Steven G. Boxer (born 1947) is an American biophysical chemist and the Camille Dreyfus Professor of Chemistry at Stanford University. He is a member of the National Academy of Sciences (elected 2008) and a fellow of the American Academy of Arts and Sciences, the American Association for the Advancement of Science, the Biophysical Society, and the Royal Society of Chemistry.
Education and Career
Boxer earned a B.S. with Honors in Chemistry from Tufts University in 1969 and a Ph.D. in Physical and Physical-Organic Chemistry from the University of Chicago in 1976 under the supervision of Gerhard L. Closs. He joined the faculty at Stanford University, where he has spent his entire academic career. He became Chair of the Department of Chemistry at Stanford in 2020.
Research
Boxer's laboratory investigates the structure and function of biological systems with a strong physical perspective, spanning three interconnected themes:
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Photosynthetic Reaction Centers: He has studied light-driven long-distance electron transfer in photosynthetic reaction centers using femtosecond fluorescence and transient absorption spectroscopy, electric field manipulation, site-specific mutagenesis, and Stark spectroscopy.
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Electrostatics in Proteins and Enzyme Catalysis: Boxer developed the use of vibrational Stark effect (VSE) spectroscopy to measure electric fields inside proteins with high spatial and field resolution. This work has enabled quantitative measurement of electrostatic contributions to enzyme catalysis, notably in ketosteroid isomerase (KSI), where his group demonstrated that extreme electric fields in the active site power catalysis. His group has also developed nitrile and carbonyl vibrational probes to map electrostatic fields in proteins, including studies of drug-target interactions in live cells using quantum cascade laser spectroscopy.
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Green Fluorescent Protein (GFP): His group was the first to demonstrate that the GFP chromophore exists in two protonation states interconvertible by ultrafast excited-state proton transfer. They have developed split GFP systems as optogenetic elements and studied photoisomerization pathways in fluorescent proteins.
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Model Membranes: Boxer's group has developed supported lipid bilayers as mimics for cell surfaces, including methods for patterning, electric field manipulation, and characterization using imaging mass spectrometry (NanoSIMS). They have studied membrane fusion mechanisms, including influenza and Zika virus fusion, and developed DNA-lipid conjugates as tools for mediating vesicle fusion.
Honors and Awards
- ACS Nakanishi Prize (2025)
- Honorary Doctor of Science, Tufts University (2025)
- Ellis R. Lippincott Award, Optica (2024)
- Founders Award, Biophysical Society (2024)
- Murray Goodman Memorial Prize, American Chemical Society (2014)
- E. Bright Wilson Award in Spectroscopy, American Chemical Society (2013)
- Earle K. Plyler Prize for Molecular Spectroscopy, American Physical Society (2008)
- Elected Member, National Academy of Sciences (2008)
- Arthur Cope Scholar Award, American Chemical Society (1995)
- Elected Fellow, American Academy of Arts and Sciences (1997)
Selected Publications
Boxer has authored over 400 research publications and has an h-index exceeding 100. Notable publications include:
- "Ultra-fast excited state dynamics in green fluorescent protein: multiple states and proton transfer" (Chattoraj, King, Bublitz, Boxer, PNAS, 1996) — over 1,000 citations
- "Extreme electric fields power catalysis in the active site of ketosteroid isomerase" (Fried, Bagchi, Boxer, Science, 2014)
- "Electric fields and enzyme catalysis" (Fried, Boxer, Annual Review of Biochemistry, 2017)
- "Measuring electric fields and noncovalent interactions using the vibrational Stark effect" (Fried, Boxer, Accounts of Chemical Research, 2015)
- "Stark spectroscopy: applications in chemistry, biology, and materials science" (Bublitz, Boxer, Annual Review of Physical Chemistry, 1997)
Personal
Boxer is married to Linda M. Boxer, the Stanley McCormick Memorial Professor in the School of Medicine at Stanford, Chief of Hematology, and Vice-Dean of the School of Medicine. They have two children.
