Geometric Singular Perturbation Theory Research Articles

Mixed-mode oscillations for slow-fast perturbed systems

This article concerns the dynamics of mixed-mode oscillations (MMOs) emerging from the calcium-based inner hair cells (IHCs) model in the auditory cortex. The paper captures the MMOs generation mechanism based on the geometric singular perturbation theory (GSPT) after exploiting the average analysis for reducing the full model. Our analysis also finds that the critical manifold and folded surface are central to the mechanism of the existence of MMOs at the folded saddle for the perturbed system. The system parameters, such like the maximal calcium channels conductance, controls the firing patterns, and many new oscillations occur for the IHCs model. Tentatively, we conduct dynamic analysis combined with dynamic method based on GSPT by giving slow-fast analysis for the singular perturbed models and bifurcation analysis. In particular, we explore the two-slow-two-fast and three-slow-one-fast IHCs perturbed systems with layer and reduced problems so that differential-algebraic equations are obtained. This paper reveals the underlying dynamic properties of perturbed systems under singular perturbation theory.

On the Validity of the Stochastic Quasi-Steady-State Approximation in Open Enzyme Catalyzed Reactions: Timescale Separation or Singular Perturbation?

The quasi-steady-state approximation is widely used to develop simplified deterministic or stochastic models of enzyme catalyzed reactions. In deterministic models, the quasi-steady-state approximation can be mathematically justified from singular perturbation theory. For several closed enzymatic reactions, the homologous extension of the quasi-steady-state approximation to the stochastic regime, known as the stochastic quasi-steady-state approximation, has been shown to be accurate under the analogous conditions that permit the quasi-steady-state reduction in the deterministic counterpart. However, it was recently demonstrated that the extension of the stochastic quasi-steady-state approximation to an open Michaelis-Menten reaction mechanism is only valid under a condition that is far more restrictive than the qualifier that ensures the validity of its corresponding deterministic quasi-steady-state approximation. In this paper, we suggest a possible explanation for this discrepancy from the lens of geometric singular perturbation theory. In so doing, we illustrate a misconception in the application of the quasi-steady-state approximation: timescale separation does not imply singular perturbation.

Traveling waves in a generalized Camassa–Holm equation involving dual-power law nonlinearities

This article focuses on traveling waves including peakons, periodic peakons, compactons and solitary waves as well as the persistence of some of these waves in a generalized Camassa–Holm (gCH) equation involving dual-power law nonlinearities. Firstly, we analyze the global phase portraits of the associated traveling wave system by a dynamical system-based approach. All the traveling waves are classified and their corresponding parameter conditions are obtained. It is found that the gCH equation with dual-power law nonlinearities can admit different peakons under different parameter conditions. Secondly, we pay attention to the persistence of solitary waves, i.e., the existence of solitary waves in a perturbed gCH equation by using geometric singular perturbation theory (GSPT) and an “explicit” Melnikov method. An elementary method to calculate the unperturbed homoclinic orbits and the associated Melnikov integral in an “explicit” way is developed in this article.

Geometric singular perturbation analysis to Camassa-Holm Kuramoto-Sivashinsky equation

We analyze a singularly Kuramoto-Sivashinsky perturbed Camassa-Holm equation with methods of the geometric singular perturbation theory. Especially, we study the persistence of smooth and peaked solitons. Whether a solitary wave of the original Camassa-Holm equation is smooth or peaked depends on whether there is linear dispersion, i.e. whether 2k=0. If 2k>0, then a unique smooth solitary wave persists with selected wave speed under singular Kuramoto-Sivashinsky perturbation just as what happens in the KS-KdV equation. On the other hand, we show that if there is no linear dispersion, i.e. 2k=0, then any observable peaked soliton fails to persist. This case is non-typical since the related slow manifold blows up and the classical geometric singular perturbation theory is not available.

