yale university phd biology

Graduate Studies in Chemical Biology

Thank you for your interest in the Chemical Biology Graduate Studies at Yale. Students with backgrounds in chemistry, biochemistry or biology are encouraged to apply directly to one of the four participating graduate programs:

Molecular Cell Biology, Genetics & Development

Biochemistry, Biophysics and Structural Biology

Molecular Medicine, Pharmacology & Physiology

Applicants should indicate their interest the Chemistry/Biology Interface Training Program in their statement of purpose.

Yale Computational Biology and Bioinformatics

Graduate program.

CBB is part of the Yale combined program in the  Biological and Biomedical Sciences (BBS) . BBS is an umbrella program that incorporates a number of different areas of biological research. Students apply to the CBB track of the BBS program and are part of this track for their first year of studies. After their first year, students join the CBB interdepartmental graduate program and work towards a degree in Computational Biology and Bioinformatics. Alternatively, students may join any of the other departments associated with BBS if they find that another department better fits their interest after the first year.

CBB offers a Ph.D. with an en-route Masters’ degree; a terminal Masters’ program is offered with a Computational Biology or Biomedical Informatics concentration as well. For more information about these degrees, visit the Curriculum  page. For more information about applying to these programs, visit the Application Information  page.

There are a variety of social and career events held throughout the year, including Research in Progress (RIP) talks, journal clubs and seminars. Check the Calendar page for upcoming events.

There are also a wide range of initiatives at Yale that relate to CBB, including (but not limited to):

• Biomedical Informatics Research Training Grant from the National Library of Medicine
• Biomedical Informatics & Data Science (BIDS)
• “West Campus” Initiatives including the Yale Center for Genome Analysis (Shrikant Mane) and Yale Systems Biology Institute (Andre Levchenko)
• ENCODE & PsychENCODE projects (Mark Gerstein, Nenad Sestan)
• Department of Pathology Informatics Group (Steven Kleinstein, Yuval Kluger)
• MCDB informatics faculty (Thierry Emonet, Damon Clark)
• Integrated Graduate Program in Physical and Engineering Biology (Corey O’Hern, Lynne Regan, and Simon Mochrie)

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• Biomaterials & Bioinspired Design
• Cell Shape & Motion
• Force Generation & Sensing
• Informatics & Data Mining
• Modeling Biological Processes
• Systems & Synthetic Biology
• Method Development
• Graduate Students
• Benefits of PEB

NEWS FLASH - First graduate student to apply to PEB defends his thesis!

Welcome to Yale’s Integrated Graduate Program in Physical and Engineering Biology

Transforming Research Through Collaboration

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Interested in applying to PEB?

We provide a step-by-step-guide  to applying to the program via different department or tracks.

APPLY TO PEB

Department of Ecology & Evolutionary Biology

The Department of Ecology and Evolutionary Biology at Yale University is home to broad, interdisciplinary and highly collaborative research with particular expertise in population, community, ecosystem, and macroecology; in evolutionary genetics, developmental evolution, behavioral evolution, and evolutionary medicine; and in phylogenetics, systematics, and biodiversity.  We are committed to producing world-class scientists, educators and professionals through undergraduate and graduate education.

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Study of slowly evolving ‘living fossils’ reveals key genetic insights

The Department of Ecology and Evolutionary Biology was created in 1997 and currently comprises 16 primary and 14 affiliated faculty members, approximately 40 graduate students, 50 postdoctorals, lecturers and research scientists, and 100 undergraduates with an EEB concentration.  Our offices and laboratories are spread across the historic Osborn Memorial Laboratories (OML), the Environmental Science Center (ESC) and Building 31 on Yale’s West Campus. 

The mission of the Department of Ecology and Evolutionary Biology at Yale University is to achieve the highest possible quality of research, undergraduate, and graduate education in the fields of ecology, evolution, and organismal biology.

We discover, create, synthesize, and disseminate knowledge about the earth’s biodiversity, its ecological interactions, and its evolutionary history.  We pursue integrative, interdisciplinary, and global research on phenomena that range from molecules to ecosystems.

We prepare Yale College undergraduate majors for careers in biology and medicine and educate other Yale College students in ecology, evolution, and biodiversity.

We seek to attract the most capable, promising, and diverse graduate students and postdoctoral appointees from the nation and the world and to prepare them for positions of leadership in research, education, and society. 

