Timothy A. Springer
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Timothy "Tim" A. Springer (born February 23, 1948) is an immunologist and the Latham Family
Professor Professor (commonly abbreviated as Prof.) is an Academy, academic rank at university, universities and other tertiary education, post-secondary education and research institutions in most countries. Literally, ''professor'' derives from Latin ...
at
Harvard Harvard University is a private Ivy League research university in Cambridge, Massachusetts, United States. Founded in 1636 and named for its first benefactor, the Puritan clergyman John Harvard, it is the oldest institution of higher lear ...
Medical School A medical school is a tertiary educational institution, professional school, or forms a part of such an institution, that teaches medicine, and awards a professional degree for physicians. Such medical degrees include the Bachelor of Medicine, ...
. He is also a professor at the Department of
Biological Chemistry Biochemistry, or biological chemistry, is the study of chemical processes within and relating to living organisms. A sub-discipline of both chemistry and biology, biochemistry may be divided into three fields: structural biology, enzymology, a ...
and Molecular
Pharmacology Pharmacology is the science of drugs and medications, including a substance's origin, composition, pharmacokinetics, pharmacodynamics, therapeutic use, and toxicology. More specifically, it is the study of the interactions that occur betwee ...
and of the Division of
Medical Sciences Medicine is the science and practice of caring for patients, managing the diagnosis, prognosis, prevention, treatment, palliation of their injury or disease, and promoting their health. Medicine encompasses a variety of health care pract ...
, and a Senior Investigator at the Research Program in Cellular and Molecular Medicine of the
Boston Children's Hospital Boston Children's Hospital (formerly known as Children's Hospital Boston until 2013) is the main pediatric training and research hospital of Harvard Medical School, Harvard University. It is a nationally ranked, freestanding acute care children ...
. Springer is best known for discovering the first
integrin Integrins are transmembrane receptors that help cell–cell and cell–extracellular matrix (ECM) adhesion. Upon ligand binding, integrins activate signal transduction pathways that mediate cellular signals such as regulation of the cell cycle, o ...
s,
LFA-1 Lymphocyte function-associated antigen 1 (LFA-1) is an integrin found on lymphocytes and other leukocytes. LFA-1 plays a key role in emigration, which is the process by which leukocytes leave the bloodstream to enter the tissues. LFA-1 also mediates ...
, and
intercellular adhesion molecule In molecular biology, intercellular adhesion molecules (ICAMs) and vascular cell adhesion molecule-1 (VCAM-1) are part of the immunoglobulin superfamily. They are important in inflammation, immune responses and in intracellular signalling event ...
s (ICAMs), and for elucidating how these
cell adhesion molecule Cell adhesion molecules (CAMs) are a subset of cell surface proteins that are involved in the binding of cells with other cells or with the extracellular matrix (ECM), in a process called cell adhesion. In essence, CAMs help cells stick to each ...
s function in the
immune system The immune system is a network of biological systems that protects an organism from diseases. It detects and responds to a wide variety of pathogens, from viruses to bacteria, as well as Tumor immunology, cancer cells, Parasitic worm, parasitic ...
. In recent years, Springer's research interest has expanded to
malaria Malaria is a Mosquito-borne disease, mosquito-borne infectious disease that affects vertebrates and ''Anopheles'' mosquitoes. Human malaria causes Signs and symptoms, symptoms that typically include fever, Fatigue (medical), fatigue, vomitin ...
,
transforming growth factor beta Transforming growth factor beta (TGF-β) is a multifunctional cytokine belonging to the transforming growth factor superfamily that includes three different mammalian isoforms (TGF-β 1 to 3, HGNC symbols TGFB1, TGFB2, TGFB3) and many other ...
(TGF-β) signaling, and
von Willebrand factor Von Willebrand factor (VWF) () is a blood glycoprotein that promotes primary hemostasis, specifically, platelet adhesion. It is deficient and/or defective in von Willebrand disease and is involved in many other diseases, including thrombotic thro ...
.


