1. Dr. GUNARETNAM (GUNA)
RAJAGOPAL
PERSONAL DETAILS
Citizenship: British (United States Permanent Resident)
Marital Status: Married (Dr. Khaw Lake Ee)
CONTACT DETAILS
Global Head – Computational Sciences, Discovery Science, Janssen Pharma R&D
410 George Street,
New Brunswick, NJ 08901
Office: 908-927-2773
grajagop@its.jnj.com
The Simons Center for Systems Biology
Institute for Advanced Study
Einstein Drive, Princeton, NJ 08540
Office: (609) 734 8325
grajagopal@ias.edu
EMPLOYMENT HISTORY
VP & Global Head of Computational Sciences, Discovery Science, 10/2013, Janssen Pharma R&D
(a J&J company).
VP & CIO – Research, Informatics and External Innovation, 6/2012 – 9/2013, Janssen Pharma
R&D (a J&J company)
Executive Director (Bioinformatics), Chief Information Officer, 9/2007 – 6/2012 – Cancer
Institute of New Jersey, Robert Wood Johnson Medical School, USA
Adjunct Professor, Radiation Oncology, 9/2009 – present, Robert Wood Johnson Medical School

2. Executive Director, 6/2006 – 8/2007 – Bioinformatics Institute, Singapore
Deputy Executive Director, 7/2001 - 5/2006 – Bioinformatics Institute, Singapore
Assistant Director of Research, 7/1996 – 6/2001, Cavendish Laboratory, University of Cambridge
Fellow & Director of Studies (Physics, Mathematics), 9/1997 – 6/2001, Jesus College, Cambridge
Postdoctoral Research Associate, Theory of Condensed Matter Group, 4/1994 – 6/1996,
Cavendish Laboratory, University of Cambridge
Visiting Research Fellow, 9/1993 – 3/1994, School of Physics, Georgia Tech, Atlanta, Georgia,
USA
Postdoctoral Research Associate, 9/1991 – 8/1993, Cavendish Laboratory, University of
Cambridge
Director's Research Assistant, 9/1987 – 8/1991, School of Physics, Georgia Tech USA
Lecturer and Head of Physics, 6/ 1986 – 8/1987, Prime College, Kuala Lumpur, Malaysia
Lecturer in Physics, 1/1984 – 5/1986, Damansara College, Kuala Lumpur, Malaysia
Physics Instructor, 6/1981 – 12/1983, University of Malaya, Kuala Lumpur, Malaysia
EDUCATION
Ph.D. Physics, Georgia Institute of Technology, December 1991
M.S. Physics, Georgia Institute of Technology, December 1989
M.Sc. in High Energy Physics, University of Malaya, August 1984
B.Sc.Ed. Physics/Mathematics/Education, University of Malaya, August 1981
EXPERTISE
 Total Informatics Strategy – helping bring better therapies to market for the benefit of
patients –At Janssen, I have developed and am implementing a integrative strategy
encompassing capabilities in IT/HPC, Informatics, Bioinformatics, Computational &
Systems Biology and Genomics/Genetics to support and advance the R&D agenda
throughout the R&D pipeline. To this end we have an overarching role across ALL of our
therapeutic areas (Neuroscience, Oncology, Immunology, Cardiovascular Disease and
Metabolism, Infectious Disease and Vaccines as well as supporting the small and large
molecule centers of excellence. This involves collecting, storing, managing, annotating

3. and integrating data generated within and outside our company during our discovery,
biomarker, pre/post PoC, drug safety, systems/clinical pharmacology and clinical trials.
Our team is leading an effort to build a patient data warehouse to facilitate effective,
efficient and secure use of our clinical data coupled with data from research to guide
patient stratification, biomarker discovery and personalized therapies. The massive
data sets are analyzed using a global network of HPC and cloud computing resources to
help our researchers make meaningful use of the derived information and informed
decisions on optimal strategies to maximize productivity.
 Integrative Genomics – advancing Personalized and Precision Medicine – In
partnership with basic and clinical researchers at CINJ/the Robert Wood Johnson
Medical School, Princeton University, Rutgers University, the University of Medicine and
Dentistry of New Jersey and the Institute for Advanced Study, I helped develop the
computational and data analytical infrastructure to support genomic analysis of
microarray and next generation sequencing data. The goal was to identify and
characterize molecular factors whose deregulation give rise to the disease phenotype.
As part of an international collaboration, I am involved in using WGS of multi-
generational family exhibiting Li-Fraumani Syndrome to identify the relevant genetic
structural variants that give rise to ‘genetic anticipation” in these families. My team has
also executed WES, RNA-Seq and CHIP-Seq studies mainly in the areas of
Skin/Breast/Prostate/Pancreatic Cancer, Leukemia stem cells, G-quad inhibition on cell
lines, autism and auto-immune diseases such as IBD/Chron’s disease.
 Translational Informatics – Linking Bench & Bedside - I lead a team working at the
interface of quantitative biology, disease biology, medicine and population sciences to
integrate & analyze data generated from diverse sources to address fundamental
problems in discovering and developing therapies and bring them to market for the
benefit of patients.. Before joining industry, in the Biopolis and New Jersey, I have been
a PI/co-PI in a number of research grants linking experimental and quantitative
approaches to address basis and translational research problems focused on Cancer
Biology, Cancer Biomarker Discovery and Stem Cell Biology at the Biopolis/Singapore,
the Institute for advanced Study in Princeton and the Cancer Institute of New Jersey.
 Computational/Systems Biology - Large-scale Systems-Level Computational Modeling
Of signaling pathways - I have applied the theoretical/computational tools and
techniques to study signaling pathway in cancer, stem cell fate decisions, and linking
aging and cancer. A quantitative understanding the disruption of cellular processes is
critical if one wants to obtain a deeper conceptual understanding of the mechanisms
that can give rise to disease as it allows an efficient way to guide the search of better
drug targets and associated mechanism of action This work is done in partnership with
colleagues within our therapeutic areas, and the Simons Center for Systems Biology at
the Institute for Advanced Studies in Princeton.
 Cyber-infrastructure Development, Deployment, Management & Application to Real-
world Problems - I have been responsible for planning, building and managing such a
complex cyber-infrastructure at the University of Cambridge and the Biopolis in
Singapore. In Cambridge, I helped set up and run the Cambridge High Performance
Computing Facility – a multi-department funded shared facility with IBM, Hitachi and

