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Selected students from spring 2024 courses in creative writing read from their work in fiction, poetry, screenwriting and literary translation as part of the Althea Ward Clark W’21 Reading Series.

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Seniors in Princeton’s renowned Program in Creative Writing read from the screenplays or collections of poems or translations written as their senior independent work. No tickets required.

2024 Senior Readings in Fiction

Seniors in Princeton’s renowned Program in Creative Writing read from the novels and collections of short stories written as their senior independent work. No tickets required.

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Reimagining writing seminar for all writers

Entryway to New South.

Tiffany tsai / the daily princetonian.

Few Princeton students can forget their writing seminars. From the stress and confusion over the D1 to the feeling of accomplishment unique to finally submitting the R3, the mandatory first-year course provides a shared introduction to all the mixed emotions that will define every Princeton student’s future academic pursuits. However, while writing seminars teach helpful research and argumentation tactics, they often ultimately fall short of helping students with their titular skill: writing. Currently, writing seminar utilizes a one-size-fits-all approach to writing, neglecting the teaching of how different forms, types of sources, and modes of analysis vary in different disciplines. It also does not teach its students how to make the best possible use of language, especially in regards to form and style. In other words, writing seminar fails at its most fundamental task: it does not teach its students how to write, nor how to do so well.

While the time limits of Princeton’s short semester — significantly briefer than that of other schools like Harvard — could limit the ability of instructors to teach broader concepts, there still exists a structural issue with writing seminar that makes its programming fundamentally ineffective. Namely, none of the three major assignments in the writing program are especially dedicated to form or style. The first paper is dedicated to learning how to analyze a single work in detail, while the second essay teaches students how to effectively compare arguments and build their own. The third paper is completely focused on fostering independent research techniques and strong engagement with a variety of primary, secondary, and tertiary sources. While these assignments are invaluable for teaching students how to form strong research papers guided with sufficient evidence, none provide the essential skills of form & style required of a good writer. 

Rarely does the writing seminar curriculum explicitly teach students how a purposeful, well-thought-out structure can effectively communicate ideas. After all, without structure, arguments simply don’t make sense. Future professors will evaluate lab scientific evaluations, close readings, and other varying assignments with their discipline-individual structures in mind because, in academic writing, coherency to a certain audience is key. The writing seminar program should set students up for success at Princeton — and in the wider world of publishing — by placing a greater emphasis on the many different kinds of idea organization in analytical writing. 

Form — which goes beyond the organization of ideas, and encompasses the text type and media used to convey an argument — also does serious work to get an author’s point across clearly. Teaching only one form of the “academic paper” neglects how helpful it can be to expose freshmen to the many forms analysis can take across fields. A lab report, a poetic analysis, and a policy memo are all forms of academic writing whose form will diverge far from the source-critique-conclusion style of essay taught in writing seminars.

The present design of writing seminar also prevents students from expanding their idea of what sources from different disciplines may look like. It focuses mainly on engaging with scholarly secondary and tertiary sources and how a thesis can be formed on external research . Although the D1/R1 does focus on primary source analysis, students spend the majority of their time simply figuring out how to write an essay instead of engaging deeply with their primary sources of choice. Additionally, because many writing seminars focus on sociological or anthropological scholarship without incorporating the works favored in other disciplines, which call for different sourcing and analytical techniques.

Humanities disciplines that involve rigorous analysis of various texts may require skills that call for a different type of analysis beyond what is offered in the writing seminar curriculum. When you analyze a novel strictly through the skills taught in writing seminar  you tend to miss out on the creative and cultural nuances of  authors’ intents and perspectives. While some seminars certainly incorporate these creative works, it’s often done at a limited, marginal level.

Furthermore, in fictional works, when one only engages with the arguments made about a character in a novel can exclude the literary context in which the character lives, and the fact that their written existence in relation to a larger cultural moment influences the author’s purpose in creating such a character at all. Analyzing other writers’ style and purpose — especially in literary works — becomes much more difficult when you haven’t learned that those are viable elements for academic investigation. Journalistic scholarly pieces are not the only argumentative works. To focus almost exclusively on published academic sources is to ignore entire fields of work ripe for analysis and leads to the exclusion or minimization of other forms of writing, and accordingly subjective, creative analysis of one body of work. 

Lastly, writing seminar also ignores the importance of style — a part of writing can greatly enhance the ability of a piece to read fluidly and engagingly. Except for passing warnings about personal style, style is excluded from the curriculum. But style is important: clarity and concision make pieces easy to read, and personal voice can make it interesting. If you don’t understand how to effectively develop and adapt your own style to different disciplines, your work will be far less engaging. This might mean professional consequences down the line, like low readership and decreased citability. 

A significant part of understanding the proper use of style comes from the ability to write and convey meaning — especially through subtle manners. The focus on directly stated arguments during writing seminars neglects the various ways that an author might further more subtle points through tone, syntax, and figurative language. Developing these skills in your own writing makes it far easier to critically examine them in other people’s work.

The very specific style of academic writing favored in writing seminars makes abundant use of the passive voice, and favors long, clause-heavy sentences with complex grammatical structures. I call this “dissertation-ese”: a language which exists to remind the reader that the author is smart. But consequently, it can be difficult to read, and the clarity of the argument can get easily lost in excessive vocabulary. This is not to imply that complex ideas don’t require idiosyncratic constructions and polysyllabic words, but rather that first-years ought to focus more on clarity as they learn the basics of argumentative writing. By holding up wordy papers as paragons of great style encourages students to mimic that same wordiness in their own work, but often at the expense of clarity. If a student is struggling to understand the fundamental points an article is making, the addition of a functional language barrier via forced loquacity does not help the situation. A simply-written paper that can be clearly understood will be much more comprehensive than one that “sounds impressive.”

Writing seminar doesn’t need to be completely overhauled, and we don’t need to compromise the high curriculum standards of the writing program. However, there are certainly key modifications that could fix these problems and make our writing and research education more well balanced. 

First, there should be a stronger focus on forming independent arguments that are informed by, but not wholly reliant on, existing scholarly discourse around a topic. In particular, an enthusiastic promotion of literary analysis can help expand students’ horizons and ideas of what analytical writing can look like. Second, there must be a greater emphasis on structural analysis as opposed to purely argument critiques. A curriculum that is designed to teach students about writing structures in different academic fields will benefit students in every discipline. Lastly, style should be emphasized in the curriculum: how to develop and hone both the subtle, creative aspects of academic writing, and the power of simplicity in service of a more interesting, widely accessible paper. Any of the above will help the writing seminar program best prepare students for larger and more complex assignments — both now, and in future academic endeavors. 

