Musculoskeletal Research Training Program (T32)

The University of Minnesota has a rich tradition of training individuals to study musculoskeletal biology and disease. This NIH training program boasts a group of highly qualified mentors in state-of-the-art research environments, providing an ideal modern setting for training young scientists. The research strengths of the training program are in the areas of animal models for bone cancer research and osteoarthritis, outcomes-based research for orthopedic and bone cancer treatments, adult stem cell research, molecular biology, and genomics research.

Program Goals

Our Goals:

  • Train predoctoral students and postdoctoral fellows for research careers as basic scientists or clinical investigators studying the causes and treatments of musculoskeletal diseases

  • Develop an educational environment for students and postdoctoral fellows which encompasses a modern spectrum of research in musculoskeletal diseases, including basic discovery, investigation of musculoskeletal disease models, and study of outcomes of standard treatments

Focus Groups

These primary focus groups allow for and encourage cross-disciplinary interactions between molecular biologists, (stem) cell biologists, orthopedic surgeons, veterinary researchers, and bioengineers. The primary interests of each faculty member guide placement into the groups; however, many faculty members make significant contributions to more than one focus group.

These primary focus groups allow for and encourage cross-disciplinary interactions between molecular biologists, (stem) cell biologists, orthopedic surgeons, veterinary researchers, and bioengineers. The primary interests of each faculty member guide placement into the groups; however, many faculty members make significant contributions to more than one focus group.

Musculoskeletal Cancer Biology and Therapeutics
(D.R. Clohisy,* M. Garwood, D. Largaespada, J. Modiano)

The collective focus of this group is to understand the etiology, progression, and clinical sequelae of musculoskeletal cancers and to develop new therapies for treatment and prevention. Specific areas of this research include:

  • Cellular mechanisms regulating osteolysis in cancer
  • Bone cancer-associated pain and tumor metastasis to bone 
  • Animal models of sarcoma
  • Clinical epidemiology research in osteosarcoma and rhabdomyosarcoma
  • Development of new techniques in MR imaging and oncology 
Osteoarthritis and Cartilage Biology Group
(C. Carlson,* J. Bechtold, P. Ludewig, G.E. Pluhar)

This group has a common interest in understanding the etiology and treatments of osteoarthritis. Specific areas of research include:

  • Development and validation of animal models of osteoarthritis
  • Bone fracture repair
  • Biomechanics of the shoulder joint
  • Pathophysiology of degenerative osteoarthritis of the spine
Musculoskeletal Development Group
(R. Gopalakrishnan, D. Kaufman, K. Mansky, L. Potter, M. O'Connor)

This group is unified by the common interest of understanding the molecular mechanisms that regulate skeletal development, maturation, and differentiation. Specific areas of research include:

  • Osteoblast and osteoclast biology and the role of these cells in the physiologic and pathophysiologic formation of the skeleton
  • Mechanisms involved in limb regeneration in xenopus tadpoles
  • Skeletal aberrations occurring in dwarfism
  • Estrogen signal transduction cascades in bone metabolism and cancer biology
Epidemiology and Clinical Outcomes Research Group
(L. Spector*, M. Swiontkowski)

This group of investigators applies state-of-the-art epidemiologic and clinical outcomes research methodologies to critically evaluate a wide variety of musculoskeletal treatments. This group has multiple strengths but the two most important are their access to large patient/sample populations through cooperative multi-institutional trials and their international reputation in orthopedic-based clinical outcomes research. Specific areas of research include:

  • Assessment of functional outcomes for the treatment of musculoskeletal conditions
  • Etiology and behavior of musculoskeletal cancers
Leadership & Faculty


  • Denis R. Clohisy, MD


  • Cathy Carlson, DVM, PhD
  • Kim C. Mansky, PhD

Advisory Council

  • Paula Ludewig, PhD
  • Lincoln Potter, PhD
  • Logan Spector, PhD


  • Victor Barocas, PhD
  • Joan Bechtold, PhD
  • Bryce Binstadt, MD, PhD
  • Michael Garwood, PhD
  • Raj Gopalakrishnan, DDS, PhD
  • Eric Jensen, PhD
  • Yasuhiko Kawakami, PhD
  • David Largaespada, Ph
  • Dawn Lowe, Ph
  • Jaime Modiano, VMD, PhD
  • Branden S. Moriarity, PhD
  • G. Elizabeth Pluhar, PhD
  • Lincoln Potter, PhD
  • Subree Subramanian, PhD
  • Troy Trumble, DVM, PhD


