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Biomedical subjects

Y de Ribaupierre

Publications and source records attributed to Y de Ribaupierre.

At least 19 recordsLinked to original sources

Inhibition of bicarbonate transport protects embryonic heart against reoxygenation-induced dysfunction.

It has not been well established whether the mechanisms participating in pH regulation in the anoxic-reoxygenated developing myocardium resemble those operating in the adult. We have specially examined the importance of Na+/H+ exchange (NHE) and HCO3-dependent transports in cardiac activity after changes in extracellular pH (pHo). Spontaneously contracting hearts isolated from 4-day-old chick embryos were submitted to single or repeated anoxia (1 min) followed by reoxygenation (10 min). The chronotropic, dromotropic and inotropic responses of the hearts were determined in standard HCO3- buffer at pHo 7.4 and at pHo 6.5 (hypercapnic acidosis). In distinct experiments, acidotic anoxia preceded reoxygenation at pHo 7.4. NHE was blocked with amiloride derivative HMA (1 micro mol/l) and HCO3-dependent transports were inactivated by replacement of HCO3 or blockade with stilbene derivative DIDS (100 micro mol/l). Anoxia caused transient tachycardia, depressed mechanical function and induced contracture. Reoxygenation temporarily provoked cardiac arrest, atrio-ventricular (AV) block, arrhythmias and depression of contractility. Addition of DIDS or substitution of HCO3 at pHo 7.4 had the same effects as acidosis per se, i.e. shortened contractile activity and increased incidence of arrhythmias during anoxia, prolonged cardioplegia and provoked arrhythmias at reoxygenation. Under anoxia at pHo 6.5/reoxygenation at pHo 7.4, cardioplegia, AV block and arrhythmias were all markedly prolonged. Interestingly, in the latter protocol, DIDS suppressed AV block and arrhythmias during reoxygenation, whereas HMA had no effect. Thus, intracellular pH regulation in the anoxic-reoxygenated embryonic heart appears to depend predominantly on HCO3 availability and transport. Furthermore, pharmacological inhibition of anion transport can protect against reoxygenation-induced dysfunction.

4,4'-Diisothiocyanostilbene-2,2'-Disulfonic Acid↗

Effects of verapamil and ryanodine on activity of the embryonic chick heart during anoxia and reoxygenation.

Perturbations of the trans-sarcolemmal and sarcoplasmic Ca2+ transport contribute to the abnormal myocardial activity provoked by anoxia and reoxygenation. Whether Ca2+ pools of the extracellular compartment and sarcoplasmic reticulum (SR) are involved to the same extent in the dysfunction of the anoxic-reoxygenated immature heart has not been investigated. Spontaneously contracting hearts isolated from 4-day-old chick embryos were submitted to repeated anoxia (1 min) followed by reoxygenation (5 min). Heart rate, atrioventricular propagation velocity, ventricular shortening, velocities of contraction and relaxation, and incidence of arrhythmias were studied, recorded continuously. Addition of verapamil (10 nM), which blocks selectively sarcolemmal L-type Ca2+ channels, was expected to protect against excessive entry of extracellular Ca2+, whereas addition of ryanodine (10 nM), which opens the SR Ca2+ release channel, was expected to increase cytosolic Ca2+ concentration. Verapamil (a) had no dromotropic effect by contrast to adult heart, (b) attenuated ventricular contracture induced by repeated anoxia, (c) shortened cardioplegia induced by reoxygenation, and (d) had remarkable antiarrhythmic properties during reoxygenation specially. On the other hand, ryanodine potentiated markedly arrhythmias both during anoxia and at reoxygenation. Thus despite its immaturity, the SR seems to be functional early in the developing chick heart and involved in the reversible dysfunction induced by anoxia-reoxygenation. Moreover, Ca2+ entry through L-type channels appears to worsen arrhythmias especially during reoxygenation. These findings show that the Ca2+-handling systems involved in irregular activity in immature heart, such as the embryonic chick heart, may differ from those in the adult.

Animals↗

Glucose is arrhythmogenic in the anoxic-reoxygenated embryonic chick heart.