• Microbiology

The Graduate Program in Microbiology is a multidepartmental, interdisciplinary Ph.D. program in training and research in the study of microorganisms and their effects on their hosts. The faculty of the program share the view that understanding the biology of microorganisms requires a multidisciplinary approach; therefore, the Microbiology graduate program emphasizes the need for strong multidisciplinary training. The program is designed to provide individualized education in modern microbiology and to prepare students for independent careers in research and teaching. Students can specialize in various areas, including bacteriology, virology, microbe-host interactions, microbial pathogenesis, cell biology and immunobiology of microbial infections, microbial genetics and physiology, structural biology, parasitology, microbiome, and microbial ecology and evolution.

• Programs of Study
• PhD - Doctor of Philosophy
• Department of Microbial Pathogenesis

Director of Graduate Studies

• [email protected]

Jennifer Atchley

Departmental Registrar (on leave February - August 2024)

• [email protected]
• 203-737-1087

Marneshia Bullock

Departmental Registrar (Temporary, February - August 2024)

• [email protected]

Admission Requirements

Standardized testing requirements.

GRE is not accepted.

English Language Requirement

TOEFL iBT or IELTS Academic is required of most applicants whose native language is not English. BBS requires a score of at least 600 on the paper version, 250 on the computer-based exam, and 100 on the internet-based exam. Please take the test no later than November and no earlier than 24 months prior to submitting your application. Use institution code 3987 when reporting your scores; you may enter any department code.

You may be exempt from this requirement if you have received (or will receive) an undergraduate degree from a college or university where English is the primary language of instruction, and if you have studied in residence at that institution for at least three years.

Admission Information

Microbiology participates in the Combined Program in the Biological and Biomedical Sciences (BBS) , and applicants interested in pursuing a degree in cell biology should apply to the Microbiology Track within BBS.

Academic Information

Program Advising Guidelines

GSAS Advising Guidelines

Academic Resources

Academic calendar.

The Graduate School's academic calendar lists important dates and deadlines related to coursework, registration, financial processes, and milestone events such as graduation.

Featured Resource

Registration Information and Dates

https://registration.yale.edu/

Students must register every term in which they are enrolled in the Graduate School. Registration for a given term takes place the semester prior, and so it's important to stay on top of your academic plan. The University Registrar's Office oversees the systems that students use to register. Instructions about how to use those systems and the dates during which registration occurs can be found on their registration website.

Financial Information

Phd stipend & funding.

PhD students at Yale are normally full-funded for a minimum of five years. During that time, our students receive a twelve-month stipend to cover living expenses and a fellowship that covers the full cost of tuition and student healthcare.

• PhD Student Funding Overview
• Graduate Financial Aid Office
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Alumni Insights

Below you will find alumni placement data for our departments and programs.

• Contact Us!

Department of Physics

You are here.

Yale’s Integrated Graduate Program in Physical and Engineering Biology (PEB for short) was established to train a new generation of scientists who are skilled at applying physical and engineering approaches to biological research, and who are also sufficiently sophisticated in their biological training that they can readily identify and tackle cutting-edge problems in the life sciences. The PEB leadership is committed to a new paradigm that transcends traditional, artificial, boundaries both between disciplines and also between research, teaching, and learning.

Physics PEB students benefit from

• Frequent interdisciplinary interactions. Being part of PEB means that you will meet and interact with graduate students, postdocs and faculty affiliated with many different departments, and spread across the Science Hill area, the Yale School of Medicine, West Campus, and the School of Engineering and Applied Science.
• Courses co-taught by faculty from biology, engineering and physics PEB courses are taught by pairs of faculty with complimentary expertise, many of whom also collaborate.
• Working with peers from a different background than yours . Several PEB courses incorporate peer teaching,  helping physicists learn biology and how to communicate effectively with other disciplines.
• PEB Discussion Group is a monthly seminar, focusing on research-in-progress talks. Two PEB students with complimentary backgrounds organize the talks. Click here for more info.
• Outreach opportunities . PEB students participate in outreach activities including devising and implementing science modules for Breakthough New Haven, a program that prepares middle school students from disadvantaged backgrounds for high school; working with 4-7th grade summer campers; and participating in activities planned by the Yale Science Diplomats.
• Professional development activities include a PEB Career Series and participation in a “Short Course on Communicating Science to the Public” led by the former chief science and health correspondent of NBC News, Bob Bazell.
• Physics of Living Systems Student Research Network (PoLS SRN) serves to connect physics students, working at the physics-biology interface, from institutions across the world. All first-year PEB students attend the annual meeting of the PoLS SRN, held at one of the institutions in the network, and present. More senior students give oral or poster presentations on their research. The PoLS SRN also funds short research exchanges.
• Physics PEB students receive a Physics Ph.D. and also a Certificate of Completion from the program.