Anna Ferris is a sophomore in the English department. She can be reached at [email protected].

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Princeton Correspondents on Undergraduate Research

creative writing

Writing lessons from my creative writing workshop.

This semester, I took my first fiction workshop in Princeton’s Creative Writing Program. I had taken two poetry courses in previous semesters and wanted to try something new. (Pro-tip: if you haven’t yet taken a CWR course, definitely consider applying for one before graduating).

Creative writing is, in many ways, a break from academic writing. It does not center on data, analysis, or argumentation. Instead, workshops focus on developing compelling images, characters, stories. Creative writing also has access to a wider variety of forms than academic writing, which tends to adhere to a narrow set of relatively conservative styles.

However, some of my workshop instructor’s writing advice has translated well to my academic writing. After all, writing is writing, and many of the same challenges confront both creative and academic writers. Below I’ve collected five of her best pieces of writing advice:

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Post Princeton Life: Interview with Isabelle Laurenzi ’15

For this year’s Spring Seasonal Series, entitled Post-Princeton Life: The Experiences of  PCUR Alumni, each correspondent has selected a PCUR alum to interview about what they have been up to. We hope that these interviews will provide helpful insight into the many different paths Princeton students take after graduation. Here, Raya shares her interview.

Teaching, travel, Congress, the Writing Center, political theory, Yale! Former PCUR chief correspondent Isabelle Laurenzi graduated from Princeton in 2015 with a degree in Religion. She has since gone on to pursue an array of adventures and projects. Most recently, Isabelle completed her first year of a Ph.D. program at Yale in political theory. For our seasonal spring series, I caught up with Isabelle to learn more about her time at Princeton and explorations after. In our conversation, Isabelle and I connected over our shared interest in interdisciplinary studies and the joy of pursuing one’s interests through varied avenues.

Continue reading Post Princeton Life: Interview with Isabelle Laurenzi ’15

Writing for Fun? (Part 2): Journalism and Academic Writing

In my last post, I started an exploration of writing on campus to understand how students approach the writing process outside the classroom in their own work and in extracurriculars. In that post, I considered creative writing and the ways academic writing can present a similar opportunity for expression and creativity.

In this post, I interview Sam Shapiro ’21 who is a Features Editor and writer for the Daily Princetonian. In my interview with Sam, we discussed the differences and similarities between journalism and academic writing and how to bring the thrill one feels when chasing a story for a publication to a term paper in class. Continue reading Writing for Fun? (Part 2): Journalism and Academic Writing

Writing for Fun? Part 1: Creativity and Academic Writing

We are constantly writing––composing emails, blackboard posts, essays, and dean’s date papers. In this two-part series, I am interested in understanding the different forms of writing students explore on campus. Specifically, I interview students who write for campus publications to see how they approach the writing process in their extracurriculars.

In this post, I Interview Serena Alagappan ’20, the Editor-in-Chief and a writer for Nassau Weekly. Serena is a comparative literature major who, for three years now, has shared poetry, cultural critiques, profiles, and fiction through the Nass. In my interview with Serena, we discuss creative writing and the connection she has experienced between her academic and personal writing. Serena encourages students to explore writing through the Creative Writing program and shares advice on how students can carry over the freedom and expression of creative writing into more formal and rigid academic subjects.

Continue reading Writing for Fun? Part 1: Creativity and Academic Writing

Unconventional Research: World Building in Creative Writing

When one thinks about research, there are certain images that come to mind: a student hunched over an old book in an empty library, or a solitary scientist in a lab coat mixing chemicals or observing animals. Emily McLean ’20 has done her fair share of the first type, as a potential Anthropology major with a strong interest in American History.

Emily is also my suitemate, and one day I heard her talking about an unfamiliar, unexpected genre of research: the research required for successful world building–the creation of a believable reality–in creative writing. Continue reading Unconventional Research: World Building in Creative Writing

Writing a Creative Thesis: An Interview with Edric Huang ’18

A couple of weeks ago, I interviewed Kristin Hauge about her independent work in the Music Department to highlight creative independent work in the arts. This week, I got in touch with Edric Huang, a senior in the Anthropology Department with certificates in Urban Studies and Creative Writing. Unlike most students on campus, he will be writing two theses this year. One is the classic research-based thesis that seniors in the sciences and humanities are familiar with, but the second will be a collection of poems for his Creative Writing Certificate. If you are unfamiliar with the kind of work that goes into creative theses, here’s what Edric had to share about his personal experience: Continue reading Writing a Creative Thesis: An Interview with Edric Huang ’18

Celebrating Senior Theses: An Interview with Claire Ashmead ‘17

In our spring series,  Senior Theses: A Celebration , we take a moment in the interlude between thesis deadlines and graduation to appreciate the diverse, personal, and impactful work of seniors’ capstone research projects.

Claire Ashmead completed two theses this spring: for her History concentration, a comparative study of McCarthyism and the Chinese Cultural Revolution, and, for her Creative Writing certificate, a novella entitled The Camel-Hair Coat . Here she reflects on writing and revising, family and loss, and the completion of her first book.  

What is your novella, The Camel-Hair Coat, about? It follows a girl, Daphne, who, four years ago, under mysterious circumstances, lost her mother in a terrible accident. Her intense grief over her mother’s death and profound yearning to know why her mother was taken away from her summons the ghost of her mother back from the dead. Daphne is faced with this choice: she can bring her mother back to reality, which she has wanted more desperately than she’s ever wanted anything before, but with terrible consequences for the rest of her family – her sister and grandmother. The price of bringing somebody back might be the exchange of other people you love, and even yourself. But if you really miss somebody – what wouldn’t you do?

How would you distill the book’s themes into a few words? Grief, wishes, and growing up.

What was the hardest part of writing the book? How much I had to revise. The book underwent a dramatic transformation, in part because I had two advisers: Joyce Carol Oates in the fall and then Jeffrey Eugenides in the spring. They’re completely different writers, which for me I think ended up being great: Joyce Carol Oates really tries to pull your creativity out of you, and she encouraged me to envision an alternative reality that ended up being a little confusing. Jeffrey Eugenides, on the other hand, is a very linear storyteller. When I gave him my draft, he told me he didn’t really understand what was going on, and that I needed to rewrite.

