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  • Open Access

Mutual Linearity Is a Generic Property of Steady-State Markov Networks

Robin Bebon and Thomas Speck*

  • *Contact author: thomas.speck@itp4.uni-stuttgart.de

Phys. Rev. Lett. 136, 137401 – Published 31 March, 2026

DOI: https://doi.org/10.1103/jcm3-57d8

Abstract

Understanding and predicting how complex systems respond to external perturbations is a central challenge in nonequilibrium statistical physics. Here, we consider continuous-time Markov networks, which we subject to perturbations along a single edge. We find that in steady state the probabilities of any two states are linearly related to one another. We show that this mutual linearity of probabilities extends to a broad class of observables, including currents but also generic counting and state-dependent observables. Moreover, we derive an exact relation between the relative response of any state’s probability and the ratio of two steady-state probabilities. Leveraging the Markov chain tree theorem, we further show that probabilities and the considered observables are constrained by the topological and kinetic properties of the network and provide analytical expressions in terms of spanning tree polynomials. Our results are general, holding for arbitrary rate parametrizations and extending far from equilibrium.

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References (95)

  1. R. Kubo, Statistical-mechanical theory of irreversible processes. I. General theory and simple applications to magnetic and conduction problems, J. Phys. Soc. Jpn. 12, 570 (1957).
  2. P. Hänggi and H. Thomas, Stochastic processes: Time evolution, symmetries and linear response, Phys. Rep. 88, 207 (1982).
  3. S. Baroni, P. Giannozzi, and A. Testa, Green’s-function approach to linear response in solids, Phys. Rev. Lett. 58, 1861 (1987).
  4. R. Kubo, The fluctuation-dissipation theorem, Rep. Prog. Phys. 29, 255 (1966).
  5. U. Seifert, Stochastic thermodynamics, fluctuation theorems and molecular machines, Rep. Prog. Phys. 75, 126001 (2012).
  6. L. Peliti and S. Pigolotti, Stochastic Thermodynamics: An Introduction (Princeton University Press, Princeton: Oxford, 2021).
  7. U. Seifert, Stochastic Thermodynamics, 1st ed. (Cambridge University Press, Cambridge, England, 2025).
  8. R. Rao and M. Esposito, Nonequilibrium thermodynamics of chemical reaction networks: Wisdom from stochastic thermodynamics, Phys. Rev. X 6, 041064 (2016).
  9. P. Gaspard, Fluctuation theorem for nonequilibrium reactions, J. Chem. Phys. 120, 8898 (2004).
  10. D. A. McQuarrie, Stochastic approach to chemical kinetics, J. Appl. Probab. 4, 413 (1967).
  11. C. Y. Mou, J.-l. Luo, and G. Nicolis, Stochastic thermodynamics of nonequilibrium steady states in chemical reaction systems, J. Chem. Phys. 84, 7011 (1986).
  12. T. Schmiedl, T. Speck, and U. Seifert, Entropy production for mechanically or chemically driven biomolecules, J. Stat. Phys. 128, 77 (2007).
  13. S. Dal Cengio, D. Levis, and I. Pagonabarraga, Linear response theory and Green-Kubo relations for active matter, Phys. Rev. Lett. 123, 238003 (2019).
  14. L. Caprini, A. Puglisi, and A. Sarracino, Fluctuation–dissipation relations in active matter systems, Symmetry 13, 81 (2021).
  15. R. Bebon, J. F. Robinson, and T. Speck, Thermodynamics of active matter: Tracking dissipation across scales, Phys. Rev. X 15, 021050 (2025).
  16. T. R. Kirkpatrick and D. Belitz, Fluctuation-response relation in nonequilibrium systems and active matter, Phys. Rev. E 111, 014102 (2025).
  17. L. K. Davis, K. Proesmans, and É. Fodor, Active matter under control: Insights from response theory, Phys. Rev. X 14, 011012 (2024).
  18. G. S. Agarwal, Fluctuation-dissipation theorems for systems in non-thermal equilibrium and applications, Z. Phys. 252, 25 (1972).
  19. T. Speck and U. Seifert, Restoring a fluctuation-dissipation theorem in a nonequilibrium steady state, Europhys. Lett. 74, 391 (2006).