Tipping Cascades in a Multi-patch System with Noise and Spatial Coupling

Forecasting tipping points in spatially extended systems is a key area of interest to ecologists. A slowly declining spatially distributed population is an important example of an ecological system that could exhibit a cascade of tipping points. Here, we develop a spatial two-patch model with environmental stochasticity that is slowly forced through population collapse, in the presence of changing environmental conditions. We begin with a basic spatial model, then introduce a fast–slow version of the model using geometric singular perturbation theory, followed by the inclusion of stochasticity. Using the spectral density of the fluctuating subpopulation in each patch, we derive analytic expressions for candidate indicators of population extinction and evaluate their performance through a simulation study. We find that coupling and spatial heterogeneity decrease the magnitude of the proposed indicators in coupled populations relative to isolated populations. Moreover, the degree of coupling dictates the trends in summary statistics. We conclude that this theory may be applied to other contexts, including the control of invasive species.

Singularly perturbed boundary-equilibrium bifurcations

Boundary equilibria bifurcation (BEB) arises in piecewise-smooth (PWS) systems when an equilibrium collides with a discontinuity set under parameter variation. Singularly perturbed BEB refers to a bifurcation arising in singular perturbation problems which limit as some ϵ → 0 to PWS systems which undergo a BEB. This work completes a classification for codimension-1 singularly perturbed BEB in the plane initiated by the present authors in [], using a combination of tools from PWS theory, geometric singular perturbation theory and a method of geometric desingularization known as blow-up. After deriving a local normal form capable of generating all 12 singularly perturbed BEBs, we describe the unfolding in each case. Detailed quantitative results on saddle-node, Andronov–Hopf, homoclinic and codimension-2 Bogdanov–Takens bifurcations involved in the unfoldings and classification are presented. Each bifurcation is singular in the sense that it occurs within a domain which shrinks to zero as ϵ → 0 at a rate determined by the rate at which the system loses smoothness. Detailed asymptotics for a distinguished homoclinic connection which forms the boundary between two singularly perturbed BEBs in parameter space are also given. Finally, we describe the explosive onset of oscillations arising in the unfolding of a particular singularly perturbed boundary-node bifurcation. We prove the existence of the oscillations as perturbations of PWS cycles, and derive a growth rate which is polynomial in ϵ and dependent on the rate at which the system loses smoothness. For all the results presented herein, corresponding results for regularised PWS systems are obtained via the limit ϵ → 0.

Persistence of solitary wave solutions to a singularly perturbed generalized mKdV equation

The existence of solitary wave solutions for a perturbed generalized mKdV equation with a cubic evolution term is investigated. All possible solitary waves for the corresponding unperturbed equation are firstly explored by dynamical system analysis. Then by using geometric singular perturbation theory and Melnikov’s method, we prove that two solitary wave solutions of the unperturbed generalized mKdV equation with particularly chosen wave speeds will persist under small singular perturbation. The results of numerical simulations are consistent with our theoretical analysis.

Canard solutions in neural mass models: consequences on critical regimes

Mathematical models at multiple temporal and spatial scales can unveil the fundamental mechanisms of critical transitions in brain activities. Neural mass models (NMMs) consider the average temporal dynamics of interconnected neuronal subpopulations without explicitly representing the underlying cellular activity. The mesoscopic level offered by the neural mass formulation has been used to model electroencephalographic (EEG) recordings and to investigate various cerebral mechanisms, such as the generation of physiological and pathological brain activities. In this work, we consider a NMM widely accepted in the context of epilepsy, which includes four interacting neuronal subpopulations with different synaptic kinetics. Due to the resulting three-time-scale structure, the model yields complex oscillations of relaxation and bursting types. By applying the principles of geometric singular perturbation theory, we unveil the existence of the canard solutions and detail how they organize the complex oscillations and excitability properties of the model. In particular, we show that boundaries between pathological epileptic discharges and physiological background activity are determined by the canard solutions. Finally we report the existence of canard-mediated small-amplitude frequency-specific oscillations in simulated local field potentials for decreased inhibition conditions. Interestingly, such oscillations are actually observed in intracerebral EEG signals recorded in epileptic patients during pre-ictal periods, close to seizure onsets.

