expression profiling of human brain tissue as a means of ... md-phd class book.pdfanalysis of...

Soluble Amyloid-beta Oligomers and Synaptic Dysfunction in Alzheimer’s Disease Kruppel-like factor 2 as a Central Link Between Blood Flow and Vascular Endothelial Function

Non-Genetic Heterogeneity in Mammalian Cell Fate Determination Genetic Dissection of the Dscam Pathway Identification of an Xist RNA binding protein and a novel genetic element at the X inactivation center Molecular Mechanisms of Erythropoiesis and Globin Gene Regulation The ApplicATion of Decision AnAlyTic

MeThoDs To Diverse public heAlTh probleMs in unDerserveD populATions The Role of Polo-like kinase 2 in synaPTic funcTion and PlasTiciTy TranscripT mapping

wiTh genome Tiling microarrays Combating Biofilms and Antibiotic Resistance with Synthetic Biology Programming and Reprogramming Cellular Identity Ikaros and Mi-2beta mediated hematopoietic lineage determination Mechanotransduction via Airway Epithelial Cells: The Effect of Compressive Stress The

Roles of Myosin ii and Rap2 in synapTic sTRucTuRe and funcTion Analysis of Cerebrospinal Fluid in Brain Development ExprEssion profiling of Human Brain TissuE as a mEans of idEnTifying

novEl CandidaTE gEnEs for sCHizopHrEnia IntegratIng VIsual InformatIon through Parallel Pathways Molecular Imaging of Pancreatic Cancer Microenvironment: Implications

on Therapy The enterohepatic microenvironment modulates immune responses to vaccination Soluble Amyloid-beta Oligomers and Synaptic Dysfunction in Alzheimer’s Disease Kruppel-like factor 2 as a Central Link Between Blood Flow and Vascular Endothelial Function Non-Genetic Heterogeneity in Mammalian Cell Fate Determination

Genetic Dissection of the Dscam Pathway Identification of an Xist RNA binding protein and a novel genetic element at the X inactivation center Molecu-lar Mechanisms of Erythropoiesis and Globin Gene Regulation The ApplicATion of Decision AnAlyTic MeThoDs To Diverse public heAlTh probleMs in unDerserveD populATions The

Role of Polo-like kinase 2 in synaPTic funcTion and PlasTiciTy TranscripT mapping wiTh genome Tiling microarrays Combating Biofilms and Anti-

biotic Resistance with Synthetic Biology Programming and Reprogramming Cellular Identity Ikaros and Mi-2beta mediated hematopoietic lineage determination Mechanotransduction via Airway Epithelial Cells: The Effect of Compressive Stress The Roles of Myosin ii and Rap2 in synapTic sTRucTuRe and funcTion

Analysis of Cerebrospinal Fluid in Brain Development ExprEssion profiling of Human Brain TissuE as a mEans of idEnTifying novEl CandidaTE gEnEs for sCHizopHrEnia IntegratIng VIsual Informa-

tIon through Parallel Pathways Molecular Imaging of Pancreatic Cancer Microenvironment: Implications on Therapy The enterohepatic microenvironment modulates im-

mune responses to vaccination Soluble Amyloid-beta Oligomers and Synaptic Dysfunction in Alzheimer’s Disease Kruppel-like factor 2 as a Central Link Between Blood Flow and

Vascular Endothelial Function Non-Genetic Heterogeneity in Mammalian Cell Fate Determination Genetic Dissection of the Dscam Pathway

Identification of an Xist RNA binding protein and a novel genetic element at the X inactivation center Molecular Mechanisms of Erythropoiesis and Globin Gene Regulation The

ApplicATion of Decision AnAlyTic MeThoDs To Diverse public heAlTh probleMs in unDerserveD populATions The Role of Polo-like kinase 2 in synaPTic funcTion and Plas-

TiciTy TranscripT mapping wiTh genome Tiling microarrays Combating Biofilms and Antibiotic Resistance with Synthetic Biology Programming and Reprogramming Cellular Identity Ikaros and Mi-2beta mediated hematopoietic lineage determination Mechanotransduction via Airway Epithelial Cells: The

Effect of Compressive Stress The Roles of Myosin ii and Rap2 in synapTic sTRucTuRe and funcTion Analysis of Cerebrospinal Fluid in Brain Development ExprEssion profiling

of Human Brain TissuE as a mEans of idEnTifying novEl CandidaTE gEnEs for sCHizopHrEnia IntegratIng VIsual InformatIon through Parallel Pathways Molecular Imaging of Pancreatic

Cancer Microenvironment: Implications on Therapy The enterohepatic microenvironment modulates immune responses to vaccination Soluble Amyloid-beta Oligomers and Synaptic Dysfunction in Alzheimer’s Disease Kruppel-like factor 2 as a Central Link Between Blood Flow and Vascular Endothelial Function Non-Genetic Heterogeneity in Mam-

malian Cell Fate Determination Genetic Dissection of the Dscam Pathway Identification of an Xist RNA binding protein and a novel genetic element at the X inactivation center Molecular Mechanisms of Erythropoiesis and Globin Gene Regulation The ApplicATion of Decision AnAlyTic MeThoDs To Diverse public heAlTh probleMs

in unDerserveD populATions The Role of Polo-like kinase 2 in synaPTic funcTion and PlasTiciTy TranscripT mapping wiTh genome Tiling microarrays

Combating Biofilms and Antibiotic Resistance with Synthetic Biology Programming and Reprogramming Cellular Identity Ikaros and Mi-2beta medi-ated hematopoietic lineage determination Mechanotransduction via Airway Epithelial Cells: The Effect of Compressive Stress The Roles of Myosin ii and Rap2 in synapTic

Harvard Medical ScHool

2 5 t H a n n u a l

S p r i n g d i n n e r

May 25 , 2010

M.D.-Ph.D.