Early life and education

Springer was born in 1948 in
Fort Benning Fort Benning (named Fort Moore from 2023–2025) is a United States Army post in the Columbus, Georgia area. Located on Georgia's border with Alabama, Fort Benning supports more than 120,000 active-duty military, family members, reserve compone ...
,
Georgia Georgia most commonly refers to: * Georgia (country), a country in the South Caucasus * Georgia (U.S. state), a state in the southeastern United States Georgia may also refer to: People and fictional characters * Georgia (name), a list of pe ...
. His father was a
physician A physician, medical practitioner (British English), medical doctor, or simply doctor is a health professional who practices medicine, which is concerned with promoting, maintaining or restoring health through the Medical education, study, Med ...
. Springer attended the
University of California, Berkeley The University of California, Berkeley (UC Berkeley, Berkeley, Cal, or California), is a Public university, public Land-grant university, land-grant research university in Berkeley, California, United States. Founded in 1868 and named after t ...
, majoring in
biochemistry Biochemistry, or biological chemistry, is the study of chemical processes within and relating to living organisms. A sub-discipline of both chemistry and biology, biochemistry may be divided into three fields: structural biology, enzymology, a ...
and graduating with a BA in 1971. He went on to pursue a
PhD A Doctor of Philosophy (PhD, DPhil; or ) is a terminal degree that usually denotes the highest level of academic achievement in a given discipline and is awarded following a course of graduate study and original research. The name of the deg ...
under
Jack Strominger Jack Leonard Strominger (born August 7, 1925) is the Higgins Professor of Biochemistry at Harvard University, specializing in the structure and function of human histocompatibility proteins and their role in disease. He won the Albert Lasker Awa ...
at
Harvard University Harvard University is a Private university, private Ivy League research university in Cambridge, Massachusetts, United States. Founded in 1636 and named for its first benefactor, the History of the Puritans in North America, Puritan clergyma ...
, completing it in 1976.


