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NIH Institutional Training Grants/Other Training Grants
  • Please contact the Research Administrator for specific DBBS data requests.
  • Three months advance contact is needed for Insitutional Grant or other award application/renewals in order to allow adequate time to meet all internal and external requirements.
Grants Management
Merit Award for Individual National Competitive Fellowships
  • Any PhD or MSTP student who obtains competitive external (fellowship/scholarship) funding awarding at least $25,836 annually in stipend will receive a merit award per year for the duration of the fellowship as long as they remain in good academic standing. At the end of the fellowship the student will return to the current DBBS stipend at that time.
  • The merit award is the full responsibility of the Mentor/Department for advanced students.
Grants Management
Module 3 - Immunotherapy

Module Leaders:  
Nathan Singh, MD, MS
   Assistant Professor of Medicine, Division of Oncology
​​Todd Fehniger, MD, PhD   Professor of Medicine, Division of Oncology

Tue: October 25
Thu: October 27
Tue: November 1
Thu: November 3
Tue: November 8
Thu: November 10
Tue: November 15
Thu: November 17 (Wrap-up) 

Clinical Mentorship Orientation:

Tue: November 22, 2022  2:00 pm - 3:00 pm*

Pathobiology of Human Disease States Course
Module 1 - Diabetes

Module Leader: 
Tom Baranski, MD, PhD
Professor of Medicine, Division of Endocrinology

Tue: August 30
Thu: September 1
Tue: September 6
Thu: September 8
Tue: September 13
Thu: September 15
Tue: September 20
Thu: September 22 (Wrap-up)

Pathobiology of Human Disease States Course
Module 2 - Kidney Disease

Module Leader: 
Carmen Halabi, MD, PhD
Assistant Professor of Pediatrics, Division of Nephrology

Tue: September 27
Thu: September 29
Tue: October 4
Thu: October 6
Tue: October 11 (No Class-Fall Break)
Thu: October 13
Tue: October 18
Thu: October 20 (Wrap-up)​

Pathobiology of Human Disease States Course
Application Process
Students applying in year 1 must be good academic standing. 
Students applying in year 2 must have completed the Program-specific qualifying exam. 
Students are encouraged to have completed one or more of the following courses: Genomics (Bio 5488), Advanced Genetics (Bio 5491) or Fundamentals of Mammalian Genetics (Bio 5285), however all interested students should apply.    
- Name, birth date, address, academic program and year and citizenship.  
- Contact information (e-mail and telephone)
- PI in which thesis work is being conducted
- CV or resume (include a list of graduate courses taken and grade)
- Paragraph description of why the student is interested in the Pathway
- Paragraph description of thesis research.* 
- Assemble these components into a single PDF, and send to
- Two letters of recommendation, one of which is from the thesis advisor*, should be sent directly to
*If you have not yet joined a thesis laboratory then a rotation lab project and a rotation advisor can be used as a substitute.
All application material should be submitted by June 15, 2022, to
Applications will be reviewed by the Pathway Co-directors, Tim Schedl (Genetics), Chris Gurnett (Neurology), Dave Spencer (Medicine) and Gary Stormo (Genetics).
Genetics & Genomics Pathway Application Process

​Washington University U-Pass Program

Washington University in St. Louis offers a comprehensive transportation program through Metro St. Louis, the region's public transportation agency.  The U-Pass program provides Metro passes for full-time students, benefits-eligible faculty and staff, and full-time employees of qualified service providers.  This program provides students and other members of the University community unlimited access to the St. Louis Metropolitan region on public transit.  In addition, there are three bus routes that specifically target the connection of Washington University's campuses to nearby residential and shopping areas.  For more information on the U-Pass program, please call (314) 935-5601 or visit​.  Metro route and schedule information is available online at  Faculty, staff, and students who use the U-Pass as the primary mode of transportation to and from campus may enroll in the Occasional Parking program and are also eligible for the Citizens for Modern Transit Guaranteed Ride Home program.  See your DBBS Program Coordinator to register for the fall semester to become eligible for the U-Pass.

For information on parking & shuttle routes on the Med School campus, visit the Washington University School of Medicine(WUSM) Transportation Office online or call 314-362-6824.

For information on parking and shuttle routes on the Danforth (main) campus, visit the Washington University in St. Louis (WUSTL) Transportation Office​ online or call 314-935-5601.​

Relocating Resources

Washington University's Off-Campus Housing Office provides apartment listings and renting information. The Apartment Referral Service (ARS) ( provides comprehensive information on renting an apartment and also maintains lists of off-campus housing options. ARS maintains lists of non-University owned housing options and works with Quadra​ngle Housing​, the management company for all University-owned properties, many of which are on the MetroBus line.  The HomeShare St. Louis program ( WashU graduate students seeking off-campus housing with homeowners who are older adults (65 years of age or older).​

Most people choose to live within a few miles of either campus in neighborhoods such as the Central West End, Clayton, Richmond Heights, Dogtown or the Loop. There are many safe areas around the campuses, but some areas do vary from street to street. We encourage you to check out your neighborhood before signing a lease, and if you are in doubt, contact Washington University’s Apartment Referral Service for more information. 

Additional rental resources:
Student Renter's Guide - Best Colleges
St. Louis Post-Dispatch On-line Classified Ads
The Riverfront Times Classified Ads (St. Louis’ alternative newsweekly)
Apartment Finder​

Because of the low cost of living in St. Louis and the availability of affordable housing, many people choose to buy property rather than rent. If you are interested in purchasing a house, the St. Louis Association of Realtors is a good place to start as well as the St. Louis Post-Dispatch.

Washington University does not investigate or endorse specific properties or landlords. Tenants are responsible to inspect a rental address and negotiate the lease terms with any landlord.

Relocating Resources
Money Matters

The 2022-2023 annual stipend is $34,500. 

 - Stipend payments are disbursed the last working day of each month. 

 - If direct deposit is not set up in time, your paycheck will be mailed to your local home address.  We encourage direct deposit as the US postal service can have a delayed delivery time which would cause you to receive your paychecks late.

