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We consider a periodically forced 1D Langevin equation that possesses two stable periodic solutions in the absence of noise. We ask the question: is there a most likely noise-induced transition path between these periodic solutions that allows us to identify a preferred phase of the forcing when tipping occurs? The quasistatic regime, where the forcing period is long compared to the adiabatic relaxation time, has been well studied; our work instead explores the case when these time scales are comparable. We compute optimal paths using the path integral method incorporating the Onsager-Machlup functional and validate results with Monte Carlo simulations. Results for the preferred tipping phase are compared with the deterministic aspects of the problem. We identify parameter regimes where nullclines, associated with the deterministic problem in a 2D extended phase space, form passageways through which the optimal paths transit. As the nullclines are independent of the relaxation time and the noise strength, this leads to a robust deterministic predictor of the preferred tipping phase in a regime where forcing is neither too fast nor too slow.
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Banded patterns consisting of alternating bare soil and dense vegetation have been observed in water-limited ecosystems across the globe, often appearing along gently sloped terrain with the stripes aligned transverse to the elevation gradient. In many cases, these vegetation bands are arced, with field observations suggesting a link between the orientation of arcing relative to the grade and the curvature of the underlying terrain. We modify the water transport in the Klausmeier model of water-biomass interactions, originally posed on a uniform hillslope, to qualitatively capture the influence of terrain curvature on the vegetation patterns. Numerical simulations of this modified model indicate that the vegetation bands arc convex-downslope when growing on top of a ridge, and convex-upslope when growing in a valley. This behaviour is consistent with observations from remote sensing data that we present here. Model simulations show further that whether bands grow on ridges, valleys or both depends on the precipitation level. A survey of three banded vegetation sites, each with a different aridity level, indicates qualitatively similar behaviour.
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Ecossistema , Modelos Biológicos , Plantas , Simulação por Computador , Demografia , ÁguaRESUMO
Climate change, amplified in the far north, has led to rapid sea ice decline in recent years. In the summer, melt ponds form on the surface of Arctic sea ice, significantly lowering the ice reflectivity (albedo) and thereby accelerating ice melt. Pond geometry controls the details of this crucial feedback; however, a reliable model of pond geometry does not currently exist. Here we show that a simple model of voids surrounding randomly sized and placed overlapping circles reproduces the essential features of pond patterns. The only two model parameters, characteristic circle radius and coverage fraction, are chosen by comparing, between the model and the aerial photographs of the ponds, two correlation functions which determine the typical pond size and their connectedness. Using these parameters, the void model robustly reproduces the ponds' area-perimeter and area-abundance relationships over more than 6 orders of magnitude. By analyzing the correlation functions of ponds on several dates, we also find that the pond scale and the connectedness are surprisingly constant across different years and ice types. Moreover, we find that ponds resemble percolation clusters near the percolation threshold. These results demonstrate that the geometry and abundance of Arctic melt ponds can be simply described, which can be exploited in future models of Arctic melt ponds that would improve predictions of the response of sea ice to Arctic warming.
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In many dryland environments, vegetation self-organizes into bands that can be clearly identified in remotely-sensed imagery. The status of individual bands can be tracked over time, allowing for a detailed remote analysis of how human populations affect the vital balance of dryland ecosystems. In this study, we characterize vegetation change in areas of the Horn of Africa where imagery taken in the early 1950s is available. We find that substantial change is associated with steep increases in human activity, which we infer primarily through the extent of road and dirt track development. A seemingly paradoxical signature of human impact appears as an increase in the widths of the vegetation bands, which effectively increases the extent of vegetation cover in many areas. We show that this widening occurs due to altered rates of vegetation colonization and mortality at the edges of the bands, and conjecture that such changes are driven by human-induced shifts in plant species composition. Our findings suggest signatures of human impact that may aid in identifying and monitoring vulnerable drylands in the Horn of Africa.