4. SGI. In Singapore, I chaired the IT cyber-infrastructure steering committee that worked
with various government agencies and multi-national companies to develop, deploy and
manage the cyber-infrastructure and services that support 3000 researchers located in 5
public and 8 private/pharma research laboratories collocated in the Biopolis. At CINJ, I
built and supported the cyber-infrastructure in partnership with the University of
Medicine and Dentistry of New Jersey to support basic, clinical and translational
research to facilitate and support collaborations within CINJ and with external partners
in academia, hospitals and the pharmaceutical industry in New Jersey.
RESEARCH ACTIVITY
At Janssen Pharma R&D
Drug Discovery - I lead a multi-disciplinary global team of physicists, mathematicians, HPC
experts, computational biologists, statisticians, bioinformatics specialists and wet lab scientists.
Our mission is to work with researchers within and outside our company to generate, evaluate
and translate biomedical data into drug targets that lead to transformational, safe and effective
medicines. To this end, my team is heavily engaged in deploying bioinformatics, machine
learning and cutting-edge computational modeling and simulations techniques. Our in silico
activities are complemented and experimentally validated by wet lab capabilities within the
team utilizing stem cell, molecular genetics, gene editing, Imaging, Next Generation Sequencing
and Omics technologies.
Big Data Analytics – My team is responsible for supporting the computational analytics needs of
partners within our five therapeutic areas within Janssen and with external pre-competitive
national/international initiatives involving Janssen scientists. This includes deploying our
bioinformatics, computational and systems biology and machine learning expertise in support of
basic, clinical and translational programs aligned with and addressing the evolving needs of
Janssen scientists
High Performance Computing – we have developed and deployed HPC capabilities in support of
our drug discovery and development mission by using internal expertise and building strong
partnerships with INTEL and major supercomputing facilities in the U.S.A., the EU and Asia.
Before joining Janssen
My research was focused on three areas: (a) the understanding of how individual genetic
profiles influence response to pathogens and therapies (b) identifying better targets for multi-
factorial diseases by using systems and network pharmacology and (s) using NGS to study the
genetics of complex diseases such as cancer, auto-immune and neurodegenerative diseases to
guide the search for more effective biomarkers and therapies. To this end, my efforts were
focused on developing integrative, quantitative techniques that help identify and characterize
the genetic variations that distinguish the normal and diseased state.

5. Projects at the Institute for Advanced Study (Princeton) & The Cancer Institute of New
Jersey - We have initiated a number of projects involving collaborations with academia,
the biomedical community and the pharmaceutical industry:
 transMED - aims to develop and deploy the latest web technologies, cyber-
infrastructure and informatics capabilities to deploy an interoperable EMR to link
physicians, patients within CINJ, the RWJ Medical School and the RWJ Hospital in New
Brunswick with the ultimate goal of linking to CINJ’s Network hospitals throughout New
Jersey. The goal is to collect, share and analyze population data (including, but not
limited to, genomic and clinical data) to aid our ongoing efforts in cancer prevention,
control and treatment. In addition, the data collected could be used (after appropriate
ethical and IRB review) as a basis for collaborations and/or partnerships with
pharmaceutical and biotech companies located within the state of New Jersey.
 Computational Analysis of Genetic Variation: (With Prof. Arnold Levine, IAS; Dr. Chang
Chan CINJ; Dr. Elke Market, CINJ; Dr. Gurinder Agrwal, CSHL) –
Statistical inference of haplotypes from genotypes is a highly non-trivial task, which has
spawned intensive recent research. The group has be on deploying an entropy-based
method based on mutual information theory on massively parallel computers for
efficient selection of structural variations and functionally characterize their role in
disease initiation and progression. We are also working on developing new tools and
techniques to (a) the analysis of differential gene expression arising as a result of genetic
variation and (b) how polymorphisms influence signal transduction.
 Network Pharmacology (Prof. Arnold Levine, Dr. Alexi Vazquez, CINJ, IAS, Dr. Eric
Perakslis (J&J) – Using tools and techniques from the statistical physics of networks, we
will probe data sets collected in the course of the drug discovery process to identify
combination of drugs that can target a number of nodes in signaling and regulatory
pathways that collectively are more sensitive and specific to control that actions of
these pathways.
 Next Generation Sequencing for Transcriptomics and Human Genetic Variation Study
(with Profs. Arnold Levine, Lorna Rodriguez & The CINJ Precision Medicine Initiative) –
we are leading the bioinformatics effort to design appropriate experiments and analyze
data generated from next generation sequencing technologies in projects related to
differential gene expression, whole genome/exome sequencing to map
driver/passenger mutations, copy number variation etc. The goal is to identify key
molecular players in cancer initiation/progression leading to optimal selection of
treatment protocols and facilitate practice of personalized medicine. We use the entire
range of NGS experiments to analyze data collected from normal/disease and germline
tissue to elucidate disease causing driver mutations and susceptibility genes.
Projects Completed at the Bioinformatics Institute, Singapore
 Developing Mathematical & Computational techniques for Stochastic Chemical Kinetics:
(With Postdoctoral associate Dr. The Yu Kai and research associates Mr. Li Bin). A crucial