Entering Class 2022 frontload details - PhD.pdf

Download the Frontload form.pdf

 - After you confirmed your start date with your Program Coordinator, please send an email to​ with the following information:

  • Start Date
  • Email address
  • Frontload form

Payroll deadlines to receive information are the following:

  • July 8th
  • August 15th

Once DBBS Finance has collectively received these three items of information, we will begin to process your stipend disbursement.  

You will receive email notifications from MyDay/WorkDay prompting you for required information, which you will enter directly into the system. Once you have entered the required information then the processing moves on to the next steps in setting up your stipend disbursements. 

For more information regarding stipend payments and possible tax implications please visit

International Students:

All new International Students must complete the Check-In Form as soon as possible after their arrival in the U.S.,​​.  Please make certain to have all appropriate original documentation when you complete the Check-In Form.​​

Please visit the WUSTL Office for International Students and Scholars (OISS) webpage for more information information and resources,

For questions concerning your VISA prior to orientation please contact the OISS Office.

Entering Students
Past Course Modules
Year Modules
1992 Sickle Cell Anemia
1993 Cystic Fibrosis
Coronary Artery Disease
1994 Sickle Cell Anemia
Acute Leukemia
1995 Multiple Endocrine Neoplasia
Undue Susceptibility to Infection
Alzheimer’s Disease
1996 Sickle Cell Disease
Multiple Sclerosis
Epstein Barr Virus
1997 Osteoporosis
Congenital Malformations
1998 Sickle Cell Disease
Hepatitis C
Chronic Myelogenous Leukemia
1999 Rheumatoid Arthritis
Prostate Cancer
2000 Sickle Cell Disease
Epstein Barr Virus
Heart Failure
2001 Alzheimer’s Disease
Chronic Myelogenous Leukemia
SLE (Lupus)
2002 Heart Failure
Mellitus Preeclampsia
2003 SLE/Rheumatoid Arthritis
Chronic Myelogenous Leukemia
2004 AIDS
Heart Failure
2005 Alzheimer’s Disease
Breast Cancer
Rheumatoid Arthritis
2006 Sickle Cell Anemia
Congestion Heart Failure
High Risk Pregnancy
2007 Rheumatoid Arthritis
Chronic Myelogenous Leukemia
Alzheimer’s Disease
2008 Cardio Disease/Heart Failure
Diabetes & Metabolism
Osteoporosis/Other Bone Disorders
2009 Sickle Cell Anemia
Major Depressive Disorders
2010 Alzheimer’s Disease
Cardio Disease/Heart Failure
2011 Chronic Myelogenous Leukemia
Cystic Fibrosis
High Risk Pregnancy
2012 Rheumatoid Arthritis
Cardiovascular Disease
2013 Diabetes
High Risk Pregnancy
2014 Translating Global Health
Cardiovascular Disease
Chronic Myelogenous Leukemia
2015 Immunotherapy
High Risk Pregnancy
Alzheimer’s Disease
2016 Evidence Based Medicine/Sickle Cell Disease
Cardiovascular Disease
2017 Immunotherapy
Alzheimer’s Disease
Hearing Loss
2018 High Risk Pregnancy
Pain Management
Cardiovascular Disease
2019 Immunotherapy
2020 Alzheimer’s Disease
Hearing Loss
Cardiovascular Disease
2021 Sickle Cell Disease
High Risk Pregnancy
Pain Management
Pathobiology of Human Disease States Course
Past Course Modules
1. Sickle Cell Anemia
2. Malaria​
3. Diabetes
1. Cystic Fibrosis
2. Coronary Artery Disease
1. Sickle Cell Anemia
2. Emphysema
3. Acute Leukemia 
1. Multiple Endocrine Neoplasia
2. Undue Susceptibility to Infection
3. Alzheimer’s Disease
1. Sickle Cell Disease
2. Multiple Sclerosis
3. Epstein Barr Virus
1. Osteoporosis
2. Thrombophilia
3. Congenital Malformations
1. Sickle Cell Disease
2. Hepatitis C
3. Chronic Myelogenous Leukemia
1. Rheumatoid Arthritis
3. Prostate Cancer
1. Sickle Cell Disease
2. Epstein Barr Virus
3. Heart Failure
1. Alzheimer’s Disease
2. Chronic Myelogenous Leukemia
3. SLE (Lupus)
1. Heart Failure
2. Diabetes Mellitus
3. Preeclampsia
1. SLE/Rheumatoid Arthritis
2. Chronic Myelogenous Leukemia
3. Asthma
2. Heart Failure
3. Preeclampsia
1. Alzheimer’s Disease
2. Breast Cancer
3. Rheumatoid Arthritis
1. Sickle Cell Anemia
2. Congestion Heart Failure
3. High Risk Pregnancy
1. Rheumatoid Arthritis
2. Chronic Myelogenous Leukemia
3. Alzheimer’s Disease
1. Cardio Disease/Heart Failure
2. Diabetes & Metabolism
3. Osteoporosis/Other Bone Disorders
1. Sickle Cell Anemia
3. Major Depressive Disorders
1. Alzheimer’s Disease
2. Cardio Disease/Heart Failure
3. Diabetes
1. Chronic Myelogenous Leukemia
2. Cystic Fibrosis
3. High Risk Pregnancy
1. Rheumatoid Arthritis
2. Cardiovascular Disease
3. Malaria
1. Diabetes
2. High Risk Pregnancy
3. Depression
1. Translating Global Health
2. Cardiovascular Disease
3. Chronic Myelogenous Leukemia
1. Immunotherapy
2. High Risk Pregnancy
3. Alzheimer’s Disease
1. Evidence Based Medicine/Sickle Cell Disease
2. Diabetes
3. Cardiovascular Disease
​2017 ​​1. Immunotherapy
2. Alzheimer's Disease
3. Cariovascular Disease
2018​ ​1. High Risk Pregnancy
2. Pain Management
3. Cardio
​2019 ​​1. Immunotherapy
2. Diabetes
3. Retina
Individualized Clinical Mentorship

An integral component of the Markey Pathway is the Clinical Mentorship. Each student meets on a regular basis with a clinically involved member of the faculty to observe and participate in the clinical enterprise, and to discuss current topics and emerging ideas in clinical medicine. This mechanism provides a continuing exposure to the pathobiology of human disease through ongoing individual interactions with a clinical mentor.