  • Laura Antos, PhD
    • Characterizing ATP’s effect on natriuretic peptide receptors in relation to long bone growth using fibroblast cell cultures
    • Mentor: Lincoln Potter, PhD
  • Emily Bermel
    • Developing a theoretical/computational model of the degeneration process using growth and remodeling theories in conjunction with realistic models of the tissue geometry and properties
    • Mentor: Victor Barocas, PhD


Charles Billington, PhD

  • Examining the spectrum of craniofacial malformations resulting from a mutation of the gene Twisted Gastrulation
    • Mentor: Anna Petryk, MD

Nicholas Blixt 

  • Providing mechanistic insight into controlling osteoclast differentiation as a potential new therapy for osteoporosis or other diseases characterized by uncontrolled bone loss
    • Mentor: Kim Manksy, PhD

Christina Brakken-Thal, PhD

  • Understanding how Decapentapolegic (Dpp), a BMP protein, is regulated to pattern the dorsal surface of the embryo
    • Mentor: Michael O’Connor, PhD

Aaron Broege, PhD

  • Determining how delta EF1 regulates osteoblast development 
    • Mentor: Michael Sanders, PhD

Amy Claeson, PhD

  • Developing a multi-scale understanding of the structure and mechanical properties of the lumbar facet capsular ligament (FCL) at the spinal, segmental, and tissue levels
    • Mentor: Victor Barocas, PhD

Brandon Diessner, PhD

  • Identifying genetic variants that predispose individuals to osteosarcoma and Ewing sarcoma development
    • Mentor: Logan G. Spector, PhD

Aaron Edmund, PhD

  • Discovering the mechanisms of activation and inactivation of natriuretic peptide receptors
    • Mentor: Lincoln Potter, PhD

Arin Ellingson, PhD 

  • Measuring the helical axis of motion of lumbar spinal motion segments using a human cadaveric model to establish factors that are distinguishable and correlated by the location and/or direction of the helical axis throughout motion
    • Mentor: David Nuckley, PhD

Bora Faulkner, PhD

  • Understanding the mechanism by which histone deacetylase 4 ( HDAC4 ) regulates osteoclasts gene expression and differentiation.
    • Mentor: Kim Mansky, PhD

Sarah Frommer, PhD 

  • Investigating the use of stem cells as treatment for Duchenne Muscular Dystrophy.
    • Mentor: Catherine Verfaillie, MD

Liz Gacek, PhD

  • Mechanically characterizing and computationally modeling the lumbar facet capsular ligament (FCL)
    • Mentor: Victor Barocas, PhD

Craig Kage

  • Cervical spine kinematics
    • Mentor: Arin Ellingson, PhD

Rachel Kahler, PhD 

  • Defining the intentions of the transcription factors, Runx2 and LEF1, and determining the role of LEF1 in osteoblast differentiation.
    • Mentor: Jennifer Westendorf, PhD

Rebekah Lawrence, PT, DPT, OCS

  • Developing a biomechanical model based on kinematic data that will help establish a foundation for a clinical examination of movement impairments for people with shoulder pain
    • Mentor: Paula Ludewig, PhD, PT

Margaret McNulty, PhD 

  • Developing a histological grading scheme for osteoarthritis in mice.
    • Mentor: Cathy Carlson, DVM, PhD

Branden Moriarity, PhD 

  • Identify osteosarcoma (OS) development and metastasis genes by using candidate genes from a forward genetic screen
    • Mentor: David Largaespada, PhD

Christopher Peters, PhD 

  • Identifying unique cellular interactions that occur within the bone microenvironment in both osteolytic and osteoblastic models of bone cancer
    • Mentor: Patrick Mantyh, PhD, JD

Emily Pope, PhD

  • Characterizing adaptations in the extracellular matrix (ECM) remodeling and tumor microenvironment in response to chemotherapy
    • Mentor: David Largaespada, PhD

Jerid Robinson, PhD 

  • The natriuretic peptide receptor B (NPR-B), which has relevance in musculoskeletal research via its effects on long bone growth.
    • Mentor: Lincoln Potter, PhD

