 1 ,
• Qingqing Yang 1 ,
• Bingliang Liu 2 ,
• Hayderbinkhalid Muhammad 3 ,
• Yingge Wang 1 ,
• Fengling Fu 1 ,
• Wanchen Li 1 &
• Haoqiang Yu 1  

BMC Genomics volume  25 , Article number:  465 ( 2024 ) Cite this article

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The early 2 factor (E2F) family is characterized as a kind of transcription factor that plays an important role in cell division, DNA damage repair, and cell size regulation. However, its stress response has not been well revealed.

In this study, ZmE2F members were comprehensively identified in the maize genome, and 21 ZmE2F genes were identified, including eight E2F subclade members, seven DEL subfamily genes, and six DP genes. All ZmE2F proteins possessed the DNA-binding domain (DBD) characterized by conserved motif 1 with the RRIYD sequence. The ZmE2F genes were unevenly distributed on eight maize chromosomes, showed diversity in gene structure, expanded by gene duplication, and contained abundant stress-responsive elements in their promoter regions. Subsequently, the ZmE2F6 gene was cloned and functionally verified in drought response. The results showed that the ZmE2F6 protein interacted with ZmPP2C26, localized in the nucleus, and responded to drought treatment. The overexpression of ZmE2F6 enhanced drought tolerance in transgenic Arabidopsis with longer root length, higher survival rate, and biomass by upregulating stress-related gene transcription.

Conclusions

This study provides novel insights into a greater understanding and functional study of the E2F family in the stress response.

Peer Review reports

Environmental stimuli, including drought, salinity, and high temperature, have frequently occurred in recent decades and led to yield loss of crops in agricultural production. Drought stress is a main misfortune in these abiotic stressors and will be a more severe challenge and threat to agriculture and humanity by 2050 [ 1 ]. In adaption to drought, plants activate a series of physiological, morphological, and molecular changes [ 1 , 2 , 3 ]. Among them, many transcription factors (TFs) can be dominated by stress to regulate gene expression and coordinate plant antagonism to adverse factors [ 4 , 5 , 6 ].

The early E2 factor (E2F) family proteins are initially discovered as TFs of the E2 gene and play key roles in cell proliferation control in adenoviruses [ 7 ]. E2Fs can be classified into typical and atypical E2Fs according to protein structure. Typically, E2Fs have only one DNA-binding domain (DBD) and form heterodimeric complexes with dimerization proteins (DPs) to bind the promoters of downstream genes, but atypical E2Fs possess duplicated two DBD [ 8 ]. In higher plants, E2Fs are also categorized into three subclades, including E2F, DP, and DEL (DP-E2F-like), due to the difference in the composition of conserved domains [ 9 , 10 ]. Over the past decade, E2Fs have well been revealed to play significant roles in the cell cycle and DNA damage repair [ 11 , 12 , 13 , 14 , 15 , 16 ]. For instance, ectopic expression of DcE2F1 of Daucus carota promotes cell proliferation in Arabidopsis seedlings [ 14 ]. In Arabidopsis , eight E2F members exhibit antagonistic roles in cell proliferation, such as E2Fa/b, which acts as a positive regulator, but E2Fc is a negative regulator [ 17 , 18 , 19 ]. E2Fa/b can also activate the DNA damage response and cell cycle progression by differentially regulating the expression of genes [ 11 ]. Additionally, AtE2Fa/DPa also inhibits growth, and AtE2Fa/DPa -overexpressing plants show an abnormal phenotype owing to ectopic cell division or enhanced DNA endoreduplication [ 12 , 20 ].

In addition to the crucial role in the cell cycle, few reports show that atypical E2F inhibits the accumulation of salicylic acid to balance growth and defense [ 21 , 22 ]. Furthermore, AtE2Fa acts downstream of ERECTA kinase, which is involved in cell size and stomatal density [ 23 ]. Via expression analysis, it is suggested that TaE2F-DP of wheat, PheE2F/DPs of Moso bamboo, E2F/DP genes of Medicago truncatula , and PvE2F/DPs of Phaseolus vulgaris respond to drought or salt stress, suggesting their potential roles in regulating stress tolerance [ 9 , 24 , 25 , 26 ]. To date, however, the role of E2F in plant stress tolerance remains obscure.

