- Open Access
Alternating pattern formation in oscillatory media with spatially uniform and nonuniform global feedback
Phys. Rev. E 114, 014211 – Published 14 July, 2026
DOI: https://doi.org/10.1103/r3d8-2p3x
Abstract
Alternating (temporally period-2) stripes, clusters, and labyrinths have been observed in chemical and biochemical reactions, in Min-protein dynamics in E. coli and in vitro systems, and in calcium alternans in cardiac myocytes. However, the mechanisms underlying the formation of these spatiotemporal patterns and the transitions among them remain incompletely understood. In this article, we use an amplitude-equation (AE) approach to perform theoretical analysis and numerical simulations to elucidate the mechanisms of the alternating pattern formation and selection, and we then use reaction-diffusion models to validate the predictions of the AE model. It was shown previously that in chemical reactions and cardiac myocytes, a global feedback loop plays a key role in the formation of alternating clusters. Based on previous AE models developed to describe cardiac alternans dynamics, we first develop an AE model that incorporates either a spatially uniform global feedback (UGF) loop or a spatially nonuniform global feedback (NUGF) loop. The NUGF is represented by a spatially Gaussian-weighted function. We then use the AE model to conduct nonlinear stability analysis to reveal the mechanisms underlying the emergence of alternating patterns under different feedback conditions. Under positive UGF, no patterns can form. When the UGF is negative, the stable patterns are only multiple stripes and circular clusters. For stripe patterns, they may be spatially periodic or random as long as they satisfy the requirements for pattern selection. For circular clusters, a medium can support one or multiple circular alternating clusters of the same radius depending on the size of the medium. Under NUGF, in addition to stripe and circular-cluster patterns, lateral instability arises when the width of the Gaussian function falls within an intermediate range, leading to transitions from stripes or circular clusters to labyrinths. Other initial-condition-dependent patterns are also observed under NUGF. Under UGF, the width of the stripe can be periodic or random, but under NUGF, the patterns become less random in space as the feedback strength increases. Finally, we incorporate the UGF and NUGF loops into several reaction-diffusion models, including the Belousov-Zhabotinsky reaction model, the carbon monoxide oxidation model, and the periodically paced FitzHugh-Nagumo model. These models successfully reproduce the alternating pattern dynamics predicted by the AE model, validating the theoretical predictions.
Physics Subject Headings (PhySH)
Article Text
References (52)
- P. Ball, The Self-Made Tapestry: Pattern Formation in Nature (Oxford University Press, Oxford, 1999).
- M. Cross and H. Greenside, Pattern Formation and Dynamics in Nonequilibrium Systems (Cambridge University Press, Cambridge, 2009).
- A. M. Turing, The chemical basis of morphogenesis, Philos. R. Soc. London B 237, 37 (1952).
- Q. Ouyang and H. L. Swinney, Transition from a uniform state to hexagonal and striped Turing patterns, Nature (London) 352, 610 (1991).
- S. Kondo and T. Miura, Reaction-diffusion model as a framework for understanding biological pattern formation, Science 329, 1616 (2010).
- P. K. Maini, R. E. Baker, and C. M. Chuong, The Turing model comes of molecular age, Science 314, 1397 (2006).
- V. K. Vanag and I. R. Epstein, Pattern formation mechanisms in reaction-diffusion systems, Int. J. Dev. Biol. 53, 673 (2009).
- V. K. Vanag and I. R. Epstein, Pattern formation in a tunable medium: The Belousov-Zhabotinsky reaction in an aerosol OT microemulsion, Phys. Rev. Lett. 87, 228301 (2001).
- L. Yang and I. R. Epstein, Oscillatory turing patterns in reaction-diffusion systems with two coupled layers, Phys. Rev. Lett. 90, 178303 (2003).
- L. Yang, A. M. Zhabotinsky, and I. R. Epstein, Stable squares and other oscillatory turing patterns in a reaction-diffusion model, Phys. Rev. Lett. 92, 198303 (2004).
- V. K. Vanag and I. R. Epstein, Stationary and oscillatory localized patterns, and subcritical bifurcations, Phys. Rev. Lett. 92, 128301 (2004).