6. component of our project is the ability to quantitatively replicate the chemical kinetics
occurring in mesoscopic systems such as that within the cell. We are developing
mathematical and computational techniques to incorporate stochasicity into our
pathway models, to quantify the effects of noise on system-level properties. The
modeling framework resulting from this effort is being applied to the projects below.
 Program Driver for Cancer Biology Initiative - Control and regulation of p53: (a)
Imaging and Modeling of the p53-Hdm2 Feedback Loop: Role of Activator-Inhibitor
Motifs in regulation of cellular processes; (b) A Systems Biology Approach to
Elucidating the Role of the P38 MAPK Pathway and its Interactions with the p53
Mediated Cell Cycle Control Checkpoints in Tumor Suppression. (With Ph.D. student Mr.
Mark Phong of Eli Lilly Systems Biology Singapore, Dr. Christopher Taylor of Lilly System
Biology Singapore, Dr. Xiang Yi at Eli Lilly USA and Prof. Sir David Lane’s Group at IMCB,
Singapore.). (a) Recently, Lahav et. al. (Nature Genetics, 36, 147, 2004) reported results
of damped oscillations of p53/mdm2 proteins in individual living cells when subject to
stress. They found that p53 was expressed in a series of discrete pulses after DNA
damage. Their main conclusion is that the p53-MDM2 feedback loop generates a
“digital” clock that releases well-timed quanta of p53 proteins until damage is repaired
or the cell dies. Our aim, through a combination of experiments and quantitative
modeling, is to verify these observations and to elucidate the mechanism that gives rise
to the “digital pulses”. (b) To study the role of p38 MAPK as a tumor suppressor (a
collaboration with Eli Lilly) we plan to: (1) To collect and annotate data related to
developing a comprehensive map of the p38 MAPK pathway and its role in p53
mediated cell cycle control for the purposes of building a computational model that
accurately describes the signal transduction process; (2) To formulate an appropriate
mathematical and computational framework describing the simplified network topology
of the p38 MAPK pathway and the p53 cell cycle response to genotoxic shock and DNA
damage; (3) To validate the accuracy and veracity of the quantitative model through
bench biology experiments on model cell lines enhancing the model iteratively; (4)
Expansion from single cell model to cell population model. The experimental work will
involve lab work in Lilly facilities in Singapore and Indianapolis. Furthermore, together
with experimental collaborators within and outside Singapore (specifically Prof. Sir
David Lane’s and Prof. Arnold Levine’s Groups) we have built and maintain a
curated p53 KnowledgeBase (p53.bii.a-star.edu.sg) that is available to the whole
community
 Program Driver for Clinical Biomarker Initiative – Discovering and validating
biomarkers for gastric cancer: (With Prof. Sir David Lane (IMCB), Prof. Yeoh KH
(Medicine/NUS), Prof. S. Itoh (Oncology Research Institute), Prof. Edison Liu (GIS), Dr.
Jeffery Hill(BII), Dr. Henry Yang(BII)) The primary focus of the Clinical Biomarkers
Discovery Group is to collaborate with the international community in the International
Cancer Biomarker Consortium (ICBC). This international consortium, led by the Prof. Lee
Hartwell (Fred Hutchinson Cancer Research Institute in Seattle, USA) involves teams
from Singapore, China, USA, France, Mexico, Korea and Taiwan etc. The goal is to
discover biomarkers, which will be validated by the clinical collaborators that are
invaluable for their potential discriminatory power in molecular classification of disease
(diagnosis), in predicting clinical outcome (prognosis) and response to drugs (therapy).
The Singapore national effort, with an initial focus on gastric cancer, is led by Prof. Sir

7. David Lane of the Institute of Molecular and Cell Biology, and will involve scientists from
the Genome Institute of Singapore, Singapore Oncology Research Institute, and the
Proteomics Facility at the Department of Biological Sciences (DBS) in the National
University of Singapore and the Clinical School at the National University Hospital. I am
responsible for the planning, deployment and management of the cyber-infrastructure
and informatics/analysis platform that integrate data generated from the different
institutions involved in an efficient, cost-effective and secure manner. To aid the
coordination of this national/international effort, we are building a Gastric Cancer
Knowledgebase that will contain actively curated biomedical information on all aspects
of gastric cancer.
 Program Driver for Stem Cell Biology Initiative - Identifying Intestinal Adult Stem Cells
and Imaging Epithelial Migration: (Flagship project within the Singapore-MIT Alliance
(SMA) in collaboration with Prof. Paul Matsudaira,Whitehead Institute (WI), Biology and
BioEng, MIT, Dr. Walter Hunziker IMCB, Dr. Mahatavan GIS, Prof. Bing Lim Harvard
Medical School, Prof. Birgit Lane at the Center for Molecular Medicine and SMA-MIT
Scholars Wang Zhengyuan and Lee Seok Yoon) - To work within the SMA Computational
and Systems Biology Initiative to pursue the following aims: (1) to use a combination of
expression/proteomics and bioinformatics to identify specific stem cell markers which
will facilitate their in vivo study via high-resolution real-time imaging of live tissues; (2) to
identify the factors that regulate aspects of the behavior of stem cells such as the process
of asymmetric division and differentiation etc. ultimately leading to a deeper
understanding of the nature of “stemness”; (3) to develop quantitative models to explore
how a gradient of Wnt morphogens along the crypt-villus axis in the small intestine can
regulate proliferation and differentiation status as epithelial cells migrate from the crypt
to the tip of the villus; (4) to elucidate the mechanism of epithelial cell migration in the
epithelium of the small intestine of the Zebra fish using real-time high-resolution in vivo
imaging of live tissue. To this end we will work closely with our experimental
collaborators to obtain and analyze the data they collect to build quantitative models.
This work is funded by a three year S$450K Singapore-MIT Alliance Grant jointly awarded
with Prof. Matsudaira. The data obtained, together with all publicly available data on
adult stem cells, will be collected and curated in a Stem Cell KnowledgeBase
(stemcell.bii.a-star.edu.sg) that will be available to the whole community as part of
theSingapore Stem Cell Consortium.
 Modeling Mechanotransduction - A Systems Biology Approach to Elucidate the Role of
Adhaerens Junctions and Tight Junctions as Mechanotransducers in Regulating Cell
Proliferation. (With AGS Scholar Lim Tong Seng, Prof. Lim Chwee Teck (BioEng, NUS), Prof.
Paul Matsudaira (WI, Biology and BioEng, MIT) and Prof. B. Lane, Center for Molecular
Medicine, Singapore) – Mechanotransduction or the translation of mechanical forces
into biochemical signals; plays an important role in the regulation of cellular activities.
How cells sense external forces and integrate these forces at the molecular level to
produce coordinated responses is an open question. In particular, the mechanism by
which mechanical stimuli regulate cell proliferation is a subject of great controversy and
a resolution to this issue has wider implications both for the fundamental understanding
of cell biology and within a clinical context. To this end, we are investigating specific
classes of proteins found in tight junctions and adhaerens junctions, which are
specialized cell-cell contacts that play an important role in regulating cellular activities.