Unlike in adult heart, embryonic myocardium works at low PO2 and depends preferentially on glucose. Therefore, activity of the embryonic heart during anoxia and reoxygenation should be particularly affected by changes in glucose availability. Hearts excised from 4-d-old chick embryos were submitted in vitro to strictly controlled anoxia-reoxygenation transitions at glucose concentrations varying from 0 to 20 mmol/L. Spontaneous and regular heart contractions were detected optically as movements of the ventricle wall and instantaneous heart rate, amplitude of contraction, and velocities of contraction and relaxation were determined. Anoxia induced transient tachycardia and rapidly depressed contractile activity, whereas reoxygenation provoked a temporary and complete cardioplegia (oxygen paradox). In the presence of glucose, atrial rhythm became irregular during anoxia and chaotic-periodic during reoxygenation. The incidence of these arrhythmias depended on duration of anoxia, and no ventricular ectopic beats were observed. Removal of glucose or blockade of glycolysis suppressed arrhythmias. These results show similarities but also differences with respect to the adult heart. Indeed, glucose 1) delayed and anoxic contractile failure, shortened the reoxygenation-induced cardiac arrest, and improved the recovery of contractile activity; 2) attenuated stunning at 20 mmol/L but worsened it at 8 mmol/L; and 3) paradoxically, was arrhythmogenic during anoxia and reoxygenation, especially when present at the physiologic concentration of 8 mmol/L. The last named phenomenon seems to be characteristic of the young embryonic heart, and our findings underscore that fluctuations of glycolytic activity may play a role in the reactivity of the embryonic myocardium to anoxiareoxygenation transitions.

Animals↗

Morphology and spatial distribution of corticothalamic terminals originating from the cat auditory cortex.

In this paper we studied the morphology and spatial distribution of corticothalamic axons and terminals originating from the auditory cortical fields of the cat. The anterograde tracer biocytin was injected at electrophysiologically characterized loci in the primary (AI) (N = 2), anterior (AAF) (N = 1), posterior (PAF) (N = 1) and secondary (AII) (N = 2) auditory fields. In all cases, two different types of labeled terminals were found in the auditory thalamus: small spherical endings (1-2 microns) and giant, finger-like endings (5-10 microns). After biocytin injections in AI and AAF, the majority of anterogradely labeled axons terminated in the rostral half of the pars lateralis (LV) of the ventral division of the medial geniculate body (vMGB). In LV, the corticothalamic axons ramified profusely, giving rise to dense terminal fields forming well delineated curved stripes, with small spherical endings. Additional terminal fields formed by small endings were observed in the medial division of the medial geniculate body (mMGB). Giant endings were observed in a small area in the dorsal nucleus (D) of the dorsal division of the medial geniculate body (dMGB), near its border with the vMGB. PAF projections were located in the caudal half of vMGB and in mMGB, where only small terminals were found. Giant endings were seen in the superficial part of dMGB emerging from labeled corticothalamic axons oriented in parallel to the dorsal surface of the MGB. Projections from AII gave rise to a main terminal field of small endings in D; a second terminal field consisting of giant endings intermingled with small endings was found in the deep dorsal nucleus (DD) of dMGB. We conclude that small terminals serve the feedback projection to the thalamic nucleus from which the injected cortical field receives its main input, whereas giant terminals cross the borders between the parallel ascending auditory pathways.

Acoustic Stimulation↗

Changes of single unit activity in the cat's auditory thalamus and cortex associated to different anesthetic conditions.

Single unit spike trains were recorded in the auditory cortex (n = 78) and in the auditory thalamus (n = 55) of nitrous oxide anesthetized cats. The electrophysiological activity was studied before and during the application of pentobarbital (P, 7 mg/kg), ketamine (K, 12 mg/kg) and a mixture of these anesthetics (KP). The units were characterized during the spontaneous and acoustically driven activity ('white' noise and pure tone bursts). For the majority of cortical (61%) and thalamic (83%) units both drugs tended to decrease the spontaneous firing rate, but affected differently its time structure: P tended to increase the average size of burst discharges, whereas K and KP tended to decrease it. In the cortex the peak firing rate evoked by 'white' noise tended to be decreased, whereas stronger excitatory responses were observed in the thalamus after injection of K or KP. The overall effect of the anesthetics during stimulation by pure tones was an increase in tonal selectivity due to a decrease in the response bandwidth. The response pattern to tones was also sometimes affected by the drugs. The direct evidence reported here for significant alterations of the discharge properties of auditory neurons in the thalamus and cortex resulting from low dose administration of K and/or P emphasizes difficulties in comparing data derived from experiments conducted in various conditions of anesthesia or in the awake state.