For further information see,  Physical Engineering and Biology (PEB)

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Yale Cancer Biology Institute

Each YCBI member is affiliated with both a Research Program and one or more Clinical Research Team at Yale Cancer Center, with the goal of the Institute being a science driver for clinical and therapeutic innovation at Yale.

Mark Lemmon, PhD

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The Yale Cellular, Molecular, and Quantitative Biology Training Program presents INSPIRE A series of talks for Yale graduate students

Career Options for Ph.D's in Science: Opening the Doors of Opportunity

University of Virginia

Host Organization

• Molecular Biophysics and Biochemistry

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Award-Winning Statistician Joins Yale School of Public Health Leadership

Bhramar mukherjee appointed inaugural senior associate dean of public health data science and data equity, award-winning statistician joins ysph.

From left to right: YSPH Dean Megan L. Ranney, Bhramar Mukherjee, and Steven (Shuangge) Ma.

Award-winning statistician Bhramar Mukherjee, PhD, has been appointed Yale School of Public Health’s inaugural Senior Associate Dean of Public Health Data Science and Data Equity, an important new leadership position that reflects the school’s focus on data science as a critical pillar for the future of public health. She joins YSPH on Aug. 1, 2024.

Mukherjee currently holds a number of distinguished academic positions at the University of Michigan School of Public Health where she is the John D. Kalbfleisch Distinguished University Professor, Siobán D. Harlow Collegiate Professor of Public Health, and chair of the Department of Biostatistics. She is also a professor of epidemiology, and of global public health. In addition, Mukherjee serves as associate director for quantitative data Sciences at the University of Michigan’s Rogel Cancer Center and, last year, she was appointed assistant vice president for research, designing a comprehensive research data analysis service infrastructure for all three U-M campuses.

YSPH Dean Megan L. Ranney, MD, MPH, said Mukherjee’s appointment aligns with the school’s strategic goal of advancing public health data science, a critical part of YSPH’s leadership of the transformation of public health.

“This new role symbolizes our commitment to both rigor and equity in data science as the foundation of the future of public health,” Ranney said in an April 8 schoolwide announcement. “Dr. Mukherjee’s arrival will allow us to coordinate across public health disciplines and across the university in achieving our shared, future-focused, consequentialist vision of public health.”

At YSPH, Mukherjee will also hold the positions of Anna M.R. Lauder endowed professor of biostatistics and professor of chronic disease epidemiology. She will be a member of the Yale Cancer Center director’s cabinet. Additionally, she will hold a secondary appointment in the Department of Statistics and Data Science and will have affiliations with the MacMillan Center and the Institute for the Foundations of Data Science.

This new role symbolizes our commitment to both rigor and equity in data science as the foundation of the future of public health. Dean Megan L. Ranney

As senior associate dean, Dr. Mukherjee will work with Dean Ranney, the school’s senior leadership team, and the broader university community to develop and realize a strategic plan for public health data science and data equity that will enable transformational and impactful research in the face of quickly changing methods, technology, and societal shifts. In doing so, she will lead the development of resources to support the school’s public health data science and data equity research enterprise. She will also work to elevate YSPH’s data science and data equity education programs, with a goal of enhancing the pipeline of diverse scholars in this field. Additionally, representing YSPH’s commitment to global public health, she will support and create international collaborations that enhance the global practice, teaching, and science of public health data science and data equity.

She also plans to continue her own research and teaching.

Mukherjee said she felt “a strong intellectual resonance” with the Yale community when she visited YSPH in August 2023 as the inaugural speaker for the Dean’s Leaders in Public Health lecture series.

“I was inspired by the transformative vision Dean Ranney has for YSPH and more broadly for academic public health,” Mukherjee said. “Data is a quintessential quantum of research, but we often forget to ask the fundamental question: who is in my study? I believe everyone should have equal opportunity of benefitting from data resources and data products.”

“Data equity, both in terms of quantity and quality of data, is a critical pillar of health equity, climate equity and more broadly, societal equity,” Mukherjee continued. “I am deeply honored to have this inaugural leadership position underscoring the importance of data to improve the human condition. I am grateful for the opportunity to work with the Yale community to advance our collective pursuit of creative, impactful, fair, and equitable data science.”

Mukherjee said the decision to leave the University of Michigan after 18 years was “exceptionally hard”. She thanked her students, friends, and colleagues at U-M for “shaping my vision for public health data science.”