Maize is a crucial crop worldwide and is widely used as food and livestock feed. During its growth, maize plants show sensitivity to water deficit due to its high water demand, leading to maize yield being greatly affected by drought stress [ 27 , 28 , 29 , 30 ]. Therefore, it becomes imperative to identify and explore drought tolerance-related genes that can be used to enhance maize resilience through molecular breeding [ 31 , 32 , 33 , 34 , 35 ]. In our previous study, we found that ZmPP2C26 regulated drought tolerance [ 36 ] and targeted maize ZmE2F (Zm00001d048412, data not shown), indicating that ZmE2F might be involved in drought response. Hence, in this study, we comprehensively investigated ZmE2F genes in the maize genome. Thereafter, phylogenetic relationships, conserved motifs and domains, gene structures and duplication, and protein-protein interaction networks were analyzed. Additionally, the ZmE2F6 (Zm00001d048412) was functionally verified by performing subcellular localization, expression patterns in drought treatment, and ectopic expression in Arabidopsis under drought stress. This study will significantly contribute to a better understanding of E2Fs in stress response.

Identification of ZmE2Fs in the maize genome

The genome and amino acid data of maize B73 were downloaded from the MaizeGDB database ( https://download.maizegdb.org/Zm-B73-REFERENCE-GRAMENE-4.0/ ). Meanwhile, the coding sequences and amino acid sequences of 8 AtE2Fs and 9 OsE2Fs of Arabidopsis and rice were downloaded from the Arabidopsis Information Resource (TAIR) ( https://www.arabidopsis.org/ ) and the Rice Genome Annotation Project (RGAP) database ( http://rice.uga.edu/ ) and were used as queries to perform local BLASTp with an E-value of 1e − 10 in the maize protein database for maize E2F searching, respectively. After removing the redundant sequences manually, the candidate sequences were further analyzed for the presence of the E2F_DP domain (PF02319) by using PFAM ( http://pfam.xfam.org/ ). The candidates possessing the E2F domain were identified as maize ZmE2F members. The secondary structure and physicochemical properties, including molecular weights, isoelectric point (PI), stability coefficient, and grand average of hydropathicity (GRAVY), of ZmE2Fs were analyzed using SOPMA ( https://npsa.lyon.inserm.fr/cgi-bin/npsa_automat.pl?page=/NPSA/npsa_sopma.html ) and EXPASY ( https://www.expasy.org/ ). The subcellular localization of ZmE2Fs was predicted using cNLS Mapper ( https://nls-mapper.iab.keio.ac.jp/cgi-bin/NLS_Mapper_form.cgi ).

Conserved motifs, domains, and phylogenetic analysis

To further identify the conserved motifs and domains, the amino acid sequences of ZmE2Fs were analyzed using MEME ( http://meme-suite.org/tools/meme ) and NCBI-CDD ( https://www.ncbi.nlm.nih.gov/cdd ), respectively. The motif and domain composition of each ZmE2F was visualized by TBtools [ 37 ]. The protein sequences of all ZmE2F, AtE2F, and OsE2F were multiple-aligned using ClustalW with default parameters. The maximum likelihood tree was built with 1000 bootstrap replications by MEGA11 ( https://www.megasoftware.net/ ). Meanwhile, protein-protein interaction (PPI) analysis among ZmE2F members was performed using the STRING tool [ 38 ].

Gene structure, promoter, duplication, and synteny analyses

The chromosomal location of each ZmE2F gene was obtained from the maizeGDB database. The coding sequences and genomic DNA sequences of every ZmE2F were downloaded and used to analyze exon-intron composition using Gene Structure Display Server 2.0 (GSDS) ( http://gsds.gao-lab.org/ ). The 2000 bp upstream sequence of the transcription start site of each ZmE2F gene was retrieved from maizeGDB and used for cis -acting element analysis using PlantCARE ( https://bioinformatics.psb.ugent.be/webtools/plantcare/html/ ). Meanwhile, the gene duplication events and the synteny relationship between ZmE2F , AtE2F , and OsE2F gene members were analyzed using MCScanX with default parameters. The chromosomal location, duplications, and synteny relationships of the ZmE2F , AtE2F , and OsE2F genes were visualized using TBtools [ 37 ]. The non-synonymous (Ka) and synonymous (Ks) substitution rates per site of the duplicated gene pairs were calculated using TBtools [ 37 ]. Then, the divergence time in millions of years (Mya) was also calculated using the following formula: T = Ks/2λ × 10 − 6 Mya (λ = 6.5 × 10 − 9 for grasses) [ 39 ].