It was February, and I was hearing from my adviser that the 140 pages I’d generated needed to go. But in another sense that actually felt great.

At first that was very scary. It was February, and I was hearing from my adviser that the 140 pages I’d generated needed to go. But in another sense that actually felt great. I’d had the sneaking suspicion that the story needed a major change, and I wasn’t sure in what way. So I decided to structure it like a four-act play, where each act takes place in one day of one season. Once I had that structure, it was like, bingo! I know how this is going to develop.

In the second writing, I only kept maybe 3% of the words I’d previously written. But because I’d already created the spaces and characters in my head, writing the story the second time around actually took almost no effort, and required much less editing.

How did you juggle writing two separate theses? It was all about time management. I also really believe that just getting words on a page is so crucial. Often people feel like writing needs to be perfect when it comes out onto the page. My experience writing for Princeton Triangle Club has taught me that actually the hardest part is just starting, and as soon as you begin to write, your thoughts become clearer.

Also, the processes of the two theses were very different, which was helpful – if they were the same it probably would’ve been much more difficult for me to do the two. For my creative thesis, I wrote almost every day. With creative writing, I want to explore characters and change dialogue, so the earlier I get it out, the better. In history, it’s a very different beast: I like getting all of my secondary source reading and research done, getting it all into my head, and then putting it out onto the page. I would research for months and months, and then sit down and write a chapter in a day or two.

Describe your happy place as a writer. I have a two-room single in Edwards, so I have a room with my bed, and then another with a desk, facing the window. I wake up early, at 6 or 7 a.m., and would either go for a run or just go get coffee at Rojo’s, and then come to my desk in my little monastic sanctuary and write.

The first time I write something, I write in pen, by hand, on blank sheets of unlined paper. The great thing about writing by hand is that it’s physically exhausting, so you only say what you need to say, and the words you pick are more exact and intentional.

I’d sit down and write for about an hour and a half every morning, which would be about five double-spaced typed pages, sometimes more. The first sentence can sometimes be difficult, but as you start writing, you reenter the world. It’s like learning to ride a bike: you remember it, and you just push forward on the momentum of describing the scene.

The first sentence can sometimes be difficult, but as you start writing, you reenter the world. It’s like learning to ride a bike: you remember it, and you just push forward on the momentum of describing the scene.

Do you have a favorite section of the book? A lot of the book was plot that I just had to get through, and then there were a few scenes that made me feel that they were exactly why I wrote the book. Here are two paragraphs, after Daphne’s mother has come back and she is able to speak with her mother again.

Her mother took her hands. “At least you’ll have your father to walk you down the aisle. Mine was gone by the time I was your age. I missed him so much.”

“What’s it like?” Daphne asked. “Missing somebody?”

“You know, I’ve never thought about it.” Her mother frowned. “I’d say missing somebody is like remembering to pick up milk at the grocery store. Most of the time, you don’t think about it at all. And then all of a sudden the thought will just occur to you. I’ve got to pick up milk at the grocery store. And the thought will occur to you once a week, every month, every year, for forever. I’ve got to pick up milk at the grocery store. There, that’s it. I think about my father every day.”

If you’d like to read more of The Camel-Hair Coat, you can find it archived – with all senior theses – on the senior thesis digital archive , where it will be available starting after graduation on June 6 th , 2017.

— Zoe Sims, Natural Sciences Correspondent

The Imp Walks in the Door: Creativity in the Research Process

Staring at my computer screen, I blink. The black cursor, a vertical slit of a pupil, blinks back.

Uh-oh. I am trying to write the first essay for my environmental nonfiction class. But, sitting down to write, I can already feel the despondent haze of writer’s block descending. I swivel in my chair. I check my email but have no new messages. I type fdsajkl; on the first line of the page, and then delete it. What’s wrong with me? I think. Am I a writer or not?

Continue reading The Imp Walks in the Door: Creativity in the Research Process

A writer’s window: How poetry is changing how I see the world

In honor of National Poetry Month, my professor, Marie Howe, suggested writing a poem every day for the month of April. “Who’s up for it?” she asked our Advanced Poetry class . “It can be just a few lines. I’ll do it if you do it.”

I decided to write a poem right when I wake up each morning – figuring this is the only way I’d consistently get it done – and to forego my computer (and its associated, infinite distractions) in favor of a pencil and notebook. Every morning, I roll out of bed, perch myself on the wide windowsill of my ground-floor room, and write a poem.

I was shocked by how easily I could reshape my early-morning habits, and how much doing so affected the rest of my day. With this new routine has come a kind of freedom: my first thought of the day is no longer my calendar or breakfast or to-do list, but something creative and unlimited. I bring this creative lens with me through the rest of the day: watching milk gush over my cereal, stepping out into the April air, listening to a lecture about respiration across the animal kingdom. Continue reading A writer’s window: How poetry is changing how I see the world

Optional Arts Supplement

The optional Arts Supplement is for students who have devoted a significant amount of time and energy to one or more art forms and who wish to have their work considered as part of their application. If you've excelled in  architecture, creative writing, dance, music, music theater, theater or visual arts , and would like us to consider your talent as part of your Princeton application, you are welcome to submit an Arts Supplement. 

Submission Guidelines

Princeton values the arts in all forms. We cannot, however, evaluate areas outside those in which our faculty have expertise. Please read the descriptions for each area for further guidelines on how to submit your supplementary materials.

Single-Choice Early action optional Arts Supplement is due on or before Nov. 6 at 11:59 p.m. (applicant's local time).

Regular decision optional Arts Supplement is due on or before Jan. 8 at 11:59 p.m. (applicant's local time).

Transfer optional Arts Supplement is due on or before March 8 at 11:59 p.m. (applicant's local time).

Instructions

On the Common or QuestBridge* Application, please indicate your intention to submit an Arts Supplement in Princeton’s member questions. You will be able to access the link to submit an Optional Arts Supplement in your Princeton Applicant Status Portal.

Please keep in mind that you need to submit your application to Princeton University before you can submit your optional Arts Supplement on your Princeton Applicant Status Portal. 

*For students who are participating in the QuestBridge College Match, we will be unable to review the optional Arts Supplement as part of the Match application review process, given the early timeline.