Acoustic Stimulation↗

Several neuronal and axonal types form long intrinsic connections in the cat primary auditory cortical field (AI).

Intrinsic connections in the cat primary auditory field (AI) as revealed by injections of Phaseolus vulgaris leucoagglutinin (PHA-L) or biocytin, had an anisotropic and patchy distribution. Neurons, labelled retrogradely with PHA-L were concentrated along a dorsoventral stripe through the injection site and rostral to it; the spread of rostrally located neurons was greater after injections into regions of low rather than high characteristic frequencies. The intensity of retrograde labelling varied from weak and granular to very strong and Golgi-like. Out of 313 Golgi like retrogradely labelled neurons 79.6% were pyramidal, 17.2% multipolar, 2.6% bipolar, and 0.6% bitufted; 13.4% were putatively inhibitory, i.e. aspiny or sparsely spiny multipolar, or bitufted. Individual anterogradely labelled intrinsic axons were reconstructed for distances of 2 to 7 mm. Five main types were distinguished on the basis of the branching pattern and the location of synaptic specialisations. Type 1 axons travelled horizontally within layers II to VI and sent collaterals at regular intervals; boutons were only present in the terminal arborizations of these collaterals. Type 2 axons also travelled horizontally within layers II to VI and had rather short and thin collateral branches; boutons or spine-like protrusions occurred in most parts of the axon. Type 3 axons travelled obliquely through the cortex and formed a single terminal arborization, the only site where boutons were found. Type 4 axons travelled for some distance in layer I; they formed a heterogeneous group as to their collaterals and synaptic specializations. Type 5 axons travelled at the interface between layer VI and the white matter; boutons en passant, spine-like protrusions, and thin short branches with boutons en passant were frequent all along their trajectory. Thus, only some axonal types sustain the patchy pattern of intrinsic connectivity, whereas others are involved in a more diffuse connectivity.

Animals↗

Auditory corticocortical interconnections in the cat: evidence for parallel and hierarchical arrangement of the auditory cortical areas.

The origin and laminar arrangement of the homolateral and callosal projections to the anterior (AAF), primary (AI), posterior (PAF) and secondary (AII) auditory cortical areas were studied in the cat by means of electrophysiological recording and WGA-HRP tracing techniques. The transcallosal projections to AAF, AI, PAF and AII were principally homotypic since the major source of input was their corresponding area in the contralateral cortex. Heterotypic transcallosal projections to AAF and AI were seen, originating from the contralateral AI and AAF, respectively. PAF received heterotypic commissural projections from the opposite ventroposterior auditory cortical field (VPAF). Heterotypic callosal inputs to AII were rare, originating from AAF and AI. The neurons of origin of the transcallosal connections were located mainly in layers II and III (70-92%), and less frequently in deep layers (V and VI, 8-30%). Single unit recordings provided evidence that both homotypic and heterotypic transcallosal projections connect corresponding frequency regions of the two hemispheres. The regional distribution of the anterogradely labeled terminals indicated that the homotypic and heterotypic auditory transcallosal projections are reciprocal. The present data suggest that the transcallosal auditory interconnections are segregated in 3 major parallel components (AAF-AI, PAF-VPAF and AII), maintaining a segregation between parallel functional channels already established for the thalamocortical auditory interconnections. For the intrahemispheric connections, the analysis of the retrograde tracing data revealed that AAF and AI receive projections from the homolateral cortical areas PAF, VPAF and AII, whose neurons of origin were located mainly in their deep (V and VI) cortical layers. The reciprocal interconnections between the homolateral AAF and AI did not show a preferential laminar arrangement since the neurons of origin were distributed almost evenly in both superficial (II and III) and deep (V and VI) cortical layers. On the contrary, PAF received inputs from the homolateral cortical fields AAF, AI, AII and VPAF, originating predominantly from their superficial (II and III) layers. The homolateral projections reaching AII originated mainly from the superficial layers of AAF and AI, but from the deep layers of VPAF and PAF. The laminar distribution of anterogradely labeled terminal fields, when they were dense enough for a confident identification, was systematically related to the laminar arrangement of neurons of origin of the reciprocal projection: a projection originating from deep layers was associated with a reciprocal projection terminating mainly in layer IV, whereas a projection originating from superficial layers was associated with a reciprocal projection terminating predominantly outside layer IV.(ABSTRACT TRUNCATED AT 400 WORDS)

Acoustic Stimulation↗

Corticofugal modulation of the information processing in the auditory thalamus of the cat.