Cloning and subcellular localization of the ZmE2F6 gene

The specific primers (Table S1 ) were designed by Primer 5.0, synthesized at Tsingke Biotech (Beijing, China), and used to amplify the sequence of the ZmE2F6 gene from maize B73 cDNA using Phanta Max Super-Fidelity DNA Polymerase (Vazyme, Nanjing). After amplification, the PCR product was purified by a gel recovery kit subcloned, and inserted into the pMD19-T vector to generate pMD19-T- ZmE2F6 and verified by sequencing. The sequencing result was aligned with the candidate sequence of the ZmE2F6 gene using DNAMAN. The open reading frame (ORF) sequence of ZmE2F6 without stop codon was amplified from the pMD19-T- ZmE2F6 plasmid using the specific primers designed by CE Design V1.04 (Table S1 ) with the Xba I and Spe I recognition sites. The PCR products and pCAMBIA2300- 35 S-eGFP plasmids were digested using Xba I and Spe I. Subsequently, they were inserted into the Xba I and Spe I sites of pCAMBIA2300- 35 S-eGFP to produce the fusion expression vector 35 S-ZmE2F6-eGFP using the ClonExpress II One Step Cloning Kit (Vazyme, Nanjing). Each construct was introduced into Agrobacterium tumefaciens strain GV3101 and then used for transient expression in the leaves of Nicotiana benthamiana . As described by Sun et al. [ 40 ], the constructs were infiltrated into the leaves of five-week-old N. benthamiana . The GFP fluorescence was observed and imaged using a confocal laser scanning microscope (Zeiss 800). The empty vector 35 S-eGFP was served as the positive control.

Plant treatment, RNA extraction, and quantitative real-time PCR (qRT‒PCR) analysis

The seeds of maize B73 lines were soaked in 10% H 2 O 2 for 15 min, rinsed twice using sterile water, soaked in distilled water for 8 h, then wrapped in filter paper and cultured at 28 °C until germination. Subsequently, the seedlings of the same size were transferred to the hydroponic cassette containing hoagland nutrient solution and cultured at 16 h light at 28 °C /8 h dark at 24 °C. Three-leaf-stage seedlings were subjected to 16% PEG-6000 treatment mimicking drought stress, and then shoots containing leaf, stem and leaf sheath, and roots were sampled at 0, 3, 6, 12, and 24 h of treatment, respectively. The total RNA of each sample was extracted using an RNAison plus kit (Takara, Dalian), examined for quality using NanoDrop OneC (ThermoFisher Scientific), treated with DNase to remove DNA contamination, reverse-transcribed into cDNA using a PrimeScript™ RT Regent Kit (Takara, Dalian), and used to perform qRT-PCR. The specific primers of ZmE2F6 were designed using Primer-BLAST ( https://www.ncbi.nlm.nih.gov/tools/primer-blast/index.cgi?LINK_LOC=BlastHome ), synthesized at TsingkeBiotech (Beijing, China), and listed in Table S1 . The ZmGAPDH gene was amplified using specific primers (Table S1 ) and used as an internal reference. The qRT-PCR was performed in the Bio-Rad CFX96™ Real-Time PCR system using 2 × Universal SYBR Green Fast qPCR Mix (ABclonal, Wuhan). The 20 µL reaction mixture contained 10 µL of 2 × Universal SYBR Green Fast qPCR Mix, 0.4 µL of each forward and reverse primer, 1.0 µL of each cDNA as template, and 8.2 µL of ddH 2 O. The reaction protocol was set as a two-step temperature cycle including 95 °C for 3 min, followed by 40 cycles at 95 °C for 5 s and 60 °C for 30 s. The relative expression level of ZmE2Fs was calculated and normalized using the 2 −ΔΔCt method [ 41 ].

Y2H and GST pull-down analysis

The ORF of ZmE2F6 was amplified using specific primers designed by CE Design V1.04 with the Nde I and EcoR I recognition sites (Table S1 ) and inserted into the pGADT7 plasmid to generate AD -ZmE2F6 as described above. The BD- ZmPP2C26 plasmid was constructed in our previous study [ 36 ]. The AD- ZmE2F6 and BD- ZmPP2C26 plasmids were cotransformed into the yeast strain Y2H Gold using a yeast transformation kit (Coolaber, Beijing). Subsequently, yeast cells were cultured on synthetic dropout (SD) medium without Trp and Leu (SD/-Trp/-Leu) at 30℃ for 2 days, and then positive clones were transferred onto SD/-Trp/-Leu/-His/-Ade plates with X-α-gal and cultured at 30℃ for 2 days. Meanwhile, the ORF of ZmE2F6 was amplified and inserted into the EcoR I and Xho I sites of pGEX-6P-1 to generate GST - ZmE2F6 and used for the GST pull-down assay. The His - ZmPP2C26 plasmids were produced in our previous study. The GST pull-down was performed as described by Lu et al. [ 36 ].

