We explore mode-locking of spontaneous oscillations of saccular hair cell bundles to periodic mechanical deflections. A simple dynamic systems framework is presented that captures the main features of the experimentally observed behavior in the form of an Arnold Tongue. We propose that the phase-locking transition can proceed via different bifurcations. At low stimulus amplitudes F, the transition to mode-locking as a function of the stimulus frequency ω has the character of a saddle-node bifurcation on an invariant circle. At higher stimulus amplitudes, the mode-locking transition has the character of a supercritical Andronov-Hopf bifurcation.

The inner ear constitutes a remarkably sensitive mechanical detector. This detection occurs in a noisy and highly viscous environment, as the sensory cells—the hair cells—are immersed in a fluid-filled compartment and operate at room or higher temperatures. We model the active motility of hair cell bundles of the vestibular system with the Adler equation, which describes the phase degree of freedom of bundle motion. We explore both analytically and numerically the response of the system to external signals, in the presence of white noise. The theoretical model predicts that hair bundles poised in the quiescent regime can exhibit sporadic spikes—sudden excursions in the position of the bundle. In this spiking regime, the system exhibits stochastic resonance, with the spiking rate peaking at an optimal level of noise. Upon the application of a very weak signal, the spikes occur at a preferential phase of the stimulus cycle. We compare the theoretical predictions of our model to experimental measurements obtained in vitro from individual hair cells. Finally, we show that an array of uncoupled hair cells could provide a sensitive detector that encodes the frequency of the applied signal.

The amphibian sacculus (AS) is an end organ that specializes in the detection of low-frequency auditory and vestibular signals. In this paper, we propose a model for the AS in the form of an array of phase oscillators with long-range coupling, subject to a steady load that suppresses spontaneous oscillations. The array is exposed to significant levels of frequency dispersion and intrinsic noise. We show that such an array can be a sensitive and robust subthreshold detector of low-frequency stimuli, though without significant frequency selectivity. The effects of intrinsic noise and frequency dispersion are contrasted. Intermediate levels of intrinsic noise greatly enhance the sensitivity through stochastic resonance. Frequency dispersion, on the other hand, only degrades detection sensitivity. However, frequency dispersion can play a useful role in terms of the suppression of spontaneous activity. As a model for the AS, the array parameters are such that the system is poised near a saddle-node bifurcation on an invariant circle. However, by a change of array parameters, the same system also can be poised near an emergent Andronov-Hopf bifurcation and thereby function as a frequency-selective detector.

Hair cells of the inner ear exhibit an active process, believed to be crucial for achieving the sensitivity of auditory and vestibular detection. One of the manifestations of the active process is the occurrence of spontaneous hair bundle oscillations in vitro. Hair bundles are coupled by overlying membranes in vivo; hence, explaining the potential role of innate bundle motility in the generation of otoacoustic emissions requires an understanding of the effects of coupling on the active bundle dynamics. We used microbeads to connect small groups of hair cell bundles, using in vitro preparations that maintain their innate oscillations. Our experiments demonstrate robust synchronization of spontaneous oscillations, with either 1:1 or multi-mode phase-locking. The frequency of synchronized oscillation was found to be near the mean of the innate frequencies of individual bundles. Coupling also led to an improved regularity of entrained oscillations, demonstrated by an increase in the quality factor.