PubMed Health⌕ Search

Biomedical subjects

M Kössl

Publications and source records attributed to M Kössl.

At least 19 recordsLinked to original sources
PubMed2004

Specializations for aerial hawking in the echolocation system of Molossus molossus (Molossidae, Chiroptera).

While searching for prey, Molossus molossus broadcasts narrow-band calls of 11.42 ms organized in pairs of pulses that alternate in frequency. The first signal of the pair is at 34.5 kHz, the second at 39.6 kHz. Pairs of calls with changing frequencies were only emitted when the interpulse intervals were below 200 ms. Maximum duty cycles during search phase are close to 20%. Frequency alternation of search calls is interpreted as a mechanism for increasing duty cycle and thus the temporal continuity of scanning, as well as increasing the detection range. A neurophysiological correlate for the processing of search calls was found in the inferior colliculus. 64% of neurons respond to frequencies in the 30- to 40-kHz range and only in this frequency range were closed tuning curves found for levels below 40 dB SPL. In addition, 15% of the neurons have double-tuned frequency-threshold curves with best thresholds at 34 and 39 kHz. Differing from observations in other bats, approach calls of M. molossus are longer and of higher frequencies than search calls. Close to the roost, the call frequency is increased to 45.0-49.8 kHz and, in addition, extremely broadband signals are emitted. This demonstrates high plasticity of call design.

Acoustic Stimulation↗
PubMed2003

Postnatal development of cochlear function in the mustached bat, Pteronotus parnellii.

Postnatal development of the mustached bat's cochlea was studied by measuring cochlear microphonic and compound action potentials. In adults, a cochlear resonance is involved in enhanced tuning to the second harmonic constant frequency component (CF2) of their echolocation calls at approximately 61 kHz This resonance is present immediately after birth in bats that do not yet echolocate. Its frequency is lower (46 kHz) and the corresponding threshold minimum of cochlear microphonic potentials is broader than in adults. Long-lasting ringing of the cochlear microphonic potential after tone stimulus offset that characterizes the adult auditory response close to CF2 is absent in newborns. In the course of the first 5 postnatal weeks, there is a concomitant upward shift of CF2 and the frequency of cochlear threshold minima. Up to the end of the third postnatal week, sensitivity of auditory threshold minima and the Q value of the cochlear resonance increase at a fast rate. Between 2 and 4 wk of age, two cochlear microphonic threshold minima are found consistently in the CF2 range that differ in their level-dependent dynamic growth behavior and are 1.5-5.7 kHz apart from each other. In older animals, there is a single minimum that approaches adult tuning in its sharpness. The data provide evidence to show that during maturation of the cochlea, the frequency and the sensitivity of the threshold minimum associated with CF2 increases and that these increases are associated with the fusion of two resonances that are partly dissociated in developing animals.

Acoustic Stimulation↗
PubMed2003

Development of echolocation calls in the mustached bat, Pteronotus parnellii.