Single unit activity of 355 cells was recorded in the auditory thalamus of anesthetized cats before, during, and after the inactivation by cooling of the ipsilateral primary auditory cortex (AI). Most of the units (n = 288) showed similar functional characteristics of firing before and after the cryogenic blockade of AI. The spontaneous firing rate remained unchanged by cooling in 20% of the units and decreased in the majority of them (60%). In some regions, i.e. dorsal division of the medial geniculate body (MGB), lateral part of the posterior group of the thalamus, and auditory sector of the reticular nucleus of the thalamus, the maximum firing rate evoked by white noise bursts was generally affected by cooling in the same direction and to the same extent as the spontaneous activity. Units in the ventral division of MGB showed a characteristic increase of signal-to-noise ratio during cortical cooling. The corticofugal modulation led to the appearance or disappearance of the best frequency of tuning in 51 units and changed it by more than 0.5 octave in 34 units. The bandwidths of different response patterns to pure tones stimulation were used to define a set of functional properties. During cryogenic blockade of AI, two cortically modulated sub-populations of units were usually distinguished that exhibited changes for a given functional property. The complexity and diversity of the effects of cortical inactivation suggest that the corticothalamic projection may be the support for selective operations such as an adaptive filtering of the incoming acoustic signal at the thalamic level adjusted as a function of cortical activity.

Acoustic Stimulation↗

Discharge properties of single units in auditory part of reticular nucleus of thalamus in cat.

1. Extracellular recordings from 153 single units were obtained in the auditory part of the reticular nucleus (RE) of the thalamus of anesthetized cats. 2. In absence of acoustic stimulation, the majority of RE units (75%) had a spontaneous activity characterized by bursts of spikes lasting over 30 ms, alternating with isolated discharges; in contrast, only 30% of units in the medial geniculate body (MGB) showed these long bursts during spontaneous activity. 3. For a majority of RE units (65%), activity evoked by noise bursts consisted in complex response patterns with multiple excitatory and/or inhibitory components. For this stimulus, most units (79%) showed nonmonotonic rate-intensity functions, and median response latency to the first excitatory component was 16 ms, which is slightly longer than that obtained in the anterior part of the ventral division of the MGB for the same conditions. 4. Frequency tuning to pure tones in RE was usually broad: the median value of the width of response ranges was five octaves in RE, as compared to two octaves in the ventral division of the MGB and three octaves in the medial division of the MGB. Responses to tones were complex, usually characterized by frequent changes of response pattern with frequency. No preferential arrangement of units as a function of their best frequency was found along a rostrocaudal or a dorsolateral to ventromedial axis. 5. The present study demonstrates that units in the auditory part of RE have discharge properties clearly different from the main auditory thalamic nucleus, the MGB.

Acoustic Stimulation↗

Tonotopic organization in the medial geniculate body (MGB) of lightly anesthetized cats.

In the medial geniculate body (MGB) of nitrous oxide anesthetized cats, the pars lateralis (PL) was the only nucleus to show a clear topographic arrangement of its neurons according to their characteristic frequency (CF). When compared to barbiturate anesthetized cats (Imig and Morel 1985a), the tonotopic organization in PL appeared less strict and was characterized by a significant local CF disparity. Furthermore, the degree of tonotopic organization varied along the rostrocaudal axis of the nucleus: it was lower in its caudal than in its rostral half. In the pars ovoidea, the rostral half of the pars magnocellularis (PM) and the suprageniculate nucleus, CF sequences and quantitative evaluations of the tonotopicity indicated the presence of some degree of tonotopic organization which was lower than in PL. No such organization was observed in the caudal part of PM nor in the ventrolateral nucleus, while in the dorsal nucleus, the proportion of tone-responding units was too low for a significant analysis.

Acoustic Stimulation↗

Functional properties and interactions of neuron pairs simultaneously recorded in the medial geniculate body of the cat.