Career

After obtaining his
PhD A Doctor of Philosophy (PhD, DPhil; or ) is a terminal degree that usually denotes the highest level of academic achievement in a given discipline and is awarded following a course of graduate study and original research. The name of the deg ...
, Springer took a
postdoctoral A postdoctoral fellow, postdoctoral researcher, or simply postdoc, is a person professionally conducting research after the completion of their doctoral studies (typically a PhD). Postdocs most commonly, but not always, have a temporary acade ...
position under
César Milstein César Milstein, CH, FRS (8 October 1927 – 24 March 2002) was an Argentine biochemist in the field of antibody research. Milstein shared the Nobel Prize in Physiology or Medicine in 1984 with Niels Kaj Jerne and Georges J. F. Köhler for ...
at the
University of Cambridge The University of Cambridge is a Public university, public collegiate university, collegiate research university in Cambridge, England. Founded in 1209, the University of Cambridge is the List of oldest universities in continuous operation, wo ...
and the
MRC Laboratory of Molecular Biology The Medical Research Council (MRC) Laboratory of Molecular Biology (LMB) is a research institute in Cambridge, England, involved in the revolution in molecular biology which occurred in the 1950–60s. Since then it has remained a major medical r ...
. A year later, he returned to the
United States The United States of America (USA), also known as the United States (U.S.) or America, is a country primarily located in North America. It is a federal republic of 50 U.S. state, states and a federal capital district, Washington, D.C. The 48 ...
and became an
assistant professor Assistant professor is an academic rank just below the rank of an associate professor used in universities or colleges, mainly in the United States, Canada, Japan, and South Korea. Overview This position is generally taken after earning a doct ...
at
Harvard Harvard University is a private Ivy League research university in Cambridge, Massachusetts, United States. Founded in 1636 and named for its first benefactor, the Puritan clergyman John Harvard, it is the oldest institution of higher lear ...
Medical School A medical school is a tertiary educational institution, professional school, or forms a part of such an institution, that teaches medicine, and awards a professional degree for physicians. Such medical degrees include the Bachelor of Medicine, ...
. Springer was promoted to
associate professor Associate professor is an academic title with two principal meanings: in the North American system and that of the ''Commonwealth system''. In the ''North American system'', used in the United States and many other countries, it is a position ...
in 1983 and was appointed Latham Family
Professor Professor (commonly abbreviated as Prof.) is an Academy, academic rank at university, universities and other tertiary education, post-secondary education and research institutions in most countries. Literally, ''professor'' derives from Latin ...
in 1989. Outside of Harvard, Springer was the Chief of the Laboratory of Membrane
Immunochemistry Immunochemistry is the study of the chemistry of the immune system. This involves the study of the properties, functions, interactions and production of the chemical components of the immune system. It also include immune responses and determina ...
at the
Dana Farber Cancer Institute Dana may refer to: Businesses and organisations * Dana (company), a Slovenian beverage company * Dana (payment service), in Indonesia * Dana Air, a Nigerian airline * Dana College, formerly in Nebraska, U.S. * Dana Energy, an Iranian oil and ga ...
between 1981 and 1988, and a Senior Investigator at the Immune Disease Institute of
Boston Boston is the capital and most populous city in the Commonwealth (U.S. state), Commonwealth of Massachusetts in the United States. The city serves as the cultural and Financial centre, financial center of New England, a region of the Northeas ...
from 1988 until its 2012 merge into
Boston Children's Hospital Boston Children's Hospital (formerly known as Children's Hospital Boston until 2013) is the main pediatric training and research hospital of Harvard Medical School, Harvard University. It is a nationally ranked, freestanding acute care children ...
. Since 2012, Springer has been a Senior Investigator at the Research Program in Cellular and Molecular Medicine of the Boston Children's Hospital. Springer is involved in a number of business ventures. He founded LeukoSite in 1993, which went
public In public relations and communication science, publics are groups of individual people, and the public (a.k.a. the general public) is the totality of such groupings. This is a different concept to the sociology, sociological concept of the ''Öf ...
in 1997 and was acquired by
Millennium Pharmaceuticals Takeda Oncology (originally Millennium Pharmaceuticals) is a biopharmaceutical company based in Cambridge, Massachusetts. It is a fully owned subsidiary of Takeda Pharmaceutical. Takeda Oncology's research, development and commercialization ac ...
the next year. He co-founded biotechnology companies Scholar Rock in 2012 and Morphic Therapeutic in 2015. He was also an early investor in Selecta Biosciences and Editas Medicine. Springer was a founding investor of the then-
startup company A startup or start-up is a company or project undertaken by an Entrepreneurship, entrepreneur to seek, develop, and validate a scalable business model. While entrepreneurship includes all new businesses including self-employment and businesses tha ...
Moderna Moderna, Inc. ( ) is an American pharmaceutical and biotechnology company based in Cambridge, Massachusetts, that focuses on RNA therapeutics, primarily mRNA vaccines. These vaccines use a copy of a molecule called messenger RNA (mRNA) to carry inst ...
after investing
USD The United States dollar (symbol: $; currency code: USD) is the official currency of the United States and several other countries. The Coinage Act of 1792 introduced the U.S. dollar at par with the Spanish silver dollar, divided it int ...
$5 million in 2010. He was the company's fourth-largest shareholder and made USD$400 million when the company launched its
initial public offering An initial public offering (IPO) or stock launch is a public offering in which shares of a company are sold to institutional investors and usually also to retail (individual) investors. An IPO is typically underwritten by one or more investm ...
(IPO) in 2018. During the
COVID-19 pandemic The COVID-19 pandemic (also known as the coronavirus pandemic and COVID pandemic), caused by severe acute respiratory syndrome coronavirus 2 (SARS-CoV-2), began with an disease outbreak, outbreak of COVID-19 in Wuhan, China, in December ...
, ''
Forbes ''Forbes'' () is an American business magazine founded by B. C. Forbes in 1917. It has been owned by the Hong Kong–based investment group Integrated Whale Media Investments since 2014. Its chairman and editor-in-chief is Steve Forbes. The co ...
'' estimated Springer's net worth as USD$1 billion after in share price of biotechnology companies surged. In 2017, Springer co-founded the
501(c)(3) organization A 501(c)(3) organization is a United States corporation, Trust (business), trust, unincorporated association or other type of organization exempt from federal income tax under section 501(c)(3) of Title 26 of the United States Code. It is one of ...
Institute for Protein Innovation, which focuses on
antibody An antibody (Ab) or immunoglobulin (Ig) is a large, Y-shaped protein belonging to the immunoglobulin superfamily which is used by the immune system to identify and neutralize antigens such as pathogenic bacteria, bacteria and viruses, includin ...
research, with him providing a $10 million foundational grant.