Class of 2010

MAY 25, 2010

The Joseph B. Martin Conference Center

6:00 PM

Cocktail ReceptionClassic Jazz by Tal Shalom-Kobi Trio

7:00 PM

Seating for Dinner

Director’s Welcome and introduction of graduates and mentorsDr. Stephen C. Blacklow

Dean’s Champagne Toast to the GraduatesDr. Jeffrey S. Flier

Special RemarksDr. Stephen C. Blacklow

Ms. Linda Burnley

Graduate SpeakersDr. Hannah Han-Chun Chang

Dr. Onyinye Ijeoma IwealaDr. Carlos Ramon Ponce

Formal Group Photo of Graduates(in dining room at conclusion of remarks)

The M.D.-Ph.D. Program welcomes the families and friends of the graduates

M.D.-Ph.D.Class of 2010

HArvArd MedicAl ScHool

AnnuAl SPring dinner in Honor of THeWelcome to the M.D.-Ph.D. Program’s Annual Spring Dinner in honor of the M.D.-Ph.D. Class of 2010 at

Harvard Medical School (HMS)! We are especially delighted to welcome the family members and significant

others who are joining the graduates, faculty, students, and staff to recognize our graduates tonight.

This year, nineteen students will graduate from our program with both M.D. and Ph.D. degrees. This book

showcases the accomplishments of all the individuals among this select group. Together, these students, who

matriculated at HMS between 1997 and 2006, collectively spent about 240 years of academic study, including

undergraduate and graduate degrees. While at Harvard they spent 8.46 years on average per student, to

complete 19 Ph.D. degrees and 19 M.D. degrees. This year’s class of five women and fourteen men reflects

the diversity of graduate training available to M.D.-Ph.D. students at Harvard Medical School. In all, they

carried out their graduate studies in 8 different programs within Harvard University’s Graduate School of Arts

& Sciences and Massachusetts Institute of Technology. One graduate pursued graduate study at Cambridge

University (UK) and the NIH. While the majority of students pursued their dissertations in the basic sciences,

two of this year’s graduates completed their dissertations within the MIT/HST Medical Engineering Medical

Physics program and another two completed their doctoral programs in Health Policy and Biostatistics.

Please spend a moment to read the individual biographies written by each of the students. Many spent

their early years in cities and towns across the United States: Arizona, California, Colorado, Massachusetts,

Washington, West Virginia, and Washington, D.C. Other students have come from Iran, Mexico, Serbia,

and Taiwan. They went on to complete their undergraduate degrees at 13 different colleges and universities

including Duke, Harvard, Massachusetts Institute of Technology, Princeton, Stanford, Universities of Arizona,

California (Berkeley and Los Angeles), Chicago, Pennsylvania, Utah, and Wisconsin. While at HMS, 10 were

enrolled in the London Health Sciences and Technology (HST) curriculum, and 9 joined the New Pathway,

representing the other four societies (4 Cannon, 1 Castle, 2 Holmes, and 1 Peabody).

While these students are meeting the joint challenges of graduate and medical study, the M.D.-Ph.D. Program

endeavors to provide a nurturing and cohesive environment throughout the course of their studies. The program

is fortunate to be able to provide financial support for the majority of the graduates under the sponsorship of the

NIH-Medical Scientist Training Program (MSTP) Grant (T32 GM07753) and other sources, and wishes it could

provide full funding for all. The graduates participated in our special courses, advising sessions, retreats, dinners,

symposia, lunches, and poster sessions. Thousands of emails also helped us to bring this diverse group of

students together in fulfilling our mission to “educate and inspire the leading physician-scientists of the future.”

We congratulate the 2010 graduates on their numerous achievements and accomplishments towards the

completion of the M.D. and Ph.D. degrees and send our most heartfelt wishes to all for continued discovery,

success and happiness in the future.

Best wishes,

The Students, Faculty, and Staff of the M.D.-Ph.D. Program

©2010 PRESIDENT AND FELLOWS OF HARVARD COLLEGE

Welcome

conTenTS

PAgeS 2–3

Photo Gallery

PAgeS 4–7

The Class of 2010

PAgeS 8–26

Graduates’ Bios

PAgeS 27–29

Photo Galleries

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(3rd row): Mauro Zappaterra, Jubin Ryu, Samuel Ng, Stephen Huffaker; (back row): Daniel Seeburg, Ganesh Shankar, Alexander Marson.

7. Simeon Kimmel and Dr. Anne E. Becker, director of the MD-PhD Program in the social sciences.

8. Dr. Elizabeth Blackburn and Dr. Stephen Blacklow share a moment at the reception following her Leaders in Biomedicine Lecture sponsored by the MD-PhD Program in 2008.

9. Dr. Stephen Blacklow, director of the MD-PhD Program since 1997.

10. Dr. Christopher A. Walsh, director of the MD-PhD Program 2003–2007.

11. Dr. Jules Dienstag, HMS dean of medical education, at the program’s graduation dinner in 2009.

12. Hannah Chang, Salil Garg, and Kyle Farh.

1. At new student orientation in 2006, Dr. Joseph Martin, Ms. Carla Fujimoto, and Dr. Nancy Oriol.

2. Dr. Jeffrey Flier, Dean of the Faculty of Medicine, at HMS Commencement in 2009.

3. Dr. Judy Lieberman at MD-PhD Program dinner.

4. Dr. Marcia Goldberg, senior associate director, and Dr. Joel Hirschhorn, associate director of the MD-PhD Program.

5. Dr. Helen Shields instructs students at their first white coat ceremony.

6. Graduates at Match Day in March 2010 with Ms. Linda Burnley (front center), executive director of the MD-PhD Program; front (L): Hannah Chang, Rebecca Spencer, Carlos Ponce, Ken Lin, Timothy Lu; (2nd row): Kush Parmar, Chara Rydzak, Onyi Iweala;

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4

3

5

6

7 8

9 10

11

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rachel Anne BortnickB.A., University of Chicago (2001)

M.Phil., University of Cambridge (UK) (2002)

Ph.D., Harvard University in Neuroscience (2009)

Dissertation: Genetic Dissection of the Dscam Pathway

M.D., Harvard Medical School (2010)

Pediatrics training in Freiburg, Germany

PAge 8

Hannah Han-chun changA.B., Princeton University (2003)

Ph.D., Harvard University in Biophysics (2008)

Dissertation: Non-Genetic Heterogeneity in Mammalian

Cell Fate Determination


resiDency: Preliminary Medicine at Beth Israel Deaconess

Medical Center, Boston, MA

Ophthalmology at Mass Eye & Ear Infirmary, Boston, MA

PAge 9

Stephen Jeffrey HuffakerB.S., University of Wisconsin (2002)