Mukherjee joins a strong team of data scientists at YSPH. Faculty in the Department of Biostatistics are considered international leaders in their field and have been extensively recognized for their pioneering work. YSPH’s acclaimed public health modeling unit , which exists across multiple departments in the school, is particularly acclaimed for its deep and direct impact in Connecticut and internationally during the COVID-19 pandemic, providing policymakers with timely data-driven insights to help guide emergency response. YSPH’s core data science centers such as the Yale Center for Analytical Sciences , the Center for Methods in Implementation and Prevention Science , and Center for Perinatal, Pediatric and Environmental Epidemiology , bring researchers and scholars together to spur development of innovative statistical tools and techniques that are used to address today’s most pressing public health and biomedical challenges.

Biostatistics Department Chair Steven (Shuangge) Ma said Mukherjee’s leadership and research strengths will elevate YSPH data science work to a new level.

“Bhramar is a remarkable researcher and a true leader in the field,” Ma said. “She has made profound methodological contributions to the foundation of biostatistics and equally importantly public health practice. We are thrilled to have her join us.”

Mukherjee’s methodological research interests include statistical methods for analysis of electronic health records, studies of gene-environment interactions, shrinkage estimation, data integration, and analysis of multiple pollutants. Her collaborative contributions have focused on cancer, cardiovascular diseases, COVID-19, exposure science, environmental epidemiology, and reproductive health. She has co-authored more than 380 publications in statistics, biostatistics, medicine, and public health and has served as a principal investigator on multiple NSF and NIH grants. She is the founding director of the University of Michigan’s undergraduate Big Data Summer Institute and has been leading the program for the past nine years.

Mukherjee’s innovation and leadership in data science has been widely recognized. She is a fellow of the American Statistical Association and the American Association for the Advancement of Science. She was elected to the National Academy of Medicine in 2022. She is the recipient of many awards for her scholarship, service, and teaching at the University of Michigan and beyond, including the Gertrude M. Cox award, the L. Adrienne Cupples award, the Janet L. Norwood award, the Karl E. Peace Award for Outstanding Statistical Contributions for the Betterment of Society, the Jerome Sacks Award for Outstanding Cross-Disciplinary Research in statistics and, most recently, the 2024 Marvin Zelen Leadership Award. She also is a recipient of the Outstanding Statistics Alumnus award, Distinguished Woman Scholar award, and Distinguished School of Science Alumnus award from her alma mater, Purdue University.

During her distinguished academic career at the University of Michigan, Mukherjee received many prestigious institutional awards including the School of Public Health Excellence in Teaching Award, Rackham Distinguished Faculty Achievement Award, the Sarah Goddard Power award for improving the work environment for women, and the highly coveted Distinguished University Professorship. Mukherjee and her team took an active role in modeling the SARS-CoV-2 virus trajectory in India during the COVID-19 pandemic. She has been featured in many major media outlets including the BBC, Der Spiegel, Forbes, NPR, the New York Times, Reuters, Wall Street Journal, and the Times of India.

• Data Science

Featured in this article

• Megan L. Ranney, MD, MPH Dean of Yale School of Public Health and C.-E. A. Winslow Professor of Public Health (Health Policy) and Professor of Emergency Medicine
• Shuangge Steven Ma, PhD Department Chair and Professor of Biostatistics; Affiliated Faculty, Yale Institute for Global Health; Director, Biostatistics and Bioinformatics Shared Resource

School of Medicine M.D./Ph.D. Training Program

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M.D./Ph.D. Training Program

Elias (eli) wisdom.

Current Program Year: Grad 1 Current Student, Neuroscience Graduate Program, School of Medicine M.D./Ph.D. Program Students, School of Medicine

Research interests: Neuroscience, neurodegeneration, proteomics, Alzheimer’s Disease, Parkinson’s Disease, aging

Clinical interests: ​​​​​​​​​​​​​​ Neurology, dementia, movement disorders, geriatrics

Elias (Eli) Wisdom is an MD-PhD student studying the molecular nature of Alzheimer’s Disease and Parkinson’s Disease under mentorship from Dr. Vivek Unni, MD, Ph.D., and Andrew Emili, PhD. He grew up in La Grande, OR, where he developed a strong appreciation for the natural world, and a deep passion for baseball. Following these, he attended Pacific University, in Forest Grove, OR, where he studied Biology while competing on the varsity baseball team all four years. During the summer before his senior year, he completed an internship at Fred Hutchinson Cancer Research Center in Seattle, WA, under mentorship from Harmit Malik, Ph.D., where he first learned that MD-PhD programs existed. Knowing that he could combine careers in medicine and scientific research with this path, he sought out further research and clinical training and moved to New Haven, CT, where he joined the lab of Daniel Colón-Ramos at Yale School of Medicine to study the cell biology of the synapse. His time in the Colón-Ramos Lab and volunteering at Yale New Haven Hospital affirmed his desire to have a career at the intersection of medicine and research. He was fortunate to matriculate into the MSTP program at Oregon Health & Science University in 2021, where he has been able to have outstanding training in his research and clinical interests while living close to family.