Plant transformation, phenotyping, and RNA-seq

The ORF of ZmE2F6 was amplified without a stop codon and inserted into the Xba I and Nde I sites of pRI201- 35 S - GUS to produce 35 S - ZmE2F6 - GUS as described above. The 35 S - ZmE2F6 - GUS construct was transformed into Agrobacterium tumefaciens strain GV3101 and then used to transform Arabidopsis thaliana (Col-0) by the floral-dip method [ 42 ]. According to the method of Sun et al. [ 40 ], the positive transformants were screened on 1/2 MS plates with 50 mg/L kanamycin, used for harvesting seeds individually. The homozygous lines without segregation on 1/2 MS plates with 50 mg/L kanamycin were screened and used for PCR detection. Meanwhile, the leaves of homozygous lines were sampled and used to perform GUS staining using the GUS Staining Kit (Coolaber, Beijing).

According to the methods described by Sun et al. [ 40 ] with minor modifications, for drought stress, the seeds of homozygous lines and wild type (WT) were surface-sterilized, planted on 1/2 MS plates supplemented with 0 (control), 150, and 250 mM mannitol, vernalized for 2 days in the dark at 4 °C, and vertically cultured in a chamber under 10 h light/14 h dark at 22 °C with 60–70% humidity. At 14 days of treatment, the seedlings were photographed and measured for root length. Moreover, another batch of overexpressed lines and WT were sown in soil and incubated in a greenhouse under the same conditions. After 2 weeks, the seedlings were subjected to withholding water for two weeks, then rewatered for at least 2 days, and monitored for phenotyping. Subsequently, the survival number of each line was counted and used to calculate the survival rate. The leaf of each line was sampled, dried at 80 °C for 3 days, and used for biomass measurement.

Meanwhile, three-week-old seedlings of homozygous lines and WT were sampled and used for RNA sequencing at Sanshubio Company (Jiangsu, China). As described by Sun et al. [ 40 ], the total RNA of each sample was extracted, qualified for quality and integrity, and used to construct a sequencing library. Then, library sequencing was conducted using the NovaSeq 6000 system. The sequencing adapters and low-quality reads of raw data were removed to generate clean data, which were mapped to the Arabidopsis genome by hisat2 [ 43 ] and used for assembling transcripts of every gene using StringTie [ 44 ]. The differentially expressed genes (DEGs) were confirmed with a p-value < 0.05 and |FoldChange| > 2 using DESeq2 [ 45 ]. The GO analysis of DEGs was performed using KOBAS [ 46 ].

Data analysis

All assays were performed with three replicates. The data are shown as the mean values ± standard error (SE). The significance was analyzed by Student’s t-test at the p  < 0.05 or p  < 0.01 level.

ZmE2F members in maize

To identify the maize ZmE2F family, a local BLASTp search against the maize protein database was performed using the amino acid sequences of 8 AtE2F and 9 OsE2F members as query references [ 10 , 47 ]. As shown in Table  1 , a total of 21 ZmE2F members were identified and designated as ZmE2F1 to ZmE2F21 . The CDS length of the ZmE2F genes ranged from 648 ( ZmE2F18 ) to 1608 bp ( ZmE2F14 ), encoding 215 to 535 amino acids with molecular weight varying from 23.39 to 59.21 kDa. The theoretical isoelectric point (pI) of ZmE2F proteins ranged from 4.71 (ZmE2F11) to 9.41 (ZmE2F13), and the grand average hydropathicity (GRAVY) of all ZmE2F proteins was less than 0 and ranged from − 0.062 (ZmE2F1) to -0.603 (ZmE2F5). The instability indices of ZmE2F proteins varied from 31.45 (ZmE2F20) to 62.39 (ZmE2F5), and 19 ZmE2F members contained the highest random coil in their secondary structure, implying that they were unstable hydrophilic proteins. All ZmE2F proteins were predicted to be localized in the nucleus.

