- Open Access
Hamilton-Jacobi theory of quantized biological function
APS Open Sci. 1, 000140 – Published 17 September, 2026
DOI: https://doi.org/10.1103/x2fd-pjlb
Abstract
Living systems display organized, context-dependent dynamics through a restricted repertoire of persistent functional states. At mesoscopic scales, where biological motion is classical, dissipative, and noisy, such discreteness cannot be attributed to microscopic quantum effects alone. Here, we formulate a mesoscopic theory of functional closure for thermodynamically open episodes in which regulatory organization confines the dynamics to an approximately invariant informational class. Within a constrained open Hamilton-Jacobi framework, exact dissipative composability requires the complete irreversible realization of a biological act—including bulk, interfacial, regulatory, composite, and unresolved internal contributions—to admit an exhaustive, nonredundant thermodynamic partition. In an approximately isothermal regime, causal accumulation of the total irreversible power defines a unique positive dissipative-action scale, . This act-dependent scale sets the resolution of distinguishable functional realizations, while closure defines the domain on which a global additive action invariant can be constructed. If the nontrivial closure spectrum has a least positive action scale, its admissible values form an exact lattice. Quantization in biological dissipative-action units occurs only when this primitive closure scale saturates the independently constructed thermodynamic resolution, so that the spectrum is resolved in integer multiples of . Coherent functional modes are the closure-compatible, persistent, and action-resolved realizations of these discrete classes. The theory predicts clustering of completed trajectories in actionlike observables and separates state correlation, hidden dissipation, and information-geometric distinguishability from the exact composability of irreversible physical cost.
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References (48)
- S. Huang, G. Eichler, Y. Bar-Yam, and D. E. Ingber, Cell fates as high-dimensional attractor states of a complex gene regulatory network, Phys. Rev. Lett. 94, 128701 (2005).
- J. Wang, K. Zhang, L. Xu, and E. Wang, Quantifying the Waddington landscape and biological paths for development and differentiation, Proc. Natl. Acad. Sci. USA 108, 8257 (2011).
- L. H. Hartwell, J. J. Hopfield, S. Leibler, and A. W. Murray, From molecular to modular cell biology, Nature (London) 402, C47 (1999).
- R. Milo, S. Shen-Orr, S. Itzkovitz, N. Kashtan, D. Chklovskii, and U. Alon, Network motifs: Simple building blocks of complex networks, Science 298, 824 (2002).
- U. Alon, Network motifs: Theory and experimental approaches, Nat. Rev. Genet. 8, 450 (2007).
- M. Mossio and A. Moreno, Organisational closure in biological organisms, Hist. Philos. Life Sci. 32, 269 (2010).
- M. Montévil and M. Mossio, Biological organisation as closure of constraints, J. Theor. Biol. 372, 179 (2015).
- G. Engel, T. R. Calhoun, E. L. Read, T. K. Ahn, T. Mancal, Y. C. Cheng, R. E. Blankenship, and G. R. Fleming, Evidence for wavelike energy transfer through quantum coherence in photosynthetic systems, Nature (London) 446, 782 (2007).
- A. Chenu and G. D. Scholes, Coherence in energy transfer and photosynthesis, Annu. Rev. Phys. Chem. 66, 69 (2015).
- N. Lambert, Y.-N. Chen, Y.-C. Cheng, C.-M. Li, G.-Y. Chen, and F. Nori, Quantum biology, Nat. Phys. 9, 10 (2013).
- S. F. Huelga and M. B. Plenio, Vibrations, quanta and biology, Contemp. Phys. 54, 181 (2013).
- J. P. Klinman, Linking protein structure and dynamics to catalysis: The role of hydrogen tunnelling, Philos. Trans. R. Soc. B 361, 1323 (2006).
- L. Masgrau, A. Roujeinikova, L. O. Johannissen, P. Hothi, J. Basran, K. E. Ranaghan, A. J. Mulholland, M. J. Sutcliffe, N. S. Scrutton, and D. Leys, Atomic description of an enzyme reaction dominated by proton tunneling, Science 312, 237 (2006).
- I. Prigogine and G. Nicolis, Self-Organization in Nonequilibrium Systems: From Dissipative Structures to Order through Fluctuations (Wiley, New York, 1977).
- I. Prigogine and I. Stengers, Order Out of Chaos: Man's New Dialogue with Nature (Bantam Books, New York, 1984).
- U. Seifert, Stochastic thermodynamics, fluctuation theorems and molecular machines, Rep. Prog. Phys. 75, 126001 (2012).
- C. Nardini, É. Fodor, E. Tjhung, F. van Wijland, J. Tailleur, and M. E. Cates, Entropy production in field theories without time-reversal symmetry: Quantifying the non-equilibrium character of active matter, Phys. Rev. X 7, 021007 (2017).
- D. J. Skinner and J. Dunkel, Improved bounds on entropy production in living systems, Proc. Natl. Acad. Sci. USA 118, e2024300118 (2021).
