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Dolores Bozovic

Publications and source records attributed to Dolores Bozovic.

18 recordsLinked to original sources

Noise-induced stability of insect swarms

Flying insect swarms exhibit cohesion without the local velocity alignment observed in flocks of birds or schools of fish. The interaction rules and channels of communication between insects that lead to collective behavior have not yet been determined. We propose a theoretical model based on acoustic communication between swarm members, where each individual is attracted to a weighted mean field of its neighbors. We demonstrate that this simple framework can describe a wide range of swarming dynamics, including a phenomenon where pairs of insects break free from the swarm in a helical orbit around each other. Counterintuitively, our numerical model suggests that stochastic noise enhances the stability of an insect swarm, as it interferes with pair formation. Finally, we demonstrate that a specific species of malarial mosquito produces swarms that reside on the edge of a transition to instability. Our study shows that fairly simple local interaction rules can be sufficient to describe the collective behavior of swarming biological systems.

physics.bio-ph

Sound Masking Strategies for Interference with Mosquito Hearing

The use of auditory masking has long been of interest in psychoacoustics and for engineering purposes, in order to cover sounds that are disruptive to humans or to species whose habitats overlap with ours. In most cases, we seek to minimize the disturbances to the communication of wildlife. However, in the case of pathogen-carrying insects, we may want to maximize these disturbances as a way to control populations. In the current work, we explore candidate masking strategies for a generic model of active auditory systems and a model of the mosquito auditory system. For both models, we find that masks with all acoustic power focused into just one or a few frequencies perform best. We propose that masks based on rapid frequency modulation are most effective for maximal disruption of information transfer and minimizing intelligibility. We hope that these results will serve to guide the avoidance or selection of possible acoustic signals for, respectively, maximizing or minimizing communication.

physics.bio-ph

Estimating Free Parameters in Stochastic Oscillatory Models Using a Weighted Cost Function

In this study, we estimate parameters in stochastic oscillatory systems by developing a novel cost function. This function incorporates power spectral density, analytic signal, and position crossings, each weighted to capture distinct oscillatory characteristics such as amplitude, frequency, and shape. By minimizing this cost via differential evolution, we estimate parameters in two stochastic systems given measured datasets. We validate this procedure by recovering known parameters from a test dataset. We then apply it to a biophysical model for auditory mechanics. Thus, we establish a general methodology for fitting stochastic oscillatory systems.

physics.comp-ph

Antennal-Based Strategies for Sound Localization by Insects

Insects rely on their hearing in order to communicate, identify and locate potential mates, and avoid predators. Due to their small sizes, many insect species are not able to utilize the interaural time and intensity differences employed by vertebrates for the localization of sound, but have instead evolved other mechanisms to perform this task. One such mechanism is the antenna, which provides directionally sensitive acoustic information. In the current work, we discuss the physical limitations imposed by the Gabor limit and the nature of acoustic radiation as small length scales. We then propose mechanisms that antennal insects may use in order to localize sound and extract precise frequency information from transient signals, thereby circumventing these physical limitations.

physics.bio-ph

A Mosquito-Inspired Theoretical Framework for Acoustic Signal Detection

Distortion products are tones produced through nonlinear effects of a system simultaneously detecting two or more frequencies. These combination tones are ubiquitous to vertebrate auditory systems and are generally regarded as byproducts of nonlinear signal amplification. It has previously been shown that several species of infectious-disease-carrying mosquitoes utilize these distortion products for detecting and locating potential mates. It has also been shown that their auditory systems contain multiple oscillatory components within the sensory structure, which respond at different frequency ranges. Using a generic theoretical model for acoustic detection, we show the signal-detection advantages that are implied by these two detection schemes: distortion product detection and cascading a signal through multiple layers of oscillator elements. Lastly, we show that the combination of these two schemes yields immense benefits for signal detection. These benefits could be essential for male mosquitoes to be able to identify and pursue a particular female within a noisy swarm environment.

physics.bio-ph

Developing Parameter-Reduction Methods on a Biophysical Model of Auditory Hair Cells