Adult mustached bats employ Doppler-sensitive sonar to hunt fluttering prey insects in acoustically cluttered habitats. The echolocation call consists of 4-5 harmonics, each composed of a long constant frequency (CF) component flanked by brief frequency modulations (FM). The 2nd harmonic CF component (CF2) at 61 kHz is the most intense, and analyzed by an exceptionally sharply tuned auditory system. The maturation of echolocation calls and the development of Doppler-shift compensation was studied in Cuba where large maternity colonies are found in hot caves. In the 1st postnatal week, infant bats did not echolocate spontaneously but could be induced to vocalize CF-FM signals by passive body motion. The CF2 frequency emitted by the smallest specimens was at 48 kHz (i.e., 0.4 octaves lower than the adult signal). CF-FM signals were spontaneously produced in the 2nd postnatal week at a CF2 frequency of 52 kHz. The CF2 frequencies of induced and spontaneous calls shifted upward to reach a value of 60.5 kHz in the 5th postnatal week. Standard deviations of CF2 frequency were large (up to +/-1.5 kHz) in the youngest bats and dropped to values of +/-250 Hz at the end of the 3rd postnatal week. Some individuals in the 4th and 5th postnatal weeks emitted with adultlike frequency precision of about +/-100 Hz. In the youngest bats, the 1st harmonic CF component (CF1) was up to 22 dB stronger than CF2. Adultlike relative levels of CF1 (-28 dB relative to CF2) were reached in the 5th postnatal week. In spontaneously emitted CF-FM calls, the duration of the CF2 component gradually increased with age from 5 ms to maximum values of 18 ms. Durations of the CF2 component in induced calls averaged 7 +/- 2.6 ms in the 1st postnatal week and 8.2 +/- 1.5 ms in the 5th postnatal week. There were no age-related changes in duration of the terminal FM sweep (3 +/- 0.4 ms) in both induced and spontaneous calls. The magnitude of the terminal FM sweep in spontaneous calls was not correlated with age (mean 13.5 +/- 2 kHz). Values for induced calls slightly increased with age from 11 +/- 2 to 13 +/- 2 kHz. The emission rate of induced CF-FM signals increased with age from values of 2.5 +/- 2 to 17 +/- 5 pulses/s. Values for spontaneously emitted calls were 4.4 +/- 3 and 9 +/- 4.5 pulses/s, respectively. Doppler-shift compensation, as tested in the pendulum task, emerged during the 4th postnatal week in young bats that were capable of very brief active flights, but before the time of active foraging outside the cave.

Acoustic Stimulation↗
PubMed2001

Laminar analysis of inhibition in the gerbil primary auditory cortex.

Physiological and immunocytochemical evidence indicates a layer-dependent organization of inhibitory circuits in the neocortex. To investigate the contribution of GABAergic inhibition to frequency tuning in the different cortical layers, we recorded single and multiple units in near-radial penetrations before and during iontophoretic application of the GABA(A)-receptor antagonist bicuculline in the auditory cortex of the lightly anesthetized gerbil. Bicuculline generally increased the spontaneous rate and enhanced and prolonged onset activity. Application of bicuculline often resulted in a shift of best frequency and a decrease of threshold (5.5 dB). A broadening of the frequency tuning evident by lower Q40dB values was observed in 63% of the units. In units with multipeaked tuning curves or clearly separated response areas, bicuculline application removed the inhibitory regions and created single-peaked tuning curves. The influence of bicuculline on the receptive field size was not significantly layer-specific but tended to be most pronounced in layers V and VI. In layer VI, we frequently found "silent" neurons that responded to sound only when GABAergic inhibition was antagonized. From the analysis of postembedding GABA immunocytochemistry, the proportion of GABAergic neurons was found to be maximal in layers I and V, and the number of GABAergic perisomatic puncta (axon terminals) on cell somata peaked in layer V.

Animals↗
PubMed2001

Components of the 2f(1)-f(2) distortion-product otoacoustic emission in a moth.

The noctuoid moth Empyreuma affinis has a simple tympanal organ with only two receptor cells attached to the eardrum. As in vertebrates, the growth of distortion-product otoacoustic emissions (DPOAE) with increasing stimulus level is characterized by two distinct components. An initial increase of DPOAE level for f(2) levels in the range of 30-65 dB SPL is followed by a second steep growth of the DPOAE at f(2) levels above 65 dB SPL. Both components increase at a slope of about 2 dB/dB and the difference between both components was used to assess a mechanical gain of the tympanal organ of 17 dB (n=23). At around 65 dB SPL, a notch in the level function coincides with an abrupt phase change of up to 180 degrees. The sensitive component induced by f(2) levels below 65 dB SPL is selectively affected by application of ethyl ether and disappears more quickly than the high-level component during ongoing deterioration of the moth preparation.

Aging↗
PubMed2000

Mechanical adaptations for echolocation in the cochlea of the bat Hipposideros lankadiva.

