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Timothy A. Springer

Timothy A. Springer is an immunologist, , and biophysicist renowned for his pioneering discoveries of molecules, including the identification of and their roles in immune cell interactions and leukocyte migration. His work has advanced understanding of physiological processes like and immunity, leading to therapeutic developments such as the FDA-approved drugs for and for dry eye disease. Springer holds the position of Latham Family Professor of Biological Chemistry and Molecular Pharmacology at , where he has been a faculty member since 1977, and he is also affiliated with . Springer earned a B.A. in biochemistry from the , in 1971, followed by a Ph.D. in and biochemistry from in 1976, and completed a postdoctoral fellowship with at the in , . He joined as an assistant professor in 1977 and was appointed the Latham Family Professor in 1989, while also serving as a senior investigator at Boston Children's Hospital's Department of Cellular and Molecular Medicine. Throughout his career, Springer has authored over 500 publications with a Hirsch index of 197 (as of 2025) and holds more than 30 patents related to adhesion molecules and protein structures. His research has focused on activation mechanisms, protein conformational changes, and applications to diseases including and autoimmune disorders. In 2022, Springer shared the Albert Lasker Basic Medical Research Award with Richard O. Hynes and Erkki Ruoslahti for their foundational work on as mediators of in health and disease. He is a member of the (since 1996) and the (since 2023) and has received the Crafoord Prize from the Royal Swedish Academy of Sciences, the American Association of Immunologists Meritorious Career Award, the Henry M. Stratton Medal from the American Society of , the Biophysical Society Founders Award (2022), the Prize (2023), and the Protein Society Stein & Moore Award (2025). As an entrepreneur, Springer founded LeukoSite in 1993, which was acquired by Millennium Pharmaceuticals in 1999 and contributed to the development of for ; he has also been a founding investor in companies including Moderna Therapeutics, Scholar Rock, and Morphic Therapeutic. In philanthropy, he established the nonprofit Institute for Protein Innovation in 2017 and pledged $210 million to it in 2023 to advance open-access antibody research for biomedical discovery.

Early Life and Education

Early Life

Timothy A. Springer was born on February 23, 1948, in , , to a . The family's frequent moves, driven by his father's military postings, shaped Springer's early childhood in a dynamic environment that exposed him to diverse settings across the . Springer's interest in science emerged during his youth, largely influenced by personal health challenges and his father's medical profession. As a child, he suffered from severe hay fever, which necessitated regular visits to an allergist for desensitization treatments using progressively stronger injections; these experiences ignited his curiosity about and the workings of the . Additionally, growing up around his father's practice provided early exposure to pharmaceuticals and medical concepts, further fostering his fascination with biology and medicine. The family eventually settled in Sacramento, California, where Springer spent much of his formative years and developed his initial academic pursuits in biology and medicine during high school. This period solidified his passion for scientific inquiry, leading him to pursue higher education at the University of California, Berkeley.

Education

After one year at Yale University, where he questioned his Ivy League path and volunteered with VISTA on a Native American reservation, Timothy A. Springer transferred to the University of California, Berkeley, earning a Bachelor of Arts degree in biochemistry in 1971. During his undergraduate studies, he gained early exposure to immunology through hands-on research involving the use of antibodies. Springer then pursued graduate studies at , where he received a Ph.D. in and biochemistry in 1976. His doctoral research, supervised by Jack L. Strominger, focused on and , with his thesis titled "Studies on the Structure of the Human Ia-like Antigens." Following his Ph.D., Springer conducted postdoctoral training as a fellow at the Medical Research Council Laboratory of Molecular Biology in Cambridge, England, from 1976 to 1977, under the guidance of . This period marked the beginning of his research interests in leukocyte function and the development of techniques.

Academic and Research Career

Academic Positions

Timothy A. Springer began his academic career at as an of in 1977, following his postdoctoral fellowship. He advanced to in 1983 and was promoted to full Professor of in 1989, later changing to Professor of Biological Chemistry and Molecular in 2011. That same year, he was appointed the Latham Family Professor, a position he has held continuously at . In parallel with his Harvard faculty roles, Springer joined the Center for Blood Research (CBR) in 1983 as a Senior Investigator, where he established and led a research laboratory focused on cellular and molecular biology. The CBR integrated into in 1992 and later became part of the Immune Disease Institute in 2006, during which time Springer continued his Senior Investigator role and laboratory leadership within the Program of Cellular and Molecular Medicine. Springer also holds an adjunct position as Resident Professor at , Inc., serving in a scientific advisory capacity since 2011.