Pathobiology of Human Disease States course

This course focuses on 3 important human diseases each year (topics including cardiovascular disease/heart failure, Alzheimer's disease, diabetes, sickle cell anemia, high risk pregnancy, AIDS, depression, rheumatoid arthritis, chronic myeloid leukemia, and cystic fibrosis).  All pathophysiology is an integrated function.  Each organ system, cell type, physiological state, and pathological stress involves complex interactions which can be dissected and studied in isolation, but must also be evaluated as a whole.  Thus, a single disease state provides a paradigm for students to experience the principal areas in human pathobiology.  The Washington University course faculty includes investigators active in the clinical sciences as well as established basic scientists.


Intent to Graduate Form must be filed:

August 1, 2021 for August 19, 2021 Graduation

October 1, 2021 for a Fall 2021 degree​

Final Dissertation must be electronically submitted to the Graduate School (in its final form, all edits corrected):

August 23, 2021 for August 19, 2021 Graduation

January 7, 2022 for a Fall 2021 degree​

Getting Ready to Graduate
Are meals provided?

A meal card will be provided to each scholar to help supplement food costs

How much is the stipend?

The stipend of $4200 will be paid in installments. The first disbursement will be after the first week of the program; another will be made at the end of June and the final installment will be at the end of the program prior to your departure.

How many summer positions are available in the Amgen Scholars Program at Washington University in St. Louis?

There are 21 positions

When will I know if I am accepted into the WU Amgen Scholars Program?

Notifications will be sent out by April 1st.

Application, Procedures and Criteria for PROGRAM Membership in the DBBS
Faculty on the Investigator/Tenure track at the assistant professor level or above from any Department of Washington University may be nominated for DBBS Program membership by their Department Head/Chair. Rare exceptions to the tenure track policy can be considered for conditional membership by the DBBS Executive Council and, at the School of Medicine, the Academic Affairs Committee.
Candidates for new Program membership in the DBBS will be evaluated using the following criteria, reflecting the importance of training and mentoring. It is expected that DBBS faculty members will have strong credentials in many/most of the following areas:
  • A high quality, independent research program in an area of foundational biological or biomedical research that is congruent with existing DBBS programs and evidence of sustainable research funding or (in the case of new investigators) a strong prognosis for securing funding.
  • A strong record of training and mentoring trainees or (for new investigators) a clear interest and potential to do so. Examples include a record of past trainees and their current professional status and achievements, a list of publications authored by trainees, formal recognition/awards for teaching or mentoring, and other activities that foster the professional development of mentees. Program members are encouraged to craft a mentorship and commitment to diversity statement for posting on the DBBS or lab website.
  • Past and ongoing teaching contributions to graduate and/or medical education. A list should be provided of course titles with a summary of the subject matter taught and the number of contact hours in the classroom.
  • For recently appointed junior faculty who have not had the opportunity to meet these criteria, the basis for a strong teaching, foundational research, and mentorship outlook should be addressed in the nomination letter from the Department Head/Chair.
The process for evaluating candidates that meet the above criteria will include the following steps.
  • The DBBS Associate Dean will screen all candidates for DBBS Program membership and convey their recommendation on the candidate's membership application to the Director(s) of the primary program which the candidate wishes to join.
  • The Program Director(s) will evaluate candidate credentials and suitability of the research program and make a recommendation to the Associate Dean. If admission is declined, a new request can be considered after one year. Appeals can be heard by the DBBS Executive Council.​
Applications must include the six components described below, attached to the DBBS membership application form, which can be downloaded below. Applications must provide evidence of mentoring skills and financial ability to support graduate students (costs include stipend, fees, and any bonus for competitive fellowship awards). 
Required documents:

Recommendation Letter or Supporting Statement from Applicant’s Department Head/Chair or Division Chief, whoever is financially responsible in the event of a lapse of funding by the faculty member (1-2 pages)
Biographical Sketch of Applicant
Current and Pending Support (include information regarding start-up funds, if applicable (1-2 pages)
Research Statement (1-2 pages)
List of Current and Past Trainees, if applicable, and other evidence of mentoring experience and skills (Past 5 Years – 1-2 pages)


See your student coordinator at least THREE - SIX months prior to thesis examination (defense).

If you are an International student, YOU MUST see the International Office prior to setting up a defense date to discuss your Visa status and its implication.

Intent to Graduate Form - The Office of Student Records requires that you complete the Intent to Graduate Form on-line through WebSTAC, see below for graduation deadlines. If you have any problems locating or completing the form in WebSTAC, please contact the Office of Student Records at (314) 935-5959.
Read through the Doctoral Dissertatio​n Guide for formatting guidelines and other important information (found at
One Month Before Thesis Examination:
  • Ask your coordinator for the Dissertation Committee Form, have your program director sign the form and return form to your coordinator.
  • Submit your CV and the dissertation abstract.  (Be sure to follow the guidelines in the Doctoral Dissertation Guide booklet). Each should be initialed by the thesis advisor.
  • Email your coordinator the dissertation title, defense date, time and location of thesis examination. It is the student’s responsibility to reserve a room for the thesis examination.  (Thesis Examination information will be published in the DBBS seminar calendar online.)
  • Submit Defense Payroll Health Form​ to your coordinator.

Two Weeks Before Thesis Examination:
  • Distribute copies of dissertation and a copy of your CV to committee members electronically.  If your PDF file is too large to send by email, we advise you to use the Washington University Large File Transfer System,​.  It is a secure and encrypted tool for transferring large files between people and works in a similar manner to Dropbox.  Members of the Wash U community can access this system with their WUSTL Key.
    Note: Your thesis advisor should be listed as the chairperson on your title page and the date would be your degree date (May, August or December are the only options). If you have questions about permission to use published papers in your thesis, you may wish to visit for help with this topic or you may wish to consult your subject librarian, or email
After Defense:
  • Submit your Thesis Examination Approval form to your coordinator.
  • Submit your finalized dissertation online.  Log on to to create an account. Once your dissertation has been successfully submitted you will receive an email informing you that everything has been received. Final submission must be completed by the deadline, indicated in deadline section below.  Plan to submit your dissertation and paperwork a few days before the deadline. You may need to make corrections to your formatting or fill out additional paperwork.
  • Fill out the Post-Graduation Job​ Survey a​
  • Fill out the Survey of Earned Docto​rates Form at
  • Check in with your coordinator to make sure you have submitted everything you need to submit. 
Binding Dissertation
  • If you would like to have your dissertation bound, visit  to upload your thesis and order bound copies. No copies should be ordered until after the electronic submission of a dissertation to ProQuest has been approved by the Graduate School; the pdf uploaded at should be identical to the approved pdf previously submitted to ProQuest.
  • Should your PI want a copy of your thesis, they can also use the Thesis on Demand site at their own expense. You will need to provide them with an electronic copy of your thesis to utilize this site-
  • Questions regarding dissertation binding should be directed to Andrew Richards, DBBS Director of Information Systems at​.
Getting Ready to Graduate
Defense Packet

Below are all the forms that need to be completed. You still must see your Coordinator for additional items and forms.