Metacommunities, fitness and gradual evolution

We analyze the evolution of a multidimensional quantitative trait in a class-structured focal species interacting with other species in a wider metacommunity. The evolutionary dynamics in the focal species as well as the ecological dynamics of the whole metacommunity is described as a continuous-time process with birth, physiological development, dispersal, and death given as rates that can depend on the state of the whole metacommunity. This can accommodate complex local community and global metacommunity environmental feedbacks owing to inter- and intra-specific interactions, as well as local environmental stochastic fluctuations. For the focal species, we derive a fitness measure for a mutant allele affecting class-specific trait expression. Using classical results from geometric singular perturbation theory, we provide a detailed proof that if the effect of the mutation on phenotypic expression is small (“weak selection”), the large system of dynamical equations needed to describe selection on the mutant allele in the metacommunity can be reduced to a single ordinary differential equation on the arithmetic mean mutant allele frequency that is of constant sign. This invariance on allele frequency entails the mutant either dies out or will out-compete the ancestral resident (or wild) type. Moreover, the directional selection coefficient driving arithmetic mean allele frequency can be expressed as an inclusive fitness effect calculated from the resident metacommunity alone, and depends, as expected, on individual fitness differentials, relatedness, and reproductive values. This formalizes the Darwinian process of gradual evolution driven by random mutation and natural selection in spatially and physiologically class-structured metacommunities.

The influence of autotoxicity on the dynamics of vegetation spots

Plant autotoxicity has proved to play an essential role in the behaviour of local vegetation. We analyse a reaction–diffusion-ODE model describing the interactions between vegetation, water, and autotoxicity. The presence of autotoxicity is seen to induce movement and deformation of spot patterns in some parameter regimes, a phenomenon which does not occur in classical biomass–water models. We aim to analytically quantify this novel feature, by studying travelling wave solutions in one spatial dimension. We use geometric singular perturbation theory to prove the existence of symmetric, stationary and non-symmetric, travelling pulse solutions, by constructing appropriate homoclinic orbits in the associated 5-dimensional dynamical system. In the singularly perturbed context, we perform an extensive scaling analysis of the dynamical system, identifying multiple asymptotic scaling regimes where (travelling) pulses may or may not be constructed. We show that, while the analytically constructed stationary pulse corresponds to its numerical counterpart, there is a discrepancy between the numerically observed travelling pulse and its analytical counterpart. Our findings indicate how the inclusion of an additional ODE may significantly influence the properties of classical biomass–water models of Klausmeier/Gray–Scott type.

Singularly perturbed boundary-focus bifurcations

We consider smooth systems limiting as ϵ → 0 to piecewise-smooth (PWS) systems with a boundary-focus (BF) bifurcation. After deriving a suitable local normal form, we study the dynamics for the smooth system with sufficiently small but non-zero ϵ , using a combination of geometric singular perturbation theory and blow-up . We show that the type of BF bifurcation in the PWS system determines the bifurcation structure for the smooth system within an ϵ −dependent domain which shrinks to zero as ϵ → 0 , identifying a supercritical Andronov-Hopf bifurcation in one case, and a supercritical Bogdanov-Takens bifurcation in two other cases. We also show that PWS cycles associated with BF bifurcations persist as relaxation oscillations in the smooth system, and prove existence of a family of stable limit cycles which connects the relaxation oscillations to regular cycles within the ϵ −dependent domain described above. Our results are applied to models for Gause predator-prey interaction and mechanical oscillation subject to friction.

Reduced Models of Cardiomyocytes Excitability: Comparing Karma and FitzHugh\u2013Nagumo

Since Noble adapted in 1962 the model of Hodgkin and Huxley to fit Purkinje fibres, the refinement of models for cardiomyocytes has continued. Most of these models are high-dimensional systems of coupled equations so that the possible mathematical analysis is quite limited, even numerically. This has inspired the development of reduced, phenomenological models that preserve qualitatively the main feature of cardiomyocyte’s dynamics. In this paper, we present a systematic comparison of the dynamics between two notable low-dimensional models, the FitzHugh–Nagumo model (FitzHugh in Bull Math Biophys 17:257–269, 1955, J Gen Physiol 43:867–896, 1960, Biophys J 1:445–466, 1961) as a prototype of excitable behaviour and a polynomial version of the Karma model (Karma in Phys Rev Lett 71(7):16, 1993, Chaos 4:461, 1994) which is specifically developed to fit cardiomyocyte’s behaviour well. We start by introducing the models and considering their pure ODE versions. We analyse the ODEs employing the main ideas and steps used in the setting of geometric singular perturbation theory. Next, we turn to the spatially extended models, where we focus on travelling wave solutions in 1D. Finally, we perform numerical simulations of the 1D PDE Karma model varying model parameters in order to systematically investigate the impact on wave propagation velocity and shape. In summary, our study provides a reference regarding key similarities as well as key differences of the two models.

