Class Book - Harvard Medical School

heimer’s Disease  Kruppel-like factor 2 as a Central Link Between Blo
ermination  Genetic Dissection of the Dscam
 Molecular Mechanisms of Erythropoiesis and Globin Gene Re
ons  The Role of Polo-like kinase 2 in synaPTic funcTion
and Antibiotic Resistance with Synthetic Biology  Program
Harvard Medical School
lineage determination  Mechanotransduction via Airway Epith
n  Analysis of Cerebrospinal Fluid in Brain Development  ExprEs
matIon through Parallel Pathways  Molecular Imaging of
mmune responses to vaccination  Soluble Amyloid-beta Oligom
and Vascular Endothelial Function  Non-Genetic Heterogeneit
way  Identification of an Xist RNA binding protein and a novel genet
 The ApplicATion of Decision AnAlyTic MeThoDs To Diverse public heA
asTiciTy  TranscripT mapping wiTh genome Tiling micr
nd Reprogramming Cellular Identity  Ikaros and M
25th Annual
s: The Effect of Compressive Stress  The Roles of Myosin ii and R
Spring Dinner
rofiling of Human Brain TissuE as a mEans of idEnTifying novEl CandidaTE gEnEs for sCH
M ay 2 5 , 2 0 1 0
ancreatic Cancer Microenvironment: Implications on Therapy  The
mers and Synaptic Dysfunction in Alzheimer’s Disease  Kruppel-lik
n Mammalian Cell Fate Determination  Genetic Dis
ment at the X inactivation center  Molecular Mechanisms of Eryth
robleMs in unDerserveD populATions  The Role of Polo-like kin
oarrays  Combating Biofilms and Antibiotic Resistance with
2beta mediated hematopoietic lineage determination  Me
n synapTic sTRucTuRe and funcTion  Analysis of Cerebrospinal F
Enia  IntegratIng VIsual InformatIon through Paralle
M.D.-Ph.D.
Class of 2010
hepatic microenvironment modulates immune responses to vaccin
2 as a Central Link Between Blood Flow and Vascular Endothelial Funct
Welcome
Welcome 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
Harvard Medical School
Annual Spring Dinner In Honor Of The
M.D.-Ph.D.
Class of 2010
MAY 25, 2010
The Joseph B. Martin Conference Center
6:00 PM
Cocktail Reception
Classic Jazz by Tal Shalom-Kobi Trio
Contents
pages 2–3
Photo Gallery
pages 4–7
The Class of 2010
7:00 PM
Seating for Dinner
Director’s Welcome and introduction of graduates and mentors
Dr. Stephen C. Blacklow
Dean’s Champagne Toast to the Graduates
Dr. Jeffrey S. Flier
Special Remarks
Dr. Stephen C. Blacklow
Ms. Linda Burnley
Graduate Speakers
Dr. Hannah Han-Chun Chang
Dr. Onyinye Ijeoma Iweala
Dr. 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
pages 8–26
Graduates’ Bios
pages 27–29
Photo Galleries
oluble 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 RN
inding protein and a novel genetic element at the X inactivation center  Molecular Mechanisms of Erythropoiesis and Globin Gene Regulation  The ApplicATion of Decision AnAlyT
MeThoDs To Diverse public heAlTh probleMs in unDerserveD populATions  The Role of Polo-like kinase 2 in synaPTic funcTion and PlasTiciTy  TranscripT mappin
6
wiTh genome Tiling microarrays  Combating Biofilms and Antibiotic Resistance with Synthetic Biology  Programming and Reprogramming Cellular
dentity 1 Ikaros and Mi-2beta mediated hematopoietic lineage determination  Mechanotransduction via Airway Epithelial Cells: The Effect of Compressive Stress 
9
of Human Brain10
TissuE as a mEans of idEnTify
oles of Myosin ii and Rap2 in synapTic sTRucTuRe and funcTion  Analysis of Cerebrospinal Fluid in Brain Development  ExprEssion profiling
ovEl CandidaTE gEnEs for sCHizopHrEnia  IntegratIng VIsual InformatIon through Parallel Pathways  Molecular Imaging of Pancreatic Cancer Microenvironment: Implicati