Helpful Tips

As one of your media uploads, we encourage you to include a résumé if you think that will add to an understanding of your training and experiences. Please keep in mind, information you provide in your arts supplement will only  be read by the arts faculty evaluating your supplement unless you submit it as additional information along with your application to Princeton.

Changes cannot be made to the optional Arts Supplement after submission. If you wish to include an arts letter of recommendation, it must be requested prior to submission and an arts résumé must be included as one of the media uploads, if you wish to submit one.

We only accept one submission per program.

Your completed arts supplement will be reflected on your Princeton Applicant Status Portal within 24 hours.

Documents (for example, additional academic information, research papers, etc.) should  not  be uploaded with your arts supplement. You may upload those materials through the additional information section of the application or on your Princeton Applicant Status Portal.

If you have excelled in an area of the arts not covered by our supplement, we are still interested in seeing how your talent and expertise might enhance our community. You may submit additional materials or links to your work through your application or on your Princeton Applicant Portal. Although we cannot guarantee that the Admission Office will have time to review your work, we will include it in your file.

For questions related to the Princeton optional Arts Supplement, please email  [email protected] .

Acceptable Formats:

We support media files as large as 5GB, but please be advised that larger files will take longer to upload from your Internet connection and may stall if you are on a wireless connection or one that cannot sustain a connection for the necessary period of time. We support the following file formats:

• .3g2, .3gp, .avi, .m2v, .m4v, .mkv, .mov, .mpeg, .mpg, .mp4, .mxf, .webm, .wmv
• .aac, .m4a, .mka, .mp3, .oga, .ogg, .wav
• .bmp, .gif, .jpg, .jpeg, .png, .tif, .tiff
• .doc, .docx, .odg, .odp, .odt, .pdf, .ppt, .pptx, .rtf, .wpd

You may also include external media from YouTube, Vimeo and SoundCloud. Please do not upload links to personal websites.

Supplement Type, Category and Description of Acceptable Submissions:

Architecture.

Submit between 3-10 samples of your work in architecture or design. These can include blueprints/plans, drawings, renderings, 3D models, photographs or video.

Creative Writing

Categories: Fiction, Nonfiction, Poetry, Translations

Include a portfolio of your work in any or all of the following media, totaling no more than five pages: poetry, short stories, novel excerpts and translations. Please do not submit journalism samples or complete books.

Categories: Choreography, Performance

Include a digital video of one or more solo performances of contrasting styles (modern, improvisation, hip hop, contemporary, ballet) and/or of your choreography (no more than five minutes total). Please do not submit group work unless you have a prominently featured solo or you are the choreographer. Art Supplements for dance should be accompanied by a personal statement related to your interest in the arts, and how you would like to integrate dance into your studies at Princeton (no more than 250 words). Please contact the Dance program office during business hours at (609) 258-3017 for more information or questions about the program.

Categories: Classical Instrument, Classical Voice, Composition, Electronic Music, Jazz, Piano

The Princeton musical community is rich in many kinds of music. This includes non-Western classical and popular music of many types. The Department of Music, however, can provide evaluations only of the performance genres for which formal instruction is offered. For further information or questions about the program, please direct emails to Greg Smith at [email protected] . 

Please attach a résumé as one of your media uploads that details your music training and accomplishments. You may send submissions in the form of digital audio or video files. Additionally please refer to the "Acceptable Formats" section above. 

Classical Instrument (except piano) — Include a 10- to 15-minute submission of advanced solo work that shows contrasting examples of expression and technique. Do not submit orchestral or ensemble work unless you have a prominently featured solo.

Classical Voice — Whether you prefer solo repertoire, small group or large choral settings, Princeton offers countless learning and performing opportunities, and we are eager to hear you sing. Your supplement submission should contain at least two songs, totaling no more than 10 minutes, that best demonstrate your level of technical and musical accomplishment. At least one submission should be from western classical art song or operatic repertoire, and at least one submission should be in a language other than English. You may additionally submit small ensemble work if this is an area of interest.

Composition — Include a submission of one to three works that demonstrate your interests and abilities in composition. The Department of Music has a broad definition of composition, one that includes concert music, improvisational music, un-notated music, and while they don’t have the expertise to fully evaluate non-Western compositional traditions, these are also welcome. Submissions can consist of PDF scores or recordings; recordings or MIDI mock-ups of scored works are welcome if you have them but not required.

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• Vavilovskii Zhurnal Genet Selektsii
• v.26(4); 2022 Jul

Language: English | Russian

Investigation of genetic polymorphism of Russian rape and turnip rape varieties using SSR and SRAP markers

Изучение генетического полиморфизма российских сортов рапса и сурепицы с использованием ssr- и srap-маркеров, i.a. klimenko.

Federal Williams Research Center of Forage Production and Agroecology, Lobnya, Moscow region, Russia

V.T. Volovik

A.a. antonov, v.a. dushkin, a.o. shamustakimova, yu.m. yu.m. mavlyutov.

Rapeseed (Brassica napus L.) and turnip rape (B. rapa L. subsp. campestris (L.)) are important agricultural plants widely used for food, fodder and technical purposes and as green manure. Over the past decades, a large number of perspective varieties that are being currently cultivated in every region of Russia have been developed. To increase the breeding eff iciency and facilitate the seed production, modern molecular-genetic techniques should be introduced as means to estimate species and varietal diversity. The objective of the presented research study was to investigate DNA polymorphism of the rapeseed and turnip rape varieties developed at Federal Williams Research Center of Forage Production and Agroecology and detect informative markers for varietal identif ication and genetic certif ication. To genotype 18 gDNA samples, 42 and 25 combinations of respective SSR and SRAP primers were used. The results obtained demonstrate that SRAP markers were more effective for polymorphism analysis: 36 % of the tested markers revealed genetic polymorphism compared with only 16.7 % of microsatellite loci. Molecular markers to detect differences at interspecif ic and intervarietal levels have also been found. For the investigated set, such microsatellite loci as Na12A02, Ni2C12, Ni02-D08a, Ra02-E01, Ni03H07а and SRAP-marker combinations as F13-R9, Me4- R7, F11-Em2, F10-R7, F9-Em2 and F9-R8 proved to be informative. Application of the two marker techniques made it possible to detect a higher level of DNA polymorphism in plants of different types (spring and winter varieties) if compared against the intervarietal differences within a species or a group. According to Nei’s genetic diversity index, in the cluster of winter rapeseed, VIK 2 and Gorizont varieties had the longest genetic distance, and in the spring cluster, these were Novosel and Veles. A high level of similarity was found between Vikros and Bizon winter rapeseed varieties. The results obtained have a high practical value for varietal specif ication of seed material and genetic certif ication of rapeseed and turnip rape varieties.