Biophysical models describing complex, cellular phenomena typically include systems of nonlinear differential equations with many free parameters. While experimental measurements can fix some parameters, those describing internal cellular processes frequently remain inaccessible. Hence, a proliferation of free parameters risks overfitting the data, limiting the model's predictive power. In this study, we develop robust methods, applying statistical analysis and dynamical-systems theory, to reduce a biophysical model's complexity. We demonstrate our techniques on an elaborate computational model designed to describe active, mechanical motility of auditory hair cells. Specifically, we use two statistical measures, the total-effect and PAWN indices, to rank each free parameter by its influence on selected, core properties of the model. With the resulting ranking, we fix most of the less influential parameters, yielding a low-parameter model with optimal predictive power. We validate the theoretical model with experimental recordings of active hair-bundle motility, specifically by using Akaike and Bayesian information criteria after obtaining maximum-likelihood fits. As a result, we determine the system's most influential parameters, which illuminate its key biophysical elements of the cell's overall features. While we demonstrated our techniques on a concrete example, they provide a general framework, applicable to other biophysical systems.

physics.comp-ph

Criticality and Chaos in Auditory and Vestibular Sensing

The auditory and vestibular systems exhibit remarkable sensitivity of detection, responding to deflections on the order of Angstroms, even in the presence of biological noise. Further, these complex systems exhibit high temporal acuity and frequency selectivity, allowing us to make sense of the acoustic world around us. As this acoustic environment of interest spans several orders of magnitude in both amplitude and frequency, these systems rely heavily on nonlinearities and power-law scaling. The behavior of these sensory systems has been extensively studied in the context of dynamical systems theory, with many empirical phenomena described by critical dynamics. Other phenomena have been explained by systems in the chaotic regime, where weak perturbations drastically impact the future state of the system. We first review the conceptual framework behind these two types of detectors, as well as the detection features that they can capture. We then explore the intersection of the two types of systems and propose ideal parameter regimes for auditory and vestibular systems.

nlin.CD

Vestibular Drop Attacks and Meniere's Disease as Results of Otolithic Membrane Damage -- A Numerical Model

BACKGROUND: Meniere's Disease (MD) is a condition of the inner ear with symptoms affecting both vestibular and hearing functions. Some patients with MD experience vestibular drop attacks (VDAs), which are violent falls caused by spurious vestibular signals from the utricle and/or saccule. Recent surgical work has shown that patients who experience VDAs also show distrupted utricular otolithic membranes. OBJECTIVE: The objective of this study is to determine if otolithic membrane damage alone is sufficient to induce spurious vestibular signals, thus potentially eliciting VDAs and the vestibular dysfunction seen in patients with MD. METHODS: We use a previously developed numerical model to describe the nonlinear dynamics of an array of active, elastically coupled hair cells. We then reduce the coupling strength of a selected region of the membrane to model the effects of tissue damage. RESULTS: As we reduce the coupling strength, we observe large and abrupt spikes in hair bundle position. As bundle displacements from the equilibrium position have been shown to lead to depolarization of the hair-cell soma and hence trigger neural activity, this spontaneous activity could elicit false detection of a vestibular signal. CONCLUSIONS: The results of this numerical model suggest that otolithic membrane damage alone may be sufficient to induce VDAs and the vestibular dysfunction seen in patients with MD. Future experimental work is needed to confirm these results in vitro.

q-bio.QM

Chimera states and frequency clustering in systems of coupled inner-ear hair cells

Coupled hair cells of the auditory and vestibular systems perform the crucial task of converting the energy of sound waves and ground-borne vibrations into ionic currents. We mechanically couple groups of living, active hair cells with artificial membranes, thus mimicking in vitro the coupled dynamical system. We identify chimera states and frequency clustering in the dynamics of these coupled nonlinear, autonomous oscillators. We find that these dynamical states can be reproduced by our numerical model with heterogeneity of the parameters. Further, we find that this model is most sensitive to external signals when poised at the onset of synchronization, where chimera and cluster states are likely to form. We therefore propose that the partial synchronization in our experimental system is a manifestation of a system poised at the verge of synchronization with optimal sensitivity.