Ph.D., University of Cambridge (UK) and NIH in

Neuroscience (2007)

Dissertation: Expression Profiling of Human Brain Tissue as a

Means of Identifying Novel Candidate Genes for Schizophrenia


resiDency: Orthopaedic Surgery at Harvard Combined

Orthopaedic Residency Program, Boston, MA

PAge 10

onyinye ijeoma iwealaA.B., Harvard University (2002)

Ph.D., Harvard University in BBS-Experimental Pathology (2009)

Dissertation: The enterohepatic microenvironment modulates

immune responses to vaccination


resiDency: Internal Medicine at Massachusetts General

Hospital, Boston, MA

PAge 11

nikola KojicB.S., University of California, Berkley (2000)

M.S., Massachusetts Institute of Technology (2003)

Ph.D., Massachusetts Institute of Technology in HST MEMP-

Biomedical Engineering (2007)

Dissertation: Mechanotransduction via Airway Epithelial

Cells: The Effect of Compressive Stress


Postdoctoral Associate at Tufts University, Boston, MA

PAge 12

Ken Young linB.A, B.S., Stanford Univeristy (2003)

Ph.D., Harvard University in Biophysics (2008)

Dissertation: Molecular Imaging of Pancreatic Cancer

Microenvironment: Implications on Therapy


resiDency: Preliminary Medicine at University of California

Irvine Medical Center, Orange, CA

Ophthalmology at University of California, Irvine, Orange, CA

PAge 13

Timothy Kuan-Ta luS.B., M.S., Massachusetts Institute of Technology (2003)

Dissertation: Combating Biofilms and Antibiotic Resistance

with Synthetic Biology

Ph.D., Massachusetts Institute of Technology in HST MEMP-

Electrical and Biomedical Engineering (2008)

Assistant Professor of EECS at Massachusetts Institute of

Technology, Cambridge, MA

PAge 14

Kaveh MaghsoudiB.A., Stanford University (1997)

M.S., Stanford University (1998)

Ph.D., Harvard University in Biostatistics (2008)

Dissertation: Transcript mapping with Genome Tiling

Microarrays


resiDency: Preliminary Medicine at Santa Clara Valley

Medical Center, San Jose, CA

Radiation Oncology at University of California, San

Francisco, San Francisco, CA

PAge 15

Alexander MarsonA.B., Harvard University (2001)


Ph.D., Massachusetts Institute of Technology in Biology (2008)

Dissertation: Programming and Reprogramming Cellular

Identity


resiDency: Internal Medicine at Brigham & Women’s


PAge 16

Samuel Yao-Ming ngB.S., B.A., University of Arizona (1998)

Ph.D., Harvard University in Immunology (2009)

Dissertation: Ikaros and Mi-2beta mediated hematopoietic

lineage determination


resiDency: Internal Medicine at Brigham & Women’s


PAge 17

Kush M. ParmarB.A., Princeton University (2002)

Ph.D., Harvard University in BBS-Pathology (2008)

Dissertation: Kruppel-like factor 2 as a Central Link

Between Blood Flow and Vascular Endothelial Function


Associate at 5AM Ventures, Boston, MA

PAge 18

carlos ramon PonceB.S., University of Utah (2001)


Dissertation: Integrating Visual Information Through

Parallel Pathways


resiDency: Pathology / Neuropathology at Massachusetts

General Hospital, Boston, MA

PAge 19

chara elaine rydzakB.A., B.A., Stanford University (2000)

Ph.D., Harvard University in Health Policy (2010)

Dissertation: The Application of Decision Analytic

Methods to Diverse Public Health Problems in Underserved

Populations


resiDency: Preliminary Medicine at Beth Israel Deaconess

Medical Center, Boston, MA

Radiology at Hospital of the University of Pennsylvania,

Philadelphia, PA

PAge 20

continued on next page

Class of 2010HA rvArd Medic A l Sc Hool M.d.-Ph.d. ProgrAM

Key

BBS: BiologicalandBiomedicalSciencesHST: HealthSciencesandTechnologyniH-gPP: NationalInstitutesofHealth–GraduatePartnershipsProgram

6 7
























Jubin Wonsun ryuB.A., B.S., Stanford University (2001)


Dissertation: The Roles of Myosin II and Rap2 in Synaptic

Structure and Function


resiDency: Transitional at Santa Clara Valley Medical Center,

San Jose, CA

Dermatology (Molecular Medicine Research Program) at

University of California, San Francisco, San Francisco, CA

PAge 21

vijay ganesh SankaranB.A., M.S., University of Pennsylvania (2002)


Ph.D., Harvard University in BBS-Genetics (2009)

Dissertation: Molecular Mechanisms of Erythropoiesis and

Globin Gene Regulation


resiDency: Pediatrics at Children’s Hospital, Boston, MA

PAge 22

daniel Philip SeeburgB.S., Duke University (2001)


Dissertation: The Role of Polo-like kinase 2 in Synaptic

Function and Plasticity


resiDency: Preliminary Medicine at University of Maryland

Mercy Medical Center, Baltimore, MD

Radiology at John Hopkins University, Baltimore, MD

PAge 23

ganesh Mani ShankarB.A., Stanford University (2002)


Dissertation: Soluble Amyloid-beta Oligomers and Synaptic

Dysfunction in Alzheimer’s Disease


resiDency: Neurological Surgery at Massachusetts General


PAge 24

rebecca Joy SpencerB.S., Washington University (1997)


Dissertation: Identification of an Xist RNA binding protein

and a novel genetic element at the X inactivation center


resiDency: Preliminary Surgery at Brigham & Women’s


Anesthesiology at Massachusetts General Hospital, Boston, MA

PAge 25

Mauro Walsh ZappaterraB.S., University California, Los Angeles (1998)


Dissertation: Analysis of Cerebrospinal Fluid in Brain

Development


resiDency: Preliminary Medicine at University of California

Irvine Medical Center, Orange, CA

Physical Medicine & Rehabilitation at VA Greater Los

Angeles Healthcare System, Los Angeles, CA

PAge 26

The Md-Phd Program Wishes to Acknowledge the Mentors of the 2010 graduates

g r A d u A T e S M e n T o r S

Rachel Anne Bortnick, M.D., Ph.D. Dietmar Schmucker, Ph.D.