MSTP students - please see the MSTP office for instructions.

  1. Visit to access:
    1. Dissertation and Thesis Template
    2. Doctoral Dissert​ation Guide
  2. Visit to access:
    1. Dissertation Defense Committee Form​
    2. ​Survey of Earned Doctorates Form
    3. Job Survey
  3. Intent to Graduate Form - filed via WEBSTAC
  4. Defense Payroll Health Form​
  5. Graduating Student Survey

Getting Ready to Graduate
Do students have access to monies/support for career, training, and professional development?

The Division provides up to $600 toward career, training, and professional development expenses for DBBS students during their graduate training (research meeting, conference or career, training and development). Unallowable expenses are: computer, computer accessories, Wi-Fi charges, software, etc.  Students may request use of the Division funds after the thesis proposal has been approved by the Thesis Advisory Committee.  Students are to complete the Career, Training, and Professional Development Funds Form then submit to the Graduate Student Coordinator for signature, and to confirm available balance. Reimbursements are based on actual expenses incurred and an original, detailed receipt for each expense must be provided. 

Automobile expenses will be reimbursed at current IRS mileage rate; gas receipts cannot be reimbursed. If mileage reimbursement exceeds the cost of airfare, the amount of the reimbursement will be the lesser expense Students must provide a simulation for airfare expense, to be included with corresponding financial documents, if requesting mileage reimbursement. If the cost is greater than $600, the additional expense must be paid by a source other than DBBS. Travel on United States Flag carriers should be used to the maximum extent possible and is required for all international travel. Alcoholic beverages will NOT be reimbursed. 

FOR INTERNATIONAL TRAVELERS: When traveling internationally, DBBS asks students to review the WashingtonUniversity International Travel Policy and strongly encourages the completion of the International Travel Registry​. Both documents can be found on the Washington University Global Opportunities website:

PLEASE NOTE: Expense request is to be entered directly into Workday by student or mentor’s administration (will depend on individual dept accounting protocol).​

DBBS Finance Group
  • Coordinates Executive Council approval of annual stipend level for students and communication to students and faculty of exact levels of financial support for each fiscal year
  • Manages initial payroll appointment and processes student payroll for first 16 months of support for PhD students, in addition to all Medical School years and GR1 level for MSTP students
  • Processes all tuition, resident candidate and student health fees for DBBS students
  • Ensures that all internal and external funds are managed in compliance with institutional and agency regulations
  • Handles all reporting and analysis of historical and projected financial support data for DBBS and University management purposes
  • Manages all aspects of DBBS Training Grant administration and reporting, in addition to providing support for individual student fellowship/award applications and other departmental training grants
Financial Support

Under normal circumstances, the mentor will attend the meeting but allow the student to take the lead in the discussion. Under exceptional circumstance, the chair may call a meeting of the thesis committee absent the student, the research mentor, or both.  However, the principal role of the committee is to facilitate discussion and provide constructive criticism to the student and the mentor.  A quorum of 4 faculty is necessary for a meeting to take place.  The chair is responsible for:

  • Facilitating thesis update meetings every 6 - 12 months
  • Submitting the Thesis Committee Report electronically after each meeting to
  • Confirm that the IDP prepared by the student, including career exploration, has been reviewed.
  • Inquiring whether the student is engaged in research subject to a confidentiality agreement and, if appropriate, the chair will direct the thesis mentor to submit a Conflict of Interest Statement: 
      • Research funding from sources that have intellectual property interests in the research, or in which the PI has personal financial interest, may create a real or perceived conflict of interest, given the dual roles of the principal investigator in obtaining funding for the lab and as a mentor for graduate students. Issues of paramount importance are (i) the ability to publish results in a timely fashion; (ii) the ability to communicate research results openly, especially to members of the thesis committee; and (iii) academic rights to publish and speak freely, especially as related to a graduate student’s thesis and defense. 

Every semester you are required (as a PI/Mentor) to evaluate DBBS students rotating, or doing thesis work in your lab. You will receive an email requesting an evaluation to be completed in the DBBS Portal​.

Test Scores

The Graduate Record Examination (GRE) General test is no longer required by DBBS Programs. Please refer to the individual program web page ( to see if submission GRE scores is optional.  If so, scores must be from tests taken in the last five years. The Subject test is not required. Applicants who need to submit scores are strongly encouraged to schedule the exam early so the official scores will reach DBBS before the December 1st deadline.

ETS Institution Code - 6929

Proficiency in English is required of all applicants. Applicants whose native language is not English must demonstrate English proficiency and are required to provide an official score report verifying such.

For the 2021-2022 Admissions season accepted ELP tests are:

1) TOEFL - Test of English as a Foreign Language (TOEFL ITP Plus and iBT Home will not be accepted)

2) International English Language Testing System (IELTS)

3) Doulingo English Test

Only scores from tests taken within the last two years will be accepted. To be eligible for an automatic (no request needed) English language proficiency waiver the applicant must have completed a full-time bachelor's or master's degree from a regionally accredited university located in the United States or an institution where English is the primary language of instruction.  PLEASE NOTE: The US Immigration service may require a English language proficiency exam score if the entire program of study is less than 3 years in duration.

ETS Institution Code - 6929

Outstanding recommendations and higher quality research experience may make up for lower test scores, so do not let lower scores discourage you from applying.

Admissions- What Makes An App Strong?
I am an international student, how do I know if I am exempt from the TOEFL or IELTS test?