8. This project aims to test the hypothesis that specific members of the tight and adhaerens
junction proteins act as mechanotransducers for regulating cell proliferation when
quiescent epithelial cells in culture are subjected to mechanical stimuli. This work is
funded by a three year S$600K Singapore Biomedical Research Council Grant.
 Modeling Differential Gene Expression: A Systems Biology Approach to Elucidate the
Frequency Decoding of GnRH Signals that Governs the Differential Expression of
Mammalian Gonadotropin Subunit Genes. (With AGS Scholar Stefan Lim and Dr. Philippa
Melmlud (DBS, NUS)): Luteinizing and follicle-stimulating hormones (LH and FSH) are
gonadotropic hormones synthesized in the pituitary gonadotropes. They share a common
a-subunit but contain a unique b-subunit that confers physiological specificity. LH and
FSH directly stimulate gonadal growth and steroidogenesis. Therefore regulation of the
precise levels of each hormone is critical to correct reproductive functioning. The
hypothalamic GnRH regulates both hormones, and variations in its pulse frequency are
associated with the differential effects on each of the three subunits. The mechanism by
which gonadotropes decode varying gonadotropin-releasing hormone (GnRH) pulse
frequencies and translate them into differential gene expression is unknown. It is the
goal of this project to resolve this issue through an integrative approach involving a
fusion of biological, genetic, computational and mathematical techniques. Understanding
the molecular mechanisms that control gonadotropin biosynthesis holds great potential
for the clinical treatment of a variety of reproductive disorders. The knowledge gained in
the resolution of this problem will also contribute to a deeper conceptual understanding
of the mechanisms governing differential expression of genes in general, which is the
cornerstone of developmental biology. This work is funded by a three year S$800K
Singapore Biomedical Research Council Grant.
 Modeling Cross-talk between RhoGTPase and MAP Kinase Signaling Pathways: (With
Research Associate Ang Boon Keong and Dr. Ong Siew Hwa (IMCB)) – The objective of
this experimental/theoretical collaboration is to elucidate the function of Rho proteins
and their regulators in various physiological processes given the importance of Rho
GTPase signaling networks in development. We have initially focused on experimentally
characterizing a specific Rho GAP protein, ArhGAP9, and its interactions with MAP
kinases such as Erk2 and p38. We aim to show how actin remodeling can be regulated
through the interaction of ArhGAP9 and the MAP kinases, cross-talking between the Rho
GTPase and MAP kinase signaling pathways. The long term goal is to study
experimentally and via bioinformatics and structural modeling how the formation of
multi-protein complexes through the different domains present in GTPase regulators
such as ArhGAP9 allows for the spatial and temporal coordination of different signaling
pathways.
TEACHING EXPERIENCE
At the Robert Wood Johnson Medical School – supervised MD/PhD and PhD candidates,
MD/PhD graduate lectures in bioinformatics and Genomics, short course on the use of
bioinformatics and computing in biomedical research (to researchers and clinicians). With my
team of computational biologists and bioinformatics specialists and biostatisticians conducted
weekly “clinics” for clinicians and researchers on effective use of genomic and statistical
techniques in experimental design of Omics studies in support of their research programs.

9. At the Bioinformatics Institute (BII), Singapore: supervised five PhD Scholars in Systems Biology
and Bioinformatics as part of the Singapore-MIT Alliance; conducted postgraduate
lectures/training for Tsinghua/Beijing University; at the national University f Singapore, gave
lecture graduate courses in Computational Biology, Molecular Modeling, Mathematical/Systems
Biology, Systems Biology of Cancer/Stem Cell; supervise MSc/PhD research projects in
bioinformatics and computational/systems biology; lead study sessions on mathematical
methods applied to biological problems; designed and implemented a MSc graduate program in
Bioinformatics/Computational Biology. I was the external examiner for Physics PhD at the
University of Cambridge.
At the University of Cambridge, UK: supervised a M. Phil. student, co-supervised seven Ph.D.
students at the Cavendish Laboratory and three PhD students at Imperial College, London;
lectured a Part II undergraduate physics course (Theoretical Physics I/II), and Part III course,
``Supercomputing in Physics"; lectures on high performance computing, biological physics and
electronic structure theory; supervised Literature Reviews and Part III projects, Part II Quantum
Mechanics, Solid State Physics, Thermal Physics, Systems, Part III Solid State Physics, Part IB
Advanced Physics and Mathematics for IA and IB (Natural Science).
At the Georgia Institute of Technology, USA: Lectured freshman and sophomore Engineering
Physics courses.
At the University of Malaya, Prime College and Damansara College, Malaysia: Physics
Instructor/lecturer.
At the Robert Wood Johnson Medical School, University of Medicine and Dentistry of New
Jersey, USA, Rutgers University: Graduate lectures in Cancer Systems Biology, Bioinformatics and
Computational Biology.
MANAGEMENT RESPONSIBILITIES
In the United States
At Janssen Pharma R&D (a J&J company): I am responsible for overseeing a team divided into
five groups: Translational Informatics, Computational & Systems Biology, High Performance
Computing, External Innovation and R&D IT. We support the evolving cyber-
infrastructure/eData capture, Informatics, Big Data analytics, needs of researchers and clinicians
throughout Janssen. I am a member of the therapeutic, functional area, Pharma R&D and R&D
Operations senior leadership team. I am also a member of CINJ’s External Advisory Board for
their Precision Medicine Initiative.
At the Cancer Institute of New Jersey (CINJ): I am responsible for all matters concerning
computing – both for research, clinic and for corporate needs. I am setting up the Cancer
Informatics Core facility from scratch. As Executive Director (Bioinformatics), I am responsible
for building up Clinical and Bioinformatics capabilities to supporting ongoing basic, clinical,
population sciences, translational research as well as postgraduate education for the CINJ
community and its partners. I am responsible for the development, deployment and smooth
operation of cyber-infrastructure and services to meet the evolving needs of the CINJ user