  20. V. Blickle, T. Speck, C. Lutz, U. Seifert, and C. Bechinger, Einstein relation generalized to nonequilibrium, Phys. Rev. Lett. 98, 210601 (2007).
  21. T. Speck and U. Seifert, Extended fluctuation-dissipation theorem for soft matter in stationary flow, Phys. Rev. E 79, 040102(R) (2009).
  22. U. Seifert, Generalized Einstein or Green-Kubo relations for active biomolecular transport, Phys. Rev. Lett. 104, 138101 (2010).
  23. U. Seifert and T. Speck, Fluctuation-dissipation theorem in nonequilibrium steady states, Europhys. Lett. 89, 10007 (2010).
  24. J. Prost, J.-F. Joanny, and J. M. R. Parrondo, Generalized fluctuation-dissipation theorem for steady-state systems, Phys. Rev. Lett. 103, 090601 (2009).
  25. B. Altaner, M. Polettini, and M. Esposito, Fluctuation-dissipation relations far from equilibrium, Phys. Rev. Lett. 117, 180601 (2016).
  26. M. Baiesi, C. Maes, and B. Wynants, Fluctuations and response of nonequilibrium states, Phys. Rev. Lett. 103, 010602 (2009).
  27. A. Dechant and S.-i. Sasa, Fluctuation–response inequality out of equilibrium, Proc. Natl. Acad. Sci. U.S.A. 117, 6430 (2020).
  28. T. Aslyamov, K. Ptaszyński, and M. Esposito, Nonequilibrium fluctuation-response relations: From identities to bounds, Phys. Rev. Lett. 134, 157101 (2025).
  29. K. Ptaszynski, T. Aslyamov, and M. Esposito, Nonequilibrium fluctuation-response relations for state observables, Phys. Rev. E 113, 024130 (2026).
  30. H.-M. Chun, Q. Gao, and J. M. Horowitz, Nonequilibrium Green-Kubo relations for hydrodynamic transport from an equilibrium-like fluctuation-response equality, Phys. Rev. Res. 3, 043172 (2021).
  31. J. Zheng and Z. Lu, Universal Non-equilibrium response theory beyond steady states, arXiv:2403.10952.
  32. A. C. Barato and U. Seifert, Thermodynamic uncertainty relation for biomolecular processes, Phys. Rev. Lett. 114, 158101 (2015).
  33. J. M. Horowitz and T. R. Gingrich, Proof of the finite-time thermodynamic uncertainty relation for steady-state currents, Phys. Rev. E 96, 020103(R) (2017).
  34. J. M. Horowitz and T. R. Gingrich, Thermodynamic uncertainty relations constrain non-equilibrium fluctuations, Nat. Phys. 16, 15 (2020).
  35. C. Dieball and A. Godec, Direct route to thermodynamic uncertainty relations and their saturation, Phys. Rev. Lett. 130, 087101 (2023).
  36. J. A. Owen, T. R. Gingrich, and J. M. Horowitz, Universal thermodynamic bounds on nonequilibrium response with biochemical applications, Phys. Rev. X 10, 011066 (2020).
  37. G. Fernandes Martins and J. M. Horowitz, Topologically constrained fluctuations and thermodynamics regulate nonequilibrium response, Phys. Rev. E 108, 044113 (2023).
  38. K. Ptaszyński, T. Aslyamov, and M. Esposito, Dissipation bounds precision of current response to kinetic perturbations, Phys. Rev. Lett. 133, 227101 (2024).
  39. K. Ptaszynski, T. Aslyamov, and M. Esposito, Nonequilibrium fluctuation-response relations for state-current correlations, Phys. Rev. E 113, 024131 (2026).
  40. T. Aslyamov and M. Esposito, Nonequilibrium response for Markov jump processes: Exact results and tight bounds, Phys. Rev. Lett. 132, 037101 (2024).
  41. J. A. Owen and J. M. Horowitz, Size limits the sensitivity of kinetic schemes, Nat. Commun. 14, 1280 (2023).
  42. P. E. Harunari, S. Dal Cengio, V. Lecomte, and M. Polettini, Mutual linearity of nonequilibrium network currents, Phys. Rev. Lett. 133, 047401 (2024).
  43. F. Khodabandehlou, C. Maes, and K. Netočný, Affine relationships between steady currents, J. Phys. A 58, 155002 (2025).
  44. S. Dal Cengio, P. E. Harunari, V. Lecomte, and M. Polettini, Mutual multilinearity of nonequilibrium network currents, SciPost Phys. 19, 111 (2025).
  45. S. M. Ross, Introduction to Probability Models, 11th ed. (Academic Press is an imprint of Elsevier, Amsterdam, 2014).
  46. T. Aslyamov and M. Esposito, General theory of static response for Markov jump processes, Phys. Rev. Lett. 133, 107103 (2024).
  47. Probability conservation enforces ∑n=1|N|pn,m(0)=1 and ∑n=1|N|χn,m=0.