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Modelos Teóricos , Plantas , África , Ecossistema , HumanosRESUMO
A particular sequence of patterns, 'gapsâlabyrinthâspots', occurs with decreasing precipitation in previously reported numerical simulations of partial differential equation dryland vegetation models. These observations have led to the suggestion that this sequence of patterns can serve as an early indicator of desertification in some ecosystems. Because parameter values in the vegetation models can take on a range of plausible values, it is important to investigate whether the pattern sequence prediction is robust to variation. For a particular model, we find that a quantity calculated via bifurcation-theoretic analysis appears to serve as a proxy for the pattern sequences that occur in numerical simulations across a range of parameter values. We find in further analysis that the quantity takes on values consistent with the standard sequence in an ecologically relevant limit of the model parameter values. This suggests that the standard sequence is a robust prediction of the model, and we conclude by proposing a methodology for assessing the robustness of the standard sequence in other models and formulations.
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A feature common to many models of vegetation pattern formation in semiarid ecosystems is a sequence of qualitatively different patterned states, "gaps â labyrinth â spots," that occurs as a parameter representing precipitation decreases. We explore the robustness of this "standard" sequence in the generic setting of a bifurcation problem on a hexagonal lattice, as well as in a particular reaction-diffusion model for vegetation pattern formation. Specifically, we consider a degeneracy of the bifurcation equations that creates a small bubble in parameter space in which stable small-amplitude patterned states may exist near two Turing bifurcations. Pattern transitions between these bifurcation points can then be analyzed in a weakly nonlinear framework. We find that a number of transition scenarios besides the standard sequence are generically possible, which calls into question the reliability of any particular pattern or sequence as a precursor to vegetation collapse. Additionally, we find that clues to the robustness of the standard sequence lie in the nonlinear details of a particular model.
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Clima Desértico , Ecossistema , Modelos Biológicos , Reconhecimento Automatizado de Padrão/métodos , Desenvolvimento Vegetal/fisiologia , Plantas/anatomia & histologia , Simulação por ComputadorRESUMO
Designing genetic networks with desired functionalities requires an accurate mathematical framework that accounts for the essential mechanistic details of the system. Here, we formulate a time-delay model of protein translation and mRNA degradation by systematically reducing a detailed mechanistic model that explicitly accounts for the ribosomal dynamics and the cleaving of mRNA by endonucleases. We exploit various technical and conceptual advantages that our time-delay model offers over the mechanistic model to probe the behavior of a self-repressing gene over wide regions of parameter space. We show that a heuristic time-delay model of protein synthesis of a commonly used form yields a notably different prediction for the parameter region where sustained oscillations occur. This suggests that such heuristics can lead to erroneous results. The functional forms that arise from our systematic reduction can be used for every system that involves transcription and translation and they could replace the commonly used heuristic time-delay models for these processes. The results from our analysis have important implications for the design of synthetic gene networks and stress that such design must be guided by a combination of heuristic models and mechanistic models that include all relevant details of the process.
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Redes Reguladoras de Genes , Modelos Genéticos , Endorribonucleases/metabolismo , Escherichia coli/genética , Escherichia coli/metabolismo , Biossíntese de Proteínas , Estabilidade de RNA , RNA Mensageiro/metabolismo , Ribossomos/metabolismo , Fatores de TempoRESUMO
Time-delays are common in many physical and biological systems and they give rise to complex dynamic phenomena. The elementary processes involved in template biopolymerization, such as mRNA and protein synthesis, introduce significant time delays. However, there is not currently a systematic mapping between the individual mechanistic parameters and the time delays in these networks. We present here the development of mathematical, time-delay models for protein translation, based on PDE models, which in turn are derived through systematic approximations of first-principles mechanistic models. Theoretical analysis suggests that the key features that determine the time-delays and the agreement between the time-delay and the mechanistic models are ribosome density and distribution, i.e., the number of ribosomes on the mRNA chain relative to their maximum and their distribution along the mRNA chain. Based on analytical considerations and on computational studies, we show that the steady-state and dynamic responses of the time-delay models are in excellent agreement with the detailed mechanistic models, under physiological conditions that correspond to uniform ribosome distribution and for ribosome density up to 70%. The methodology presented here can be used for the development of reduced time-delay models of mRNA synthesis and large genetic networks. The good agreement between the time-delay and the mechanistic models will allow us to use the reduced model and advanced computational methods from nonlinear dynamics in order to perform studies that are not practical using the large-scale mechanistic models.