Key Research Contributions

Timothy A. Springer discovered (LFA-1), a key on the surface of cytotoxic T cells essential for immune and target cell interactions, in the early 1980s through studies that identified its role in T cell-mediated and mixed reactions. His work demonstrated that antibodies targeting LFA-1 could block T cell adhesion to target cells, establishing it as a critical mediator of immune responses. This discovery, detailed in a 1981 Journal of Immunology paper, laid the foundation for understanding leukocyte adhesion deficiencies. In 1982–1985, Springer pioneered the identification of the integrin family by linking LFA-1 to other leukocyte adhesion molecules like Mac-1 and p150,95 through their shared β2 subunit, revealing a superfamily of transmembrane receptors that mediate cell-cell and cell-matrix interactions crucial for leukocyte trafficking and . His 1983 Journal of Experimental Medicine publication characterized these β2 , showing their conserved structure and function in immune . By 1986–1987, Springer's research expanded the integrin concept to include both and signaling roles, influencing fields from to . Springer's development of intercellular adhesion molecule 1 () research highlighted its role as the primary endothelial ligand for LFA-1, facilitating during inflammation and immune responses; a 1988 paper confirmed this interaction, showing how it enables firm under shear flow. This work elucidated the stepwise adhesion cascade—rolling, activation, and firm arrest—and its dysregulation in autoimmune diseases. In the 1990s–2000s, Springer contributed to malaria research by identifying ICAM-1 as an endothelial receptor for Plasmodium falciparum-infected erythrocytes, a finding from his 1992 Cell study that linked cytoadherence to cerebral malaria pathogenesis. Subsequent structural studies on PfEMP1 domains binding ICAM-1, following the 1995 identification of PfEMP1 expression from var genes, advanced understanding of antigenic variation and immune evasion, informing vaccine strategies targeting conserved binding sites to prevent severe malaria. Springer's investigations into transforming growth factor-β (TGF-β) signaling pathways revealed mechanisms of and , including the role of αv in releasing active TGF-β from its prodomain, with implications for , cancer, and immune regulation; key structures from 2011–2016 publications in and detailed the TGF-β/ complex and evolutionary conservation. His work on (vWF) structure elucidated its multimeric assembly and shear-dependent in and , using cryo-EM to show force-induced unfolding of the A2 domain for cleavage, as reported in 2014–2021 and PNAS papers. These contributions underscore vWF's role in platelet adhesion under high shear. With over 500 publications (exceeding 600 as of 2025), Springer's seminal works, such as the 1985 review on adhesion molecules and the 1987 paper on structure, have been highly cited and shaped modern .

Biotechnology Entrepreneurship

Founded Companies

Timothy A. Springer has founded several companies, leveraging his expertise in protein interactions and to advance therapeutic development in areas such as , neuromuscular disorders, and antibody engineering. In 1993, Springer founded LeukoSite, Inc., a company focused on discovering and developing therapeutics targeting leukocyte molecules to treat inflammatory diseases. The company went public in 1998 and was acquired by Pharmaceuticals in 1999 for $635 million in stock, which later appreciated significantly as 's value grew. LeukoSite's work contributed to the development of key drugs like (Entyvio), approved for . Springer co-founded Scholar Rock Holding Corporation in 2012 with Harvard colleague Leonard I. Zon, aiming to develop protein therapeutics that selectively modulate TGF-β signaling for neuromuscular and cardiometabolic diseases. The company raised initial funding through a $20 million Series A in 2014, led by ARCH Venture Partners with participation from Springer and others. Scholar Rock completed its on in May 2018, raising $75 million to advance its pipeline, including candidates for and other rare diseases. In 2015, Springer established Morphic Therapeutic, Inc. (now part of Morphic Holding), based on discoveries from his laboratory regarding structures, to develop oral small-molecule therapies for chronic inflammatory conditions like . Co-founded with Partners, the company secured a $51.5 million Series A financing in 2016 to support preclinical and clinical development. Morphic went public in 2019 and was acquired by in 2024 for up to $3.2 billion, highlighting the impact of its alpha4beta7 inhibitor program. Springer has also been involved in founding early-stage ventures, such as Biotherapeutics in 2015, which applies computational and to design novel for immune-mediated diseases. As a lead founder and board member, he has guided its focus on platforms. In 2019, Springer co-founded Tectonic Therapeutic with Kruse, focusing on using computational design and to target challenging membrane proteins for and . The company launched with $80 million in Series A financing in 2021. In 2022, Springer co-founded Seismic Therapeutic, integrating with to develop precision immunomodulators for autoimmune diseases and . Seismic emerged with $101 million in Series A .