10. community. I was the Chair of the Electronic Medical Record Implementation Committee and
Co-Chair of the Steering Committee: responsible for deploying and integrating a Electronic
Medical Record system across the Cancer Institute of New Jersey, the Robert Wood Johnson
Hospital and the Robert Wood Johnson Medical School. I am also member of the Clinical
Enterprises Committee that oversees all clinical operations at the CINJ: responsible for providing
advise on Informatics and IT/security issues relating to clinical operations.
I am a member of various internal advisory committees for overseeing CINJ Core facilities:
Biostatistics Core, Microarray Core, Sequencing Core and Tissue Analytic Services.
Member of the Executive Council for the Cancer Institute of New Jersey: this is the key
leadership committee that is chaired by the Director which meets weekly to manage the affairs
of the Institute.
Member of the Scientific Council for the CINJ/Rutgers University/Princeton University Cancer
Consortium: responsible for providing advise on scientific computing, biomedical informatics for
inter-institutional collaborations.
Member of the DIMACS Executive Council, (2008 - 2012 ): Efficient, effective and secure access
to and analysis of health data is THE central problem in health care informatics with serious
implications for the economics of health care delivery, basic, clinical and translational research
and in the prevention of disease. We are working with DIMACS to provide to tools and
expertise to address one of the key challenges in the medical arena and am co-organizer of the
“DIMACS/CINJ Working Group on Electronic Medical Records”
(http://dimacs.rutgers.edu/Workshops/WGMedicalRecords/) and “Algorithmic Medical
Decision Making: Leveraging Heterogeneous Data Sources for Quantifying Risk” Workshop
()http://dimacs.rutgers.edu/Workshops/QuantifyingRisk/).
At the Institute for Advanced Study: Conduct research in the areas of Computational/Systems
Biology focused on the genetic variations in the human genome that give rise to complex
diseases (such as cancer) and provide advice on all matters concerning computing needs of the
Institute.
At the American Association for Cancer Research (AACR): chair the Bioinformatics
Subcommittee for the AACR – NCI - FDA Cancer Biomarkers Collaborative. The subcommittee
will explore current thinking from academia (i.e. the Cancer Bioinformatics Grid, caBIG) and
industry on the role of informatics in cancer biomarker discovery. Its mandate is to map out and
where possible, make recommendations on “best practices” in the use of bioinformatics tools
and cyber-infrastructure resources and services in facilitating the discovery, validation,
qualification and clinical deployment of diagnostic biomarkers for cancer,
In Singapore:
At the Bioinformatics Institute: set up the Institute from scratch in July 2001. Mission of the BII
encompasses research in bioinformatics and computational biology, post-graduate education, IT
and application support for the biomedical user community in Singapore and cyber-
infrastructure development at the national level. The BII is the central national resource for

11. biomedical computing for the biomedical research community. My responsibilities include
setting up and overseeing the management, research, biomedical computing support and post-
graduate education activities of the Institute. Program driver for the Cancer Biology, Clinical
Biomarkers and Stem Cell Initiatives that links experimental and quantitative modeling involving
researchers within and beyond the Biopolis to address key biomedical questions in cancer and
stem cell biology and to coordinate the building of a Web based knowledgebase to advance R&D
in these areas. I was the Executive Director, responsible for providing the scientific and
administrative leadership for the Institute.
Cross Institution responsibilities: Chairman of the Joint Management Committee overseeing the
BII-NUS Postgraduate Program in Bioinformatics. Planned and implemented a graduate program
in Bioinformatics involving 2 years of course work and academic and industrial attachment.
Research Council Responsibilities: Chairman of the IT Steering Committee for the Biomedical
Research Council and Co-Chair of the One-north IT steering Committee responsible for planning,
implementing and managing the outsourced cyber-infrastructure and services for the new
science park, the Biopolis (Project SELECT). The Biopolis (www.one-north.com) is a physical co-
location of public and private research institutes and biotech companies that will leverage on
each other to advance biomedical research and development and exploit the results of the
research for wealth creation. I direct the Project Team which is responsible for formulating,
managing the tender process and the implementation of infrastructure and services. Member
of the BMRC/NMRC Grant Evaluation Committee (2006-2009).
National Cyber-infrastructure development responsibilities: member of the Steering Committee
for the Singapore National Grid Initiative whose goal is to aggregate national computing
resources to effectively and efficiently address the needs of academia, industry and the medical
community. To build up the expertise to accomplish this challenging task, I initiated the
Singapore BioMed Grid (BMG) Project (in collaboration with similar Grid projects in the US and
Europe). The main focus is to develop and implement grid-based middleware and application
software of interest to the biomedical community to advance research and development in the
public and private sector. We work very closely with companies such as Intel Sun and HP as well
as the US national supercomputing centers (NCSA, PNNL and SDSC) to share expertise in
developing grid computing for the biomedical user community. We are founding members of
international collaborations in grid computing such as the PRAGMA Consortium and the GELATO
Federation.
At the University of Cambridge, United Kingdom:
Cross-Department Responsibilities: Coordinator and member of management committee of the
Cambridge High Performance Computing Facility (1994-2001); management of the day to day
operation of the University of Cambridge High-Performance Computing Facility; providing
technical advice to the user community within the University on all aspects of high performance
computing; enhancement of supercomputing resources by preparing grant applications to
funding bodies.
Cavendish Laboratory, University of Cambridge: Lecturer in Part II/Part III courses in Theoretical
Physics and Computational Physics; Part III Physics Examiner for the 1998/1999 and 1999/2000
academic years.

12. EPSRC Research Council Responsibilities: Member of the EPSRC "Computational Partial
Differential Equations" Panel (1998-2001). I refereed grant applications for EPSRC and
HEFCE/JREI, the Welcome Trust and for joint MRC/DFG projects involve advanced computing in
Biomedical research.
Jesus College: Elected member of Jesus College Governing Council (1998). Elected to the
Mastership Committee (April 2000).Elected to the College Development Committee (July 2000).
Director of Studies for Physics and Mathematics for Natural Scientists in Jesus College (1997-
2001).
CONSULTANTCIES/SCIENTIFIC BOARDS
Grant reviewer for NIH, NSF, AACR, EU (ESF), UK (MRC, EPSRC, CRUK), Germany (DFG),
Singapore (ASTAR) and Japan (MOE, JSI). Served as a member of external review boards for
governments of Singapore, Ireland, UK and Malaysia.
Informatics Consultant for the Johnson & Johnson Centacor (Pharma arm of J&J):
Informatics Consultant for the Scientific Board for Novartis AG: Reviewed ongoing informatics
activities throughout Novartis’s global R&D organizations, 2009 – 2012
Informatics Consultant for Daiichi Sankyo: Reviewed ongoing informatics activities throughout
their global R&D organizations, 2010 – 2012
Program Grant Reviewer for Science Foundation Ireland, 2009 – present
Advisor, University of Malaya Medical Faculty, 2010 – present
Advisor, Cancer Research Foundation , Malaysia, 2008 - present
External Review Board, Personalized Medicine, Rutgers Cancer Institute of New Jersey, 2012 –
present.
Scientific Advisory Board, IMEC, Belgium 2014 - present
Member of the European Commission Panel of Experts, Digital Agenda for Europe, 2014 -
present
COMPUTATIONAL EXPERTISE
I have published research that used large scale computing to address challenge problems in
computational chemistry, biology and physics. I am now focused on transferring the techniques
and algorithms developed to understand problems in computational/systems biology and
genetics (especially the use of NGS to advance drug discovery and development). As part of our
Biomedical Grid initiative in Singapore I have been directing the year long training of grid
engineers (53 trained with government funding), building a operational test-bed for biomedical