n 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 Determinati
 Genetic Dissection of the Dscam Pathway  Identification of an Xist RNA binding protein and a novel genetic element at the X inactivation center  Molec
ar Mechanisms of Erythropoiesis and Globin Gene Regulation  The ApplicATion of Decision AnAlyTic MeThoDs To Diverse public heAlTh probleMs in unDerserveD populATions  Th
ole of Polo-like kinase 2 in synaPTic funcTion and PlasTiciTy  TranscripT mapping wiTh genome Tiling microarrays  Combating Biofilms and Anti
iotic 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 
nalysis 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
Ion through Parallel Pathways  Molecular Imaging of Pancreatic Cancer Microenvironment: Implications on Therapy  The enterohepatic microenvironment modulates
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 a
2
3
ascular Endothelial Function  Non-Genetic Heterogeneity in Mammalian Cell Fate Determination  Genetic Dissection of the Dscam Pathw
 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  Th
2
3
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8
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Reprogramming Cellular Identity  Ikaros and Mi-2beta mediated hematopoietic lineage determination  Mechanotransduction via Airway Epithelial Cells: T
ffect of Compressive Stress  The Roles of Myosin ii and Rap2 in synapTic sTRucTuRe and funcTion  Analysis of Cerebrospinal Fluid in Brain Development  ExprEssion profilin
f Human Brain TissuE as a mEans of idEnTifying novEl CandidaTE gEnEs for sCHizopHrEnia  IntegratIng VIsual InformatIon through Parallel Pathways  Molecular Imaging of Pancrea
Cancer Microenvironment: Implications on Therapy  The enterohepatic microenvironment modulates immune responses to vaccination  Soluble Amyloid-beta Oligomers an
5
ynaptic Dysfunction in Alzheimer’s Disease  Kruppel-like factor 2 as a Central Link Between Blood Flow and Vascular Endothelial Function  Non-Genetic Heterogeneity in Ma
1. At new student orientation in 2006, Dr. Joseph Martin, Ms. Carla
(3 row): Mauro Zappaterra, Jubin Ryu, Samuel Ng, Stephen Huffaker;
malian Cell Fate Determination  Genetic
Dissection
of the Dscam Pathway
Identification
of an Xist RNA
binding protein and a novel genetic eleme
Fujimoto,
and Dr. Nancy Oriol.
(back row): Daniel Seeburg,
Ganesh
Shankar, Alexander Marson.
12
2. Dr. Jeffrey Flier, Dean of the Faculty of Medicine, at HMS
7. Simeon Kimmel and Dr. Anne E. Becker, director of the MD-PhD
t the
of Erythropoiesis and Globin Gene Regulation
 The ApplicATion of Decision AnAlyTic MeThoDs To Diverse public heAlTh prob
4 X inactivation center  Molecular Mechanisms
Commencement in 2009.
Program in the social sciences.
3. Dr. Judy kinase
Lieberman at MD-PhD
dinner.
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Blackburn and Dr.
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at the
n unDerserveD populATions  The Role of Polo-like
2 inProgram
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the MD-PhD
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Synthetic
Biology
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Hirschhorn,
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of the MD-PhD
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5. D
r. Helen Shields instructs students at theirvia
first white
coat
ted hematopoietic lineage determination 
Mechanotransduction
Airway
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rd
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;
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.