nlin.AO

Chaos stabilizes synchronization in systems of coupled inner-ear hair cells

Hair cells of the auditory and vestibular systems display astonishing sensitivity, frequency selectivity, and temporal resolution to external signals. These specialized cells utilize an internal active amplifier to achieve highly sensitive mechanical detection. One of the manifestations of this active process is the occurrence of spontaneous limit-cycle motion of the hair cell bundle. As hair bundles under in vivo conditions are typically coupled to each other by overlying structures, we explore the role of this coupling on the dynamics of the system, using a combination of theoretical and experimental approaches. Our numerical model suggests that the presence of chaotic dynamics in the response of individual bundles enhances their ability to synchronize when coupled, resulting in significant improvement in the system's ability to detect weak signals. This synchronization persists even for a large frequency dispersion and a large number of oscillators comprising the system. Further, the amplitude and coherence of the active motion is not reduced upon increasing the number of oscillators. Using artificial membranes, we impose mechanical coupling on groups of live and functional hair bundles, selected from in vitro preparations of the sensory epithelium, allowing us to explore the role of coupling experimentally. Consistent with the numerical simulations of the chaotic system, synchronization occurs even for large frequency dispersion and a large number of hair cells. Further, the amplitude and coherence of the spontaneous oscillations are independent of the number of hair cells in the network. We therefore propose that hair cells utilize their chaotic dynamics to stabilize the synchronized state and avoid the amplitude death regime, resulting in collective coherent motion that could play a role in generating spontaneous otoacoustic emissions and an enhanced ability to detect weak signals.

q-bio.NC

Violation of generalized fluctuation theorems in adaptively driven steady states: Applications to hair cell oscillations

The spontaneously oscillating hair bundle of sensory cells in the inner ear is an example of a stochastic, nonlinear oscillator driven by internal active processes. Moreover, this internal activity is adaptive -- its power input depends on the current state of the system. We study fluctuation dissipation relations in such adaptively-driven, nonequilibrium limit-cycle oscillators. We observe the expected violation of the well-known, equilibrium fluctuation-dissipation theorem (FDT), and verify the existence of a generalized fluctuation-dissipation theorem (GFDT) in the non-adaptively driven model of the hair cell oscillator. This generalized fluctuation theorem requires the system to be analyzed in the co-moving frame associated with the mean limit cycle of the stochastic oscillator. We then demonstrate, via numerical simulations and analytic calculations, that the adaptively-driven dynamical hair cell model violates both the FDT and the GFDT. We go on to show, using stochastic, finite-state, dynamical models, that such a feedback-controlled drive in stochastic limit cycle oscillators generically violates both the FDT and GFDT. We propose that one may in fact use the breakdown of the GFDT as a tool to more broadly look for and quantify the effect of adaptive, feedback mechanisms associated with driven (nonequilibrium) biological dynamics.

cond-mat.stat-mech

Chaotic Dynamics Enhance the Sensitivity of Inner Ear Hair Cells

Hair cells of the auditory and vestibular systems are capable of detecting sounds that induce sub-nanometer vibrations of the hair bundle, below the stochastic noise levels of the surrounding fluid. Hair bundles of certain species are also known to oscillate without external stimulation, indicating the presence of an underlying active mechanism. We propose that chaotic dynamics enhance the sensitivity and temporal resolution of the hair bundle response, and provide experimental and theoretical evidence for this effect. By varying the viscosity and ionic composition of the surrounding fluid, we are able to modulate the degree of chaos observed in the hair bundle dynamics in vitro. We consistently find that the hair bundle is most sensitive to a stimulus of small amplitude when it is poised in the weakly chaotic regime. Further, we show that the response time to a force step decreases with increasing levels of chaos. These results agree well with our numerical simulations of a chaotic Hopf oscillator and suggest that chaos may be responsible for the sensitivity and temporal resolution of hair cells.

q-bio.NC

Noise-induced distortion of nonequilibrium oscillator mean limit cycle

Under general conditions imposed on an active stochastic oscillator, we study change in the size and shape of its average limit cycle as a function of temperature. Such dynamics occur in a multitude of nonequilibrium systems, including the spontaneous oscillations of hair cells of the inner ear. We demonstrate one of the many mechanisms through which distortion of the mean curve may occur and identify regions in the driven oscillator phase space susceptible to corner-cutting due to noise. This we infer causes rounding of certain sharp features in the noiseless curve given by the underlying complex theoretical model, making them inaccessible under any amount of averaging of the experimentally obtained finite-temperature trajectory.

