Welcome¶
Principal Investigator
Santiago Schnell
Santiago Schnell is the Provost, Professor of Mathematics at Dartmouth, and Adjunct Professor of Biochemistry & Cell Biology and Biomedical Data Science in the Geisel School of Medicine at Dartmouth. His research focuses on developing methods and models to obtain high-quality measurements with rigor, reproducibility, and robustness in the biomedical sciences.
Santiago uses mathematical, computational, and statistical approaches to tackle complex problems in the biomedical sciences. Our theoretical models serve as powerful tools for hypothesis testing and data interpretation, driving discovery in partnership with experimental research groups worldwide.
Selected Projects¶
Mathematical Biology
Principles of Enzyme and Biochemical Kinetics
We are deriving rate equations and developoing standard based approaches to measure the physical constants of biopchemical reactions.
Scientometrics
Measuring and Improving Research University Performance
We are developing new metrics and models to measure and improve performance of scientists, research universities, and academic policies.
Team¶
Postdoctoral Fellow
Abhishek Mallela
Abhishek works on parameter estimation of enzyme-catalyzed experiments. He is also exploring lumping methods in reaction networks, and how AI systems face fundamental social dilemma with game theory.
Postdoctoral Fellow
Jakub Svoboda
Jakub works on evolutionary dynamics on graphs, studying how population structure influences the spread of cooperative strategies. He is also investigating optimal decision-making in stochastic games on graphs.
Latest Cover Features¶
Biophysical Chemistry
July 1, 2021
Diabetes
September 1, 2019
The Journal of Physiology
March 31, 2019
See the rest of our cover features →
Latest Publications¶
J. D. Calhoun, C. Wang, C. Biar, J. R. Gunti, J. Lee, A. M. Geller, J. H. Hong, S. Schnell, L. T. Dang, Y. Wang, J. Parent, L. L. Isom, M. D. Uhler, H. C. Mefford, M. E. Ross, V. Pulido, G. Carvill (2026). EpiPred: A gene-specific machine learning model for classifying missense variants in the epilepsy-related gene STXBP1. Journal of Clinical Investigation, in press.
S. Schnell (2026). The stairs of reconciliation: A mathematical tourist in Graz. The Mathematical Intelligencer, in press.
S. Schnell (2026). Academic freedom and Catholic mission: The procedural overlap zone in university governance. Journal of Catholic Higher Education, in press.
S. Schnell (2026). From Signals to measurands: A measurement-science roadmap for reproducible analytical biochemistry. ACS Measurement Science Au, in press [Preprint ChemRxiv, DOI: 10.26434/chemrxiv.15005518]. (perspective) code
I. V. Gimón, C. Sandefur, S. Schnell (2026). Reproducible agent-based simulations of protein aggreation: A FAIR implementation. Bulletin of Mathematical Biology 88, art. 181. code
S. Schnell (2026). Reaction acceleration: Reviving the second derivative in chemical kinetics. ChemSystemsChem 8, e70047. code
J. Eilertsen, S. Schnell, S. Walcher (2026). Ehrlich occupancy time: Beyond \(k_{\rm off}\) to a complete residence time framework. Journal of Pharmacokinetics and Pharmacodynamics 53, art. 46. code
S. Pal, A. Mallela, L. Pracher, C. Wei, F. Fu, S. Schnell, M. A. Nowak (2026). Large language models instantiate evolutionarily robust strategies of cooperation. PNAS Nexus 5, pgag210. data code
H. V. Westerhoff, B. M. Bakker, A. S. Bommarius, M. L. Cárdenas, A. Cornish-Bowden, P. Fitzpatrick, P. Halling, V. Hatzimanikatis, C. Kettner, Y. Liu, A. G. McDonald, E. Noor, J. Pleiss, F. Raushel, J. Rohwer, S. Schnell, K. F. Tipton, M.-D. Tsai, U. von Stockar, U. Wittig, R. Wohlgemuth, J. M. Woodley (2026). Metabolic thermodynamics: Pertinent reference state and energy potentials FEBS Journal 293, 3111-3126.