13. applications and developing new code to run over the grid cyber-infrastructure that links
geographically distributed computing and data resources.
HONORS & AWARDS
Distinguished Visiting Professor, Ministry of Science and Technology, Malaysia (2010);
Informatics Consultant (Advisor) for Norvatis (2009 -201 ); Johnson & Johnson (2008 - ); Daiichi
Sankyo (2010 -2012 ); Special Merit Pay Allowance, University of Cambridge, (1991-1996, 1999-
2004); Elected Fellow of Jesus College (1997); Awarded National Science Foundation (USA)
scholarship to attend Supercomputing Summer School at the Pittsburgh Supercomputing Centre
(June 1990); Director's Research Assistantship, School of Physics, Georgia Tech, (1987-1991);
Appointed Junior Associate of the ICTP Trieste, Italy, (1986-1989).
GRANTS AWARDED
1. US$11.7M support from the Robert Wood Johnson University Hospital to design,
deploy and operate/manage an Oncology based EMR, 2009-2011.
2. Jointly awarded US$1.500M for Bioinformatics support for NIH Challenge grants,
2009 as co-PI in grants awarded to scientists at CINJ and RWJMS.
3. US$3M grant from State of New Jersey for building Computational, Systems Biology
and Bioinformatics capability at the Cancer Institute of New Jersey, 2007-2010
4. US$12M grant from the Robert Wood Johnson Foundation, as co-investigator in
population sciences study of cancer prevention, control and survivorship.
5. S$1.800M BMRC grant with Dr. Lim Chwee Teck (BioEng/NUS) to study biochemical
and mechanical perturbation regulating cellular mechanotransduction in 2006-2009.
6. S$1.360M Grant with Dr. Henry Yang for the Stem Cell Knowledgebase, 2006-2009
7. S$1.80M BMRC grant with Dr. Phillipa Melamad for study of frequency-dependent
differential gene expression (DBS/NUS) 2005-2008.
8. S$3.5M to plan and build a test bed for the Singapore Biomedical Grid (BMG) 2001-
2004.
9. S$3.4M to build up of expertise in Grid Computing by the InfoCom Development
Authority of Singapore 2002-2003.
10. £1.250M Hitachi Japan Ltd. to collaborate with Hitachi's Central Research Lab in
Tokyo in the project "Atomic Level Simulations of Submicron Devices" 1996-2001.
11. Three successful awards from the Joint Research Equipment Initiative (JREI) for
enhancing supercomputing hardware and technical support in the Cambridge High

14. Performance Computing Facility. Total of £15.5M of funding awarded
1996/1997/1998.
12. ROPA grant (£179,729) by the United Kingdom Engineering and Physical Sciences
Research Council (jointly with Dr. R J Needs and Dr. M C Payne) 1996-1999.
13. NATO Collaborative Research grant of 215,000 BF (Grant number CRG 951105jointly
with Dr. RJ Needs (Cavendish) and Prof. M-Y Chou (Georgia Tech) 1996-1998
AFFILIATIONS
American Association for Cancer Research (AACR), the American Society of Clinical Oncology
(ASCO), The American Society of Human Genetics (ASHG), The New York Academy of Science
(NYAS), American Association for the Advancement of Science (AAAS), American Society for
Clinical Pharmacology and Therapeutics (ASCPT) and the American Medical Informatics
Association (AMIA).
REFEREES
Prof. Arnold J Levine, Director, Simons Center for Systems Biology, Institute for Advanced Study,
Princeton. alevine@ias.edu
Dr. William Hait, VP & Global Head of R&D, Johnson & Johnson, WHait@ITS.JNJ.COM
Prof. Robert DiPaola, Director, The Cancer Institute of New Jersey, Robert Wood Johnson
Medical School, DiPaolRS@umdnj.edu
Prof. Sidney Brenner, Chairman, Biomedical Research Council, sidney_brenner@a-star.edu.sg
Prof. Sir David Lane, Executive Director, IMCB, d.p.lane@imcb.a-star.edu.sg
Prof. Robert Mair, The Master, Jesus College, Cambridge, UK, rjm50@eng.cam.ac.uk
Prof. Michael Levitt, Stanford University, Stanford, CA, michael.levitt@stanford.edu
Prof. Michael Waterman, University of Southern California, Los Angeles, msw@usc.edu
PUBLICATIONS
REVIEWS
S. Vedula, C.T. Lim, T.S. Lim, G. Rajagopal, W. Hunziker, B. Lane and M. Sokabe, "Role of external
mechanical forces in cell signal transduction", Biomechanics At Micro And Nanoscale Levels. H.
Wada (ed), (World Scientific, Singapore, 2007).

15. W. M. C. Foulkes, L. Mitas, R.J. Needs and G. Rajagopal, "Quantum Monte Carlo Simulations of
Solids", Reviews of Modern Physics. 73 , 33-81, (2001).
M. Nekovee, W.M.C. Foulkes, A.J. Williamson, G Rajagopal, and R.J. Needs, "A quantum Monte
Carlo approach to the adiabatic connection method", 33, 189, Advances in Quantum Chemistry,
Thematic volume on DFT, edited by J.M. Seminario, (1999).
W. M. C. Foulkes, R.J. Needs and G. Rajagopal, "Quantum Monte Carlo Simulations of Real
Solids", High Performance Computing. R.J. Allen et. el. (eds), (Kluwer Academic Publishers,
Dordrecht, 1999).
R.J. Needs, G. Rajagopal, A.J. Williamson, L.M. Fraser, S.D. Kenny, W.M.C. Foulkes, A.J. James,
and P. Maccallum ``Quantum Monte Carlo studies of electronic systems '', J. Korean Physical
Society 29, S116-S120 (1996).
U. Landman, R. Barnett, G. Rajagopal and A. Nitzan, ``Energetics and Dynamics of Solvation and
Fission in Clusters", Proceedings of the European Physical Society, Topical Conference on
Nuclear and Atomic Clusters, Finland, (1992).
R. Barnett, U. Landman, G. Rajagopal and C. Cleveland, ``Evolutionary Patterns of Structure and
Stability of Clusters", NATO Advanced Research Workshop on Physics and Chemistry of Finite
Systems : From Clusters to Crystals, Vol. 374, 165, edited by P. Jena, S. Khanna and B. Rao
(Kluwer Academic Publishers, Dordrecht, 1992).
R. Barnett, U. Landman, G. Rajagopal and A. Nitzan, ``Dynamics, Spectra and Relaxation
phenomena of excess electrons in Clusters", Israel Journal of Chemistry, 30, (1990).
Peer-reviewed Journal Articles
S.P. Chia and G. Rajagopal, ``An exact calculation for the flavor-changing quark-photon vertex ",
Physics Letters, 156B, 405 (1985).
G. Rajagopal, R. Barnett, A. Nitzan, U. Landman, E. Honea, P. Labistie, M.L. Homer and R.L.
Whetten, ``Optical Spectra of Localized excess electrons in Alkali Halide Clusters", Physical
Review Letters, 64, 2933 (1990).
R. Barnett, U. Landman, A. Nitzan and G. Rajagopal,``Born-Oppenheimer dynamics using Density-
Functional Theory : Equilibrium and Fragmentation of small Sodium Clusters", Journal of
Chemical Physics, 94, 608 (1991).
G. Rajagopal, R. Barnett and U. Landman, ``Metalization of Ionic Clusters ", Physical Review
Letters, 67, 727, (1991).
R. Barnett, U. Landman, and G. Rajagopal,``Patterns and Barriers for fission of charged small
metal Clusters", Physical Review Letters, 67, 3058, (1991).