Phylogenetic analysis

To investigate the evolutionary relationships of ZmE2Fs and model plants, a total of 38 amino acid sequences, including 21, 8, and 9 E2Fs from maize, Arabidopsis , and rice, respectively, were used to construct a phylogenetic tree (Fig.  1 ). It showed that the ZmE2F proteins could be classified into three subfamilies, including E2F, DP, and DEL clades [ 9 ]. Eight members, including ZmE2F2, 4, 5, 11, 14, 16, 17, and 21, were clustered into the E2F subscale with AtE2Fs and OsE2Fs. Seven proteins containing ZmE2F1, 7, 10, 12, 13, 15, and 20 were branched into the DEL subfamily with AtDELs and OsDELs. The other 6 ZmE2Fs (ZmE2F3, 6, 8, 9, 18, and 19) belonged to the DP subclade with AtDP and OsDP members. In addition, ZmE2Fs showed a closer phylogenetic relationship with OsE2Fs than with AtE2Fs, indicating more sequence similarity with OsE2Fs.

Phylogenetic tree of the E2F family. The E2Fs of Arabidopsis , rice, and maize are marked in blue, red, and black, respectively

Conserved motif and domain

Five conserved motifs were identified in the amino acid sequences of ZmE2Fs using the MEME online program (Fig.  2 A). Among them, motif 1 was highly conserved in all ZmE2F members and characterized by the RRIYD sequence that was a DNA-binding motif followed by dimerization residues DNVLE sequence [ 48 ]. Conserved domain analysis revealed that all ZmE2F members contained at least one DNA-binding domain (DBD) (Pfam ID PF02319, Table S2 ). ZmE2F7, 12, and 13 possessed two DBDs (Fig.  2 B). In addition, ZmE2F5, 6, 9, 18, and 19 also contained a dimerization domain (DD). The coiled coil (CC)-marked box (MB) domain (CC-MB) was found in ZmE2F4, 11, 16, and 17. Furthermore, all ZmE2F members contained a nuclear localization signal.

Conserved motif and domain of ZmE2Fs

ZmE2Fs protein-protein Interaction Prediction

Due to the presence of dimerization residues DNVLE sequence within motif 1 in every ZmE2F and DD or CC-MB in some ZmE2Fs, to explore the potential interactions among ZmE2F members, protein-protein interaction (PPI) analysis was performed. As shown in Fig.  3 , ten ZmE2Fs were predicted to interact with each other, which generated 23 PPI combinations. Among them, ZmE2F6, 9, 18 and 19 had the largest number of PPIs (6 interactions), while ZmE2F4, 5, 11, 14, 16, and 17 had 4 PPIs. These results imply that the DNVLE sequence, DD, and CC-MB play crucial roles during ZmE2F dimerization.

ZmE2F protein-protein interaction prediction

Gene structure, duplication, synteny, and cis -acting elements

Gene structure analysis revealed that the ZmE2F gene family showed diversity in exon and intron composition (Fig.  4 A). The number of exons among ZmE2Fs ranged from six to fourteen. Except for ZmE2F18 and ZmE2F20 , the other 19 ZmE2Fs contained 5ʹ or 3ʹ terminal untranslated regions. The ZmE2Fs were unevenly distributed on eight maize chromosomes, excluding chromosomes 3 and 7. Gene duplication analysis showed that there were eight paralogous pairs of ZmE2Fs in the maize genome, including ZmE2F4 and ZmE2F17 , ZmE2F6 and ZmE2F18 , ZmE2F6 and ZmE2F19 , ZmE2F7 and ZmE2F12 , ZmE2F7 and ZmE2F13 , ZmE2F11 and ZmE2F17 , ZmE2F12 and ZmE2F13 , and ZmE2F18 and ZmE2F19 . Likewise, gene synteny analysis revealed one and twenty-two orthologous pairs between ZmE2F and AtE2F and ZmE2F and OsE2F , respectively (Fig.  4 B; Table S3 ). Meanwhile, the Ka/Ks rates among ZmE2Fs paralogous pairs ranged from 0.19 to 0.35 and their duplication time was estimated to be 16.54–139.30 million years ago (Table S4 ). The results suggest that gene duplication contributed to E2F expansion during the evolutionary process.