Рапс (Brassica napus L.) и сурепица (B. rapa L. subsp. campestris (L.)) – важные сельскохозяйственные культуры, широко используются для продовольственных, кормовых и технических целей, а также в качестве сидератов. За последние десятилетия создано большое количество перспективных сортов, культивируемых практически во всех регионах России. Для повышения эффективности селекционного процесса и успешного развития семеноводства необходимо внедрять современные молекулярно-генетические методы оценки видового и сортового разнообразия. Цель настоящей работы заключалась в изучении ДНК-полиморфизма сортов рапса и сурепицы селекции Федерального научного центра кормопроизводства и агроэкологии им. В.Р. Вильямса и выявлении информативных маркеров для сортовой идентификации и генетической паспортизации. Для генотипирования 18 образцов геномной ДНК использовали 42 и 25 комбинаций SSR- и SRAP-праймеров соответственно. Результаты показали, что маркеры SRAP более эффективны для анализа полиморфизма изучаемого материала: 36 % от общего числа испытанных маркеров демонстрировали генетический полиморфизм, тогда как для микросателлитных локусов этот показатель равнялся 16.7 %. Определены молекулярные маркеры для выявления различий на межвидовом и межсортовом уровнях. Информативными для исследуемой выборки сортов оказались микросателлитные локусы Na12A02, Ni2C12, Ni02-D08a, Ra02-E01, Ni03H07а и комбинации SRAP-маркеров F13-R9, Me4-R7, F11-Em2, F10-R7, F9-Em2 и F9-R8. Анализ сортового материала по двум системам маркирования показал более высокий уровень ДНК-полиморфизма у образцов растений разного типа развития (яровой/озимый) в сравнении с различиями между сортами в пределах вида или группы. Согласно индексам генетического разнообразия Нея, в кластере сортов озимого рапса наибольшей генетической удаленностью выделялись ВИК 2 и Горизонт, среди яровых – Новосёл и Велес. Высокий уровень сходства обнаружен между яровыми сортами рапса Викрос и Бизон. Полученная информация имеет практическое значение для контроля сортовой принадлежности и генетической паспортизации семенного материала сортов рапса и сурепицы.

Introduction

Cabbage oilseed crops such as rapeseed (Brassica napus L.) and turnip rape (B. rapa L. subsp. campestris (L.)) are cultivated in almost every region of Russia, and, for the foreseeable future, are regarded as the main reserve for increasing the production of vegetable oil and fodder protein. These plants are widely used in food, fodder, technical purposes and as green manure that increases soil fertility thanks to the plants’ root remains containing up to 6 tons of organic maters, 80 kg of nitrogen, 60 kg of phosphorus and 90 kg of potassium per hectare. As for their food and fodder properties, rapeseed and turnip rape exceed many other cultivated crops since their seeds are 40–48 % fat and 21–33 % protein and contain a high amount of essential amino acids (Volovik, 2015). Rapeseed can provide livestock with green forage from early spring to late fall thanks to their cold hardiness and fast regrowth after mowing. They are also an excellent silage material, and their seeds and seed by-pass products are processed to produce seed cake and coarse meal. In the recent years the varieties of rapeseed and turnip rape with low or no erucic-acid content became available and seed production has increased more than 7 times to reach the world’s third place after soybeans and cotton. Russia’s short-term plans are to increase rapeseed planting acreage to 2.5 mln he.

As for Russian research institutions working intensely to select cabbage oilseed crops, the leading ones are All-Russian Research Institute of Rapeseed, All-Russian Research Institute of Oilseed Crops and All-Russian Williams Fodder Research Institute. For the two last decades, they have produced the perspective varieties of rapeseed, turnip rape, white mustard and oil radish that have been recommended for oil production, livestock and poultry green forage, combination fodder, seed cake and coarse meal production. In 2021, “State Register” of the Russian Federation included 13 varieties of rapeseed and 3 varieties of turnip rape selected by Federal Williams Research Center of Forage Production and Agroecology (Kosolapov et al., 2019; State Register…, 2021).

For preservation and rational use of newly available varieties, intensification of the selection process and protection of intellectual property, modern and effective methods to estimate species and varietal diversity at a genetic level are to be introduced. One of such techniques that has been successfully applied in the recent years is molecular DNA markers, which, if compared against the traditional morphological indicators, possess a number of advantages. These include a high level of polymorphism; even genome distribution; reliability; a possibility to automate the assay procedure that does not depend on environmental conditions or a plant development phase (Agarwal et al., 2008; Khlestkina, 2011; Chesnokov, 2018). If the most informative and convenient DNA markers are selected, their capabilities to estimate the genetic variability of selection material are regarded as unlimited.

Laboratory for Molecular and Genetic Studies in Federal Williams Research Center of Forage Production and Agroecology has been developing a system for DNA identification and genetic certification of Russian fodder crops. For the time being, the varietal identification techniques have been adapted for perennial legume grasses such as red clover and different species of alfalfa (Klimenko et al., 2020a, b). The assay uses samples of the summary total DNA obtained through a modified method from an arbitrary selected sample of every variety’s germinants. Two types of molecular markers were used: SSR (simple sequence repeats), which detect the variability of microsatellite genome sequences, and SRAP (sequence related amplified polymorphism), which is based on PCR with a pair of primers for amplification of intron/exon regions (open reading frames). The techniques have been tested on different species of fodder crops to optimize the amplification conditions, detection and analysis of results.