  48. S. Liang, P. De Los Rios, and D. M. Busiello, Thermodynamic bounds on symmetry breaking in linear and catalytic biochemical systems, Phys. Rev. Lett. 132, 228402 (2024).
  49. See Supplemental Material at http://link.aps.org/supplemental/10.1103/jcm3-57d8 for additional results, proofs, and details on the examples, which includes Refs. [50–52].
  50. C. Maes and K. Netočný, Heat bounds and the blowtorch theorem, Ann. Henri Poincaré 14, 1193 (2013).
  51. U. Çetiner and J. Gunawardena, Reformulating nonequilibrium steady states and generalized Hopfield discrimination, Phys. Rev. E 106, 064128 (2022).
  52. J. D. Mallory, A. B. Kolomeisky, and O. A. Igoshin, Kinetic control of stationary flux ratios for a wide range of biochemical processes, Proc. Natl. Acad. Sci. U.S.A. 117, 8884 (2020).
  53. T. L. Hill, Studies in irreversible thermodynamics IV. Diagrammatic representation of steady state fluxes for unimolecular systems, J. Theor. Biol. 10, 442 (1966).
  54. J. Schnakenberg, Network theory of microscopic and macroscopic behavior of master equation systems, Rev. Mod. Phys. 48, 571 (1976).
  55. W. T. Tutte and C. S. J. A. Nash-Williams, Graph Theory, Encyclopedia of Mathematics and Its Applications Vol. 21 (Cambridge University Press, Cambridge, England, 2005).
  56. Rooted trees Tnμ are defined as directed subgraphs containing all nodes and no cycles, with every node—except the root n—having outdegree 1.

  57. For a concrete example, consider Fig. 1. Here, the upper B2+ (B4−) and lower B2− (B4+) bounds of p2 and p4 are mirrored due to their different scalings with k+I [cf. Fig. 1].

  58. M. Merolle, J. P. Garrahan, and D. Chandler, Space–time thermodynamics of the glass transition, Proc. Natl. Acad. Sci. U.S.A. 102, 10837 (2005).
  59. C. Maes, Non-Dissipative Effects in Nonequilibrium Systems (Springer, Berlin Heidelberg, New York, 2017).
  60. C. Maes, Frenetic bounds on the entropy production, Phys. Rev. Lett. 119, 160601 (2017).
  61. I. Di Terlizzi and M. Baiesi, Kinetic uncertainty relation, J. Phys. A 52, 02LT03 (2019).
  62. C. Maes, Frenesy: Time-symmetric dynamical activity in nonequilibria, Phys. Rep. 850, 1 (2020).
  63. One example is currents through bridges, which are zero regardless of input rates.