Hannah Han-Chun Chang, M.D., Ph.D. Sui Huang, M.D., Ph.D. and Donald E. Ingber, M.D., Ph.D.

Stephen Jeffrey Huffaker, M.D., Ph.D. Daniel R. Weinberger, M.D. and Sabine Bahn, M.D., Ph.D.

Onyinye Ijeoma Iweala, M.D., Ph.D. Cathryn R. Nagler, Ph.D.

Nikola Kojic, M.D., Ph.D. Jeffrey M. Drazen, M.D. and Daniel Tschumperlin, Ph.D.

Ken Young Lin, M.D., Ph.D. Umar Mahmood, M.D., Ph.D.

Timothy Kuan-Ta Lu, M.D., Ph.D. James J. Collins, Ph.D.

Kaveh Maghsoudi, M.D., Ph.D. X. Shirley Liu, Ph.D. and John Quackenbush, Ph.D.

Alexander Marson, M.D., Ph.D. Richard A. Young, Ph.D. and Rudolf Jaenisch, M.D.

Samuel Yao-Ming Ng, M.D., Ph.D. Katia Georgopoulos, Ph.D.

Kush M. Parmar, M.D., Ph.D. Guillermo Garcia-Cardena, Ph.D.

Carlos Ramon Ponce, M.D., Ph.D. Richard T. Born, M.D.

Chara Elaine Rydzak, M.D., Ph.D. Sue J. Goldie, M.D., M.P.H

Jubin Wonsun Ryu, M.D., Ph.D. Morgan H. Sheng, Ph.D.

Vijay Ganesh Sankaran, M.D., Ph.D. Stuart H. Orkin, M.D.

Daniel Philip Seeburg, M.D., Ph.D. Morgan H. Sheng, Ph.D.

Ganesh Mani Shankar, M.D., Ph.D. Dennis J. Selkoe, M.D. and Bernardo L. Sabatini, M.D., Ph.D.

Rebecca Joy Spencer, M.D., Ph.D. Jeannie T. Lee, M.D., Ph.D.

Mauro Walsh Zappaterra, M.D., Ph.D. Christopher A. Walsh, M.D., Ph.D.

crediTS

The 2010 M.D.- Ph.D. Class Book was produced by the Harvard Medical School Office of the M.D.- Ph.D. Program.

EDITOR: Linda Burnley

PRODUCTION ASSISTANT: Janelle O’Rourke

WRITING: Biographical summaries were written by the graduates.

DESIGN: Kathy Sayre, Faith Hruby

PHOTOGRAPHY

Linda Burnley: page 27, photo 8 | Suzanne Camarata: pages 2–3, photos 3, 5, 8–12; page 27, photos 1, 3–7; page 28, photos 2–5; pages 8–26, individual head shots of the graduates | Steve Gilbert: page 2, photo 2 | Liza Green*: inside front cover; page 2, photos 1, 5; page 7; page 27, photo 2 | Onyi Iweala: page 29, photos 6, 9 | Moshe Jakubowski: page 2, photo 6; page 27, photo 9; pages 28–29, photos 1, 7, 8 | Marc Wein: page 29, photo 10.

Other photos courtesy of the graduates who submitted personal photos for use in this book.

*© President and Fellows of Harvard College on behalf of HMS Media Services, Photo by Liza Green, All Rights Reserved.

8 9

harvard m.d.- ph.d. program Class of 2010 harvard m.d.- ph.d. program Class of 2010

Hannah Han-Chun ChangI arrived in Boston in 2003 not knowing if I would ever be a researcher, a doctor, or both. I came because learning had been a lifelong hobby, and the opportunity to study both medicine and science seemed too good to be true. Having spent my childhood in both Taiwan and the U.S., I was comfortable with traversing between unfamiliar territories. From quantum chemistry I went into gross anatomy and human pathology, then from there to stem cell biology and biophysics, and finally back to clinical medicine. I was guided by a common intellectual thread known only to myself. Along the way, I learned how to be an inquisitive scientist and a caring doctor.

In the end, it was not the academics but the rest of life that provided the most humbling lessons over the past few years. Figuring out how to utilize my idiosyn-cratic set of talents/weaknesses towards meaningful pursuits is an ongoing project. Forming and learning to accept the loss of important relationships remain challeng-ing. Trying to trust clinical medicine again after the recent loss of my sister has been even harder. Through it all, I have been bolstered by the patience of my loving fam-ily and friends. I am also indebted to the MD-PhD Program for bringing me to this Quad beaming with brilliant minds, and so close to suffering patients. This proxim-ity provided me with not only knowledge, but opportunities to develop compassion, resilience, and humility. I could not have asked for more.

Rachel Anne BortnickAs a child growing up near Washington, D.C., I didn’t know I wanted to be a doctor. One of my first career aspirations was to sell children’s shoes, and I also thought about becoming a comedian. Then two things pivotal happened — the first was that I became increasingly fascinated with the medical journals my best friend’s father left lying around their house. Then, at the age of 17, I was handed a unique oppor-tunity to conduct bench research at the National Cancer Institute. This experience, enhanced by the guidance of fantastic mentors, set me on a path that would eventu-ally lead to academic medicine.

Of course, in true MD-PhD student spirit, I pursued other interests along the way. As an undergraduate at the University of Chicago, I studied philosophy in addi-tion to spending long hours in an immunology lab. Another incredible opportunity enabled me to do an MPhil at the University of Cambridge, solidifying my interest in neuroscience. NGO and journalism work in the Republic of Georgia rounded out my trajectory towards a career combining research, clinical medicine and interna-tional health.

MD-PhD is not without challenges, such as long nights in the lab and trying to get that one last experiment to work. There is no way I would have survived the last few years without the support of family and friends. My biggest accomplishment by far in these past 7 years was marrying my husband, Jan Pruszak. I am incredibly grateful for his unending love and support and look forward to embarking on our next step together.