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Modelos Genéticos , Biossíntese de Proteínas , RNA Mensageiro/fisiologia , Ribossomos/fisiologia , Algoritmos , Simulação por Computador , Escherichia coli/genética , Proteínas de Escherichia coli/genética , Cinética , Dinâmica não Linear , RNA Mensageiro/genética , RNA Mensageiro/metabolismo , Ribossomos/genética , Ribossomos/metabolismo , Ativação TranscricionalRESUMO
Animal behaviour arises through a complex mixture of biomechanical, neuronal, sensory and control constraints. By focusing on a simple, stereotyped movement, the prey capture strike of a weakly electric fish, we show that the trajectory of a strike is one which minimizes effort. Specifically, we model the fish as a rigid ellipsoid moving through a fluid with no viscosity, governed by Kirchhoff's equations. This formulation allows us to exploit methods of discrete mechanics and optimal control to compute idealized fish trajectories that minimize a cost function. We compare these with the measured prey capture strikes of weakly electric fish from a previous study. The fish has certain movement limitations that are not incorporated in the mathematical model, such as not being able to move sideways. Nonetheless, we show quantitatively that the computed least-cost trajectories are remarkably similar to the measured trajectories. Since, in this simplified model, the basic geometry of the idealized fish determines the favourable modes of movement, this suggests a high degree of influence between body shape and movement capability. Simplified minimal models and optimization methods can give significant insight into how body morphology and movement capability are closely attuned in fish locomotion.
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Gimnotiformes/fisiologia , Locomoção/fisiologia , Comportamento Predatório/fisiologia , Animais , Fenômenos Biomecânicos , Gimnotiformes/anatomia & histologia , Modelos Anatômicos , Percepção de Movimento/fisiologia , NataçãoRESUMO
For many years it was believed that an unstable periodic orbit with an odd number of real Floquet multipliers greater than unity cannot be stabilized by the time-delayed feedback control mechanism of Pyragas. A recent paper by Fiedler et al. Phys. Rev. Lett. 98, 114101 (2007) uses the normal form of a subcritical Hopf bifurcation to give a counterexample to this theorem. Using the Lorenz equations as an example, we demonstrate that the stabilization mechanism identified by Fiedler et al. for the Hopf normal form can also apply to unstable periodic orbits created by subcritical Hopf bifurcations in higher-dimensional dynamical systems. Our analysis focuses on a particular codimension-two bifurcation that captures the stabilization mechanism in the Hopf normal form example, and we show that the same codimension-two bifurcation is present in the Lorenz equations with appropriately chosen Pyragas-type time-delayed feedback. This example suggests a possible strategy for choosing the feedback gain matrix in Pyragas control of unstable periodic orbits that arise from a subcritical Hopf bifurcation of a stable equilibrium. In particular, our choice of feedback gain matrix is informed by the Fiedler et al. example, and it works over a broad range of parameters, despite the fact that a center-manifold reduction of the higher-dimensional problem does not lead to their model problem.
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We investigate the relationship between the linear surface wave instabilities of a shallow viscous fluid layer and the shape of the periodic, parametric-forcing function (describing the vertical acceleration of the fluid container) that excites them. We find numerically that the envelope of the resonance tongues can only develop multiple minima when the forcing function has more than two local extrema per cycle. With this insight, we construct a multi-frequency forcing function that generates at onset a nontrivial harmonic instability which is distinct from a subharmonic response to any of its frequency components. We measure the corresponding surface patterns experimentally and verify that small changes in the forcing waveform cause a transition, through a bicritical point, from the predicted harmonic short-wavelength pattern to a much larger standard subharmonic pattern. Using a formulation valid in the lubrication regime (thin viscous fluid layer) and a Wentzel-Kramers-Brillouin (WKB) method to find its analytic solutions, we explore the origin of the observed relation between the forcing function shape and the resonance tongue structure. In particular, we show that for square and triangular forcing functions the envelope of these tongues has only one minimum, as in the usual sinusoidal case.