The English proficiency testing requirement may be waived for applicants who meet any of the below criteria:

 •    You have completed an undergraduate or master’s degree in the U.S. and completed the entire program while in the U.S.

•    Is a citizen of Australia, Cameroon, Commonwealth Caribbean nations, Ghana, India, Ireland, Kenya, Liberia, New Zealand, Nigeria, Singapore, Uganda, the United Kingdom, or Zimbabwe.

•    Has completed a minimum of three years of documented study in a university-level academic program in the United States or one of the countries listed above.

•    Has completed a minimum of three years of documented study in an English-medium academic program in Canada, Hong Kong, or South Africa.


Please note that the English proficiency testing requirement is only waived for immigration if criteria in bullets above are met.

Test Scores

The Graduate Record Examination (GRE) General test is no longer required by DBBS Programs, however it is optional for some. Please refer to the individual program web page ( to see if GRE scores may be optionally submitted.  If so, scores must be from tests taken in the last five years.

ETS Institution Code - 6929

English Language Proficiency Exams:

Proficiency in English is required of all applicants. Applicants whose native language is not English must demonstrate English proficiency and are required to provide an official score report from the Test of English as a Foreign Language (TOEFL), the International English Language Testing System (IELTS) or Duolingo. Only scores from tests taken within the last two years will be accepted.  If you have not taken the test, please enter the date you plan to take the test.  Your application can be reviewed with your self-reported scores. Scores must be from tests taken within the last two years. Request TOEFL score reports to be sent to Institution code 6929. IELTS or Duolingo scores will need to be mailed to DBBS PhD Admissions, 660 S. Euclid Avenue, MSC 8226-04-13, St. Louis, MO 63110.

The English proficiency testing requirement may be waived for applicants who meet any of the below criteria: 

•    You have completed an undergraduate or master’s degree in the U.S. and completed the entire program while in the U.S.

•    Is a citizen of Australia, Cameroon, Commonwealth Caribbean nations, Ghana, India, Ireland, Kenya, Liberia, New Zealand, Nigeria, Singapore, Uganda, the United Kingdom, or Zimbabwe.

•    Has completed a minimum of three years of documented study in a university-level academic program in the United States or one of the countries listed above.

•    Has completed a minimum of three years of documented study in an English-medium academic program in Canada, Hong Kong, or South Africa.

Please note that the English proficiency testing requirement is only waived for immigration if criteria in bullets above are met.

PhD Application Instructions
Faculty Mentors


Dixit, Ram: Dr. Dixit focuses on understanding on how the microtubule cytoskeleton regulates plant cell shape. His lab uses transgenic plants and follow fluorescently tagged proteins in living cells using total internal reflection fluorescence microscopy to study dynamics and function of proteins at the single molecule level. In addition, by combining mutational analysis with live imaging of new two-color marker lines generated in the Dixit lab, they examine the way in which microtubule severing proteins are responsible for pruning unaligned cortical microtubules at crossover sites and how this activity is involved in creating ordered arrays. Collaborators: Herzog, Piston.

Herzog, Erik: Dr. Herzog studies the cellular and molecular basis for circadian rhythms, focusing on the suprachiasmatic nucleus of the hypothalamus. By combining electrophysiological and molecular imaging techniques, his lab is identifying pacemaking cells and how these cells coordinate their activities to drive behavior. The lab compares the circadian rhythms expressed behaviorally and by cells and tissues using a variety of techniques including behavioral monitoring and imaging with multielectrode recordings, bioluminescence and fluorescence from animals carrying transgenic reporters. Trainees in the Herzog lab pursue optical and digital imaging of low-light bioluminescence, fluorescence, and bright-field preparations. Dr. Herzog received an Outstanding Mentor Award in 2008. Collaborators: Holy, Culver, Taghert.


An, Hongyu: Dr. An has extensive experience in MR and PET/MR imaging and is the associate director of the Center for Clinical Imaging Research (CCIR). Her expertise includes MRI physics, MR sequence design and programming, image reconstruction, image and data analysis, PET/MR attenuation correction, and motion correction. Simultaneously acquired anatomical, physiological and metabolic MR imaging and physiological and molecular PET imaging provide unprecedented diagnostic and prognostic values in many diseases. A specialty of Dr. An’s group has been developing novel MR based PET attenuation methods. An application area is the important MR imaging challenge of quantifying cerebral oxygenation. Collaborators: Ackerman, Hershey, Woodard. 

Chen, Hong:  Dr. Chen’s research is focused on developing image-guided ultrasound drug delivery (IGUDD) techniques. A new assistant professor, Dr. Chen has a joint appointment with Radiation oncology. Her laboratory is setting up two experimental systems: an ultrasound-image-guided focused ultrasound system and an MRI-guided focused ultrasound system. The goal is to translate basic research advances in imaging and ultrasound therapy into image-guided therapy devices that can impact cancer patient care. Collaborators: Anastasio, Hallahan, Parikh. 

Raman, Barani: Dr. Raman’s research focuses on examining the spatio-temporal signals in neural systems to understand the design and computing principles of biological sensory systems using relatively simple invertebrate models (e.g., Drosophila melanogaster). His lab employ’s a variety of multi-dimensional electrophysiological recording techniques and computational modeling approaches to investigate how dynamic odor signals are encoded as neural representations (odor coding). Recent work from Dr. Raman’s lab, published in Nature Communications and Nature Neuroscience, has revealed the behavioral relevance of combinations of neurons activated by an odorant (i.e., ‘the combinatorial code’) and in the temporal structure of the neural activity (i.e., ‘the temporal code’). Collaborators: Gruev, Holy, Petersen. 


Cooper, John: The laboratory uses a variety of light and electron microscopy techniques to address questions of how cells control their shape and movement. Those techniques might include low-light level fluorescence microscopy of living cell preparations, including spinning-disk confocal and total internal reflection microscopy. Collaborators: Bayly, Piston. 