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Onyinye Ijeoma IwealaI was born in Manchester, U.K., on March 12, 1981 to a neurosurgery resident, Ikemba Iweala, and a newly minted economist, Ngozi Okonjo-Iweala. At 11 months old, I relocated with my family to Washington, DC. I spent my childhood in DC and the nearby Maryland suburbs, eventually ending up in Potomac, Maryland. By 6th grade, I knew I wanted a PhD because my mom had one. In 9th grade, I fell in love for the first time, after being introduced to proteins, carbohydrates, and lipids during Freshman Biology. Just knowing that these compounds existed made me feel so empowered, and from that day I knew I wanted to become a physician and a scientist.

I was introduced to the wonders of the wet lab during freshman and sophomore years of college under the tutelage of Peter Romanienko, then a postdoc at the NIH, in the lab of R. Dan Camerini-Otero. I studied the role of Spo11, a type II topoisomerases, in recombination and meiosis in yeast. But more importantly, I learned how to pipette, load and run gels. As a junior in Cathy Nagler’s lab, I was immersed in the wondrous world of mucosal immunology, live attenuated oral vaccines, and gastrointestinal parasites, and had the privilege of being mentored by her graduate student at the time, Don Smith. Family trips to Nigeria gave me the opportunity to watch my father in action deliver-ing primary and urgent care to fellow villagers in his one-man clinic in Umuda Isingwu, Nigeria. This experience coupled with shadowing Cambodian midwives and physicians who educated and treated female sex workers at mobile STD clinics in Phnom Penh, Cambodia, exposed me to the practice of medicine in resource-poor settings.

As an MD-PhD student, I wanted to make a scientific contribution that could potentially impact health-care delivery in these types of settings. As a graduate student, I found myself back in Cathy Nagler’s lab where, four years and several hundred mice later, I characterized the role of immunization route, an innate immune signaling mol-ecule and preexisting chronic infection with a gastrointestinal helminth on the genera-tion of antibody responses to vaccination. As I near the end of my fantastic educational voyage, I would like to thank Cathy Nagler for pushing me and advocating for me throughout my PhD and beyond, Helen Shields and Holmes Society, Steve Blacklow, Linda Burnley, Yi Shen and the MD-PhD Program. Thank you to my husband Andrew, my friends and family for surrounding me with your love and unflagging support.

Stephen Jeffrey HuffakerI was born in Boulder, Colorado, and attended the University of Wisconsin-Mad-ison. As a college freshman, I began working in the laboratory of Dr. Neal First, investigating aspects of development and tissue generation using bovine embryos. Nurturing a growing interest in development and biomedical engineering, I com-pleted my senior honor thesis in the lab of Dr. James Thomson, focusing on gen-erating cardiomyocytes from human embryonic stem cells. After graduating with a BS in genetics, I started graduate school as part of the first class of Cambridge-NIH Health Science Scholars. The program offered the incredibly unique opportunity to combine international education with NIH research. As a Cambridge Scholar, I worked in the laboratories of Daniel Weinberger (NIH) and Sabine Bahn (Univer-sity of Cambridge), where my dissertation focused on the developmental aspects of schizophrenia and, more specifically, on efforts to identify new mechanisms of dis-ease and potential drug targets. However, while I enjoyed the intellectual freedom of research, I was left unsatisfied. I eventually realized that my questioning and tinkering at the bench were rooted in the hope of eventual translation to patients. Consequently, I decided to apply to medical school. After being accepted to Harvard Medical School, two other students in the Oxford/Cambridge NIH program and I developed the idea of an MD-PhD program that would allow for integration of international research and medical training. Eventually, we were successful in developing the Ox/Cam/NIH MSTP, a program that now supports over 65 MD-PhD students.

While in medical school, my interests in development have continued but with a more direct focus on using knowledge of developmental processes to aid in medical therapies. From day one, I have been drawn to orthopaedic surgery, where I believe the processes of development offer incredible opportunities for improved methods of surgical reconstructions and tissue engineering/generation. Though my heart is still out west in Colorado, I am ecstatic to be starting the Harvard Combined Or-thopaedic Residency Program this July.

12 13


Ken Young LinI grew up in Tainan, one of the oldest cities in Taiwan. Memories from childhood feel as if they happened only yesterday. I remember traveling to many places sitting on my dad’s shoulders. He helped me see better both literally and metaphorically. My mom taught me how to play the piano and to appreciate music. Not a day has gone by that I didn’t learn something from her. I would ride on my grandfather’s motorcy-cle to play baseball with him at the park. Upon our return grandma would bring out the most quenching cold drinks and hear our stories. I was the luckiest and happiest child thanks to these people.

I came to Orange County, CA, at age 14. Learning to speak English and navigate through high school was tough but fun. I signed up for cross country thinking that I got to cross the country. Later at Stanford, my mentor John Cooke introduced me to vascular biology and inspired me to consider an MD-PhD career.

I met my wife Tina when I was invited to revisit HMS. It was love at first sight. We got married in 2008. She tirelessly supported me through my med school years de-spite her heavy schedule as a private boarding school faculty. She helped me maintain a balanced healthy life and taught me how to think outside of the box.

Our son Leonardo was born last December. Every day of his life has been a joyous learning experience for us. A smile from him and we’d forget about the pain of con-stantly waking up at night. We are very happy to head to sunny southern California to start our new life as I complete my residency.

Special thanks to my advisor Umar Mahmood, Linda Burnley at the MD-PhD Program office, my wife Tina, my son Leonardo, my parents Alex and Brenda Lin, my in-laws Nobutami and Yuka Okina and their medical foundation (Iryouhojin Shadan Sho-Oh Kai), and in loving memory of my grandparents Peter and Lilly Tu.

Nikola KojicEach reflection, in whole, or in part, that I make on my 10-year journey at HST, always brings fond memories and yields a smile on my face. It was a twisting path of wonders, masterfully bridging the seemingly distant shores of engineering and medicine. As a young student, I had thought those two shores would be nearly im-possible to bridge. But, examples at HST quickly convinced me otherwise. Once on the path, so complete was my conversion (or revelation) that I decided to transfer into the M.D. program after finishing my Ph.D. The joy of working with patients is something that I will treasure forever. Equally valuable has been the journey itself that included a tour-de-force of biology, basic science, and medicine. Growing up in Kragujevac, Serbia, I never could have dreamed of learning so much in such a wonderful environment. The scores of tremendous HST people with whom I have worked and played over the last decade have made me a better researcher and, per-haps more importantly, a better person.