16. G. Rajagopal and R.J. Needs, ``An optimized Ewald method for long-ranged potentials", Journal
of Computational Physics, 115, 399 (1994).
G. Rajagopal, R.J. Needs, S. Kenny, W.M.C. Foulkes and A. James, ``Quantum Monte Carlo
calculations for solids using special k-points methods", Physical Review Letters, 73 1959 (1994).
G. Rajagopal, R.J. Needs, W.M.C. Foulkes, W.M.C. Foulkes, S. Kenny and A. James, ``Variational
and diffusion Quantum Monte Carlo calculations at Non-zero Wavevectors : Theory and
application to Diamond-Structure Germanium", Physical Review B 51 , 10591 (1995).
S. Kenny, G. Rajagopal and R.J. Needs, ``Relativistic corrections to atomic energies using
quantum Monte Carlo calculations", Physical Review A 51 , 1898 (1995).
L. Fraser, W.M.C. Foulkes, G. Rajagopal, and R.J. Needs, ``Finite-size effects and Coulomb
interactions in quantum Monte Carlo calculations for homogeneous systems with periodic
boundary conditions", Physical Review B 53 , 1814 (1996).
A.J. Williamson, G.Rajagopal, R.J. Needs, L. Fraser, A. James, W.M.C. Foulkes and P. Maccallum,
``Optimized Wavefunctions for quantum Monte Carlo studies of Atoms and Solids", Physical
Review B 53, 9640 (1996).
S.D. Kenny, G.Rajagopal, R.J. Needs, A.J. Williamson, W-K Leung, L.M. Fraser, W.M.C. Foulkes
and M.J. Godfrey, ``Quantum Monte Carlo Calculations of Relativistic Corrections to the Energy
of the Homogeneous Electron Gas.", Physical Review Letters, 77 1099 (1996).
A.J. Williamson, G.Rajagopal, R.J. Needs, L.M. Fraser, W.M.C. Foulkes Y. Wang and M.-Y. Chou,
``Elimination of Coulomb finite size effects in quantum many-body simulations.", Physical Review
B55 R4867 (1997).
R.J. Needs and G. Rajagopal ``First-principles calculations of the adsorbate-induced surface stress
of K/Al(111) ()R30". Surface Science 372, 179-184 (1997).
R.Q. Hood, M.-Y. Chou, A.J. Williamson, G.Rajagopal, R.J. Needs, W.M.C. Foulkes ``Quantum
Monte Carlo Investigation of Exchange and Correlation in Silicon.", Phy. Rev. Letts. 78, 3350,
1997.
W.M.C Foulkes, M. Nekovee, R.Q. Hood, M.Y. Chou, R.J. Needs, G. Rajagopal, A.J. Williamson,
"Quantum Monte Carlo studies of exchange and correlation in solids", ABSTRACTS OF PAPERS OF
THE AMERICAN CHEMICAL SOCIETY, 213, 125, 1997.
R.Q. Hood, M.-Y. Chou, A.J. Williamson, G. Rajagopal, and R.J. Needs, " Exchange and correlation
in silicon", Physical Review B57 8972 (1998).
A.J. Williamson, R.Q. Hood, R.J. Needs, G. Rajagopal ``Diffusion Quantum Monte Carlo
Calculation of the excited states of Silicon", Physical Review B57 12140 (1998).

17. P.R.C. Kent, R.Q. Hood, M.D. Towler, R.J. Needs and G. Rajagopal, "Quantum Monte Carlo
calculations of the one-body density matrix and excitation energies of silicon", Physical Review
B57 15293 (1998).
P. R. C. Kent, Randolph Q. Hood, A. J. Williamson, R. J. Needs, G. Rajagopal, and W. M. C. Foulkes,
"Finite size errors in quantum many-body simulations of extended systems, Physical Review B59,
1917 (1998).
P.R.C. Kent, R.J. Needs and G. Rajagopal. "Monte Carlo energy and variance minimization
techniques for optimizing many-body wave functions", Physical Review B59, 12344, (1999).
W.K. Leung, R.J. Needs, G. Rajagopal, S. Itoh and S. Ihara "Calculations of Silicon Self-Interstitials
Defects", Physical Review Letters 83 , 2351, (1999).
K. C. Huang, R. J. Needs, and G. Rajagopal, Comment on ``Quantum Monte Carlo study of the
dipole moment of CO'' [J. Chem. Phys. 110, 11700 (1999)], J. Chem. Phys. 112, 4419, (2000).
P.R.C. Kent, M.D. Towler, R.J. Needs and G. Rajagopal, "Carbon clusters near the crossover for
fullerene stability '', Physical Review B62, 15 394 (2000).
Y. Lee, P. Kent, M. Towler, R. Needs and G. Rajagopal, "Pseudopotentials for Correlated Electron
Calculations ", Physical Review B62, 13347, (2000).
R. Gaudoin, M. Nekovee, W. M. C. Foulkes, R. J. Needs and G. Rajagopal, " Inhomogeneous
random-phase approximation and many-electron trial wave functions ", Physical Review B63,
115115, (2001).
W.K. Leung, R.J. Needs, G. Rajagopal, S. Itoh and S. Ihara "Quantum Monte Carlo study of Silicon
Self-Interstitials Defects", VLSI Design, 13(1-4): 229, (2001).
R. Gaudoin, W. M. C. Foulkes, and G. Rajagopal, " Ab Initio Calculations of the Cohesive Energy
and Bulk Modulus of Aluminium”, J. Phys.: Condens. Matter 14, 8787-8793 (2002).
R.J. Needs, P.R.C. Kent, A.R. Porter, M.D. Towler and G. Rajagopal, "Quantum Monte Carlo
Calculations for Ground and Excited States", Int. J. Quant. Chem. 86, 218, (2002).
Zhu H, Sun Y, Rajagopal G, Mondry A and Dhar P, “Facilitating arrhythmia simulation: the
method of quantitative cellular automata ”, BioMedical Engineering OnLine 2004, 3:29
(30 August 2004).
Vedula S.R.K., Tong Seng Lim., Kausalya P.J, Gunaretnam R., Hunziker W., and Lim C.T,
“Quantifying adhesion forces of tight junction proteins”, paper published in the Proceedings of
the 12th Int. Conf. On Biomedical Engineering, 2005.
Vedula S.R.K., TS Lim., Kausalya P.J, Hunziker W., G. Rajagopal and Lim C.T, “Biophysical
approaches for studying the integrity and function of tight junctions”, Molecular and Cellular
Biomechanics, 2, 105-123, 2006.