  64. Y. Tu, The nonequilibrium mechanism for ultrasensitivity in a biological switch: Sensing by Maxwell’s demons, Proc. Natl. Acad. Sci. U.S.A. 105, 11737 (2008).
  65. G. Lan, P. Sartori, S. Neumann, V. Sourjik, and Y. Tu, The energy–speed–accuracy trade-off in sensory adaptation, Nat. Phys. 8, 422 (2012).
  66. A. Murugan and S. Vaikuntanathan, Topologically protected modes in non-equilibrium stochastic systems, Nat. Commun. 8, 13881 (2017).
  67. N. Barkai and S. Leibler, Robustness in simple biochemical networks, Nature (London) 387, 913 (1997).
  68. Y. Tu, Quantitative modeling of bacterial chemotaxis: Signal amplification and accurate adaptation, Annu. Rev. Biophys. 42, 337 (2013).
  69. R. G. Endres and N. S. Wingreen, Precise adaptation in bacterial chemotaxis through “assistance neighborhoods,” Proc. Natl. Acad. Sci. U.S.A. 103, 13040 (2006).
  70. Y. V. Kalinin, L. Jiang, Y. Tu, and M. Wu, Logarithmic sensing in Escherichia Coli bacterial chemotaxis, Biophys. J. 96, 2439 (2009).
  71. M. Skoge, S. Naqvi, Y. Meir, and N. S. Wingreen, Chemical sensing by nonequilibrium cooperative receptors, Phys. Rev. Lett. 110, 248102 (2013).
  72. V. Kharbanda and B. Sabass, Sensory adaptation in a continuum model of bacterial chemotaxis—working range, cost-accuracy relation, and coupled systems, New J. Phys. 26, 023045 (2024).
  73. P. Sartori and Y. Tu, Free energy cost of reducing noise while maintaining a high sensitivity, Phys. Rev. Lett. 115, 118102 (2015).
  74. C. Tietz, S. Schuler, T. Speck, U. Seifert, and J. Wrachtrup, Measurement of stochastic entropy production, Phys. Rev. Lett. 97, 050602 (2006).
  75. H. S. Chung, K. McHale, J. M. Louis, and W. A. Eaton, Single-molecule fluorescence experiments determine protein folding transition path times, Science 335, 981 (2012).
  76. H. S. Chung and W. A. Eaton, Single-molecule fluorescence probes dynamics of barrier crossing, Nature (London) 502, 685 (2013).
  77. J. Stigler, F. Ziegler, A. Gieseke, J. C. M. Gebhardt, and M. Rief, The complex folding network of single calmodulin molecules, Science 334, 512 (2011).
  78. R. Petrosyan, A. Narayan, and M. T. Woodside, Single-molecule force spectroscopy of protein folding, J. Mol. Biol. 433, 167207 (2021).
  79. W. Ye, M. Götz, S. Celiksoy, L. Tüting, C. Ratzke, J. Prasad, J. Ricken, S. V. Wegner, R. Ahijado-Guzmán, T. Hugel, and C. Sönnichsen, Conformational dynamics of a single protein monitored for 24 h at video rate, Nano Lett. 18, 6633 (2018).
  80. L. Vollmar, R. Bebon, J. Schimpf, B. Flietel, S. Celiksoy, C. Sönnichsen, A. Godec, and T. Hugel, Model-free inference of memory in conformational dynamics of a multi-domain protein, J. Phys. A 57, 365001 (2024).
  81. S. Arrhenius, über die Reaktionsgeschwindigkeit bei der Inversion von Rohrzucker durch Säuren, Z. Phys. Chem. 4U, 226 (1889).
  82. H. Eyring, The activated complex in chemical reactions, J. Chem. Phys. 3, 107 (1935).
  83. P. Clifford and A. Sudbury, A model for spatial conflict, Biometrika 60, 581 (1973).
  84. V. Sood and S. Redner, Voter model on heterogeneous graphs, Phys. Rev. Lett. 94, 178701 (2005).
  85. M. Ángeles Serrano, K. Klemm, F. Vazquez, V. M. Eguíluz, and M. San Miguel, Conservation laws for voter-like models on random directed networks, J. Stat. Mech. (2009) P10024.
  86. N. Masuda and H. Ohtsuki, Evolutionary dynamics and fixation probabilities in directed networks, New J. Phys. 11, 033012 (2009).
  87. C. Floyd, A. R. Dinner, S. Vaikuntanathan, Local imperfect feedback control in non-equilibrium biophysical systems enabled by thermodynamic constraints, arXiv:2507.07295.
  88. R. Bebon and T. Speck, Supporting data for “Mutual Linearity is a Generic Property of Steady-State Markov Networks” (2026), 10.18419/DARUS-5742.
  89. M. R. Evans and S. N. Majumdar, Diffusion with stochastic resetting, Phys. Rev. Lett. 106, 160601 (2011).
  90. J. Fuchs, S. Goldt, and U. Seifert, Stochastic thermodynamics of resetting, Europhys. Lett. 113, 60009 (2016).
  91. A. Pal and S. Reuveni, First passage under restart, Phys. Rev. Lett. 118, 030603 (2017).
  92. M. R. Evans, S. N. Majumdar, and G. Schehr, Stochastic resetting and applications, J. Phys. A 53, 193001 (2020).
  93. R. Bebon and A. Godec, Controlling uncertainty of empirical first-passage times in the small-sample regime, Phys. Rev. Lett. 131, 237101 (2023).
  94. A. Pal, S. Reuveni, and S. Rahav, Thermodynamic uncertainty relation for first-passage times on Markov chains, Phys. Rev. Res. 3, L032034 (2021).
  95. A. Pal, S. Reuveni, and S. Rahav, Thermodynamic uncertainty relation for systems with unidirectional transitions, Phys. Rev. Res. 3, 013273 (2021).

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