Gene structure ( A ) and duplication ( B ) of ZmE2Fs. Yellow and blue boxes represent exons and untranslated regions, respectively. Black lines represent introns. Yellow, blue, and red boxes with a number represent chromosomes of maize, Arabidopsis , and rice, respectively. The green, blue, and red lines indicate duplicated E2F gene pairs among maize, as well as between maize and Arabidopsis and rice, respectively

Furthermore, cis -acting element analysis revealed that abundant stress- or hormone-responsive elements were found in the promoter regions of ZmE2Fs , such as ABREs, MBSs, AREs, and LTR elements (Table S5 ), which were responsive to abscisic acid, drought, anaerobic induction, and low temperature. For instance, ABREs were found in all ZmE2F promoters. MBS elements, the MYB binding site involved in drought response, were found in 15 ZmE2F promoters. This finding implies that the ZmE2F genes may be involved in abiotic stress responses.

ZmE2F6 interacts with ZmPP2C26

In our previous study, ZmE2F6 was identified as a potential target of two ZmPP2C26 splicing variants (ZmPP2C26L/S) via Y2H library screening. Hence, to verify whether ZmE2F6 interacts with ZmPP2C26, Y2H, and GST pull-down assays were performed. As shown in Fig.  5 , the yeast cells transformed with AD- ZmE2F6 and BD- ZmPP2C26L/S exhibited normal growth and turned blue on SD/-Leu/-Trp/-His/-Ade plates containing X-α-gal. Moreover, the GST-ZmE2F6 protein could be pulled down by His-ZmPP2C26 L/S in the GST pull-down assay. These findings confirmed the interaction between ZmE2F6 and ZmPP2C26L/S. Likewise, there were 7 potential phosphorylated sites in ZmE2F6 amino acid sequences (Table S6 ) were predicted using NetPhos-3.1 ( https://services.healthtech.dtu.dk/services/NetPhos-3.1/ ).

ZmE2F6 interacts with ZmPP2C26. ( A ) Yeast two-hybrid (Y2H). ( B ) GST pull-down. ZmPP2C26L and ZmPP2C26S represent two splicing variants of ZmPP2C26. The combination of AD-T with BD-53 and AD-T with BD-lam were used as negative and positive controls, respectively

ZmE2F6 localized to the nucleus

To further validate the cellular localization of the ZmE2F6 protein, the ORF of ZmE2F6 was cloned and inserted into the 35 S - eGFP plasmid to fuse with eGFP . The results of transient expression in tobacco leaves revealed that fluorescent signals were observed in whole cells, including the nucleus and cytoplasm, transformed by the 35 S - eGFP empty vector. However, in tobacco leaves transformed with 35 S - ZmE2F6 - eGFP , fluorescent signals were exclusively detected in the nucleus (Fig.  6 ). This observation is consistent with the bioinformatic prediction, confirming that the ZmE2F6 transcription factor is solely localized in the nucleus.

Subcellular localization of ZmE2Fs

The expression of ZmE2F6 was induced by drought stress

The qRT-PCR results demonstrated that the expression of the ZmE2F6 gene was responsive to drought stress (Fig.  7 ). After drought treatment, the expression of ZmE2F6 in maize shoots was significantly upregulated and reached approximately 11-, 8-, 100-, and 47-fold that of the control at 3, 6, 12, and 24 h of treatment. In roots, the expression of ZmE2F6 was significantly upregulated after 3 h of drought treatment and then downregulated at 6, 12, and 24 h of treatment. These results suggest that ZmE2F6 responds to drought stress.

The relative expression level of the ZmE2F gene under drought stress

Expression of ZmE2F6 enhanced drought tolerance in Arabidopsis thaliana

To assess the function of ZmE2F6 in regulating drought tolerance, we generated transgenic Arabidopsis lines overexpressing ZmE2F6 . In the T 1 generation, ten positive transgenic lines were screened by kanamycin on 1/2 MS plates, identified by PCR, and harvested to produce the next generation. Finally, in the T 3 generation, two homozygous lines (OE6-5 and OE6-10) were selected, identified by GUS staining, and used for phenotyping (Fig.  8 A). It was found that the root length of the OE6-5 and OE6-10 lines was significantly longer than that of the WT under 1/2 MS plates or supplemented with 150 mM mannitol (Fig.  8 B, C). Furthermore, natural drought treatment in soil was performed to monitor the drought tolerance of the OE6-5 and OE6-10 lines. The results showed that there was no difference between the transgenic lines and WT before treatment. Subsequently, after two weeks of drought treatment, WT plants were seriously wilting, but OE6-5 and OE6-10 showed slightly inhibited phenotypes. After 3 days of rewatering, the OE-65 and OE6-10 lines exhibited significantly higher survival rates and biomass, indicating that overexpression of ZmE2F6 contributes to enhancing drought tolerance in transgenic Arabidopsis and that ZmE2F6 positively regulates drought tolerance.

