important step towards understanding the working principles of the mammalian hearing sensor is the concept of an active cochlear amplifier. Theoretical arguments and physiological measurements suggest that the active cochlear amplifiers originate from systems close to a Hopf bifurcation. Efforts to model the mammalian hearing sensor on these grounds have, however, either had problems in reproducing sufficiently close essential aspects of the biological example (Magnasco, M.O. Phys. Rev. Lett. 90, 058101 (2003); Duke, T. & Jülicher, F. Phys. Rev. Lett. 90, 158101 (2003)), or required complicated spatially coupled differential equation systems that are unfeasible for transient signals (Kern, A. & Stoop, R. Phys. Rev. Lett. 91, 128101 (2003)). Here, we demonstrate a simple system of electronically coupled Hopf amplifiers that not only leads to the desired biological response behavior, but also has real-time capacity. The obtained electronic Hopf cochlea shares all salient signal processing features exhibited by the mammalian cochlea and thus provides a simple and efficient design of an artificial mammalian hearing sensor.

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In response to a sound stimulus, the inner ear emits sounds called otoacoustic emissions. While the exact mechanism for the production of otoacoustic emissions is not known, active motion of individual hair cells is thought to play a role. Two possible sources for otoacoustic emissions, both localized within individual hair cells, include somatic motility and hair bundle motility. Because physiological models of each of these systems are thought to be poised near a Hopf bifurcation, the dynamics of each can be described by the normal form for a system near a Hopf bifurcation. Here we demonstrate that experimental results from three–frequency suppression experiments can be predicted based on the response of an array of noninteracting Hopf oscillators tuned at different frequencies. This supports the idea that active motion of individual hair cells contributes to active processing of sounds in the ear. Interestingly, the model suggests an explanation for differing results recorded in mammals and nonmammals. Key words: Hair Cells; Inner Ear; Hopf Bifurcation; Otoacoustic EmissionsMultifrequency Forcing 2

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Active hair bundle motility has been proposed to underlie the amplification mechanism in the auditory endorgans of nonmammals and in the vestibular systems of all vertebrates, and to constitute a crucial component of cochlear amplification in mammals. We used semi-intact in vitro preparations of the bullfrog sacculus to study the effects of elastic mechanical loading on both natively coupled and freely oscillating hair bundles. For the latter, we attached glass fibers of different stiffness to the stereocilia and observed the induced changes in the spontaneous bundle movement. When driven with sinusoidal deflections, hair bundles displayed phase-locked response indicative of an Arnold Tongue, with the frequency selectivity highest at low amplitudes and decreasing under stronger stimulation. A striking broadening of the mode-locked response was seen with increasing stiffness of the load, until approximate impedance matching, where the phase-locked response remained flat over the physiological range of frequencies. When the otolithic membrane was left intact atop the preparation, the natural loading of the bundles likewise decreased their frequency selectivity with respect to that observed in freely oscillating bundles. To probe for signatures of the active process under natural loading and coupling conditions, we applied transient mechanical stimuli to the otolithic membrane. Following the pulses, the underlying bundles displayed active movement in the opposite direction, analogous to the twitches observed in individual cells. Tracking features in the otolithic membrane indicated that it moved in phase with the bundles. Hence, synchronous active motility evoked in the

We use the oscillating Min proteins of Escherichia coli as a prototype system to illustrate the current state and potential of modeling protein dynamics in space and time. We demonstrate how a theoretical approach has led to striking new insights into the mechanisms of selforganization in bacterial cells and indicate how these ideas may be applicable to more complex structure formation in eukaryotic cells.