The cochlear mechanics of bats with long constant-frequency components in their echolocation calls are sharply tuned to the dominant second harmonic constant frequency. Hipposiderid bats employ a shorter constant-frequency call component whose frequency is less stable than in long-constant-frequency bats. To investigate to what degree cochlear mechanics in hipposiderid bats are already specialized for the processing of constant frequencies, we recorded distortion-product otoacoustic emissions in Hipposideros lankadiva. Iso-distortion threshold curves for the 2f1-f2 distortion-product otoacoustic emission reveal a threshold maximum close to the second harmonic constant frequency, between 65.0 and 70.0 kHz, and a second insensitivity close to the first harmonic constant frequency. The group delay of the 2f1-f2 distortion is prolonged for both frequency ranges, indicating that a specialized cochlear resonance may act to absorb the constant-frequency call components. Compared to long-constant-frequency bats, the threshold maximum at the second harmonic constant frequency is less pronounced and the optimum cochlear frequency separation is larger. Distortion-product otoacoustic emission suppression tuning curves and neuronal tuning curves recorded from neurons in the cochlear nucleus display an increase of tuning sharpness close to the second harmonic constant-frequency range which is smaller than that reported for long-constant-frequency bats. Our data suggest that the cochlea of hipposiderid bats represents an intermediate state between that of non-specialized bats and long-constant-frequency bats.

Adaptation, Physiological↗
PubMed2000

Consequences of outer hair cell damage for otoacoustic emissions and audio-vocal feedback in the mustached bat.

The cochlea of the mustached bat is adapted to process ultrasonic echolocation signals. To assess the involvement of active sound amplification by outer hair cells in high-frequency hearing and in audio-vocal interaction, selective hair cell damage was induced by the antibiotic Amikacin. Amikacin preferentially damaged the first row of outer hair cells in the basal cochlear turn. The cochlear regions coding for the second-harmonic constant-frequency component of the echolocation call (CF2) at 61 kHz and for frequencies between 75 and 100 kHz were the most affected. This was reflected in an increase of mechanical thresholds obtained by measuring distortion-product otoacoustic emissions. During initial periods of minor hair cell damage, when thresholds had deteriorated by less than 40 dB, a sharp, mechanical, cochlear resonance that is responsible for enhanced tuning to 61 kHz was still measurable as a stimulus-frequency otoacoustic emission and its frequency decreased by 350 Hz. The persistence of the resonance suggests that passive structures like the tectorial or basilar membrane are important for generation of the resonance. Behaviorally, the bats reacted to the change in cochlear micromechanics with a decrease of their CF2 frequency by 360 Hz. After larger hair cell damage, when the cochlear resonance had disappeared, the bats vocalized only sparsely and the CF2 frequency increased by up to 2 kHz, which may correspond to a state without audiovocal feedback. This indicates that audio-vocal feedback in the nondamaged animal works to lower the call frequency.

Animals↗
PubMed2000

Evidence for multiple DPOAE components based upon group delay of the 2f(1)-f(2) distortion in the gerbil.

The cochlear delay of the 2f(1)-f(2) distortion product otoacoustic emission (DPOAE) was measured using the phase gradient method. With a constant f(2) and swept f(1), the resulting phase change of 2f(1)-f(2) was used to calculate the group delay for f(2) frequencies from 1 to 60 kHz. For f(2) frequencies between 2 and 60 kHz, the group delays were between 2.2 and 0.11 ms and continuously decreased for increasing f(2) and for increasing primary stimulus levels. For f(2) frequencies below 2 kHz, the group delay decreased to around 1 ms and was largely independent of stimulus level. The ratio curves resulting from the f(1) sweeps for high frequencies (f(2)16 kHz) displayed the typical mammalian shape with a peak in the level of 2f(1)-f(2) for a larger primary frequency separation (f(2)/f(1)1.15) and decreasing 2f(1)-f(2) level for smaller primary separation. In addition to this typical level maximum, for f(2) frequencies from about 1.8 to 16 kHz, the ratio curves displayed a second component in the form of an increase in the level of 2f(1)-f(2) for small primary separation at higher primary levels (level of f(2)30 dB SPL). For f(2) frequencies below 1.8 kHz, only the second component and no typical ratio peak as for higher f(2) could be observed and the associated group delay was always close to 0.8 ms. Several possible causes for this behavior are discussed, including different modes of DPOAE generation and modulation as well as changes in the nature of mechanical processing from base to apex in the gerbil cochlea. To evaluate the relative sensitivity of non-linear cochlear mechanics, an iso-distortion threshold curve was constructed from acoustical growth functions of the 2f(1)-f(2) DPOAE at optimum primary separation, by plotting the level of f(2) sufficient to evoke a distortion of -10 dB SPL as a function of f(2)2.5 kHz but failed to reflect the sensitivity for lower frequencies. This may be a consequence of more linear frequency processing in the apex.