The phenotype of transgenic Arabidopsis under drought stress. ( A ) PCR detection and GUS staining of transgenic lines. ( B ) The phenotype of transgenic lines on 1/2 MS plates with mannitol. ( C ) Root length. ( D ) The phenotype of transgenic lines in soil. ( E ) Survival rate and biomass of each line. OE6-1 to OE6-10 represent transgenic lines overexpressing ZmE2F6 . WT, wild type

To investigate the impact of ZmE2F6 overexpression on endogenous genes in Arabidopsis thaliana , RNA-Seq was conducted on four transgenic lines, OE6-5, OE6-10, and WT. The results showed that there were 657 DEGs in transgenic lines compared to WT. Totally, 19 DEGs were identified in two transgenic lines. Among them, 15 DEGs were upregulated in transgenic lines, including AtMYB44 (AT5G67300), AtB1L (AT1G18740), AtJAZ7 (AT2G34600), AtEXS (AT1G35350), AtIP5PII (AT4G18010), AtPATL2 (AT1G22530), AtAZI1 (AT4G12470), AtXTH23 (AT4G25810), and AtEXL5 (AT2G17230) (Fig.  9 ). GO analysis showed that these DEGs were associated with stress responses (Figure S1 ).

The expression level (Log2 Fold change) of differentially expressed genes (DEGs) in each line

In eukaryotes, the ZmE2F family is characterized as a TF that plays crucial roles in cell division, DNA repair, and differentiation [ 11 , 13 , 14 , 15 , 16 ]. However, the E2F family is only genome-wide identified in a few plants, including Arabidopsis , rice, wheat, Moso bamboo, Medicago truncatula , and Phaseolus vulgaris [ 9 , 24 , 25 , 26 ]. In the present study, 21 ZmE2F members were identified in the maize genome (Table  1 ) and classified into E2F, DP, and DEL subclades (Fig.  1 ), which was higher than the number of E2F members identified in Arabidopsis (8), rice (9), Medicago truncatula (5), and Phaseolus vulgaris (7) but close to the number in wheat (27) and Moso bamboo (23), owing to their comparable genome size and gene duplication being a major driving force of gene families [ 25 , 49 ]. Likewise, eight paralogous pairs of ZmE2Fs and twenty-three orthologous pairs of E2Fs among maize, Arabidopsis , and rice were found (Fig.  4 ). All the Ka/Ks ratios of the ZmE2Fs paralogous pairs were < 1 (Table S4 ), indicating their duplication evolved under purifying selection [ 50 ]. A similar phenomenon was consistently observed in the PheE2F/DP gene family [ 25 ]. A previous study showed that there are 12 ZmE2F genes in maize [ 51 ], which is much lower than the number identified in this study. This may be due to their preliminary BLAST search using maize genome release 5b.60 and not a comprehensive analysis [ 51 ].

All ZmE2F proteins possess at least one DBD characterized by motif 1 containing the RRIYD sequence and dimerization residue DNVLE sequence (Fig.  2 ), which contributes to the binding of E2F and DNA as a homodimer or as a heterodimer with its dimerization partner DP. In addition, some ZmE2F proteins also have DD and CC-MB domains, which promotes their formation of heterodimers to regulate downstream genes [ 52 ]. As a result, ten ZmE2Fs are predicted to interact with each other and generate 23 PPI combinations (Fig.  3 ), owing to the presence of the DNVLE residues, DD, or CC-MB domain in these proteins.

To date, the function of E2F in regulating plant stress tolerance remains unknown, although few reports have shown that the expression of some E2F genes is responsive to stress [ 9 , 24 , 25 , 26 ]. In our study, abundant stress-responsive acting elements were found in ZmE2F promoter regions, such as ABREs and MBSs, suggesting the response of ZmE2Fs to stress and the potential roles of ZmE2Fs . In our previous study, the ZmPP2C26 gene, a B clade of maize PP2C members, was found to be responsive to drought stress and negatively regulated drought tolerance in Arabidopsis , rice, and maize [ 36 ] and targeted on maize ZmE2F6 (Zm00001d048412), indicating its function in the drought response. Hence, the ZmE2F6 gene was cloned and functionally validated. It is found that the ZmE2F6 interacts with two splicing variants of ZmPP2C26 but localized in the nucleus (Figs.  5 and 6 ), suggesting that ZmPP2C26 physically targets ZmE2F6. In Arabidopsis , it has been previously reported that B clade PP2C (AP2C1) dephosphorylates the autophosphorylated form of CBL-interacting protein kinase 9 (CIPK9) to regulate root growth, seedling development, and stress tolerance (low-K + ) [ 53 ]. Previous studies showed that BES1/BZR1 TFs are phosphorylated and degraded but moved to the nucleus after dephosphorylation in the cytoplasm [ 54 , 55 ]. It is proposed that ZmPP2C26 might dephosphorylate ZmE2F6 in the cytoplasm, which needs to be further revealed in our next study.