A problem of reliable varietal identification is particularly topical for rapeseed due to its limited genetic variability conditioned by the intensive selection aimed at higher content and quality of oil. Currently, a significant number of published studies have been devoted to using different DNA markers for estimation of the genetic diversity of rapeseed varieties and hybrids (Plieske, Struss, 2001; Snowdon, Friedt, 2004; Klyachenko et al., 2018; Mozgova et al., 2019); to genetic mapping (Piquemal et al., 2005; Gao et al., 2007; Geng, 2012) and marking the genes of economically valuable traits (Chen et al., 2010; Ananga et al., 2012). However, only a few such studies have investigated Russian varieties. Four varieties of winter and spring rapeseed (Podmoskovniy, Vikros, VIK 2 and Severyanin) were studied by Byelorussian researchers to identify the gene alleles determining the concentration of oleic and linolic acids in rapeseed oil (Lemesh et al., 2015). The same varieties were investigated to detect the DNA markers of the genes responsible for erucic-acid synthesis (Amosova et al., 2014). Microsatellite markers were used to study the genetic polymorphism of Russian varieties Ratnik and SNK- 198 (Satina, 2010) as well as the genetic homogeneity of spring rapeseed varieties Bulat and Forward (Rogozhina et al., 2015). Such winter varieties as Stolychniy, Laureat, Gorizont, Nord and Severyanin were investigated to detect the quantitative trait loci (QTLs) associated with high winter hardiness (Mozgova et al., 2019).

The objective of the presented study was to investigate DNA polymorphism of rapeseed and turnip rape varieties developed by breeders of Federal Williams Research Center of Forage Production and Agroecology and to identify the informative markers for varietal differentiation and genetic certification.

Materials and methods

Plant material. The study investigated 15 varieties of winter (Severyanin, Stolychniy, VIK 2, Nord, Laureat, Gorizont, Garant) and spring (Vikros, Novik, Novosel, Veles, Grant, Podmoskovniy, Lugovskoy, Bizon) rapeseed and 3 varieties of winter (Zarya) and spring (Nadezhda, Svetlana) turnip rape.

DNA extraction and PCR analysis. The gDNA was extracted from 30 germinants of each abovementioned variety (bulk samples) using the basic SDS method (Kirby, Cook, 1967; Dellaporta et al., 1983) with some modifications (Klimenko et al., 2020b). The quality and concentration of the obtained DNA fractions were verified with agarose gel (1.5 %) electrophoresis and using a Nabi spectrophotometer (MicroDigital, South Korea).

To carry out SSR analysis, 42 markers from the database Brassica info (https://www.brassica.info) and available publications were applied. The efficiency of the primers devised for these markers had been demonstrated in the studies devoted to development of the technology of rapeseed genotyping (Satina, 2010) and selection of the samples with low erucic-acid and glucosinolate content (Hasan et al., 2008). A part of the markers included in the analysis was used for hybridization control and detection of Alternaria blight resistant genotypes in Indian mustard (B. juncea L.) (Chandra et al., 2013; Sharma et al., 2018).

The PCR-mixture of 20 μl contained 3 μl 10 × PCR buffer (Taq Turbo Buffer), 0.5 μl 50 × dNTPs mix, 0.4 μl Taq polymerase (5U), forward and reverse primers (0.1 μl each, 100 μm) and 0.1 μl of DNA sample (20 ng/μl). The amplification was performed in a T-1000 thermal cycler (Bio-Rad, USA) at two different temperature regimes. The first amplification program was an initial 3-min denaturation at 95 °C followed by 30 cycles of 30 s at 94 °C, 30 s at 55–57 °C, 30 s at 72 °C and a final 5-min elongation at 72 °C (Satina, 2010). The second program included an initial 5-min denaturation at 95 °C followed by 39 cycles of 1 min at 94 °C, 2 min at 46–51 °C (depending on the primer pair in use), 2 min at 72 °C and a final 10-min elongation at 72 °C (Chandra et al., 2013). The reproducibility of obtained results was attested in three-fold replication.

SRAP analysis was carried out using 25 primer combinations comprised from 10 single oligonucleotides: F9, F13, Me4, F10, F11, R9, R7, Em2, R14, R8 (Li, Quiros, 2001; Rhouma et al., 2017). The amplification program was an initial 4-min denaturation at 94 °C followed by 10 cycles with changing temperature and duration parameters (1 min at 94 °C, 1 min at 35 °C, 1 min at 72 °C); followed by 30 cycles (1 min at 94 °C, 1 min at 50 °C, 1 min at 72 °C) and a final 5-min elongation step run at 72 °C. The PCR-mixture composition was similar to that used for the microsatellite analysis.

PCR-products were separated using 90-min 50-V agarosegel electrophoresis (4 % MetaPhorR Agarose, Rockland or 1.6 % LE, Lonza, USA). As the reference markers, 20 bp DNA Ruler (Bio-Rad), 100 kb DNA Ladder (Thermo Fisher Scientific, USA) and 100 bp + 1.5 kb (SibEnzyme, Russia) were applied.

Analysis of the obtained results. PCR-product detection and size measurement was performed using a GelDoc XR+ imaging system (Bio-Rad) and the ImageLab software (Bio- Rad Lab., Inc.) for molecular-mass markers. The obtained results were transformed into a binary matrix, and PopGene v. 1.32 (Yeh et al., 2000) was applied to determine such genetic diversity indices as the effective number of alleles per locus; Shannon’s index; expected heterozygosity; Nei’s genetic distance (Nei, Li, 1979). Polymorphism information content (PIC) for every pair of primers was calculated by the formula presented in the study (Chesnokov, Artemyeva, 2015). To build the genetic similarity dendrogram, the unweighted pair group method with arithmetic averages was applied in NTSYSpc v 2.10 (Rohlf, 2000).

To obtain gDNA from the rapeseed and turnip rape germinants, a modified SDS method was used. The applied protocol proved more effective and less costly compared to other known protocols and commercial reagents kits. The results of electrophoresis and spectrophotometry attested to the DNA’s high concentration and purification degree from protein compounds and polysaccharides for all experimental samples (Fig. 1, 2).

Lanes 1–15 (rape varieties): Severyanin, Stolychniy, VIK 2, Nord, Laureat, Gorizont, Garant, Vikros, Novik, Novosel, Veles, Grant, Podmoskovniy, Lugovskoy, Bizon; 16–18 (turnip rape varieties): Zarya, Nadezhda, Svetlana.

SSR-analysis

For genotyping the full variety collection, out of 42 SSR primers, 7 primers providing stable and reproducible amplification were selected (Table 1).

Analysis of the amplification fragments obtained using the listed primers detected 42 alleles. Their number per locus was 6 on average, varying from 3 (Ni2C12 and Bna.M.010) to 10 (Ra02-E01a). The fragment size varied from 110 bps (Ni2C12) to 1200 bps (Ni02-D08a). The maximum allele frequency was registered for Bna.M.010 (0.83), and the minimum – for Ni03H07a (0.27); the mean value was 0.42. The primers developed for Ni03H07a, Ni02-D08a and Ra02-E01a markers made it possible to detect 8–10 alleles per locus and had the highest PIC (0.82).