Acoustic Stimulation↗
PubMed1999

Evolutionary adaptations of cochlear function in Jamaican mormoopid bats.

Mormoopid bat species have their echolocation system adapted to different hunting strategies. To study the corresponding mechanical properties of their inner ear, we measured distortion-product otoacoustic emissions to assess cochlear sensitivity and tuning. Mormoops blainvillii, Pteronotus macleayii and P. quadridens use frequency-modulated echolocation signals, sometimes preceded by a short narrowband signal component. Their distortion-product otoacoustic emission-threshold curves are most sensitive between 30 and 50 kHz and show no adaptation to the narrowband echolocation components. In contrast, the constant-frequency bat P. parnellii always uses long constant-frequency call components. Its inner ear is maximally sensitive at 62 kHz, the echo-frequency of the dominant constant-frequency component, and pronounced insensitivities at 61 and 93 kHz (CF2 and CF3 call frequency) are the major evolutionary change in comparison to its relatives. Furthermore, in P. parnellii, the optimum cochlear frequency separation is minimal at 62 and 93 kHz, associated with enhanced cochlear tuning, while for the other mormoopids there is no indication of enhanced tuning. The phylogeny of mormoopids, assessed by mitochondrial DNA analysis, shows a close relationship between the Pteronotus species. This suggests that major cochlear redesign, associated with the acquisition of echolocation-call specific cochlear processing in P. parnellii, has occurred within a relatively short evolutionary time scale.

Acoustic Stimulation↗
PubMed1999

Binaural influences on Doppler shift compensation of the horseshoe bat Rhinolophus rouxi.

The flying horseshoe bat Rhinolophus rouxi compensates for Doppler shifts in echoes of their orientation pulses. By lowering the frequency of subsequent calls the echo's constant frequency is stabilized at the so-called reference frequency centered in a narrow and sensitive cochlear filter. This audio-vocal behaviour is known as Doppler shift compensation. To investigate whether the bats depend on binaural cues when compensating, three animals were tested for compensation on a swing before and after unilateral deafening. In each case compensation was severely impaired by unilateral deafening. Individual animals' compensation amplitude was reduced to 28-48% of the preoperational compensation of a +1.8 kHz shift. Doppler shift compensation performance did not recover to control levels during the observed period of 24 h after surgery. In contrast, unilateral middle ear removal which induces a unilateral auditory threshold increase of 9-14 dB does not impair compensation performance on the swing. To mimick Doppler shifts in a fixed setup, the frequencies of recorded echolocation calls were experimentally shifted between 0 and +2 kHz and played back via earphones to six animals. The bats completely compensated the experimental shifts only as long as the interaural intensity difference of the playback did not exceed 20 dB. No animal compensated with monaural playback.

Adaptation, Physiological↗
PubMed1999

Neuronal response to cochlear distortion products in the anteroventral cochlear nucleus of the gerbil.

To receive information on the intracochlear magnitude and propagation properties of cochlear distortion products, the neuronal response of AVCN cells to distortion stimuli and the acoustical correlates in the ear canal (DPOAE) were measured for frequencies between 0.7 and 45.3 kHz. Comparison of the growth of neuronal response to a distortion stimulus and the neuronal rate-versus-level function for an externally applied pure tone of equal frequency allowed for an assessment of the intracochlear magnitude of the distortions. AVCN neurons with a characteristic frequency (CF) > 1.8 kHz started to respond to the intracochlear distortion stimulus, at primary levels for which the ear-canal level of the corresponding DPOAE was close to the pure-tone threshold of the units. This finding suggests that transmission of sound energy is comparable in the forward and reverse direction, and that mechanical distortions of the cochlea are fully encoded by neurons in the AVCN. For neurons with a CF < 1.8 kHz, the intracochlear magnitude of the distortion stimulus appeared to be about 15 to 30 dB higher than the corresponding DPOAE, at the threshold of neuronal response. This discrepancy between intracochlear magnitude of cochlear distortions and their acoustical expression may be explained by high-pass filter action of the middle ear during DPOAE re-emission from the cochlea. A contribution to the observed discrepancy of the type of distortion (cubic versus quadratic) used as stimulus, and possible differences in mechanical frequency processing between the apex and base of the gerbil cochlea, are also discussed. The delay of the neuronal response to an intracochlear distortion stimulus was on average 1.1 ms longer than the neuronal delay to an external pure tone of equal frequency and intensity, most likely stemming from the activation delay of the DPOAE generating mechanisms.