Controlling Canard Cycles

Canard cycles are periodic orbits that appear as special solutions of fast-slow systems (or singularly perturbed ordinary differential equations). It is well known that canard cycles are difficult to detect, hard to reproduce numerically, and that they are sensible to exponentially small changes in parameters. In this paper, we combine techniques from geometric singular perturbation theory, the blow-up method, and control theory, to design controllers that stabilize canard cycles of planar fast-slow systems with a folded critical manifold. As an application, we propose a controller that produces stable mixed-mode oscillations in the van der Pol oscillator.

Analysis and simulation of periodic and solitary waves in nonlinear dispersive-dissipative solids

In this paper, we consider a general realistic nonlinear elastic rod model which is embedded in an external medium with weak disturbance. We apply the geometric singular perturbation theory to prove the existence of a unique periodic traveling wave in this model. With a dynamical system approach, it is shown that the periodic wave appears near a stationary state and then amplifies and degenerates into a solitary wave. Simulations are presented to demonstrate an excellent agreement with the theoretical predictions.

Effects on I–V relations from small permanent charge and channel geometry via classical Poisson–Nernst–Planck equations with multiple cations

We analyse a quasi-one-dimensional Poisson–Nernst–Planck system for ionic flow through a membrane channel with nonzero but small permanent charge. The system includes three ion species, two cations with the same valences and one anion. The cross-sectional area of the channel is included in the system, which provides certain information of the geometry of the three-dimensional channel. This is critical for our analysis. We treat the model as a boundary value problem of a singularly perturbed system of differential equations. Under the framework of geometric singular perturbation theory, together with specific structures of the model, the existence and local uniqueness of solutions to the boundary value problem for small permanent charges is established. Furthermore, treating the permanent charge as a small parameter, via regular perturbation analysis, we are able to derive an approximation of the I–V (current–voltage) relations explicitly, from which the permanent charge and channel geometry effects on ionic flows are analysed in detail. Critical potentials are identified and their roles in characterising the ionic flow properties of interest are studied. Some critical potentials can be experimentally estimated, and this provides an efficient way to adjust the boundary pot to observe distinct dynamics of ionic flows. Mathematical analysis indicates that a short and narrow filter within which the permanent charge is confined is expected in order to optimise the effect of permanent charges. This is consistent with the typical structure of an ion channel.

Traveling wave solutions in a model for social outbursts in a tension‐inhibitive regime

AbstractIn this work, we investigate the existence of nonmonotone traveling wave solutions to a reaction‐diffusion system modeling social outbursts, such as rioting activity, originally proposed in Berestycki et al (Netw Heterog Media. 2015;10(3):443–475). The model consists of two scalar values, the level of unrest and a tension field . A key component of the model is a bandwagon effect in the unrest, provided the tension is sufficiently high. We focus on the so‐called tension‐inhibitive regime, characterized by the fact that the level of unrest has a negative feedback on the tension. This regime has been shown to be physically relevant for the spatiotemporal spread of the 2005 French riots. We use Geometric Singular Perturbation Theory to study the existence of such solutions in two situations. The first is when both and diffuse at a very small rate. Here, the time scale over which the bandwagon effect is observed plays a key role. The second case we consider is when the tension diffuses at a much slower rate than the level of unrest. In this case, we are able to deduce that the driving dynamics are modeled by the well‐known Fisher–Kolmogorov‐Petrovsky‐Piskunov (KPP) equation.