Research

Springer began his research career in
immunology Immunology is a branch of biology and medicine that covers the study of Immune system, immune systems in all Organism, organisms. Immunology charts, measures, and contextualizes the Physiology, physiological functioning of the immune system in ...
. He was studying
cytotoxic T cell A cytotoxic T cell (also known as TC, cytotoxic T lymphocyte, CTL, T-killer cell, cytolytic T cell, CD8+ T-cell or killer T cell) is a T lymphocyte (a type of white blood cell) that kills cancer cells, cells that are infected by intracellular ...
s' interaction with
antigen In immunology, an antigen (Ag) is a molecule, moiety, foreign particulate matter, or an allergen, such as pollen, that can bind to a specific antibody or T-cell receptor. The presence of antigens in the body may trigger an immune response. ...
s, and, since this interaction depends on
magnesium Magnesium is a chemical element; it has Symbol (chemistry), symbol Mg and atomic number 12. It is a shiny gray metal having a low density, low melting point and high chemical reactivity. Like the other alkaline earth metals (group 2 ...
, believed a
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residue (biochemistry), residues. Proteins perform a vast array of functions within organisms, including Enzyme catalysis, catalysing metab ...
in addition to the
T-cell receptor The T-cell receptor (TCR) is a protein complex, located on the surface of T cells (also called T lymphocytes). They are responsible for recognizing fragments of antigen as peptides bound to major histocompatibility complex (MHC) molecules. ...
is required. He found a
monoclonal antibody A monoclonal antibody (mAb, more rarely called moAb) is an antibody produced from a cell lineage made by cloning a unique white blood cell. All subsequent antibodies derived this way trace back to a unique parent cell. Monoclonal antibodie ...
that binds a new protein prevents cytotoxic T cells interacting with antigens, and named the protein
LFA-1 Lymphocyte function-associated antigen 1 (LFA-1) is an integrin found on lymphocytes and other leukocytes. LFA-1 plays a key role in emigration, which is the process by which leukocytes leave the bloodstream to enter the tissues. LFA-1 also mediates ...
. LFA-1 is a heterodimer, meaning it is made of two different
protein subunit In structural biology, a protein subunit is a polypeptide chain or single protein molecule that assembles (or "''coassembles''") with others to form a protein complex. Large assemblies of proteins such as viruses often use a small number of t ...
s. Around the same period,
Richard Hynes Richard Olding Hynes (born 29 November 1944) is a British biologist, a Howard Hughes Medical Institute Investigator, and the Daniel K. Ludwig Professor for Cancer Research at the Koch Institute for Integrative Cancer Research, Massachusetts ...
from the
United States The United States of America (USA), also known as the United States (U.S.) or America, is a country primarily located in North America. It is a federal republic of 50 U.S. state, states and a federal capital district, Washington, D.C. The 48 ...
and
Erkki Ruoslahti Erkki Ruoslahti (born 16 February 1940 in Imatra, Finland) is a cancer researcher and distinguished professor at Sanford Burnham Prebys Medical Discovery Institute. He moved from Finland to the United States in 1976. Ruoslahti made seminal contri ...
from
Finland Finland, officially the Republic of Finland, is a Nordic country in Northern Europe. It borders Sweden to the northwest, Norway to the north, and Russia to the east, with the Gulf of Bothnia to the west and the Gulf of Finland to the south, ...
were independently characterizing proteins on the
cell Cell most often refers to: * Cell (biology), the functional basic unit of life * Cellphone, a phone connected to a cellular network * Clandestine cell, a penetration-resistant form of a secret or outlawed organization * Electrochemical cell, a de ...
surface that helps attach cells to the surrounding
extracellular matrix In biology, the extracellular matrix (ECM), also called intercellular matrix (ICM), is a network consisting of extracellular macromolecules and minerals, such as collagen, enzymes, glycoproteins and hydroxyapatite that provide structural and bio ...
. They discovered