S. Marques, J. Planas-Iglesias, J. Velecký, M. Musil, Y. Asano, T. Borowski, V. Brissos, M. Cespugli, K. Chorozian, M. Dadashipour, E. Erdem, E. Ferrandi, K. Grigorakis, A. Kluza, J. Lawniczek, K. Makryniotis, D. Monti, B. Nestl, A. Ngo, E. Nikolaivits, S. Patti, C. Pentari, C. Ferro Rodrigues, T. Schopper, K. Seweryn-Ożóg, M. Szaleniec, A. Taborda, M. Tataruch, D. Tischler, E. Topakas, J. Wang, P. Wójcik, A. Wojtkiewicz, J. Woodley, O. Zastawny, L. Martins, M. Fraaije, J. Pleiss, S. Schnell, J. Damborský, S. Mazurenko, D. Bednar (2026). Mobilizing the biocatalysis community for reproducible and reusable data collection. ACS Catalysis 16, 8858–8868.
A. Mallela, S. Schnell (2026). Structural hormesis in protein aggregation: A minimal mechanistic model Journal of the Royal Society Interface 23, 20251037. code
I. V. Gimón, C. Sandefur, S. Schnell (2026). Macromolecular crowding and protein aggregation: Friend, foe or contextual force. Progress in Biophysics and Molecular Biology 199, 79-98. (review)
I. V. Gimón, C. Sandefur, S. Schnell (2025). The hidden influences of macromolecular crowding: A new frontier in cellular biology and medicine. In: Uversky, V.N. (eds) (Macro)Molecular Crowding. Subcellular Biochemistry, vol 109. Springer, Cham. (review)
S. Schnell, D. L. Gumucio (2025). Medicine and The Arts: Commentary on Thistle. Academic Medicine 100, 1153. (see the image, Thistle by Maria Morrell, PhD)
K. Srivastava, J. Eilertsen, V. Booth, S. Schnell (2025). Accuracy versus Predominance: Reassessing the validity of the quasi-steady-state approximation. Bulletin of Mathematical Biology 87, 73.
S. Schnell, P. K. Maini (2024). In Memory of Edmund John Crampin: Multi-scale and multi-physics phenomena in biology.. Mathematical Biosciences 376, 109283. (editorial)
J. Eilertsen, S. Schnell, S. Walcher (2024). The Michaelis-Menten reaction at low substrate concentrations: Pseudo-first-order kinetics and conditions for timescale separation. Bulletin of Mathematical Biology 86, 68.
M. S. DeNies, Allen P. Liu, S. Schnell (2024). Seeing beyond the blot: A critical look at assumptions and raw data interpretation in Western blotting. Biomolecular Concepts 15, 20220047.
J. Eilertsen, S. Schnell, S. Walcher (2024). The unreasonable effectiveness of the total quasi-steady state approximation, and its limitations.. Journal of Theoretical Biology 583, 111770.
J. Eilertsen, S. Schnell, S. Walcher (2024). Rigorous estimates for the quasi-steady state approximation of the Michaelis-Menten reaction mechanism at low enzyme concentrations.. Nonlinear Analysis: Real World Applications 120, e2219683120.
K. Ikami, S. Shoffner-Beck, M. Tyczynska Weh, S. Schnell, S. Yoshida, E. A. Diaz Miranda, S. Ko, L. Lei (2023). Branched germline cysts and female-specific cyst fragmentation facilitate oocyte determination in mice.. Proceedings of the National Academy of Sciences of the United States of America 120, e2219683120.
J. Eilertsen, S. Schnell, S. Walcher (2023). Natural parameter conditions for singular perturbations of chemical and biochemical reaction networks.. Bulletin of Mathematical Biology 85, 48.
M. A. Gingerich, J. Zhu, B. Chai, M. P. Vincent, N. Xie, V. Sidarala, N. A. Kotov, D. Sahu. D. J. Klionsky, S. Schnell, S. A. Soleimanpour (2023). Reciprocal regulatory balance within the CLEC16A-RNF41 mitophagy complex depends on an intrinsically disordered protein region.. Journal of Biological Chemistry 299, 103057.