cond-mat.stat-mech

Noise-Induced Chaos and Signal Detection by the Nonisochronous Hopf Oscillator

The Hopf oscillator has been shown to capture many phenomena of the auditory and vestibular systems. These systems exhibit remarkable temporal resolution and sensitivity to weak signals, as they are able to detect sounds that induce motion in the sub-nanometer regime. In the present work, we find the analytic response function of a nonisochronous Hopf oscillator to a step stimulus and show that the system is most sensitive in the regime where noise induces chaotic dynamics. We show that this regime also provides a faster response and enhanced temporal resolution. Thus, the system can detect a very brief, low-amplitude pulse. Finally, we subject the oscillator to periodic delta-function forcing, mimicking a spike train, and find the exact analytic expressions for the stroboscopic maps. Using these maps, we find a period-doubling cascade to chaos with increasing force strength.

nlin.CD

Nonequilibrium limit cycle oscillators: fluctuations in hair bundle dynamics

We develop a framework for the general interpretation of the stochastic dynamical system near a limit cycle. Such quasi-periodic dynamics are commonly found in a variety of nonequilibrium systems, including the spontaneous oscillations of hair cells in the inner ear. We demonstrate quite generally that in the presence of noise, the phase of the limit cycle oscillator will diffuse while deviations in the directions locally orthogonal to that limit cycle will display the Lorentzian power spectrum of a damped oscillator. We identify two mechanisms by which these stochastic dynamics can acquire a complex frequency dependence, and discuss the deformation of the mean limit cycle as a function of temperature. The theoretical ideas are applied to data obtained from spontaneously oscillating hair cells of the amphibian sacculus.

cond-mat.stat-mech

Molding the asymmetry of localized frequency-locking waves by a generalized forcing and implications to the inner ear

Frequency locking to an external forcing frequency is a {well} known phenomenon. In the auditory system, it results in a localized traveling wave, the shape of which is essential for efficient discrimination between incoming frequencies. An amplitude equation approach is used to show that the shape of the localized traveling wave depends crucially on the relative strength of additive vs. parametric forcing components; the stronger the parametric forcing the more asymmetric the response profile and the sharper the traveling-wave front. The analysis captures the empirically observed regions of linear and nonlinear responses and highlights the significance of parametric forcing mechanisms in shaping the resonant response in the inner ear.

nlin.PS

Chaotic Dynamics of Inner Ear Hair Cells

Experimental records of active bundle motility are used to demonstrate the presence of a low-dimensional chaotic attractor in hair cell dynamics. Dimensionality tests from dynamic systems theory are applied to estimate the number of independent variables sufficient for modeling the hair cell response. Poincare maps are constructed to observe a quasiperiodic transition from chaos to order with increasing amplitudes of mechanical forcing. The onset of this transition is accompanied by a reduction of Kolmogorov entropy in the system and an increase in mutual information between the stimulus and the hair bundle, indicative of signal detection. A simple theoretical model is used to describe the observed chaotic dynamics. The model exhibits an enhancement of sensitivity to weak stimuli when the system is poised in the chaotic regime. We propose that chaos may play a role in the hair cell's ability to detect low-amplitude sounds.

physics.bio-ph

Frequency locking in auditory hair cells: Distinguishing between additive and parametric forcing

The auditory system displays remarkable sensitivity and frequency discrimination, attributes shown to rely on an amplification process that involves a mechanical as well as a biochemical response. Models that display proximity to an oscillatory onset (a.k.a. Hopf bifurcation) exhibit a resonant response to distinct frequencies of incoming sound, and can explain many features of the amplification phenomenology. To understand the dynamics of this resonance, frequency locking is examined in a system near the Hopf bifurcation and subject to two types of driving forces: additive and parametric. Derivation of a universal amplitude equation that contains both forcing terms enables a study of their relative impact on the hair cell response. In the parametric case, although the resonant solutions are 1:1 frequency locked, they show the coexistence of solutions obeying a phase shift of $π$, a feature typical of the 2:1 resonance. Different characteristics are predicted for the transition from unlocked to locked solutions, leading to smooth or abrupt dynamics in response to different types of forcing. The theoretical framework provides a more realistic model of the auditory system, which incorporates a direct modulation of the internal control parameter by an applied drive. The results presented here can be generalized to many other media, including Faraday waves, chemical reactions, and nonlinear optics, which are known to exhibit resonant behavior.

nlin.PS