fibronectin Fibronectin is a high- molecular weight (~500-~600 kDa) glycoprotein of the extracellular matrix that binds to membrane-spanning receptor proteins called integrins. Fibronectin also binds to other extracellular matrix proteins such as col ...
and a receptor protein to which fibronectin binds. The fibronectin receptor, which Hynes named "integrin", is also a protein heterodimer. Springer found that one of the protein subunits of LFA-1 and Mac-1, a protein heterodimer found on
macrophage Macrophages (; abbreviated MPhi, φ, MΦ or MP) are a type of white blood cell of the innate immune system that engulf and digest pathogens, such as cancer cells, microbes, cellular debris and foreign substances, which do not have proteins that ...
s, has a highly similar
DNA Deoxyribonucleic acid (; DNA) is a polymer composed of two polynucleotide chains that coil around each other to form a double helix. The polymer carries genetic instructions for the development, functioning, growth and reproduction of al ...
sequence to one of the subunits of the fibronectin receptor, suggesting the three proteins belong to the same family of proteins. Springer's group also discovered
ICAM-1 ICAM-1 (Intercellular adhesion molecule, Intercellular Adhesion Molecule 1) also known as CD54 (Cluster of Differentiation 54) is a protein that in humans is encoded by the ''ICAM1'' gene. This gene encodes a cell surface glycoprotein which is ty ...
, the protein to which LFA-1 binds, and that this interaction is essential for cytotoxic T cells to recognize antigens. His innovative use of
monoclonal antibodies A monoclonal antibody (mAb, more rarely called moAb) is an antibody produced from a Lineage (evolution), cell lineage made by cloning a unique white blood cell. All subsequent antibodies derived this way trace back to a unique parent cell. Mon ...
in these discoveries paved the way for the development of
therapeutic antibodies Monoclonal antibodies (mAbs) have varied therapeutic uses. It is possible to create a mAb that binds specifically to almost any extracellular target, such as cell surface proteins and cytokines. They can be used to render their target ineffective ...
, known as selective adhesion molecule inhibitors, to treat
autoimmune disease An autoimmune disease is a condition that results from an anomalous response of the adaptive immune system, wherein it mistakenly targets and attacks healthy, functioning parts of the body as if they were foreign organisms. It is estimated tha ...
s. In addition, Springer identified the steps through which
white blood cell White blood cells (scientific name leukocytes), also called immune cells or immunocytes, are cells of the immune system that are involved in protecting the body against both infectious disease and foreign entities. White blood cells are genera ...
s move out of the
circulatory system In vertebrates, the circulatory system is a system of organs that includes the heart, blood vessels, and blood which is circulated throughout the body. It includes the cardiovascular system, or vascular system, that consists of the heart ...
towards the site of damage or
infection An infection is the invasion of tissue (biology), tissues by pathogens, their multiplication, and the reaction of host (biology), host tissues to the infectious agent and the toxins they produce. An infectious disease, also known as a transmis ...
. More recently, research at Springer's group has expanded to
malaria Malaria is a Mosquito-borne disease, mosquito-borne infectious disease that affects vertebrates and ''Anopheles'' mosquitoes. Human malaria causes Signs and symptoms, symptoms that typically include fever, Fatigue (medical), fatigue, vomitin ...
vaccine A vaccine is a biological Dosage form, preparation that provides active acquired immunity to a particular infectious disease, infectious or cancer, malignant disease. The safety and effectiveness of vaccines has been widely studied and verifi ...
,
transforming growth factor beta Transforming growth factor beta (TGF-β) is a multifunctional cytokine belonging to the transforming growth factor superfamily that includes three different mammalian isoforms (TGF-β 1 to 3, HGNC symbols TGFB1, TGFB2, TGFB3) and many other ...
(TGF-β) signaling, and how
von Willebrand factor Von Willebrand factor (VWF) () is a blood glycoprotein that promotes primary hemostasis, specifically, platelet adhesion. It is deficient and/or defective in von Willebrand disease and is involved in many other diseases, including thrombotic thro ...
starts the process of stopping bleeding.