Mecham, Robert: Dr. Mecham studies the extracellular matrix, the critical material that helps bind together and support the structures and tissues of the human body. He is a well-known leader in uncovering the structure of elastic fiber and understanding the complex process involved in producing it. His laboratory focuses on learning how cells produce elastic fibers, a major component of the extracellular matrix. His work includes live-cell imaging of extracellular matrix assembly. Collaborators: Holtzman, Taber 

Piston, David:  The main research focus of the Piston lab is the understanding of glucose-regulated hormone secretion from islets of Langerhans in the pancreas. To perform live cell measurements in situ and in vivo, his lab develops unique, state-of-the-art fluorescence imaging methods to assay responses along critical signaling pathways in both glucagon-secreting α-cells and insulin-secreting β-cells. These quantitative microscopy measurements are combined with standard biochemical and molecular biological techniques to obtain valuable information that bridges the gap between the known details of the signaling pathways in individual cells and the overall response of a whole islet. Experimental work involves 5D live cell imaging and high-content screening. Collaborators: Nichols, Urano, Gross, Lawson. 


Ackerman, Joseph: Trainees perform research in the development and application of magnetic resonance spectroscopy (MRS) and imaging (MRI) for study of intact biological systems, from cultured cells to mice to man. A major area of research is the development of MR techniques that will provide a more complete understanding of the complex structure and operating organization of mammalian tissues in the intact, functioning state. Collaborators: Bayly, Culver, Weilbaecher. 

Mirica, Liviu: Dr. Mirica uses inorganic chemistry, organic chemistry, and biological chemistry to address metal-mediated processes with energy, biological, and medical relevance. One of his projects involves investigation of the interaction of transition metal ions with Aβ peptides and study of the role of metal ions in amyloid plaque and reactive oxygen species (ROS) formation in patients with AD — whose plaques exhibit unusually high concentrations of copper, iron, and zinc. He is developing Cu-64 complexes that can be employed for PET imaging and early diagnosis of AD. Collaborators: Rath, Tai. 


Gruev, Victor:  Dr. Gruev’s research focuses on borrowing key concepts from nature to develop ultra-sensitive, compact, lightweight and conformal imaging sensors capable of recording spectral and polarization properties with high spatial resolution and to bring these new sensory devices to clinical settings. Gruev’s lab has been able to successfully mimic both the optics and underlying neural circuitry from the visual system of both Morpho butterflies and mantis shrimp by using various nanomaterials and nanofabrication techniques and monolithically integrate them with circuits fabricated with advanced CMOS technologies. The compact realization of these bio-inspired spectral-polarization imaging sensors combined with wearable goggle devices and real-time image processing implemented on FPGA platform, were recently used to translate this technology into the operating room to provide instant visual feedback to physicians. Collaborators: Achilefu, Culver, Raman. 

Pless, Robert:  Dr. Pless works on developing tools for the fundamental mathematical modeling and analysis of motion in video sequences. He co-founded the Media and Machines Laboratory, which now includes five full time faculty and is a focal point for research on Computer Vision, Robotics, Graphics, Medical Imaging and Human Computer Interaction. Driven by biological imaging applications, the primary mathematical tools are data-driven, non-parametric statistical models that represent scene-specific or patient-specific models of common motions and behaviors. These models are ignore distracting motions (e.g., breathing artifacts in CT). Collaborators: Bayly, Leuthardt, Miller, O’Sullivan, Taber. 

Ju, Tau:  Dr. Tau’ works on computer graphics and image analysis with application to biological imaging. His early works pioneered the cage-based deformation paradigm which is now widely used in both entertainment industry and academics. In collaboration with a group of image processing specialists and neuroscientists, his lab used geometric atlases to map the gene expression patterns in the mouse brain. While the prototype of the mapped database (see was initially done in 2D, his lab recently completed a 3D version (hosted on the same website) with the support of an NSF grant. His lab also is working on theoretical foundations and practical algorithms to quantify how “tubular” or “plate-like” an object (or one of its part) is. This work is mostly motivated by the analysis of biological structures in biomedical images with applications to optical and electron microscopy. Collaborators: Dacey, Zipfel, Prior. 


Lew, Mathew:  Dr. Lew, a new faculty recruit, is interested in developing imaging platforms for visualizing biomolecules in living organisms across length scales, from subcellular to whole subjects. He trained in the lab of W.E. Morner (Noble prize 2014). His work primarily focuses on super-resolution microscopy. For example he developed method simultaneous accurate measurement of the 3D position and 2D orientation of single molecules and solutions for mitigating localization errors through modified labeling or optical strategies. On the applications side, he works on labeling and imaging internal cellular structures and external cell surfaces, in 3D, with resolution beyond the diffraction limit. These techniques will enabled the mapping of protein locations and interactions in studies of developmental cell biology. Collaborator: Achilefu. 

Nehorai, AryeDr. Nehorai’s research deals with analysis of space-time data in a number of biomedical areas. In biomedicine, he is developing methods for locating electrical sources in the brain using arrays of electrodes (EEG) or magnetometers (MEG) placed around the head. His solutions are important for clinical applications such as finding origins of seizures, or in neuroscience for mapping the brain functions. He is also developing procedures that find the stiffness of the heart wall using MRI. In microscopy imaging, he is working on algorithms to quantify targets (e.g., antigens, proteins etc.) from 3D microarray-based images, and quantum-dot (q-dot) barcoded microparticle ensembles. Collaborators: Achilefu, Garbow, Song. 

O’Sullivan, Jody:  Dr. O'Sullivan was the director of the Electronic Systems and Signals Research Laboratory (ESSRL) from 1998-2007, and is now dean of the new joint engineering program between University of Missouri-St. Louis and WU. He conducts research in a wide range of science and technology for security applications, including borders, target and object recognition theory, information hiding for secure and clandestine communication, and spectral analysis for biochemical agent detection. Current imaging research includes spiral CT imaging in the presence high-density attenuators and microPET. Collaborators: Tai, Culver. 


Bayly, Phillip:  Dr. Bayly, Professor and Chair of Mechanical Engineering, uses MRI to study deformation and to infer mechanical properties of soft tissue, particularly in the brain and spinal cord. The changes in shape and mechanical properties are important both in rapid events such as brain trauma, and very slow events, such as brain morphogenesis. His students employ MR tagging and analysis of tagged images to study the deformation of the brain during linear angular acceleration of the skull. Dr. Bayly collaborates with other researchers who use MRI measurement of water diffusion to characterize the effects of trauma on the brain and spinal cord, in vivo, in animal models. Collaborators: Ackerman, Carlsson, Cooper, Garbow, Pham. 