Gaura Saini, DPT

  • Testing how anticipated patterns of muscle activation based on movement patterns identified using biplane fluoroscopy correlate to quantitatively observed muscle activity
    • Mentor: Paula Ludewig, PhD, PT

Brandon Smeester

  • Validating candidate cancer genes using the CRISPR system and testing novel therapeutic targets to treat osteosarcoma
    • Mentors: Branden Moriarty, PhD and David Largaespada, PhD


Alexandra Armstrong, DVM

  • Osteochondrosis (OC)/juvenile osteochondritis dissecans (JOCD)
    • Mentor: Cathy Carlson, DVM, PhD, DACVP

Randy Daughters, PhD

  • Understanding the origin and specification of satellite cells during embryonic development and their contribution to muscle regeneration in Xenopus Laevis tadpole tails
    • Mentor: Jonathan Slack, PhD

Dawne Dougherty, PhD

  • Understanding prostate cancer as it progresses to bone metastases, particularly with respect to the contributing hormonal signaling pathways
    • Mentor: Michel Sanders, PhD

Emily Gillespie, MD

  • Discovering underlying systemic autoimmune disease with the hope of improving disease management and generating better therapies
    • Mentor: Timothy Behrens, MD

Curtis Goreham-Voss, PhD

  • Applying finite element analysis to simulate the implant-bone fixation
    • Mentor: Joan Bechtold, PhD

Eric Jensen, PhD 

  • Studying interactions between Runx2 and histone deacetylases in the regulation of bone-specific genes
    • Mentor: Jennifer Westendorf, PhD

Kristina Kiefer, DVM 

  • Evaluating the use of canine adipose derived stem cells as a therapeutic modality for the treatment of intervertebral disc disease and degenerative myelopathy, as well as other various orthopedic conditions in canines
    • Mentor: Elizabeth Pluhar, DVM, PhD

Gengyun (Coco) Le, PhD

  • Investigating how estrogen affects the inflammatory responses and strength recovery following traumatic muscle injury
    • Mentor: Dawn Loe, PhD

Amanda Leightner, PhD

  • Understanding the role of PKD2 in osteoclast differentiation and function
    • Mentor: Eric Jensen, PhD

Carol Lin, MD

  • Comparing reoperation rates and outcomes in patients not enrolled or excluded from SPRINT to assess the influence of enrollment in the trial
    • Mentor: Marc Swiontkowski, MD

John Looft, PhD

  • Developing a high fidelity shoulder finite element model for clinical applications by combining finite element models along with MRI and fluoroscopy to develop predictive models for diagnosing shoulder pain and dysfunction
    • Mentor: Paula Ludewig, PhD

Kai Ludwig, PhD

  • Developing and evaluating advanced magnetic resonance imaging (MRI) techniques at clinical-strength and ultra-high magnetic field for noninvasive diagnosis and monitoring of musculoskeletal diseases (e.g., juvenile osteochondritis dissecans (JOCD)
    • Mentors: Jutta Ellermann MD, PhD, and Greg Metzger, PhD

Shawn Mahmud, MD, PhD

  • Understanding the breakdown of tolerance in human autoimmune diseases by identifying, tracking, and characterizing rare, autoreactive T and B cells to understand how they cause autoimmunity
    • Mentors: Marc Jenkins, PhD, and Bryce Binstadt, MD, PhD

Christiana Raymond-Pope, PhD

  • Examining the musculoskeletal system following complex orthopedic and volumetric muscle loss (VML) injury
    • Mentor: Sarah M. Greising, PhD

Frequently Asked Questions

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Frequently Asked Questions

What is the T32 training grant and what is its purpose?

The training grant is a 5-year award from the National Institute of Arthritis, Musculoskeletal and Skin Diseases (NIAMS) of the NIH. Its purpose is to provide state-of-the-art research training in musculoskeletal biology to predoctoral graduate students and postdoctoral fellows.

How many positions are available on the training grant?

Three predoctoral positions and two postdoctoral positions are supported by this training grant. Each individual may be funded up to a total of three years. After two years of support, trainees wishing to continue for a third year must reapply for the position.

Who is eligible to apply for a position on the training grant?