Looking ahead, I decided to return to research with the goal of applying my clini-cal experiences to practical biomedical engineering problems. Presently I am a post-doc at Tufts University and I would like to stay in academia long-term. Finally, I would like to thank my fiancée Ksenija, my parents Gordana and Milos, my brother Aleksandar, my thesis advisors Drs. Jeffrey Drazen and Daniel Tschumperlin, and the entire HST community for their unwavering support over the years.

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Kaveh MaghsoudiI was born in Iran, had a short stint in Fort Collins, Colorado, then back to Iran for 2nd grade, until finally swinging around and settling in Lafayette, Louisiana, the heart of Cajun Country, all without a single frequent flier mile to show for it. Here most of my friends rooted for the Saints, but I reveled in the glory years of the 49ers (my, how things have changed!).

Not surprisingly, I moved to the Bay Area for my undergraduate years to be closer to Jerry Rice, but inadvertently fell in love with college football and basketball instead. Then at HMS, I realized that Stanford games start at 10:30 PM EST and that could interfere with my education. That’s when I decided to do a PhD, so that I didn’t have to be at work until after 9AM. But then I got married and had to get on with my life.

However, since this sketch is for the MD-PhD Program, I should say something about research: I liked it a lot. What I’ve liked even more are the people I’ve had a chance to share this process with: my teachers, and especially advisors Shirley Liu and John Quackenbush who have been great role models, my classmates who have encouraged and inspired me, the HST staff who showed me how to go on this journey, and my wife Jennifer who always let me be myself. Thanks to everyone for being in the right place at the right time!

Timothy Kuan-Ta LuMy research centers on synthetic biology with the ultimate goal of developing syn-thetic circuits and organisms for medical and industrial applications. Application areas of interest to me include designing new therapeutics for infectious diseases and synthetic organisms that produce biofuels. During my time as an M.D.-Ph.D. student, I developed novel synthetic gene circuits with increasing complexity, such as biological memory and the world’s first in vivo biological counters. I also en-gineered bacteriophage and antimicrobial peptides to treat intractable infections caused by bacterial biofilms and antibiotic-resistant bacteria. These technologies are orders-of-magnitude more effective than conventional antimicrobials. I am actively working to translate these treatments into clinical therapeutics as a co-founder of Novophage Therapeutics. Additionally, my current research is focused on several topics — blocking amyloid formation in bacterial and mammalian systems, design-ing rational engineering methods for scalable genetic circuits, and investigating the human microbiome.

During my career in HST, I worked in the lab of James Collins at Boston Univer-sity and HHMI with the support of an HHMI fellowship. While an M.D.-Ph.D. stu-dent, I was awarded the 2008 Lemelson-MIT Student Prize and Grand Prize in the 2008 National Inventors Hall of Fame Collegiate Inventors Competition. I will be joining MIT as an assistant professor and the Broad Institute as an associate mem-ber in fall 2010. In addition to learning medicine and engineering biology, I married my long-time girlfriend, Sandy Wang, in 2009. I have also enjoyed playing club vol-leyball at MIT and learning how to snowboard in the mountains of New England.

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Samuel Yao-Ming NgI was born in Beth Israel Hospital here in the Longwood medical area and my earli-est memories are of western Massachusetts, but the bulk of my childhood was spent in Phoenix, Arizona. My initial attempts to wander out of the desert were thwarted by the Flinn Foundation, which convinced me to stay by picking up the tab for my undergraduate education at the University of Arizona where I majored in Molecu-lar Biology and English Literature. I was finally led out after spending the summer before my senior year working at NIH for Mike Lenardo and being sucked into the field of Immunology and ultimately all the way back to the East Coast.

A few months after starting medical school, I met Katia Georgopoulos, and be-gan a long and fruitful relationship studying gene regulation in the earliest stages of lymphocyte development. Fortunately, I was able to stay out of lab enough nights to meet my wife, Alissa. It is difficult to know which was more difficult, finishing my doctorate or convincing her to move to Boston.

During pre-clinical years in HST, I met many of the people who I now count among my closest friends, most of whom have already graduated from the MD-PhD Program at HMS. While it is somewhat lonely bringing up the tail of my medical school class, although I must credit Kaveh Maghsoudi with not leaving me completely alone, I am looking forward to being reunited in training with many of them at Brigham and Women’s Hospital where I will start my residency in Internal Medicine next year.

Alexander MarsonWhen I was 14 I abandoned my aspirations to be an archeologist—instead I would pursue something called medical research. With little understanding of what this career might hold, I found myself drawn to the mystery and power of biology.

In college, I took a circuitous path through philosophy and neuroscience to discover what would become my major academic interest, how cells enact distinct genetic programs to take on specialized cellular identities. But, only through the first two years of HMS did I come to realize that clinical medicine would be most rewarding for me in continued dialogue with basic science. So my plans came full circle; joining the MD-PhD Program during my second year, I decided once again that medical research would be my career.

During the past eight years I have been mentored by spectacularly talented and spectacularly generous members of the faculty at Harvard and MIT. Many thanks to everyone in the MD-PhD Program. Buck Strewler provided wisdom as I navigated the path from clinical training to research and back. Cliff Tabin encouraged my interest in developmental biology and genetics, introduced me to medical education in Nepal, sat on my thesis committee, and has been a wonderful role model. David Hafler adopted me into his lab and his family. Rick Young and Rudolf Jaenisch of-fered me world-class training and showed me the joys, excitement, and satisfaction of inquiry and discovery.

None of this ever would have happened without my friends, my sister, and my parents, who encouraged me and had incredible patience for the three decades of schooling I needed so that I could be here — ready to begin a medicine residency at the Brigham and embark on a career of medical research.

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Carlos Ramon PonceIn the summer of 2008, I was the humble recipient of two of the highest honors I could ever hope for: a PhD in neurobiology and American citizenship. That hap-piest of periods is now joined by this upcoming Graduation Day. This ceremony will be another reminder that the improbably fortunate path that my life has taken — from a small farm in central Mexico, through years as an undocumented worker in Utah, to the academic richness of Boston — was shaped by many caring individuals. I am deeply indebted to them. I am grateful to my mother, aunt, and uncle, who risked everything they had in the pursuit of a better life in the United States. I am thankful to the high school teachers who saw past my early broken Eng-lish and nudged me into more challenging academic courses. I am grateful to the college mentors that exposed me to the expansive and wonderful range of science, to the many medical and graduate school professors that advanced my knowledge and taught me to think like a scientist and physician, and to Harvard Medical School and the MD-PhD program for their support. I do not see Commencement Day as a testament of individual achievement, but as a reflection of the strong communi-ties that sustain us. As I begin the next phase of my training as a neuroscientist and pathologist, I hope also to contribute positively to the paths of other people’s lives.