18. CM Song, BH Yeo, E. Tantoso, Y. Yang, YP Lim, KB Li and G. Rajagopal, “iHAP – Integrated
haplotype analysis pipeline for characterizing the haplotype structures of genes”, BMC
Bioinformatics, 7:525, 2006
KH Chiam, C.M. Tan, V. Bhargava, and G. Rajagopal, “Hybrid simulations of stochastic reaction-
diffusion processes for modeling intracellular signaling pathways”, Physical Review E 74, 51910,
2006.
Lee BT1, Song CM, Yeo BH, Chung CW, Chan YL, Lim TT, Chua YB, Loh MC, Ang BK, Vijayakumar P,
Liew L, Lim J, Lim YP, Wong CH, Chuon D, Rajagopal G, Hill J. “Gastric Cancer(Biomarkers)
Knowledgebase (GCBKB):A curated and Fully Integrated Knowledgebase of Putative Biomarkers
Related to Gastric Cancer”, Biomarker Insights, 2, 1 2006.
YP Lim, TT Lim, CM Song, BH Yeoh, D. Coomber, G, Rajagopal and D. Lane, “The p53
KnowledgeBase – An Integrated Information Resource for p53 Research ”, Nature Oncogene,
September 2006.
KH Chiam and G. Rajagopal, “Oscillations in Signaling Cascades”, Physical Review E 75, 061901,
2007.
BK Ang, CY Lim, SH Koh, S. Taib, KB Lim, S. Ahmad, G, Rajagopal, G. Guy and SH Ong, “ArhGAP9,
a novel MAP Kinase docking protein, inhibits Erk and p38 activation through WW domain
binding ”, 2:1, Journal of Molecular Signaling, 2007.
Sri Ram Krishna Vedula; Tong Seng Lim; Hui Shi; Jaya P. Kausalya; Birgitte Lane; Gunaretnam
Rajagopal; Walter Hunziker “Molecular force spectroscopy of homophilic nectin-1 interactions”,
Biochem Biophys Res Commun. 2007 Nov 3;362(4):886-92.
Vedula, S. R.; Lim, T. S.; Kausalya, P. J.; Hunziker, W.; Rajagopal, G; Lim, C. T. Biophysical
approaches for studying the integrity and function of tight junctions. Mol Cell Biomech (2005)
2:105-123.
S.R.K. Vedula, T.S. Lim, P.J. Kausalya, E.B. Lane, G. Rajagopal, W. Hunziker, C.T. Lim,
“Quantifying Forces Mediated by Integral Tight Junction Proteins in Cell–Cell Adhesion”,
accepted for publication in Experimental Mechanics, 49:3-9, 2009.
Bernett T. K. Lee, LaiLing Liew, JiaHao Lim, Jonathan K. L. Tan, Tze Chuen Lee, Pardha S.
Veladandi, Yun Ping Lim, Hao Han, Gunaretnam Rajagopal, N. Leigh Anderson, “Candidate List of
yoUr Biomarkers(CLUB): a Web-based Platform to Aid Cancer Biomarker Research”, Biomarker
Insights, 4, 2008
Zhaohui Feng, Wenwei Hu, Gunaretnam Rajagopal and Arnold J. Levine, “The Tumor Suppressor
p53: Cancer and Aging”, Cell Cycle, April, 2009.
Chen, T.; Kim, S.; Zhou, J.; Metaxas, D.; Rajagopal, G; Yue, N. 3D meshless prostate segmentation
and registration in image guided radiotherapy. Med Image Comput Comput Assist Interv (2009)
12:43-50.

19. Henry Lim, S.; Pnueli, L.; Tan, J. H.; Naor, Z.; Rajagopal, G; Melamed, P. Negative feedback
governs gonadotrope frequency-decoding of gonadotropin releasing hormone pulse-frequency.
PLoS One (2009) 4:e7244.
Ting Chen, Sung Kim, Sharad Goyal, Salma Jabbour, Jinghao Zhou, Gunaretnam Rajagopal, Bruce
Haffty, Ning Yue, "Object-Constrained Meshless Deformable Algorithm for High Speed 3D Non-
rigid Registration between CT and CBCT" , Med. Phys. 37 197 (2010).
Hany Ariffin, Pierre Hainaut, Anna Puzio-Kuter, Soo Sin Choong, Adelyne Sue Li, Chan, Denis
Tolkunov, Gunaretnam Rajagopal, Wenfeng Kang, Leon Li Wen Lim, Shekhar Krishnan , Kok-
Siong Chen, Maria Isabel Achatz, Mawar Karsa, Jannah Shamsani, Arnold J Levine and Chang S
Chan, “Whole genome sequencing analysis of phenotypic heterogeneity and anticipation in Li-
Fraumeni cancer predisposition syndrome ”, PNAS, 111, 15497, 2014.