Pairs of single units were recorded simultaneously from the same microelectrode in the medial geniculate body (MGB) of nitrous oxide anaesthetized cats. Crosscorrelation histograms of each spike train pairs were computed and distributed in five main classes, interpreted in terms of functional interaction between neighbouring neurons. Overall 40% of the 950 recorded pairs showed independent firing during spontaneous activity. Histogram shapes suggestive of simple functional connectivity were found in 38% of the pairs: common inputs (20%), excitatory (15%) and inhibitory (3%) synaptic links. Complex and 'borderline' histogram shapes made up the rest of the sample (22%). Subdivisions of the MGB had the same types and proportions of interaction classes. Several of these interactions showed, in statistical terms only, preferential associations with some other descriptive features of neuronal behaviour: pattern of spontaneous firing, sensitivity to acoustic stimuli, and to a lesser extent characteristic frequency, response latency, response pattern and width of the response range. Simultaneous recording of nearby neuron pairs allowed to test if a given functional property is spatially organized or randomly distributed in the MGB. Response latency, width of the response range behaved as if randomly distributed, whereas binaural interactions and characteristic frequency as if spatially organized.

Acoustic Stimulation↗

Micromeasurement of total and regional CO2 productions in the one-day-old chick embryo.

A conductometric micromethod combined with image analysis system has been developed allowing to determine the CO2 production within 'two-dimensional' tissues, i.e., flat and thin cell layers or epithelial sheets. The preparation was mounted into an airtight chamber separated in two compartments by a thin silicone membrane permeable to gases. The lower compartment contained the nutritive medium and the preparation. The upper compartment and a conductivity measuring capillary connected in series were perfused with a solution of Ba(OH)2. The CO2 produced by the tissue precipitated as BaCO3 and the resulting decrease of electrical conductivity was linearly related to the total CO2 production. In addition, the pattern of CO2 production was directly observable as the BaCO3 crystals formed upon the silicone membrane over the regions which produced CO2. The spatial distribution of the crystals was quantified by video image processing and the regional CO2 production evaluated with a spatial resolution of 100 microns. This new microtechnique was originally developed to study the CO2 production in the early chick blastoderm which is a disc 1-5 cells thick. At the stage of young neurula the CO2 production was found to be 235 +/- 37 nmol.h-1 (mean +/- SD; n = 10) per blastoderm and large variations of local CO2 production were detected from one region to another (from 0.6 to 6.5 nmol.h-1.mm-2). These results indicate a high metabolic and functional differentiation of cells within the blastoderm. The possible applications and improvements of such a microtechnique are discussed.

Animals↗

Evoked mechanical responses of isolated cochlear outer hair cells.

Intracellular current administration evokes rapid, graded, and bidirectional mechanical responses of isolated outer hair cells from the mammalian inner ear. The cells become shorter in response to depolarizing and longer in response to hyperpolarizing currents in the synaptic end of the cell. The cells respond with either an increase or decrease in length to transcellular alternating current stimulation. The direction of the movement with transcellular stimuli appears to be frequency dependent. Iontophoretic application of acetylcholine to the synaptic end of the cell decreases its length. The microarchitecture of the organ of Corti permits length changes of outer hair cells in a manner that could significantly influence the mechanics of the cochlear partition and thereby contribute to the exquisite sensitivity of mammalian hearing.

Acetylcholine↗

Neural correlates of cubic difference tones in the medial geniculate body of the cat.

Single unit responses to the cubic difference tone CDT (2f1 - f2 = CF) and the difference tone DT (f2 - f1 = CF) were studied in the medial geniculate body (MGB) of the cat. Out of 66 units tested with CDT stimuli and having characteristic frequencies (CF) below 10 kHz, 77% gave a response to the two-tone combination stimulus. The component tones when presented alone evoked no responses, or in some cases a response pattern that was different from the one observed for the combination tone. The CDT response pattern was always similar to that seen for a pure tone at the CF. The threshold of response for the CDT was 10-70 dB higher than for a pure tone stimulus at the CF. The few units which were phase-locked could be synchronised with the CF, CDT, or DT, depending on the particular stimulus conditions. The index of synchrony was in many cases found to be higher for CDT responses than for a pure tone at CF.

Animals↗

Intensity functions of single unit responses to tone in the medial geniculate body of cat.