Kush M. ParmarI was raised in a sleepy fishing town on the Gulf Coast of Mexico. Growing up knowing nothing but lazy afternoons and the ocean’s music, I could never have imagined having the privilege to be a part of something as incredible as the Har-vard MD-PhD Program. What has made this journey so poignantly beautiful is the community of colleagues and mentors, a community that has been a gateway to phenomenal personal and intellectual growth.

Soon after arriving in Boston from Princeton, I found an inspirational advisor in Guillermo Garcia-Cardena, who was recently starting up his laboratory. My research focused on how local blood flow transforms the biology of blood vessels, conse-quently determining the location of where life-threatening plaques form. Traversing the terrain from a basic question to findings relevant to disease was a fascinating and equally humbling journey. Entering the clinic in 2008, I came to finally appreci-ate the unmatched beauty and responsibility of taking care of patients.

Some of the most profound lessons in life have made their way into my con-sciousness during these years, and I will take the invaluable lessons from this jour-ney as my future unfolds. I have many to thank for these lessons and shared experi-ences, but I must first thank my family for your unyielding love and support: Mom, Dad, Sheila, and a most special bonus of my Ph.D. years — my fiancée Padmaja. My advisor Guillermo has been a tremendous support through the entirety of my time in the Program, as have been my closest friends over the years.

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Jubin Wonsun RyuIn 1977, my parents immigrated from Seoul, Korea, to Laurel, Maryland. Having finished their graduate education, my father was seeking to become an orthopedic surgeon and my mother a librarian. I was born in the fall of 1979, in Philadelphia, PA. Since then, I have lived in many places — very cold places (Minnesota), very hot places (Texas), very hilly places (West Virginia), and very perfect places (California). Most importantly, those years gave me my brother Justin, who was born in 1986.

Boston has been as much a home to me as anywhere. I have lived here for eight years — the longest I have stayed anywhere — and I find myself exiting in middle age. As much as my training has taught me about the specifics of medical knowledge and research, I value more its intangible, internal lessons. The patients and co-work-ers I met in the hospital indelibly stretched and reshaped my imagined boundaries of human joy and sorrow. Just as valuably, my time in graduate school was absent of many of the external markers of time and progress that I had grown accustomed to in the classroom, and it taught me to measure these things on my own terms.

I am indebted to the people who have supported me through these years. I want to thank my Ph.D. advisor Dr. Morgan Sheng, who is unfailingly fair, generous, and kind. Thanks also to my best and dearest of friends — Chen, Ben, Dave, Rich, Young, Jennifer, Michael, Alex, Nancy, Alaka, Jason. Lastly, thank you to my brother, father, and mother — my family and my comfort.

Chara Elaine RydzakI grew up in the rainy climes of Seattle where I spent my childhood exploring the woods, swinging in the backyard, and searching for strawberries in the garden. Given our abundant waterways, I quickly found my athletic calling as a competi-tive synchronized swimmer and split my time between the pool and my studies. Although times were tough, my family always supported me in my academic and extracurricular pursuits, whether it was staying late for Science Olympiad, tolerat-ing hours of squeaky cello playing, or driving to 6 am swim practices. It was these formative experiences that taught me the value of hard work, perseverance, and the joys of team and individual achievement.

At Stanford, I chose to double major in Human Biology and English, two distinct but complementary fields — the first provided practical exposure to the biological sciences, while the other allowed me to ponder the rich complexities of the human condition. This led me, upon graduation, to take a health policy research position at Stanford PCOR. The experience confirmed my desire to gain advanced health policy training in conjunction with clinical training. Shortly thereafter, I enrolled in Harvard’s MD-PhD Program in Health Policy.

Looking back after eight intense years, I feel my experiences at Harvard have been invaluable, helping me develop the knowledge, skills and strength to pursue any goal. I am excited to be taking my next professional step by entering a preliminary medicine year at BIDMC followed by a radiology residency at the University of Pennsylvania.

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Daniel Philip SeeburgAfter growing up in the beautiful cities of San Francisco and Heidelberg, Germany, I returned to the USA to go to college at Duke University. There, I studied philosophy and neuroscience, and, more importantly, met my future wife Whitney. I then moved up north to Boston to enter the MD-PhD Program and she followed a year later to study law. I was lucky to quickly make wonderful friends here among the other entering MD-PhD students and quickly felt at home in Boston.

During my first years in the program, I studied the role of neuromodula-tion in synaptic transmission in the visual system with Chinfei Chen, who was a patient and wonderful mentor. I then did my PhD at MIT under another great mentor, Morgan Sheng, studying molecular pathways underlying homeostatic plasticity in the brain. When I returned to the wards, I fell in love with radiology as a clinical specialty, and in the future, I’d like to use molecular imaging tools to enhance our ability to screen for and detect neurological disease non-invasively.

I would like to thank my wonderful wife Whitney, who has been my life com-panion for almost 12 years. Her patience and encouragement over the years have helped me immeasurably and I consider myself extremely lucky to have her. This last year has been amazing for us with the birth of our daughter Katia. She is such a joy and undoubtedly represents the biggest achievement of my life so far.

Vijay Ganesh SankaranI grew up in the sheltered town of Reading, Massachusetts. As a child, I made occasional trips to India, where I witnessed the striking disparities present in the developing world, which I suspect at least subconsciously had an influence on my eventual goal of pursuing a career in medicine. After finishing college and thinking I would focus my career on basic science, I happened to see David Weatherall speak at Max Perutz’s memorial, which got me excited about the potential of combining medicine and science. As a first year medical student, I had the great fortune to work with Ellis Neufeld, with whom I met a sickle cell patient who forever altered my scientific goals. I was able to pursue my resulting interest in hemoglobin switching as a graduate student thanks to the generous support of my PhD advisor Stuart Orkin, along with invaluable guidance from my mentors Joel Hirschhorn, Sam Lux, and David Nathan. After a lot of struggle and some eventual success, I returned to medical school and realized I truly loved clinical medicine, as well. In the midst of my confusion over career goals, I was fortunate to receive guidance and encouragement from David Nathan and Harvey Lodish to keep pursuing the scientific problems that drove me. So I have continued to pursue the rather tortuous path of becoming a physician-scientist. All along the way, I have been privileged to have had numerous friends and caring mentors who have always kept me grounded. Finally, and most importantly, none of this would have been possible without inspiration from many courageous patients and the unwavering support that I have received from my parents.