Acoustic Stimulation↗
PubMed1999

Micromechanical responses to tones in the auditory fovea of the greater mustached bat's cochlea.

An extended region of the greater mustached bat's cochlea, the sparsely innervated (SI) zone, is located just basally to the frequency place of the dominant 61-kHz component of the echolocation signal (CF2). Anatomic adaptations in the SI zone are thought to provide the basis for cochlear resonance to the CF2 echoes and for the extremely sharp tuning throughout the auditory system that allows these bats to detect Doppler shifts in the echoes caused by insect wing beat. We measured basilar membrane (BM) displacements in the SI zone with a laser interferometer and recorded acoustic distortion products at the ear drum at frequencies represented in the SI zone. The basilar membrane in the SI region was tuned both to its characteristic frequency (62-72 kHz) and to the resonance frequency (61-62 kHz). With increasing stimulus levels, the displacement growth functions are compressive curves with initial slopes close to unity, and their properties are consistent with the mammalian cochlear amplifier working at high sound frequencies. The sharp basilar membrane resonance is associated with a phase lag of 180 degrees and with a shift of the peak resonance to lower frequencies for high stimulus levels. Within the range of the resonance, the distortion product otoacoustic emissions, which have been attributed to the resonance of the tectorial membrane in the SI region, are associated with an abrupt phase change of 360 degrees. It is proposed that a standing wave resonance of the tectorial membrane drives the BM in the SI region and that the outer hair cells enhance, fine tune, and control the resonance. In the SI region, cochlear micromechanics appear to be able to work in two different modes: a conventional traveling wave leads to shear displacement between basilar and tectorial membrane and to neuronal excitation for 62-70 kHz. In addition, the SI region responds to 61-62 kHz with a resonance based on standing waves and thus preprocesses signals which are represented more apically in the CF2 region of the cochlea.

Acoustic Stimulation↗
PubMed1997

Sound emission from cochlear filters and foveae--does the auditory sense organ make sense?

Sense organs filter relevant information from a broad background of physical interactions and discard possible perceptual input that has not proven useful during the course of biological evolution. Sense organs not only limit the access to physical reality, under certain conditions they have a life of their own and produce responses even in the absence of physical stimulation. As a perfect example, the inner ear, the cochlea, in addition to detecting incoming sound waves, it also is capable of producing sound energy. Such "active" processes, however, seem to be necessary to push detection thresholds close to physical limits. The price that has to be paid are "cochlear artifacts" like otoacoustic emissions. In the following, measurement of sound that is emitted by the ear will be introduced as a noninvasive means to assess cochlear function and to help to unravel the mechanical interaction between sensory cells and supporting structures that ultimately leads to sensitive and sharply tuned auditory perception. One focus will be on the cochlea of echo-locating bats that use audition as the main window of perception to their environment and therefore have highest demands on cochlear performance.

Animals↗
PubMed1997

Acoustical and electrical biasing of the cochlea partition. Effects on the acoustic two tone distortions f2-f1 and 2f1-f2.