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Parametrically-excited surface waves, forced by a repeating sequence of delta-function impulses, are considered within the framework of the Zhang-Viñals model [W. Zhang and J. Viñals, J. Fluid Mech. 336, 301 (1997)]. With impulsive forcing, the linear stability analysis can be carried out exactly and leads to an implicit equation for the neutral stability curves. As noted previously [J. Bechhoefer and B. Johnson, Am. J. Phys. 64, 1482 (1996)], in the simplest case of N=2 equally-spaced impulses per period (which alternate up and down) there are only subharmonic modes of instability. The familiar situation of alternating subharmonic and harmonic resonance tongues emerges only if an asymmetry in the spacing between the impulses is introduced. We extend the linear analysis for N=2 impulses per period to the weakly nonlinear regime, where we determine the leading order nonlinear saturation of one-dimensional standing waves as a function of forcing strength. Specifically, an analytic expression for the cubic Landau coefficient in the bifurcation equation is derived as a function of the dimensionless spacing between the two impulses and the fluid parameters that appear in the Zhang-Viñals model. As the capillary parameter is varied, one finds a parameter regime of wave amplitude suppression, which is due to a familiar 1:2 spatiotemporal resonance between the subharmonic mode of instability and a damped harmonic mode. This resonance occurs for impulsive forcing even when harmonic resonance tongues are absent from the neutral stability curves. The strength of this resonance feature can be tuned by varying the spacing between the impulses. This finding is interpreted in terms of a recent symmetry-based analysis of multifrequency forced Faraday waves [J. Porter, C. M. Topaz, and M. Silber, Phys. Lett. 93, 034502 (2004); C. M. Topaz, J. Porter, and M. Silber, Phys. Rev. E 70, 066206 (2004)].
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We use symmetry considerations to investigate control of a class of resonant three-wave interactions relevant to pattern formation in weakly damped, parametrically forced systems near onset. We classify and tabulate the most important damped, resonant modes and determine how the corresponding resonant triad interactions depend on the forcing parameters. The relative phase of the forcing terms may be used to enhance or suppress the nonlinear interactions. We compare our symmetry-based predictions with numerical and experimental results for Faraday waves. Our results suggest how to design multifrequency forcing functions that favor chosen patterns in the lab.
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We show how pattern formation in Faraday waves may be manipulated by varying the harmonic content of the periodic forcing function. Our approach relies on the crucial influence of resonant triad interactions coupling pairs of critical standing wave modes with damped, spatiotemporally resonant modes. Under the assumption of weak damping and forcing, we perform a symmetry-based analysis that reveals the damped modes most relevant for pattern selection, and how the strength of the corresponding triad interactions depends on the forcing frequencies, amplitudes, and phases. In many cases, the further assumption of Hamiltonian structure in the inviscid limit determines whether the given triad interaction has an enhancing or suppressing effect on related patterns. Surprisingly, even for forcing functions with arbitrarily many frequency components, there are at most five frequencies that affect each of the important triad interactions at leading order. The relative phases of those forcing components play a key role, sometimes making the difference between an enhancing and suppressing effect. In numerical examples, we examine the validity of our results for larger values of the damping and forcing. Finally, we apply our findings to one-dimensional periodic patterns obtained with impulsive forcing and to two-dimensional superlattice patterns and quasipatterns obtained with multifrequency forcing.
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We exploit the approximate (broken) symmetries of time translation, time reversal, and Hamiltonian structure to obtain general scaling laws governing the process of pattern formation in weakly damped Faraday waves. Using explicit parameter symmetries we determine, for the case of two-frequency forcing, how the strength of observed three-wave interactions depends on the frequency ratio and on the relative phase of the two driving terms. These symmetry-based predictions are verified for numerically calculated coefficients, and help explain the results of recent experiments.