- V. K. Vanag, L. Yang, M. Dolnik, A. M. Zhabotinsky, and I. R. Epstein, Oscillatory cluster patterns in a homogeneous chemical system with global feedback, Nature (London) 406, 389 (2000).
- V. K. Vanag, A. M. Zhabotinsky, and I. R. Epstein, pattern formation in the Belousov−Zhabotinsky reaction with photochemical global feedback, J. Phys. Chem. A 104, 11566 (2000).
- M. Kim, M. Bertram, M. Pollmann, A. v. Oertzen, A. S. Mikhailov, H. H. Rotermund, and G. Ertl, Controlling chemical turbulence by global delayed feedback: Pattern formation in catalytic CO oxidation on Pt(110), Science 292, 1357 (2001).
- V. Petrov, Q. Ouyang, and H. L. Swinney, Resonant pattern formation in a chemical system, Nature (London) 388, 655 (1997).
- A. L. Lin, M. Bertram, K. Martinez, H. L. Swinney, A. Ardelea, and G. F. Carey, Resonant phase patterns in a reaction-diffusion system, Phys. Rev. Lett. 84, 4240 (2000).
- S. Kretschmer, T. Heermann, A. Tassinari, P. Glock, and P. Schwille, Increasing MinD's membrane affinity yields standing wave oscillations and functional gradients on flat membranes, ACS Synth. Biol. 10, 939 (2021).
- A. Kaminaga, V. K. Vanag, and I. R. Epstein, Wavelength halving in a transition between standing waves and traveling waves, Phys. Rev. Lett. 95, 058302 (2005).
- Z. Qu and J. N. Weiss, Cardiac alternans: From bedside to bench and back, Circ. Res. 132, 127 (2023).
- J. Kockskamper and L. A. Blatter, Subcellular alternans represents a novel mechanism for the generation of arrhythmogenic waves in cat atrial myocytes, J. Physiol. 545, 65 (2002).
- M. E. Diaz, D. A. Eisner, and S. C. O'Neill, Depressed ryanodine receptor activity increases variability and duration of the systolic transient in rat ventricular myocytes, Circ. Res. 91, 585 (2002).
- S. A. Gaeta, G. Bub, G. W. Abbott, and D. J. Christini, Dynamical mechanism for subcellular alternans in cardiac myocytes, Circ. Res. 105, 335 (2009).
- G. L. Aistrup, Y. Shiferaw, S. Kapur, A. H. Kadish, and J. A. Wasserstrom, Mechanisms underlying the formation and dynamics of subcellular calcium alternans in the intact rat heart, Circ. Res. 104, 639 (2009).
- L. H. Xie and J. N. Weiss, Arrhythmogenic consequences of intracellular calcium waves, Am. J. Physiol. Heart Circ. Physiol. 297, H997 (2009).
- D. S. Rosenbaum, L. E. Jackson, J. M. Smith, H. Garan, J. N. Ruskin, and R. J. Cohen, Electrical alternans and vulnerability to ventricular arrhythmias, N. Engl. J. Med. 330, 235 (1994).
- J. N. Weiss, A. Karma, Y. Shiferaw, P. S. Chen, A. Garfinkel, and Z. Qu, From pulsus to pulseless: The saga of cardiac alternans, Circ. Res. 98, 1244 (2006).
- D. M. Raskin and P. A. J. de Boer, Rapid pole-to-pole oscillation of a protein required for directing division to the middle of Escherichia coli, Proc. Natl. Acad. Sci. USA 96, 4971 (1999).
- X. Fu, Y.-L. Shih, Y. Zhang, and L. I. Rothfield, The MinE ring required for proper placement of the division site is a mobile structure that changes its cellular location during the Escherichia coli division cycle, Proc. Natl. Acad. Sci. USA 98, 980 (2001).
- C. A. Hale, H. Meinhardt, and P. A. J. de Boer, Dynamic localization cycle of the cell division regulator MinE in Escherichia coli, EMBO J. 20, 1563 (2001).
- R. A. Kerr, H. Levine, T. J. Sejnowski, and W.-J. Rappel, Division accuracy in a stochastic model of Min oscillations in Escherichia coli, Proc. Natl. Acad. Sci. USA 103, 347 (2006).