Education and training

​​​​​​ Bachelors of Science, 2019, Pacific University, Oregon

Fellowship : Postbaccalaureate Research Fellowship, Yale University, 2019-2021

Memberships and associations:

• American Neurological Association
• Society for Neuroscience
• Oregon Geriatrics Society
• American College of Physicians
• American Physician Scientist Association
• Beta-Beta-Beta Biological Honor Society
• Phi-Eta-Sigma National Honor Society

Additional information

https://orcid.org/0000-0002-7549-4127

Honors and awards

2018 | Present  Pacific University Berglund Fellow 2019       Fred Hutchinson Intern Alumni Scholarship 2019       Outstanding Senior of School of Natural Sciences, Pacific University 2019       Outstanding Senior in Biology, Pacific University 2019       NCAA Northwest Conference All-Conference Team 2018       $25,000 Inspired Ideas Competition Winner, Entrepreneur Award 2018       Summer Undergraduate Research Program, Fred Hutchinson C.R.C. 2016 | 2019       NCAA Northwest Conference Scholar Athlete 2015 | 2019        Pacific University Honor’s Scholar 2015 | 2019        Oregon Student Scholar 2015 | 2019        Pacific Pacesetter Scholarship 2015 | 2019        Pacific University Dean's List (6 out of 8 semesters)

Publications

Selected publications.

2023                      Almoril-Porras, A., Calvo, A.C., Niu, L., Beagan, J., Hawk, J.D., Aljobeh, A., Wisdom, E. M ., Ren, I., Malcom Díaz-García, M., Wang, Z.W., Colón-Ramos, D.A. “Specific configurations of electrical synapses filter sensory information to drive choices in behavior”. BioRxiv. DOI: https://doi.org/10.1101/2023.08.01.551556 *Currently under review.

2021                      Ransey, E., Chesnov, K., Thomas, G.E., Wisdom, E.M. , Bowman, R., Rodriguez, T., Adamson, E., Almoril-Porras, A., Walder-Christensen, K.K., Hughes, D., Schwennesen, H., Mague, S.D., Colón-Ramos, D.A., Hultman, R., Bursac, N., Dzirasa, K. “Long-term precision editing of neural circuits using engineered gap junction hemichannels”. BioRxiv. DOI: https://doi.org/10.1101/2021.08.24.457429   *Currently under review.

2021                      Hawk, J.D., Wisdom, E.M ., Sengupta, T., Kashlan, Z.D., Colón-Ramos, D.A. “HySyn: A                 genetically encoded tool for reconstituting synthetic modulatory neurotransmission and reconnect neural circuits in vivo” . Nature Communications. DOI: https://doi.org/10.1038/s41467-021-24690-9

2018                      Clark, B., Eberhardt, M., Gift, S., Goodwin, S., Hirasa, K., Kaupiko, K., Lotomau, S., Powell, N., Shick, E., Tamanaha, K., Tang, C., Teal, M., Warren, A., Wisdom, E.M. “Analysis of Microbial Metabolic Diversity in Two Locations at Fernhill Wetlands: A Report to Clean Water Services”. Government Agency Report. 2018. Clean Water Services, Forest Grove, OR. * All authors listed in alphabetical order.

• College of Arts & Sciences
• Graduate Division
• College of Liberal and Professional Studies

• Graduate Student Awards in Microbiology

Microbiology, Virology and Parasitology program within the Cell and Molecular Biology graduate group has announced awards to three talented graduate students. We are excited to see where your research leads you! 

Priyanka Chatterjee is a 4th year PhD student in the Microbiology, Virology and Parasitology subtrack of the Cell and Molecular Biology graduate group. She received the NIH F31 Ruth L. Kirschstein National Research Service Award to fund her dissertation research in the Pohlschröder Lab.

Yirui Hong is a 4th year PhD student in the Biology graduate group. She received the Teece Dissertation Research Award to fund her dissertation research in the Pohlschröder Lab.

Jacob Cote is a 2nd  year Ph.D. student in the Microbiology, Virology and Parasitology subtrack of the Cell and Molecular Biology graduate group. He received an NSF Graduate Research Fellowship (GRFP) to fund his dissertation research in the Pohlschröder Lab.