oluble 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 RN
inding protein and a novel genetic element at the X inactivation center  Molecular Mechanisms of Erythropoiesis and Globin Gene Regulation  The ApplicATion of Decision AnAlyT
MeThoDs To Diverse public heAlTh probleMs in unDerserveD populATions  The Role of Polo-like kinase 2 in synaPTic funcTion and PlasTiciTy  TranscripT mappin
wiTh genome Tiling microarrays  Combating Biofilms and Antibiotic Resistance with Synthetic Biology  Programming and Reprogramming Cellular
dentity  Ikaros and Mi-2beta mediated hematopoietic lineage determination  Mechanotransduction via Airway Epithelial Cells: The Effect of Compressive Stress 
oles 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 idEnTify
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 Genetic Dissection of the Dscam Pathway  Identification of an Xist RNA binding protein and a novel genetic element at the X inactivation center  Molec
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Class of 2010
issertation
issertation
issertation
issertation
esidency
Page 8
Page 18
Page 12
Page 15
issertation
issertation
issertation
esidency
esidency
esidency
issertation
Page 19
Page 9
Page 13
esidency
Page 16
issertation
issertation
issertation
esidency
esidency
issertation
Page 10
Page 14
esidency
Page 20
continued on next page
Page 17
issertation
M.D., Harvard Medical School (2010)
Residency: Internal Medicine at Massachusetts General
Hospital, Boston, MA
Page 11
Key
BBS:
HST:
NIH-GPP:
Biological and Biomedical Sciences
Health Sciences and Technology
National Institutes of Health – Graduate Partnerships Program
entity  Ikaros and Mi-2beta mediated hematopoietic lineage determination  Mechanotransduction via Airway Epithelial Cells: The Effect of Compressive Stress  T
oles 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 idEnTifyin
vEl CandidaTE gEnEs for sCHizopHrEnia  IntegratIng VIsual InformatIon through Parallel Pathways  Molecular Imaging of Pancreatic Cancer Microenvironment: Implicatio
Therapy  The enterohepatic microenvironment modulates immune responses to vaccination  Soluble Amyloid-beta Oligomers and Synaptic Dysfunction in Alzheimer’s
isease  Kruppel-like factor 2 as a Central Link Between Blood Flow and Vascular Endothelial Function  Non-Genetic Heterogeneity in Mammalian Cell Fate Determinatio
Genetic Dissection of the Dscam Pathway  Identification of an Xist RNA binding protein and a novel genetic element at the X inactivation center  Molecu
r Mechanisms of Erythropoiesis and Globin Gene Regulation  The ApplicATion of Decision AnAlyTic MeThoDs ToThe
Diverse
public
heAlTh
probleMs
in unDerserveD populATions  Th
MD-PhD
Program
Wishes
to Acknowledge
Mentors
of the 2010 Graduates
ole of Polo-like kinase 2 in synaPTic funcTion and PlasTiciTy  TranscripT mapping wiTh genomethe
Tiling
microarrays
 Combating Biofilms and Antiotic Resistance with Synthetic Biology  Programming and Reprogramming Cellular Identity  Ikaros and Mi-2beta mediated hematopoietic lineage
Jubin Wonsun Ryu
Ganesh Mani Shankar
G r a d u at e s
Mentors
etermination
Mechanotransduction
via Airway
Cells:
B.A., B.S.,
Stanford
University (2001)
B.A.,Epithelial
Stanford University
(2002) The Effect of Compressive Stress  The Roles of Myosin ii and Rap2 in synapTic sTRucTuRe and funcTion 
Rachel Anne Bortnick, M.D., Ph.D.
Dietmar Schmucker, Ph.D.
Ph.D., Massachusetts Institute of Technology in Biology (2008)
Ph.D., Harvard University in Neuroscience (2008)
nalysis of Cerebrospinal
Fluid
in
Brain
Development

ExprEssion
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Brain
TissuE
as
a
mEans
of
idEnTifying
novEl
CandidaTE
gEnEs
for
sCHizopHrEnia
Hannah
Han-Chun
Chang,
M.D.,
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M.D., Harvard Medical School (2010)
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Onyinye Ijeoma Iweala, M.D., Ph.D.
Cathryn R. Nagler, Ph.D.
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Nikola Kojić, M.D., Ph.D.