All individuals supported by this grant mechanism must be a US citizen or noncitizen national (permanent resident). Students on temporary or student visas are not eligible. Predoctoral students must be enrolled in a graduate program at the University of Minnesota and be in good academic standing. Postdoctoral fellows must have a MD, DDS, DVM, and/or PhD and must have had no more than one year of previous postdoctoral support on another NIH institutional training grant or individual NIH fellowship award.

What are the criteria for acceptance into the program?

The criteria for acceptance include the following:

  1. Academic credentials
  2. Strong interest in musculoskeletal research and a desire for a research career
  3. Desirable personal characteristics, including integrity, perseverance, and communications skills

How do individuals apply to the program and what is the application deadline?

A complete application must include the following:

  1. A completed predoctoral or postdoctoral application form that is available online or from Tanya Doble
  2. A statement of career goals and interests
  3. Curriculum vitae
  4. University-certified transcripts
  5. GRE scores (if available)
  6. Reprints of a maximum of 3 peer-reviewed manuscripts
  7. Three letters of recommendation from those who are knowledgeable of the applicant’s academic performance and research skills. If the applicant is currently located at the University of Minnesota and has identified a mentor, the application package should also include a letter of support from the mentor. These materials should be submitted to Tanya Doble, Orthopedic Surgery, 2450 Riverside Av., Suite R200, Minneapolis, MN 55454; Phone 612-273-8052; FAX 612-273-8072; or by email at

What is the level of stipend support for trainees supported by this grant?

Predoctoral stipends are set by the trainee’s graduate program. The NIH provides a set stipend level; any amount over the NIH funded level to match graduate program level must be funded by faculty mentor’s non-sponsored funds.

Postdoctoral stipend levels are set by the NIH each year and vary depending on the number of years of experience.

The training grant can supplement tuition/fees, health insurance, travel, and trainee-related expenses. Exact amounts for predoctoral or postdoctoral support are available here.

Trainees (predoctorate and postdoctorate) are not considered University of Minnesota employees and thus withholding taxes are not withheld unless requested by the trainee. If appointed, you are responsible for addressing this issue, learn more here.

What are the expectations of the trainees?

Trainees are expected to be committed to full-time research training (40 hours/week). Clinical duties must be confined to those that are an integral part of the research training experience. Trainees will be expected to regularly attend program seminars and journal clubs. They will also be required to present journal club and research seminars in designated forums each year. Additionally, trainees are required to complete a research ethics course, a grant writing course, and a Skeletal Biology course (OBIO 8012/CMB 8012).

Clinical fellows may opt to participate in the Department of Orthopaedic Surgery conferences instead of the Skeletal Biology Journal Club. Postdoctoral trainees are strongly encouraged to apply for mentored career awards while being supported by the training grant. Annual reviews of trainees by the Steering Committee is required prior to re-appointment for continued support.

Who are the faculty mentors that are listed on the grant?

Denis Clohisy, MD, Cathy Carlson, DVM, PhD, and Kim Mansky, PhD are the directors of the training program. Additional training faculty are listed on the training grant website under the Leadership & Faculty tab.

Can additional faculty members participate?

Additional mentors may be added, based on the research interests of trainees (i.e. trainee has a research interest outside of the current mentor research expertise and identifies an individual he/she wishes to work with). These individuals must meet the criteria outlined below, including an extramurally funded, independent research program and a strong interest in providing musculoskeletal research training. Faculty members may apply by submitting a letter of interest and a CV to Dr. Kim Mansky. A completed application form is also required. New faculty must be approved by the training program faculty as well as at the NIH program level.

The criteria for membership are: 

  • Full member of the graduate faculty
  • Established, extramurally supported research program 
  • Commitment to strong research training in spirit of the program

Your responsibilities as a mentor will include:
When sponsoring a trainee, mentors will submit a proposal to the steering committee explaining a potential trainee’s project, its merits as a training experience, including cross-disciplinary elements and mentorship, and ancillary supporting experiences that will be given to the trainee.  The mentor will also submit an annual report to the Steering Committee, describing progress, performance, and a summary diary of mentoring of the trainee.  Reports and trainee programs will be reviewed and approved by the Steering Committee to ensure compliance with mission.  As needed, the Steering Committee will determine remedy.

To apply for membership, please complete the attached application form and provide us with the requested materials. Applications from new prospective mentors will be reviewed and approved by the Steering Committee.