SRAP-analysis

Based on the results of preliminary testing, the initial 25 combinations of SRAP primers were reduced to 10 pairs, amplifying stable polymorphic DNA fragments (Table 2). In total, 53 PCR fragments of 132–1674 nucleotide pairs in size were obtained. One combination contained from 4 (F9-R9) to 7 (F10-R8, F11-Em2, F10-R7) amplicons. A part of the markers proved to be informative to detect the amplification fragments for differentiating the type of plants (winter/ spring). Using 6 combinations made it possible to obtain the amplicons specific for varieties identification (marked with a star in the Table 2).

Fig. 3 demonstrates the electrophoregram of PCR results with the F9-R8 primer combination. Significant DNA profile differences were found between winter (I) and spring (II) rapeseed varieties (joined in curly brackets). The arrows mark the variety-specific PCR products characteristic for Stolychniy winter rapeseed (508 bps) and Nadezhda spring turnip rape (700 bps) as well as the absence of an amplicon in size of 460 bps in spring rapeseed Podmoskovniy though it was a specific characteristic for other varieties in this group. The performed analysis demonstrated that it is possible to identify rapeseed varieties Grant and Novosel with 3 marker combinations (F11-Em2, F10-R7 and Me4-R7), and Gorizont and Lugovskoy – with 2 (F13-R9 and Me4-R7). Variety VIK 2 was identified with SRAP primers F9-Em2, and spring ones Veles – with F10-R7. Specific DNA spectra for rapeseed varieties Stolychniy, Podmoskovniy and turnip rape Nadezhda were obtained with F9-R8 combination.

Winter rapeseed varieties: Severyanin (1), Stolychniy (2), VIK 2 (3), Nord (4), Laureat (5), Gorizont (6), Garant (7); spring rapeseed varieties: Vikros (8), Novik (9), Novosel (10), Veles (11), Grant (12), Podmoskovniy (13), Lugovskoy (14), Bizon (15). Winter turnip rape: Zarya (16); spring turnip rape: Nadezhda (17), Svetlana (18). H2O control (19). M – molecular weight marker (100 кb DNA Ladder).

The performed analysis demonstrated that it is possible to identify rapeseed varieties Grant and Novosel with 3 marker combinations (F11-Em2, F10-R7 and Me4-R7), and Gorizont and Lugovskoy – with 2 (F13-R9 and Me4-R7). Variety VIK 2 was identified with SRAP primers F9-Em2, and spring ones Veles – with F10-R7. Specific DNA spectra for rapeseed varieties Stolychniy, Podmoskovniy and turnip rape Nadezhda were obtained with F9-R8 combination

The obtained data were transformed into a binary matrix to calculate Nei’s genetic distances (Table 3). The lowest genetic similarity coefficient (0.7069) was found between rapeseed varieties Gorizont, Novosel and Grant, the highest – between spring varieties Vikros and Bizon (1.0) as well as Veles and Bizon (0.9655). A similarly high genetic distance (0.3228) indicated significant differences between pairs: Grant and VIK 2, and Lugovskoy and Stolychniy. Low distance values and high genetic similarity were demonstrated by spring varieties Bizon and Vikros (zero distance) and winter varieties Garant, Severyanin, Stolychniy, Nord, Laureat (0.0174).

Notе. According to the data of 1 (Rhouma et al., 2017); 2 (Сатина, 2010); 3 (Chandra et al., 2013).

The results of PCR analysis for SSR and SRAP markers were used to determine the genetic variability indices and build an UPGMA dendrogram depicting the varieties’ phylogenetic relationships. The variety material had a low degree of genetic heterogeneity, while higher values of expected heterozygosity (He) and the number of effective alleles (ne) were determined with SSR markers: 0.25 on average against 0.14 and 1.47 per locus if compared to 1.24, respectively. However, the SRAP method has enabled obtaining more PCR products applicable for varietal differentiation (Table 4).

Notе. No. 1–15 – rapeseed varieties Severyanin, Stolychniy, VIK 2, Nord, Laureat, Gorizont, Garant, Vikros, Novik, Novosel, Veles, Grant, Podmoskovniy, Lugovskoy, Bizon.

Analysis of the UPGMA dendrogram demonstrated that the winter/spring rapeseed varieties were divided into two distinguishable clusters (Fig. 4). The first one united such winter cultivars as Severyanin, Garant, Stolychniy, Nord, Laureat, Gorizont, VIK 2; the second – all the spring ones. In the winter cluster VIK 2 and Gorizont were the most distant from the other varieties. The distances between Stolychniy, Nord, Laureat as well as between Garant and Severyanin were much shorter, which was confirmed by their high genetic similarity indices being 0.9655 and 0.9828, respectively (see Table 3). The most distant among spring rapeseed were twozero varieties Novosel, Grant and Lugovskoy, which had the longest genetic distances in the cluster (0.3469 and 0.3228). Bizon and Vikros belonged to one subgroup, sharing a common branch of the dendrogram.

The bulk strategy of DNA sampling from 30 germinants per variety has significantly reduced the labor efforts and cost of the research if compared to the traditional method of individual sample genotyping. The method has proved its efficiency for different cultures especially in large-scale studies of vast populations (Liu et al., 2018). However, this approach is only justified if the analyzed set of samples is representative. For cross-pollinating species with a high level of intrapopulation variations, it should include at least 30–50 plants per variety, which significantly increases the likelihood of registering a rare alleles, the occurrence of which in the population does not exceed 10 % (Crossa, 1989; Semerikov et al., 2002). The plants of winter rapeseed are known for their high self-pollination capacity (up to 70 % of flowers) (Shpaar, 2012), many varieties are linear; while in spring rapeseed this capacity reaches 40 % (Osipova, 1998). That’s why in our study we used budk samples that combined 30 seedlings from each variety.

A significant part of SSR primers tested in our study generated monomorphic amplification fragments. They did not allow us to properly estimate the genetic variability and had low reproducibility in replicated experiments. A proportion of the markers proven effective for intervarietal DNA polymorphism detection comprised 16.7 %, being much lower than in other studies (Plieske, Struss, 2001; Hasan et al., 2008; Tian et al., 2017). It was probably due to the composition of the tested collection that had a narrow genetic basis considering the varieties’ pedigree. At the same time, such parameters of genetic variability as the number of allelic variants, singleallele frequency, PIC and He were comparable to those found in published data (Satina, 2010; Klyachenko et al., 2018).