Investments and Board Roles

Springer made a pivotal seed investment of $5 million in , Inc. in 2010 as one of its founding investors. Following Moderna's in 2018, this stake appreciated to more than $400 million, playing a key role in elevating Springer's wealth to status by 2020. In addition to his Moderna stake, Springer has actively participated as a lead and board member at Biotherapeutics, a company developing proprietary platforms for discovery in protein therapeutics. His involvement underscores his commitment to innovative approaches in biologics development. Springer holds board positions in several biotech firms beyond those he founded, including as an at Tectonic Therapeutic since 2020 and as a at Cartesian Therapeutics (formerly Selecta Biosciences) since 2016. He has also maintained board roles in Scholar Rock Holding Corporation and Morphic Holding, Inc., providing strategic oversight following their foundings in 2012 and 2015, respectively. These roles reflect his ongoing influence in advancing therapies for musculoskeletal disorders, , and other conditions. Beyond these, Springer has invested in other emerging biotechs, such as and Seismic Therapeutics, supporting advancements in gene editing and immunology-based treatments. His diversified portfolio has fueled innovation across the sector, with his estimated at $1.1 billion as of 2025.

Philanthropy

Institute for Protein Innovation

The Institute for Protein Innovation (IPI) was established in 2017 in , , as a non-profit organization dedicated to generating open-source, recombinant antibodies to support protein research and address longstanding gaps in available research tools. Co-founded by Timothy A. Springer and Kruse, IPI employs yeast surface display technology to produce high-quality, synthetic antibodies targeting human extracellular and secreted proteins, with an initial focus on creating a comprehensive library for applications. Springer, a at and , served as the primary founder and initial funder, motivated in part by challenges encountered in his own research on , where reliable antibodies were often lacking. Under his vision, IPI operates as an open-access foundry, validating antibodies through rigorous testing and sharing them freely via platforms like Addgene to accelerate biomedical discoveries without commercial restrictions. By 2025, IPI had expanded its operations significantly, incorporating and to optimize design and generate data tailored for computational modeling, marking a shift toward more efficient, precision-engineered tools. The institute now maintains advanced facilities on the campus and has forged key partnerships with academic labs, including those at , the , and initiatives funded by the , to co-develop antibodies for and other fields. IPI's contributions have notably impacted protein by distributing validated collections freely to researchers globally, including 26 recombinant antibodies against the family launched in 2024 and a 2024 set targeting nine common tags such as V5 and DYKDDDDK (FLAG), enabling reproducible studies in , structural determination, and disease modeling. These resources, accompanied by plasmids, sequences, and validation data, have supported hundreds of investigators in advancing human health research while promoting community-driven validation standards.

Major Donations

In 2023, Timothy A. Springer and his family donated $210 million to the Institute for Protein Innovation (IPI), establishing an endowment that marked one of the largest gifts to a biomedical research nonprofit at the time and enabling the organization's expansion in protein science research. This contribution built on Springer's prior support for IPI, where he provided seed funding totaling $40 million between 2017 and 2022 to launch and sustain the nonprofit's initial operations focused on open-access antibody resources. Springer has also made significant endowments to academic institutions, including chairs at and dedicated to advancing research. These philanthropic efforts have been largely funded by proceeds from his early investment in , directing resources toward initiatives and underfunded areas of biomedical discovery such as protein therapeutics.

Personal Life

Family

Timothy A. Springer is married to Chafen Lu, a structural biologist and former assistant professor at who previously worked as a researcher in his laboratory. The couple has collaborated professionally on multiple research projects, including studies on subunit folding, dimerization mechanisms, and fusogen structures, with co-authored publications spanning from the late 1990s to the . Springer and Lu have five children together, three of whom are from Springer's first marriage. The family has demonstrated unity in philanthropic efforts, jointly contributing to a $210 million donation to for Protein Innovation on March 29, 2023. Despite Springer's demanding career as a Harvard and biotech entrepreneur, the family maintains a low public profile, with Lu supporting shared interests in scientific research.