- I. Di Terlizzi, M. Gironella, D. Herraez-Aguilar, T. Betz, F. Monroy, M. Baiesi, and F. Ritort, Variance sum rule for entropy production, Science 383, 971 (2024).
- C. Bechinger, R. Di Leonardo, H. Löwen, C. Reichhardt, G. Volpe, and G. Volpe, Active particles in complex and crowded environments, Rev. Mod. Phys. 88, 045006 (2016).
- F. S. Gnesotto, F. Mura, J. Gladrow, and C. P. Broedersz, Broken detailed balance and non-equilibrium dynamics in living systems: A review, Rep. Prog. Phys. 81, 066601 (2018).
- H. Maturana and F. Varela, Autopoiesis and Cognition: The Realization of the Living (D. Reidel, Dordrecht, 1972).
- D. C. Krakauer, N. Bertschinger, E. Olbrich, J. C. Flack, and N. Ay, The information theory of individuality, Theory Biosci. 139, 209 (2020).
- M. Esposito, Stochastic thermodynamics under coarse graining, Phys. Rev. E 85, 041125 (2012).
- K. Kawaguchi and Y. Nakayama, Fluctuation theorem for hidden entropy production, Phys. Rev. E 88, 022147 (2013).
- D. M. Busiello, M. Ciarchi, and I. Di Terlizzi, Unraveling active baths through their hidden degrees of freedom, Phys. Rev. Res. 6, 013190 (2024).
- R. Bebon, J. F. Robinson, and T. Speck, Thermodynamics of active matter: Tracking dissipation across scales, Phys. Rev. X 15, 021050 (2025).
- L. Bertini, A. De Sole, D. Gabrielli, G. Jona-Lasinio, and C. Landim, Macroscopic fluctuation theory, Rev. Mod. Phys. 87, 593 (2015).
- S. Ito and A. Dechant, Stochastic time evolution, information geometry, and the Cramér–Rao bound, Phys. Rev. X 10, 021056 (2020).
- P. B. Melo, S. M. D. Queirós, and W. A. M. Morgado, Stochastic thermodynamics of Fisher information, Phys. Rev. E 111, 014101 (2025).
- P. B. Melo, F. Iemini, D. O. Soares-Pinto, S. M. D. Queirós, and W. A. M. Morgado, Thermodynamic interpretation to stochastic Fisher information and single-trajectory speed limits, Phys. Rev. E 112, 014126 (2025).
- See Supplemental Material at http://link.aps.org/supplemental/10.1103/x2fd-pjlb for the complete variational derivation, thermodynamic construction, treatment of hidden dissipation, closure-lattice theorem, and modal-coherence and breakdown criteria.
- F. Jacob and J. Monod, Genetic regulatory mechanisms in the synthesis of proteins, J. Mol. Biol. 3, 318 (1961).
- J. Monod, Chance and Necessity: An Essay on the Natural Philosophy of Modern Biology (Alfred A. Knopf, New York, 1971).
- V. I. Arnold, Mathematical Methods of Classical Mechanics, 2nd ed., Graduate Texts in Mathematics, Vol. 60 (Springer, New York, 1989)
- L. D. Landau and E. M. Lifshitz, Mechanics, 3rd ed. (Pergamon Press, Oxford, 1976).
- C. Lanczos, The Variational Principles of Mechanics, 4th ed. (Dover Publications, New York, 1986), reprint of the 1970 University of Toronto Press edition.
- J. Paulsson, Summing up the noise in gene networks, Nature (London) 427, 415 (2004).
- M. J. Dunlop, R. Sidney Cox III, J. H. Levine, R. M. Murray, and M. B. Elowitz, Regulatory activity revealed by dynamic correlations in gene expression noise, Nat. Genet. 40, 1493 (2008).
- N. Bain and D. Bartolo, Critical mingling and universal correlations in model binary active liquids, Nat. Commun. 8, 15969 (2017).
- U. M. B. Marconi, A. Puglisi, L. Rondoni, and A. Vulpiani, Fluctuation-dissipation: Response theory in statistical physics, Phys. Rep. 461, 111 (2008).
- L. F. Cugliandolo, The effective temperature, J. Phys. A: Math. Theor. 44, 483001 (2011).
- A. Puglisi, A. Sarracino, and A. Vulpiani, Temperature in and out of equilibrium: A review of concepts, tools and attempts, Phys. Rep. 709–710, 1 (2017).
- J. R. Medeiros and S. M. D. Queirós, Effective temperatures for single particle system under dichotomous noise, J. Stat. Mech. (2021) 063205.
- E. Madelung, Quantentheorie in hydrodynamischer form, Z. Phys. 40, 322 (1927).
- L. D. Landau and E. M. Lifshitz, Quantum Mechanics: Non-Relativistic Theory, 3rd ed., Course of Theoretical Physics, Vol. 3 (Pergamon Press, Oxford, 1977).
- D. Bohm, A suggested interpretation of the quantum theory in terms of “hidden” variables. I, Phys. Rev., 85, 166 (1952).
- D. Bohm, A suggested interpretation of the quantum theory in terms of “hidden” variables. II, Phys. Rev. 85, 180 (1952).