Jeffrey M. Drazen, M.D. and Daniel Tschumperlin, Ph.D.
San Jose, CA
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Ken Young Lin, M.D.,
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Kuan-Ta Lu, M.D., Ph.D.
James J. Collins, Ph.D.
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Molecular
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Kaveh Maghsoudi, M.D., Ph.D.
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Harvard
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Rebecca Joy Spencer, M.D., Ph.D.
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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
Page 26
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.
h a r va r d m.d.- ph.d. p r o g r a m
8
Class of 2010
h a r va r d m.d.- ph.d. p r o g r a m
Class of 2010
Rachel Anne Bortnick
Hannah Han-Chun Chang
As 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 opportunity 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 eventually 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 addition 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 international 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.
I 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 idiosyncratic set of talents/weaknesses towards meaningful pursuits is an ongoing project.
Forming and learning to accept the loss of important relationships remain challenging. 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 family 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 proximity provided me with not only knowledge, but opportunities to develop compassion,
resilience, and humility. I could not have asked for more.
9
h a r va r d m.d.- ph.d. p r o g r a m
10
Class of 2010
h a r va r d m.d.- ph.d. p r o g r a m
Class of 2010
Stephen Jeffrey Huffaker
Onyinye Ijeoma Iweala
I was born in Boulder, Colorado, and attended the University of Wisconsin-Madison. 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 completed my senior honor thesis in the lab of Dr. James Thomson, focusing on generating 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 (University of Cambridge), where my dissertation focused on the developmental aspects of
schizophrenia and, more specifically, on efforts to identify new mechanisms of disease 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 Orthopaedic Residency Program this July.
I 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 delivering 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 molecule and preexisting chronic infection with a gastrointestinal helminth on the generation 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.
11
h a r va r d m.d.- ph.d. p r o g r a m
12
Class of 2010
h a r va r d m.d.- ph.d. p r o g r a m
Class of 2010
Nikola Kojić
Ken Young Lin
Each 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 impossible 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, perhaps more importantly, a better person.
Looking ahead, I decided to return to research with the goal of applying my clinical experiences to practical biomedical engineering problems. Presently I am a postdoc 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.
I 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 motorcycle 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 despite 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 constantly 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.
13
h a r va r d m.d.- ph.d. p r o g r a m
14
Class of 2010
h a r va r d m.d.- ph.d. p r o g r a m
Class of 2010
Timothy Kuan-Ta Lu
Kaveh Maghsoudi
My research centers on synthetic biology with the ultimate goal of developing synthetic 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 engineered 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, designing 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 University and HHMI with the support of an HHMI fellowship. While an M.D.-Ph.D. student, 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 member 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 volleyball at MIT and learning how to snowboard in the mountains of New England.
I 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!
15
h a r va r d m.d.- ph.d. p r o g r a m
16
Class of 2010
h a r va r d m.d.- ph.d. p r o g r a m
Class of 2010
Alexander Marson
Samuel Yao-Ming Ng
When 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 offered 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.
I was born in Beth Israel Hospital here in the Longwood medical area and my earliest 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 Molecular 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 began 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 MDPhD 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.
17
h a r va r d m.d.- ph.d. p r o g r a m
18
Class of 2010
h a r va r d m.d.- ph.d. p r o g r a m
Class of 2010
Kush M. Parmar
Carlos Ramon Ponce
I 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 Harvard 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, consequently 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 appreciate the unmatched beauty and responsibility of taking care of patients.
Some of the most profound lessons in life have made their way into my consciousness during these years, and I will take the invaluable lessons from this journey as my future unfolds. I have many to thank for these lessons and shared experiences, 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.
In 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 happiest 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 English 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 communities 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.
19
h a r va r d m.d.- ph.d. p r o g r a m
20
Class of 2010
h a r va r d m.d.- ph.d. p r o g r a m
Class of 2010
Chara Elaine Rydzak
Jubin Wonsun Ryu
I 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 competitive 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, tolerating 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.