The qRT-PCR results showed that the ZmE2F6 gene was induced by drought stress (Fig.  7 ), which could be explained by the presence of three MBS and two ABREs in the ZmE2F6 promoter (Table S5 ). It’s confirmed that MYB transcription factors bind to the MYB binding sites (MBS) of nuclear gene promoters to adjust their transcription and regulate drought tolerance in plants [ 56 , 57 ]. Likewise, ABRE (ABA-responsive element), the major cis -element for ABA-responsive gene expression, is targeted by ABRE-binding protein (AREB) or ABRE-binding factor (ABF) TFs to regulate the drought response via the ABA signaling pathway [ 58 ]. These findings further imply the regulation of ZmE2F6 in drought tolerance.

After overexpressing ZmE2F6 in Arabidopsis , the transgenic lines showed longer root lengths than the WT (Fig.  8 B, C), which is consistent with the fact that E2Fs regulate root growth in Arabidopsis [ 19 , 22 , 59 ]. The elevated root growth of ZmE2F6 -overexpressing lines may confer osmotic tolerance because roots can respond to moisture and coordinate responses to drought [ 1 ]. The ectopic expression of ZmE2F6 enhances drought tolerance in transgenic Arabidopsis (Fig.  8 ). We found that some stress-related genes were significantly upregulated in the transgenic lines, such as AtMYB44 , AtB1L , AtJAZ7 , AtEXS , AtIP5PII , AtPATL2 , AtAZI1 , AtXTH23 , and AtEXL5 (Fig.  9 ), which are well known to regulate abiotic stresses [ 60 , 61 , 62 , 63 , 64 , 65 , 66 , 67 , 68 ]. The ZmE2F6 protein acts as a TF and localizes in the nucleus to regulate the expression of these genes (Fig.  6 ). However, the molecular mechanism regulated by ZmE2F6 in maize remains unknown.

Overall, in this study, a comprehensive analysis of the ZmE2F gene family was performed. In total, 21 ZmE2F TFs were identified in the maize genome and divided into three subclades. All ZmE2F proteins possessed at least one DBD characterized by the RRIYD (DNA binding motif) and DNVLE (dimerization residues) sequences. The ZmE2F genes showed diversity in gene structure, expanded by gene duplication, and contained abundant stress-responsive elements in their promoter regions. Then, the ZmE2F6 gene was cloned and functionally verified in the drought response. The ZmE2F6 protein interacted with ZmPP2C26, localized in the nucleus, and responded to drought treatment. The overexpression of ZmE2F6 enhanced drought tolerance in transgenic Arabidopsis by upregulating stress-related gene transcription. This study sheds light on the role of ZmE2F in the drought response and provides novel insights into a greater understanding of the E2F family in crops.

Data availability

The RNA-seq datasets generated during the current study are available in the NCBI repository under BioProject accession number PRJNA1028664.

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This study was supported by the National Key R&D Program of China (2021YFF1000303), the Sichuan Science and Technology Program (2022YFH0067) and the National Natural Science Foundation of China (32102226).

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Yang Cao, Kexin Wang, Fengzhong Lu, Qi Li, Qingqing Yang, Yingge Wang, Fengling Fu, Wanchen Li & Haoqiang Yu

College of Food and Biological Engineering, Chengdu University, Chengdu, 610106, China

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Y.C., K.W. and F.L. designed and carried out the experiments; Y.C., K.W., B.L., H.M. and Y.W. carried out all bioinformatics analysis and wrote the manuscript; Q. Y. provided technical support. F.F., W.L. and H.Y supervised the experiments; H.Y. directed and revised the manuscript. All authors reviewed and approved the final manuscript.

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Cao, Y., Wang, K., Lu, F. et al. Comprehensive identification of maize ZmE2F transcription factors and the positive role of ZmE2F6 in response to drought stress. BMC Genomics 25 , 465 (2024). https://doi.org/10.1186/s12864-024-10369-0

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