Low frequency acoustical biasing of the cochlear partition with 5 Hz tones produces phase correlated changes of the acoustic two-tone distortions 2f1-f2 and f2-f1. Pronounced changes of f2-f1 and only small changes of 2f1-f2 for lower bias tone levels indicate that there is a close relation between changes in the difference tone f2-f1 and changes in the operating point of the cochlear amplifier (Frank and Kössl, 1996). To further investigate this relationship, the cochlear partition was additionally biased by current injection into the scala media of the gerbil. The injection of low frequency (5 Hz) AC currents (max. 1.3 microA) has a similar effect to that caused by low frequency tones in that both produce phase correlated changes of the two distortions (so-called biasing patterns), with stronger effects on f2-f1. For bias tone levels of about 105 dB SPL and current values of 1.3 microA, the effects are approximately of the same size. A change in the f2-f1 biasing pattern that can be found for increasing bias tone levels can also be seen for increasing primary levels. Changing the setpoint of the cochlear amplifier through the injection of DC current into the scala media during acoustical biasing of the cochlear partition produces the same changes of f2-f1 biasing patterns as increasing the primary levels. This indicates that the operating point of the outer hair cells that respond to the primary tones is not only influenced by low frequency biasing stimuli but also by shifts with increasing primary levels.

Acoustic Stimulation↗
PubMed1996

The acoustic two-tone distortions 2f1-f2 and f2-f1 and their possible relation to changes in the operating point of the cochlear amplifier.

Acoustic two-tone distortions are generated during non-linear mechanical amplification in the cochlea. Generation of the cubic distortion 2f1-f2 depends on asymmetric components of a non-linear transfer function whereas the difference tone f2-f1 relies on symmetric components. Therefore, a change of the operating point and hence the symmetry of the cochlear amplifier could be strongly reflected in the level of the f2-f1 distortion. To test this hypothesis, low-frequency tones (5 Hz) were used to bias the position of the cochlear partition in the gerbil. Phase-correlated changes of f2-f1 occurred at bias tone levels where there were almost no effects on 2f1-f2. Higher levels of the bias tone induced pronounced changes of both distortions. These results are qualitatively in good agreement with the results of a simulation in which the operating point of a Boltzman function was shifted. This function is similar to those used to describe outer hair cell (OHC) transduction. To influence OHC motility, salicylate was injected. It caused a decrease of the 2f1-f2 level and an increase in the level of f2-f1. Such reciprocal changes of both distortions, again, can be interpreted in terms of a shift of the operating point of the cochlear amplifier along a non-linear transfer characteristic. To directly influence the cochlear amplifier, DC current was injected into the scala media. Large negative currents (> -2 microA) caused a pronounced decrease of 2f1-f2 (> 15 dB) and positive currents had more complex effects with increasing and/or decreasing 2f1-f2 distortion level. The effects were time and primary level dependent. Changes of f2-f1 for DC currents > magnitude of mu 2A were in most cases larger compared to 2f1-f2 and reversed for certain primary levels. The current effects probably result from a combination of changing the endocochlear potential and shifting the operating point along a non-linear transfer function.

Acoustic Stimulation↗
PubMed1996

Acoustic distortion products from the cochlea of the blind African mole rat, Cryptomys spec.

The measurement of distortion-product otoacoustic emissions is a noninvasive method that can be used for assessing the sensitivity and the frequency tuning of nonlinear cochlear mechanics. During stimulation with two pure tones f1 and f2, the acoustic 2f1-f2 distortion was recorded in the ear canal of Cryptomys spec. to study specializations in cochlear mechanics that could be associated with the presence of a frequency expanded cochlear region between 0.8-1 kHz. In addition, a distortion threshold curve was obtained which describes relative threshold of nonlinear cochlear mechanics. Sensitive distortion thresholds could be measured for stimulus frequencies between 0.4 to 18 kHz with a broad minimum between 0.75 to 2.5 kHz. The distortion threshold curve extends to higher frequencies than previous neuronal data indicated. As a measure of mechanical tuning sharpness in the cochlea, suppression tuning curves of 2f1-f2 were recorded. The tuning curves reflected the typical mammalian pattern with shallow low frequency and steep high frequency slopes. Their tuning sharpness was poor with Q10dB values between 0.3 and 1.88. In the range of the frequency expanded region, the Q10dB values were below 0.5. This finding emphasizes that the presence of frequency expansion does not necessarily lead to enhanced mechanical tuning in the cochlea and one has to consider if in certain bat species with cochlear frequency expansion and particularly sharp cochlear tuning, the two phenomena may not be interlinked.

Acoustic Stimulation↗