In 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-workers 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.
21
h a r va r d m.d.- ph.d. p r o g r a m
22
Class of 2010
h a r va r d m.d.- ph.d. p r o g r a m
Class of 2010
Vijay Ganesh Sankaran
Daniel Philip Seeburg
I 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.
After 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 neuromodulation 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 companion 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.
23
h a r va r d m.d.- ph.d. p r o g r a m
24
Class of 2010
h a r va r d m.d.- ph.d. p r o g r a m
Class of 2010
Ganesh Mani Shankar
Rebecca Joy Spencer
I 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 antioxidant 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 economics 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 teachers for shaping my world.
There 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 removed. 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 attention. 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 antitumor drug interaction with DNA. Since reading Barbara McClintock’s autobiography 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.
25
oluble 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 RN
a r va r d m.d.- ph.d. p r o g r a m Class of 2010
inding hprotein
and a novel genetic element at the X inactivation center  Molecular Mechanisms of Erythropoiesis and Globin Gene Regulation  The ApplicATion of Decision AnAlyT
MeThoDs To Diverse public heAlTh probleMs in unDerserveD populATions  The Role of Polo-like kinase 2 in synaPTic funcTion and PlasTiciTy  TranscripT mappin
wiTh genome Tiling microarrays  Combating Biofilms and Antibiotic Resistance with Synthetic Biology  Programming and Reprogramming Cellular
dentity  Ikaros and Mi-2beta mediated hematopoietic lineage determination  Mechanotransduction via Airway Epithelial Cells: The Effect of Compressive Stress 
1
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My parents
in me two very important
life lessons:
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and think with your own head. I came to Harvard with that philosophy and it has
n Therapy  The enterohepatic
microenvironment
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responses
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been a wonderful
journey through numerous
uncharted territories.
I am only
here
today2because
of the tremendous
love and support
I received
the yearsEndothelial Function  Non-Genetic Heterogeneity in Mammalian Cell Fate Determinati
Disease  Kruppel-like factor
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from my wife Cami, my parents Enzo and Stella, my brother Fabrizio, my sister Lara,
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Cami’s parents
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and Marie
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the
MD-PhD Program — John Hanna, Daniel Seeburg, and Sashank Reddy, the entire
ar Mechanisms of Erythropoiesis
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MD-PhD Program staff, and of course my PhD thesis advisor Christopher A. Walsh
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and helping me connect my mind and my heart.
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Ion through ParallelUCLA/Greater
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Los Angeles VA Healthcare System. The basic philosophy of PM&R
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26
27
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carry forward.
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9
malian Cell Fate Determination  Genetic Dissection of the Dscam Pathway
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6
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 Genetic Dissection of the Dscam Pathway  Identification of an Xist RNA binding protein and a novel genetic element at the X inactivation center  Molec
ar Mechanisms of Erythropoiesis and Globin Gene Regulation  The ApplicATion of Decision AnAlyTic MeThoDs To Diverse public heAlTh probleMs in unDerserveD populATions  Th
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iotic Resistance with Synthetic Biology  Programming and Reprogramming Cellular Identity  Ikaros and Mi-2beta mediated hematopoietic lineage
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Ion through Parallel Pathways  Molecular Imaging of Pancreatic Cancer Microenvironment: Implications on Therapy  The enterohepatic microenvironment modulates
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 a
ascular Endothelial Function  Non-Genetic Heterogeneity in Mammalian Cell Fate Determination  Genetic Dissection of the Dscam Pathw
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7
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28
29
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Waterville Valley.
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with group assembled for photo
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6. The Retreat’s traditional dance floor.
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who matriculated in MD-PhD
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Cert no. SW-COC-002508
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Analysis of Cerebrospinal Fluid in Brain Development  ExprEssion pr
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Cancer Microenvironment: Implications on Therapy  The enteroh
www.hms.harvard.edu/md_phd
Synaptic Dysfunction in Alzheimer’s Disease  Kruppel-like factor