Lake, Spence:   Dr. Lake’s research focuses on multiscale structure-function relationships of musculoskeletal soft tissues and joints. He uses various imaging techniques (e.g., quantitative polarized light imaging, two-photo microscopy, x-ray microscopy) to quantify structural organization of tissues at various length scales and correlate with region-specific compositional and mechanical properties. His work seeks to understand fundamental principles that govern how soft tissues function in healthy conditions, how these relationships change in injury/disease, and how connective tissue damage can be better prevented, treated, or replaced.


Weilbaecher, Katherine:  Dr. Weilbaecher’s laboratory investigates the molecular mechanisms of tumor metastasis to bone. They utilize luciferase/GFP labeled osteolytic cancer cell lines and evaluate tumor metastasis and bone tumor growth using in vivo bioluminescence in genetically targeted osteoclast and platelet defective mice. They also utilize MRI and PET imaging to evaluate bone tumor growth and metastasis in spontaneous metastasis tumor mouse models. Trainees gain experience in metastasis biology and host cell/tumor cell interactions using an array of in vivo imaging techniques, including PET, bioluminescence and MRI. Collaborators: Achilefu, Ackerman, Garbow, Lanza. 


Petersen, Steven:  Dr. Peterson pioneered the use of brain imaging (PET and fMRI) to identify brain regions that contribute to attention, learning, memory and language. He also investigates the effects of disease and brain damage on these cognitive processes. Currently, he has two main areas of interest. The first focus is the development of neural mechanisms underlying cognition. Methods have been developed that allow direct statistical comparison of child and adult imaging data. The second focus is identifying and characterizing fMRI signals related to task organization and executive control. Recently his lab developed a series of seminal papers on functional connectivity mapping with MRI related to the management of motion artifacts, the applications of graph theory and the mapping of network hubs. Collaborators: Barch, Culver, Hershey, Raman. 


Holy, Timothy:  Dr. Holy’s research in imaging focuses on developing new optical methods for imaging neuronal activity. He has devised a new method, called objective-couple planar illumination microscopy, for imaging neuronal activity simultaneously in large neuronal populations. This approach uses a sheet of light to provide three-dimensional resolution without point-scanning. The principal advantage of this technique is that hundreds or thousands of neurons can be imaged at high speed and high signal-to-noise ratio. Current work on this technology includes optical and algorithmic methods for enhancing resolution deeper into tissue. Collaborators: Herzog, Raman, Taghert. 

Taghert, Pau:  Dr. Taghert’s research focuses on (i) how peptidergic neurons differentiate and (ii) how neural circuits are controlled by the circadian clock to generate rhythmic behaviors. Both areas of study rely heavily on imaging methods, including standard epifluorescent and confocal microscopy, low light level imaging methods, and use of bioluminesent reporters to interrogate pacemaker neuron function and peptidergic cell secretion mechanisms. Collaborators: Hanson, Herzog, Holy. 


Barch, Deanna:  Dr. Barch’s research program is focused on developing and using a variety of neuroimaging techniques to understand the developmental interplay among cognition, emotion, and brain function to better understand the deficits in behavior and cognition found in illnesses such as schizophrenia, depression and substance abuse. She has a long history of mentoring graduate, postdoctoral fellows and junior faculty in psychology, psychiatry, and neuroscience who have gone on to productive research careers. She was the Director of Graduate Studies in Psychology 2004 to 2014 (now Chair of Psychology) and is a co-Investigator on the Human Connectome Project. Cofounder of our Cognitive, Computational and Systems Neuroscience integrative training pathway, Dr. Barch and has been actively involved in training students in cross-disciplinary neuroimaging research. Collaborators: Petersen, Hershey. 


Hershey, TamaraDr. Hershey’s research is in the fields of neuroimaging and cognitive and clinical neuroscience. Her lab uses a range of neuroimaging, pharmacological and cognitive techniques to understand the impact of metabolic and neurodegenerative conditions on the brain, particularly during development. For example, her lab explores the neural underpinnings of cognitive and mood dysfunction in disorders relevant to dopamine and the basal ganglia (e.g., Parkinson disease, Tourette syndrome), the effects of diabetes and obesity on the brain, particularly within development, and the neurodevelopmental and neurodegenerative impact of a rare monogenic diabetes. Dr. Hershey is deputy lab chief of the WUSM Neuroimaging Labs, and has mentored numerous undergraduate and graduate students, postdocs and junior faculty and co-directs a WU Peer Mentoring Program. Collaborators: Barch, Culver, Raichle. 


Zhang, Tiezhi:  Dr. Zhang’s primary research interests include the development of multi-pixel x-ray source, tetrahedron x-ray imaging systems based on scanning x-ray sources. Almost all modern x-ray imaging systems including x-ray radiography, fluoroscopy, mammography and cone beam CT, to name only a few, utilize a single x-ray source and a 2D detector to acquire 2D images. Dr. Zhang’s lab develops new linear scan x-ray sources and tetrahedron beam imaging systems that can overcome the problems in traditional x-ray imaging, including excessive x-ray scattering, suboptimal detector performance and limited detector dimension. The novel imaging system may find important uses in many medical procedures such as image guided radiotherapy (IGRT), image guided intervention, and office-based point-of-care diagnostic imaging. Besides x-ray imaging, Dr. Zhang’s lab also develops novel technologies for precise radiation (x-ray and proton) treatment of cancers.


Achilefu, Samuel:  Dr. Achilefu is interested in molecular optical imaging, the design and development of new molecular probes and nanomaterials, specific delivery of imaging agents and drugs to target cells or tissues, development of tissue-specific multi-modal imaging molecules, and tumor-specific photodynamic therapy agents. He is co-leader of the oncologic imaging program for the NCI-designated Siteman Cancer Center, and Director of WU molecular imaging center. His Optical Radiology Lab provides a multidisciplinary environment for students in a variety of disciplines, including the chemistry, physics, and biology of optical imaging of diseases. The lab is equipped with state-of-the-art instruments to train the student in all aspects of optical imaging, depending on the expressed interest level of the student. Collaborators: Culver, Gruev, Lew, Shokeen, Weilbaecher, Woodard. 