Extracellular spike activity was recorded from single units in the medial geniculate body (MGB) of nitrous oxide anaesthetized cats. The responses of 291 units to tone bursts at the characteristic frequency (CF) were studied as a function of stimulus intensity, covering a range from 10 to 100 dB SPL. The proportion of MGB units characterized by a monotonic or a non-monotonic discharge rate--intensity function was 26% and 74%, respectively. In addition, changes of response latency as a function of tone levels were demonstrated to be either monotonic (38% of units) or non-monotonic (62% of units). One third Of MGB units showed a change of response pattern with increasing intensities, in similar proportion towards either prevailing excitatory or inhibitory components. The monotonic units tended to differ from non-monotonic ones in addition to their intensity function by showing shorter response latencies, a higher response probability to broad-band stimuli and simpler response patterns. The mean dynamic range of the monotonic unit population was 60 dB, with thresholds ranging from 10 to 90 dB SPL; most discharge rate--intensity functions did not saturate at sound levels of 100 dB SPL. In the population of non-monotonic units, the 'best' intensity, defined as th intensity giving the strongest response, ranged between 10 and 100 dB SPL. The present results suggest that the intensity could be signaled by the mean firing rate of a restricted population of monotonic units or place coded by the distribution of maximally activated non-monotonic units which are broadly tuned to different intensities.

Acoustic Stimulation↗

Measurement of hand tremor induced by industrial exposure to metallic mercury.

Hand tremor due to industrial exposure to metallic mercury vapour was recorded in 26 exposed and 25 non-exposed male workers by an accelerometer attached to the dorsum of the hand. The subjects were instructed to hold the hand and the forearm in the same position first without and then with a load of 1250 g supported by the hand. Analysis of the records showed that the highest peak frequencies (HPF) (the frequency corresponding to the highest acceleration) were higher in the exposed men than in the controls and were related to the duration of exposure to mercury and to age. The changes in HPF between rest and load were again higher in the exposed men than in the controls and again related to the duration of exposure and to age. The second moment (M2) (an index taking into account the whole recorded spectrum) was similar in the exposed men and controls at rest. The changes in M2 between rest and load were higher in the exposed men than in the controls but were related to duration of exposure and to the biological measurements (loge of mercury in urine or blood) and not to age. These neurophysiological impairments might result from the tendency of metallic mercury to accumulate in the cerebellum and the basal ganglia. These results are consistent with the hypothesis that metallic mercury, even at concentrations probably below the current TLV-TWA of 0.05 mg/m3, can lead to neurological disorders.

Adult↗

In situ recording of the mechanical behaviour of cells in the chick embryo.

The changes of viscosity in different regions of the intact living early chick embryo were recorded by a new sensitive and non-invasive technique. In the heart, the very first yet invisible mechanical activity has been clearly detected and its further differentiation followed. In other regions, especially in the Hensen's node, spontaneous slow and oscillating mechanical activity has been observed. It has been shown that the undifferentiated embryonic tissue can as well respond to electrical, mechanical or chemical stimulations. The recorded changes have been interpreted as manifestations of the cell mechanical work i.e. active tension, cell shaping or movement. The technique represents a new tool for physiological and pharmacological studies of the undifferentiated embryonic cells and the early morphogenetic phenomena.

Animals↗

Neural coding of repetitive clicks in the medial geniculate body of cat.

The activity of 418 medial geniculate body (MGB) units was studied in response to repetitive acoustic pulses in 35 nitrous oxide anaesthetized cats. The proportion of MGB neurons insensitive to repetitive clicks was close to 30%. On the basis of their pattern of discharge, the responsive units were divided into three categories. The majority of them (71%), classified as "lockers', showed discharges precisely time-locked to the individual clicks of the train. A few units (8%), called "groupers', had discharges loosely synchronized to low-rate repetitive clicks. When the spikes were not synchronized, the cell had transient or sustained responses for a limited frequency range and was classified as a "special responder' (21%). Responses of "lockers' were time-locked up to a limiting rate, which varied between 10 and 800 Hz; half of the "lockers' had a limiting rate of locking equal to or higher than 100 Hz. The degree of entrainment, defined as the probability that each click evokes at least one spike, regularly decreases for increasing rates; on the other hand, the precision of locking increasing increases with frequency. The time jitter observed at 100 Hz might be as small as 0.2 ms and was 1.2 ms on average. The population of "lockers' can mark with precision the transients of complex sounds and has response properties still compatible with a temporal coding of the fundamental frequency of most animal vocalizations.

Animals↗