In general, the used markers made it possible to detect DNA polymorphism between rapeseed and turnip rape as well as between the winter and spring varieties within each species. However, Na12A02 marker turned out to be variety-specific for Bizon winter rapeseed and Zarya spring turnip rape, and Ra02-E01а – for VIK 2 winter rapeseed and Svetlana spring turnip rape. The unique alleles of Podmoskovniy and Lugovskoy rapeseed were detected using Ni02-D08a loci. The indicated markers can be used for varietal DNA identification and genetic certification.

SSR primers for the markers of Indian mustard’s Alternaria blight resistance genes (Chandra et al., 2013), such as Ni02- D08a, Ni03H07a and RA02-E01a, proved to be the most effective. Their application enabled us to detect the specific amplification fragments for linear winter rapeseed variety VIK 2. They also proved effective for Gorizont, which had been obtained on the base of VIK 2 by seed freezing followed by their selection at low-temperature stress. These two varieties share high winter hardiness and are resistant to Alternaria blight. Thereby the results of our study can be useful for further selection of perspective breeding material and QTL analysis on disease resistance.

Among the spring rapeseed, Veles variety turned out to be substantially different while Lugovskoy and Garant had many similarities in the studied microsatellite parts of regions of the genome. Veles is a new perspective variety that has been approved for use since 2021 and was selected based on Vikros using the method of chemical mutagenesis, producing a high frequency of nucleotide changes. This is possibly the reason for Veles having unique alleles in three loci: Ni2C12, Ra02- E01a, Na12A02. For Vikros variety, a specific DNA profile was also obtained with Ni2C12 marker.

Rapeseed Grant was selected using the method of interspecies and intervarietal hybridization of early-maturing foreign breeding samples and the high-yielding varieties Lugovskoy and Vikros, developed at Federal Williams Research Center of Forage Production and Agroecology. Their common origin is probably the reason for the genetic similarity found between Grant and Lugovskoy varieties.

In general, SSR analysis failed to achieve optimum effect in identification of the investigated varieties: from the total set, including 42 primers for microsatellite genome loci, only four were attested as variety-specific for rapeseed, and only one (Ni03H07а) – for Nadezhda spring turnip rape.

For further investigation of DNA polymorphism, SRAP analysis was applied. SRAP is the third generation of molecular markers that were initially designed for the genes of B. oleracea L. (Li, Quiros, 2001) and are successfully used these days for genetic variability estimation and genetic mapping in different plants (Aneja et al., 2012; Rhouma et al., 2017; Liu et al., 2018). This is a cheap, effective and highly reproducible technique

2017; Liu et al., 2018). This is a cheap, effective and highly reproducible technique

The final dendrogram of phylogenetic relations made it possible to visually estimate the degrees of genetic similarities and differences of the studied material. For instance, close placing of such rapeseed varieties as Stolychniy, Nord and Laureat was probably determined by the features of their origin: they were selected for winter hardiness from a combination, in which one of the parental forms was Promin’, a well-known winter rapeseed variety

Garant, selected for winter hardiness, and Severyanin, which was obtained by seed freezing in a climatic chamber and the following individual-family selection, turned out to be in the common subgroup and at a short genetic distance (0.0174) from each other. In addition to high winter hardiness, these varieties are resistant to lodging and to damage by pathogenic fungi

A two-zero spring variety Novosel takes a special position in his group (Nei’s distance is 0.3469). Novosel was developed based on the foreign breeding samples and Russian varieties Lugovskoy and Vikros, characterized by early maturing and high yield. Specific properties of the new breeding achievement are shorter maturation period in comparison to standard varieties and high resistance to Alternaria blight.

Spring rapeseed Bizon and Vikros take the common branch of the dendrogram. The varieties were developed using the method of interspecies hybridization but from different parental forms; characterized by high yield productivity, early maturation and low glucosinolate content.

The presented study has proved the efficiency of SSR and SRAP markers for estimation of DNA polymorphism in rapeseed and turnip rape varieties developed in Federal Williams Research Center of Forage Production and Agroecology. During the study, SRAP technique has demonstrated a higher level of informativity: 36 % of the tested markers were polymorphic, while for the microsatellite loci this rate did not exceed 16.7 %.

Both techniques of molecular analysis enabled detecting the DNA markers for identification of 10 out of 15 rapeseed varieties tested and for 2 turnip rape samples. Microsatellite loci Na12A02, Ni2C12, Ra02-E01 and Ni02-D08a allowed obtaining unique PCR products for Bizon, Veles, Vikros, VIK 2, Podmoskovniy and Lugovskoy rapeseed varieties. Marker Ni03H07а proved effective for identifying Nadezhda turnip rape. In the used SRAP test kit, such primers as F13-R9, Me4- R7, F11-Em2, F10-R7, F9-Em2 and F9-R8 proved effective for detecting variety-specific amplicons or obtaining unique DNA profiles for different types of plants (winter/spring) in rapeseed varieties Grant, Novosel, Gorizont, Stolychniy, Lugovskoy, Podmoskovniy and in spring turnip rape Svetlana.

The results of the study can be used for development of the perspective breeding samples and hybrids, for genetic certification and seed material purity control.

Conflict of interest

The authors declare no conflict of interest.

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Acknowledgments

The presented investigation was supported by the means of the federal budget, directed for performing the government assignment (project No. 0442-2019-0001АААА-А19-119122590053-0).

Contributor Information

I.A. Klimenko, Federal Williams Research Center of Forage Production and Agroecology, Lobnya, Moscow region, Russia .

V.T. Volovik, Federal Williams Research Center of Forage Production and Agroecology, Lobnya, Moscow region, Russia .

A.A. Antonov, Federal Williams Research Center of Forage Production and Agroecology, Lobnya, Moscow region, Russia .

V.A. Dushkin, Federal Williams Research Center of Forage Production and Agroecology, Lobnya, Moscow region, Russia .

A.O. Shamustakimova, Federal Williams Research Center of Forage Production and Agroecology, Lobnya, Moscow region, Russia .

Yu.M. Yu.M. Mavlyutov, Federal Williams Research Center of Forage Production and Agroecology, Lobnya, Moscow region, Russia .