Personal life

Springer is a
gongshi ''Gongshi'' (), also known as scholar's rocks or viewing stones, are naturally occurring or shaped rocks which are traditionally appreciated by Chinese scholars.Metropolitan Museum of Art "The World of Scholars' Rocks Gardens, Studios, and Pai ...
collector. Springer is married to Chafen Lu, a former
assistant professor Assistant professor is an academic rank just below the rank of an associate professor used in universities or colleges, mainly in the United States, Canada, Japan, and South Korea. Overview This position is generally taken after earning a doct ...
at
Harvard Harvard University is a private Ivy League research university in Cambridge, Massachusetts, United States. Founded in 1636 and named for its first benefactor, the Puritan clergyman John Harvard, it is the oldest institution of higher lear ...
Medical School A medical school is a tertiary educational institution, professional school, or forms a part of such an institution, that teaches medicine, and awards a professional degree for physicians. Such medical degrees include the Bachelor of Medicine, ...
and an alumnus of his lab. He has five children, three from his first marriage. Springer founded and has made contributions to the Institute for Protein Innovation, a Boston-based nonprofit biomedical research organization. He also has endowed professorships at Harvard Medical School, Boston Children's Hospital, and Berkeley.


Honors and awards

*
American Heart Association The American Heart Association (AHA) is a nonprofit organization in the United States that funds cardiovascular medical research, educates consumers on healthy living and fosters appropriate Heart, cardiac care in an effort to reduce disability ...
Basic Research Prize (1993) * William B. Coley Award for Distinguished Research in Basic and Tumor Immunology (1995) * Member of the
National Academy of Sciences The National Academy of Sciences (NAS) is a United States nonprofit, NGO, non-governmental organization. NAS is part of the National Academies of Sciences, Engineering, and Medicine, along with the National Academy of Engineering (NAE) and the ...
(1996) * Fellow of the
American Academy of Arts and Sciences The American Academy of Arts and Sciences (The Academy) is one of the oldest learned societies in the United States. It was founded in 1780 during the American Revolution by John Adams, John Hancock, James Bowdoin, Andrew Oliver, and other ...
(2001) *
Crafoord Prize The Crafoord Prize () is an annual science prize established in 1980 by Holger Crafoord, a Swedish industrialist, and his wife Anna-Greta Crafoord following a donation to the Royal Swedish Academy of Sciences. It is awarded jointly by the Acade ...
in
Polyarthritis Polyarthritis is any type of arthritis that involves 5 or more joints simultaneously. It can be associated with autoimmune conditions; it may be experienced at any age and is not sex specific. Causes Polyarthritis is often caused by an auto-immun ...
(2004) *
Guggenheim Fellowship Guggenheim Fellowships are Grant (money), grants that have been awarded annually since by the John Simon Guggenheim Memorial Foundation, endowed by the late Simon Guggenheim, Simon and Olga Hirsh Guggenheim. These awards are bestowed upon indiv ...
(2004) *
Fellow A fellow is a title and form of address for distinguished, learned, or skilled individuals in academia, medicine, research, and industry. The exact meaning of the term differs in each field. In learned society, learned or professional society, p ...
of the
American Association for the Advancement of Science The American Association for the Advancement of Science (AAAS) is a United States–based international nonprofit with the stated mission of promoting cooperation among scientists, defending scientific freedom, encouraging scientific responsib ...
(2013) * AAI-Life Technologies Meritorious Career Award (now AAI-Thermo Fisher Meritorious Career Award),
American Association of Immunologists American(s) may refer to: * American, something of, from, or related to the United States of America, commonly known as the "United States" or "America" ** Americans, citizens and nationals of the United States of America ** American ancestry, p ...
(2014) * Henry M. Stratton Medal,
American Society of Hematology The American Society of Hematology (ASH) is a professional organization representing hematologists, founded in 1958. Its annual meeting is held in December of every year and has attracted more than 30,000 attendees. The society publishes the medi ...
(2014) *
Canada Gairdner International Award The Canada Gairdner International Award is given annually by the Gairdner Foundation at a special dinner to five individuals for outstanding discoveries or contributions to medical science. Receipt of the Gairdner is traditionally considered a ...
(2019) *
Albert Lasker Award for Basic Medical Research The Albert Lasker Award for Basic Medical Research is one of the Lasker Award, prizes awarded by the Lasker Foundation for a fundamental discovery that opens up a new area of biomedical science. The award frequently precedes a Nobel Prize in Phys ...
(2022) *
Robert Koch Prize The Robert Koch Medal and Award are two prizes awarded annually by the German for excellence in the biomedical sciences. These awards grew out of early attempts by German physician Robert Koch to generate funding to support his research into the ...
(2023) He was a
Phi Beta Kappa The Phi Beta Kappa Society () is the oldest academic honor society in the United States. It was founded in 1776 at the College of William & Mary in Virginia. Phi Beta Kappa aims to promote and advocate excellence in the liberal arts and sciences, ...
graduate from the
University of California, Berkeley The University of California, Berkeley (UC Berkeley, Berkeley, Cal, or California), is a Public university, public Land-grant university, land-grant research university in Berkeley, California, United States. Founded in 1868 and named after t ...
.


References

{{DEFAULTSORT:Springer, Timothy Alan Harvard University faculty 1948 births Living people Fellows of the American Academy of Arts and Sciences Fellows of the American Association for the Advancement of Science Members of the United States National Academy of Sciences Recipients of the Albert Lasker Award for Basic Medical Research American billionaires American immunologists Harvard Medical School alumni UC Berkeley College of Letters and Science alumni University of California, Berkeley alumni Moderna people Academics of the University of Cambridge Members of the National Academy of Medicine