Interests and Hobbies

Timothy A. Springer maintains a notable collection of , or scholar's rocks, which are naturally sculpted stones valued for their abstract forms and symbolic resonance with nature. These rocks, formed over millennia by from rivers and wind, embody aesthetic principles rooted in literati , evoking landscapes, mountains, and philosophical ideals. Springer's passion for them highlights his appreciation for the interplay of natural processes and artistic contemplation. His collection began after acquiring an initial piece, prompting him to study the tradition under the guidance of Kemin Hu, a prominent expert and dealer based in . Springer has since traveled extensively to on sourcing expeditions, amassing pieces through dedicated buying trips; he once shipped seven containers back to his home, including one monumental stone weighing 23 tons. The rocks are prominently displayed in his backyard garden, integrated among specimen trees such as a cedar of , creating a contemplative outdoor space. One favored Duanxi stone serves as his personal "sitting rock," inscribed with a poem he composed. Springer describes his pursuits as his "greatest extravagance," underscoring how this offers respite and creative outlet amid his rigorous scientific and entrepreneurial endeavors. By immersing himself in the selection and arrangement of these natural artifacts, he finds equilibrium, channeling a sense of wonder derived from their organic beauty into his daily life.

Honors and Awards

Major Scientific Awards

Timothy A. Springer has received several of the world's most prestigious scientific awards for his pioneering discoveries in molecules and , which have transformed understanding of immune responses and led to therapeutic advancements. In 2004, Springer shared the Crafoord Prize in from the Royal Swedish Academy of Sciences with Eugene Butcher for their discoveries of molecules that enable to exit blood vessels, fight infections, and contribute to inflammatory diseases such as allergies, arthritis, and cancers. The award, valued at $500,000, recognized Springer's identification of key molecules like , LFA-1, and LFA-3, which underpin cellular interactions and have informed treatments for conditions including and blood cancers. In 2014, Springer received the American Association of Immunologists Meritorious Career Award for his outstanding research contributions to , particularly in the discovery and characterization of leukocyte molecules. Also in 2014, Springer was awarded the Henry M. Stratton Medal from the American Society of , shared with Geraldine P. Schechter, for basic and clinical research contributions to , recognizing his work on and their role in immune cell function. Springer received the in 2019 for his seminal discoveries of the first adhesion molecules, including elucidation of their roles in recognition, leukocyte homing, and that enables movement. This work also highlighted his contributions to translating these findings into therapeutics, such as monoclonal antibodies for autoimmune diseases like , , , and , as well as applications in . In 2022, Springer received the Biophysical Society Founders Award for his pioneering contributions to biophysical studies of immune cell rolling, activation, and adhesion, revealing force-based mechanisms in integrin function. In 2022, Springer was one of three recipients of the Albert Lasker Basic Medical Research Award, shared with Richard O. Hynes and Erkki Ruoslahti, for discoveries concerning integrins as key mediators of cell-matrix and cell-cell adhesion in physiology and disease. Springer's specific contributions included identifying transmembrane proteins such as LFA-1 and Mac-1 that facilitate immune cell interactions, linking them to a shared β-chain gene family, and revealing the ligand ICAM-1 along with the dynamic activation of integrins critical for immune cell migration. The Prize was awarded to Springer in 2023, shared with Sanchez-Madrid, for their groundbreaking research on molecules and their essential roles in immune and . Valued at €120,000, the prize honored Springer's discovery and mechanistic insights into , which drive and immune responses, and his development of monoclonal antibodies that have advanced therapies for and . In 2025, Springer received the Stein & Moore Award from The Protein Society for his groundbreaking work on and function, particularly in and their role in cellular adhesion and signaling.

Other Recognitions

Springer was elected to the in 1996, recognizing his foundational work in and related to cellular interactions. He became a of the American Academy of Arts and Sciences in 2001, honoring his interdisciplinary contributions to and . In 2023, Springer was elected to the for his pioneering research on receptor-ligand interactions and transmembrane signaling in immune responses. Since 1989, Springer has held the Latham Family Professorship at , an endowed chair that acknowledges his long-standing impact on biological chemistry, molecular , and . He has delivered several named lectureships, including the Tai-Shun Lin Memorial Lectureship at in 2025, where he discussed the and of ensembles. Additionally, in 2011, awarded him an honorary , celebrating his global influence on health sciences. Springer's entrepreneurial achievements have garnered recognition in financial media, with profiling him as a starting in 2020 due to his stakes in companies like , which surged amid the . These profiles highlight how his early discoveries in integrin biology translated into high-impact investments advancing therapeutic development.

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