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Rebecca Joy SpencerThere was always a microscope set up and a project in progress in the house where I grew up in Meridian, Idaho. My first lessons in anatomy came from dissecting mice my brother and father collected from neighboring fields when the hay bales were re-moved. My mother and father, a chemist and physicist respectively, fostered an early interest in science and provided an environment that nurtured curiosity.

As I worked toward a degree in biochemistry at the University of Washington, my uncle first brought the possibility of participating in a MD-PhD program to my at-tention. He was instrumental in helping me establish myself in a lab that conducted research on Hepatitis C, and in later years I moved on to a lab that investigated anti-tumor drug interaction with DNA. Since reading Barbara McClintock’s autobiogra-phy in high school I had been interested in genetics, and in graduate school I began research in that area. As a PhD student at Harvard I did my thesis investigating X-inactivation using mouse embryonic stem cells with Jeannie Lee as my advisor.

I am extremely thankful for the guidance of my teachers and the steadfastness of the friends I’ve meet since arriving in Boston. I’ve benefited enormously from their mentorship and generosity. I am grateful for my family and their unfailing support. I would especially like to thank my grandfather, whose wisdom and inspiration have been invaluable to me.

Ganesh Mani ShankarI have been shaped by the people who have surrounded me since I was born on that sunny December day in Thousand Oaks, CA. The early years were marked by my parents, V.J. and Devika, and my siblings, Leena and Mark. We were soon joined in the late 80’s by my mom’s parents. As I finished high school, I realized how fortunate I was to have my outlook tinted by growing up in a tri-generational home.

At Stanford I developed dual interests in biology and business. I studied anti-oxidant defenses in lipid peroxidation with Dr. David Stevenson. This was my first foray into a lab and I remain grateful to Dr. Henk Vreman for investing so much into developing me as a student of science. My studies in health policy and econom-ics were guided by Dr. Alan Garber. I finished college with an internal tug of war, not quite sure how all this would come together.

This identity crisis was addressed during my first year at HMS when I heard Dr. Dennis Selkoe speak on Alzheimer’s disease. I was completely taken aback by how willing Dennis was to give me a chance at his science. I was initially taught by Dr. Dominic Walsh, who was finishing his time as a post-doctoral fellow. I decided to do a PhD so that I would not have to prematurely stop my research on the effects of amyloid beta on synapse physiology. Dr. Bernardo Sabatini assumed a large role early in my thesis. I could not imagine a better experience for a student than the one afforded me by Dennis, Bernardo and Dominic. Space precludes me from detailing how each one has deeply contributed to my education.

I was ultimately drawn towards neurosurgery by the prospect of advancing the surgical management of complex neurological disorders. These interests were molded through my interactions with Drs. Art Day, Robert Friedlander and Chris Ogilvy.

Towards the end of my time in medical school, I met the love of my life – Kalpana Narayan. We became engaged in October 2009; I cannot imagine a happier beginning to the rest of my life. She will be heading to University of Pennsylvania for two years and will return back to Boston, where I will be a neurosurgical resident at MGH.

As the chapters turn, I come to appreciate ever more John Donne’s poem “No Man is an Island.” I find myself completely indebted to my family, friends and teach-ers for shaping my world.

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harvard m.d.- ph.d. program Class of 2010

Mauro Walsh ZappaterraMy parents instilled in me two very important life lessons: always reach for the stars and think with your own head. I came to Harvard with that philosophy and it has been a wonderful journey through numerous uncharted territories. I am only here today because of the tremendous love and support I received throughout the years from my wife Cami, my parents Enzo and Stella, my brother Fabrizio, my sister Lara, Cami’s parents Jim and Marie Walsh, Cami’s sister Ashley, my great friends from the MD-PhD Program — John Hanna, Daniel Seeburg, and Sashank Reddy, the entire MD-PhD Program staff, and of course my PhD thesis advisor Christopher A. Walsh who allowed me to study the role of cerebrospinal fluid in brain development.

These years have been quite fruitful and spectacular. The best thing to ever hap-pen to me is that I met my wife Cami in 2002. In paving our path together she has been a true inspiration by pushing me to be my best, teaching me to open my heart, and helping me connect my mind and my heart.

It is with great anticipation, joy, and excitement that we move to California. I will be completing a residency in Physical Medicine and Rehabilitation (PM&R) at the UCLA/Greater Los Angeles VA Healthcare System. The basic philosophy of PM&R is to help people improve their quality of life and to relieve suffering. Being able to work with people and help them improve their quality of life is one of the greatest honors possible and a lifelong responsibility I am excited to take on and willing to carry forward.

1. Hannah Chang.

2. Jordan Kreidberg, Mauro Zappaterra.

3. Rebecca Spencer.

4. Alice Shaw, Onyi Iweala.

5. Mauro Zappaterra, Stephen Blacklow, Zuzana Tothova.

6. Vijay Sanakaran.

7. Amy Saltzman, Steve Huffaker.

8. Kenny Lin.

9. Alex Marson.

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1. The Program’s 25th Annual Retreat in 2007.

2. Hike in the White Mountains.

3. Group is welcomed back at Waterville Valley.

4. Kyle Farh and Hannah Chang.

5. Steve Blacklow, Director (center) with group assembled for photo at retreat.

6. The Retreat’s traditional dance floor.

7. Daniel Seeberg, Ganesh Shankar, and Jubin Ryu on Match Day.

8. Rick Mitchell speaks with Ken Lin and Tim Lu.

9. Onyi Iweala and Sarah Henrickson at PhD Graduation.

10. Marlys Fassett and Kush Parmar who matriculated in MD-PhD Program in 2002.

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