Benzinger, Tammie:  Dr. Benzinger`s research focuses on translating advanced neuromagnetic resonance imaging techniques from small animal research in the Department of Radiology, to translational research in the Center for Clinical Imaging Research (CCIR), and into clinical practice. In particular, her current research focuses on using directional diffusivity measurements derived from diffusion tensor imaging (DTI) to measure axonal and myelin damage in pediatric and adult demyelination, dysmyelinating diseases, in traumatic brain injury (TBI), and as a function of aging. Diseases under study in Dr. Benzinger`s laboratory include multiple sclerosis (MS), acute disseminated encephalomyelitis (ADEM), adrenoleukodystrophy, Krabbe`s disease, Pelizaeus-Merzbacher`s disease, and head trauma. In addition, Dr. Benzinger combines advanced neuromagnetic resonance techniques, such as DTI and spectroscopy, and positron emission tomography (PET) to study interactions between normal aging, Alzherimer`s disease, depression, and delirium in older adults.  Collaborators: Achilefu, Ackerman, Hershey, Culver, Woodard

Culver, Joseph:  Dr. Culver’s Lab develops neurophotonic technology for mapping brain function in humans and animal models. With the goal of producing high-performance portable brain imaging in humans, his group has been developing a series of innovations for diffuse optical tomography (DOT) instrumentation and algorithms. Recently they presented the first DOT system capable of mapping distributed brain function and networks (Nature Photonics). Applied projects include mapping brain function in infants in the neonatal ICU, and in stroke patients in the Adult ICU. In parallel with human imaging efforts, the Culver lab is also developing mouse equivalent measurements of functional connectivity using optical intrinsic signal imaging (fcOIS) - so as to link human fcMRI with mouse models of disease (e.g., amyloid-beta models of Alzheimer’s, stroke, brain tumors, autism). Recently, to work with faster physiological signals, they have extend fcOIS to mice with genetically encoded calcium indicators and are exploring transitions between awake/sleep and anesthesia. Collaborators: Achilefu, Ackerman, Anastasio, Bruchas, Hershey, O’Sullivan, Petersen, Shokeen. 

Eggebrecht, Adam:  Dr. Eggebrecht’s lab is focused on developing new hardware and software tools for mapping human brain function beyond the reach of current technology. Current projects include optimizing and applying high density diffuse optical tomography (HD-DOT) and functional magnetic resonance imaging (fMRI) to understand how brain function underlies behavior during early childhood development and how it is altered in children with autism spectrum disorder. Additional projects include optimizing HD-DOT for bedside neuromonitoring applications in infants in acute care settings. The Eggebrecht lab also develops computational software suites for modeling, data registration, and analysis of next generation HD-DOT systems. Collaborators: Constantino, Culver, Hershey, Marrus, Said, Smyser.

Shokeen, Monica:  Dr. Shokeen’s lab has expertise in the development and evaluation of molecularly targeted small molecule and multi-functional macromolecular bio-conjugates for nuclear and optical imaging of cancer and cardiovascular diseases. Her group aspires to utilize the translational capabilities of quantitative imaging modalities (PET, SPECT, FMT and MRI) to bring the bench side discoveries into patient care. Working on the chemistry of imaging, the Shokeen lab has been evaluating high-affinity 64Cu labeled-Very Late Antigen-4 (VLA-4) targeted PET radiopharmaceuticals to assess disease progression and response to treatment in pre-clinical mouse and human models of multiple myeloma by quantitative receptor measurements. The ultimate goal of these studies is successful clinical translation. Her group is also investigating the unique metabolic pathways and metabolite fate tracking in multiple myeloma tissues by using 13C edited 1H NMR and 11C-Acetate/PET-CT imaging. Additionally, as part of a multi-PI team, the Schokeen lab is developing a high-throughput optical in vivo imaging platform for the detection of unstable plaque in carotid arteries using a novel custom built Fluorescence Molecular Tomography (FMT) system. Collaborators: Woodard, Achilefu, Culver. 

Tai, Yuan-Chuan:  Dr. Tai’s team conceived and demonstrated the feasibility of the virtual-pinhole PET insert technology for improving the image resolution of existing human and animal PET scanners. This technology is currently being evaluated for whole-body cancer staging to improve the sensitivity of metastatic cancer detection. Additionally, Tai’s lab has developed several high resolution PET and multimodality imaging systems for preclinical, clinical, and functional plant imaging applications. The plant PET imager is now used routinely for molecular plant imaging research and has brought the in vivo imaging technology to plant scientists and triggered new interdisciplinary researches across multiple universities and institutions. Collaborators: O’Sullivan, Laforest. 

Woodard, Pamela:  Dr. Woodard’s expertise is in translational imaging and clinical trials, particularly in cardiovascular MRI, CT and PET. She is Radiology’s Vice Chair of Clinical Translational Research, has an appointment in Biomedical Engineering and is the Director of the Center for Clinical Imaging Research (CCIR). She has been principal investigator (PI) or co-investigator on numerous NIH grants and subcontracts, including the PIOPED II and III Trials. Most recently, her lab has developed a receptor-targeted nanoparticle PET imaging agent for assessment of atherosclerosis, brought it through preclinical safety testing, applied for and received an FDA eIND for testing in human subjects, and have begun testing in normal volunteers and patients. New extensions of the same receptor targeted nanoparticle include optical labelling for imaging with fluorescence molecular tomography. Collaborators: Shokeen, Achilefu, Culver. 

William H. Danforth Fellowship in Plant Sciences

Natasha Bilkey - Plant and Microbial Biosciences

​Jordan Brock - Evolution, Ecology and Population Biology

Ryan Calcutt - Plant and Microbial Biosciences

Ryan Emenecker​​ - Plant and Microbial Biosciences

Emma FrawleyEvolution, Ecology and Population Biology

Wen-Hsi KuoEvolution, Ecology and Population Biology

Erin Mattoon​ - Plant and Microbial Biosciences

Kari MillerPlant and Microbial Biosciences

Angela Schlegel - Plant and Microbial Biosciences

Jennette Shoots - Plant and Microbial Biosciences

External Fellowship Awardees
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