- A. Amiranashvili, N. D. Schnellbächer, and U. S. Schwarz, Stochastic switching between multistable oscillation patterns of the Min-system, New J. Phys. 18, 093049 (2016).
- D. Fange and J. Elf, Noise-induced Min phenotypes in E. coli, PLoS Comput. Biol. 2, e80 (2006).
- H. Meinhardt and P. A. J. de Boer, Pattern formation in Escherichia coli: A model for the pole-to-pole oscillations of Min proteins and the localization of the division site, Proc. Natl. Acad. Sci. USA 98, 14202 (2001).
- K. Zieske and P. Schwille, in Methods in Cell Biology, edited by J. Ross and W. F. Marshall, (Academic, 2015), Vol. 128, p. 149.
- M. Bonny, E. Fischer-Friedrich, M. Loose, P. Schwille, and K. Kruse, Membrane binding of MinE allows for a comprehensive description of min-protein pattern formation, PLoS Comput. Biol. 9, e1003347 (2013).
- L. Wettmann and K. Kruse, The Min-protein oscillations in Escherichia coli: An example of self-organized cellular protein waves, Philos. Trans. R. Soc. London B Biol. Sci. 373, 20170111 (2018).
- Z. Ren, H. Weyer, M. Sandler, et al., Robust and resource-optimal dynamic pattern formation of Min proteins in vivo, Nat. Phys. 21, 1160 (2025).
- K. Zieske and P. Schwille, Reconstitution of self-organizing protein gradients as spatial cues in cell-free systems, eLife 3, e03949 (2014).
- M. Falcke and H. Engel, Pattern formation during the CO oxidation on Pt(110) surfaces under global coupling, J. Chem. Phys. 101, 6255 (1994).
- M. Falcke, H. Engel, and M. Neufeld, Cluster formation, standing waves, and stripe patterns in oscillatory active media with local and global coupling, Phys. Rev. E 52, 763 (1995).
- L. Yang, M. Dolnik, A. M. Zhabotinsky, and I. R. Epstein, Oscillatory clusters in a model of the photosensitive Belousov-Zhabotinsky reaction system with global feedback, Phys. Rev. E 62, 6414 (2000).
- M. Bertram and A. S. Mikhailov, Pattern formation in a surface chemical reaction with global delayed feedback, Phys. Rev. E 63, 066102 (2001).
- Y. Shiferaw and A. Karma, Turing instability mediated by voltage and calcium diffusion in paced cardiac cells, Proc. Natl. Acad. Sci. USA 103, 5670 (2006).
- Z. Song and Z. Qu, Delayed global feedback in the genesis and stability of spatiotemporal excitation patterns in paced biological excitable media, PLoS Comput. Biol. 16, e1007931 (2020).
- C. Huang, Z. Song, and Z. Qu, Transition from alternating stripes to alternating labyrinths in oscillatory media, Phys. Rev. E 111, L032201 (2025).
- B. Echebarria and A. Karma, Instability and spatiotemporal dynamics of alternans in paced cardiac tissue, Phys. Rev. Lett. 88, 208101 (2002).
- B. Echebarria and A. Karma, Amplitude equation approach to spatiotemporal dynamics of cardiac alternans, Phys. Rev. E 76, 051911 (2007).
- C. Huang, Z. Song, Z. Di, and Z. Qu, Stability of spatially discordant repolarization alternans in cardiac tissue, Chaos 30, 123141 (2020).
- C. Huang, Z. Song, and Z. Qu, Synchronization of spatially discordant voltage and calcium alternans in cardiac tissue, Phys. Rev. E 106, 024406 (2022).
- J. P. Keener and J. Sneyd, Mathematical Physiology (Springer, New York, 1998).
- M. Loose, E. Fischer-Friedrich, J. Ries, K. Kruse, and P. Schwille, Spatial regulators for bacterial cell division self-organize into surface waves in vitro, Science 320, 789 (2008).
- B. Ramm, A. Goychuk, A. Khmelinskaia, P. Blumhardt, H. Eto, K. A. Ganzinger, E. Frey, and P. Schwille, A diffusiophoretic mechanism for ATP-driven transport without motor proteins, Nat. Phys. 17, 850 (2021).