Geometric analysis of mixed-mode oscillations in a model of electrical activity in human beta-cells

Human pancreatic beta-cells may exhibit complex mixed-mode oscillatory electrical activity, which underlies insulin secretion. A recent biophysical model of human beta-cell electrophysiology can simulate such bursting behavior, but a mathematical understanding of the model’s dynamics is still lacking. Here we exploit time-scale separation to simplify the original model to a simpler three-dimensional model that retains the behavior of the original model and allows us to apply geometric singular perturbation theory to investigate the origin of mixed-mode oscillations. Changing a parameter modeling the maximal conductance of a potassium current, we find that the reduced model possesses a singular Hopf bifurcation that results in small-amplitude oscillations, which go through a period-doubling sequence and chaos until the birth of a large-scale return mechanism and bursting dynamics. The theory of folded node singularities provide insight into the bursting dynamics further away from the singular Hopf bifurcation and the eventual transition to simple spiking activity. Numerical simulations confirm that the insight obtained from the analysis of the reduced model can be lifted back to the original model.

Geometric singular perturbation theory analysis of an epidemic model with spontaneous human behavioral change.

We consider a model due to Piero Poletti and collaborators that adds spontaneous human behavioral change to the standard SIR epidemic model. In its simplest form, the Poletti model adds one differential equation, motivated by evolutionary game theory, to the SIR model. The new equation describes the evolution of a variable x that represents the fraction of the population following normal behavior. The remaining fraction 1-x uses altered behavior such as staying home, social isolation, mask wearing, etc. Normal behavior offers a higher payoff when the number of infectives is low; altered behavior offers a higher payoff when the number is high. We show that the entry–exit function of geometric singular perturbation theory can be used to analyze the model in the limit in which behavior changes on a much faster time scale than that of the epidemic. In particular, behavior does not change as soon as a different behavior has a higher payoff; current behavior is sticky. The delay until behavior changes is predicted by the entry–exit function.

Small Permanent Charge Effects on Individual Fluxes via Poisson–Nernst–Planck Models with Multiple Cations

A quasi-one-dimensional Poisson–Nernst–Planck system for ionic flow through a membrane channel is studied. Nonzero but small permanent charge, the major structural quantity of an ion channel, is included in the model. The system includes three ion species, two cations with the same valences and one anion, which provides more correlations/interactions between ions compared to the case included only two oppositely charged particles. The cross-section area of the channel is included in the system, which provides certain information of the geometry of the three-dimensional channel. This is crucial for our analysis. Under the framework of geometric singular perturbation theory, more importantly, the specific structure of the model, the existence and local uniqueness of solutions to the system for small permanent charges is established. Furthermore, treating the permanent charge as a small parameter, through regular perturbation analysis, we are able to derive approximations of the individual fluxes explicitly, and this allows us to examine the small permanent charge effects on ionic flows in detail. Of particular interest is the competition between two cations, which is related to the selectivity phenomena of ion channels. Critical potentials are identified and their roles in characterizing ionic flow properties are studied. Some critical potentials can be estimated experimentally, and this provides an efficient way to adjust/control boundary conditions (electric potential and concentrations) to observe distinct qualitative properties of ionic flows. Mathematical analysis further indicates that to optimize the effect of permanent charges, a short and narrow filter, within which the permanent charge is confined, is expected, which is consistent with the typical structure of an ion channel.

Shock-fronted travelling waves in a reaction–diffusion model with nonlinear forward–backward–forward diffusion

Reaction–diffusion equations (RDEs) are often derived as continuum limits of lattice-based discrete models. Recently, a discrete model which allows the rates of movement, proliferation and death to depend upon whether the agents are isolated has been proposed, and this approach gives various RDEs where the diffusion term is convex and can become negative (Johnston et al., 2017), i.e. forward–backward–forward diffusion. Numerical simulations suggest these RDEs support shock-fronted travelling waves when the reaction term includes an Allee effect. In this work we formalise these preliminary numerical observations by analysing the shock-fronted travelling waves through embedding the RDE into a larger class of higher order partial differential equations (PDEs). Subsequently, we use geometric singular perturbation theory to study this larger class of equations and prove the existence of these shock-fronted travelling waves. Most notable, we show that different embeddings yield shock-fronted travelling waves with different properties.