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1.
Auditory and vestibular functions of otolithic organs vary among vertebrate taxa. The saccule has been considered a major hearing organ in many fishes. However, little is known about the auditory role of the lagena in fishes. In this study we analyzed directional and frequency responses from single lagenar fibers of Dormitator latifrons to linear accelerations that simulate underwater acoustic particle motion. Characteristic frequencies of the lagenar fibers fell into two groups: 50 Hz and 80–125 Hz. We observed various temporal response patterns: strong phase-locking, double phase-locking, phase-locked bursting, and non-phase-locked bursting. Some bursting responses have not been previously observed in vertebrate otolithic nerve fibers. Lagenar fibers could respond to accelerations as small as 1.1 mm s–2. Like saccular fibers, lagenar fibers were directionally responsive and decreased directional selectivity with stimulus level. Best response axes of the lagenar fibers clustered around the lagenar longitudinal axis in the horizontal plane, but distributed in a diversity of axes in the mid-sagittal plane, which generally reflect morphological polarizations of hair cells in the lagena. We conclude that the lagena of D. latifrons plays a role in sound localization in elevation, particularly at high stimulus intensities where responses of most saccular fibers are saturated.Abbreviations BRA best response axis/axes - BS best sensitivity - CF characteristic frequency - CV coefficient of variation - DI directionality index - ISIH inter-spike interval histogram - PSTH peri-stimulus time histogram - SR spontaneous rate  相似文献   

2.
The otolithic end organs in the ears of teleost fishes play important roles in hearing. Although previous studies have shown that afferent fibers innervating otolithic organs are directionally sensitive to acoustic stimulation, no study has demonstrated that directionality of the otolithic afferent neurons derives directly from morphological polarity of the hair cells that they innervate. In this study we investigated whether or not there exists such a structure and function relationship in one of the otolithic organs, the saccule, by using intracellular and extracellular tracing, histochemistry, and confocal imaging techniques. We observed a variety of morphologies of dendritic terminals of saccular ganglion neurons. Arbor innervation areas of these saccular neurons ranged from 893 microm2 to 21,393 microm2, and the number of dendritic endings fell into a range between 10 and 54. We found that the response directionality of saccular ganglion neurons correlates significantly with the morphological polarization of the hair cells in the regions that they innervate. Therefore, we provide direct evidence to support the hypothesis that fish are able to encode directional information about a sound source, particularly in elevation, using arrays of hair cells in the otolithic organs that are oriented specifically along the sound propagation axis.  相似文献   

3.
Our previous studies have shown that the peripheral auditory system of the toadfish encodes the direction of a sound source. Here, we compare directional responses of peripheral saccular afferents, cells in the descending octaval nucleus (DON) of the medulla, and the torus semicircularis (TS) of the midbrain. Recording locations in the brain were labeled with neurobiotin to confirm the site. To compare directional responses among cells, we calculated an index [sharpening ratio (SR)] that weights the relative strength of responses to the best direction for that cell and to the adjacent stimulus angles tested. Unsharpened saccular afferents tend to have a cosinusoidal directional response pattern (DRP) with an expected SR of 0.87. In DON, more than 60% of the cells exhibited directional sharpening (defined as SR <0.8). In TS, more than 80% of the cells exhibited directional sharpening. We conclude that directional auditory sharpening first occurs in DON and some additional sharpening occurs in the ascending pathway to the midbrain, particularly in azimuth. The sharpening of directional selectivity is likely to be an important component of the neural computations underlying directional hearing.  相似文献   

4.
It is unknown whether the fish utricle contributes to directional hearing. Here, we report response properties of single utricular fibers in a teleost fish (Dormitator latifrons) to linear accelerations at various stimulus frequencies and axes. Characteristic frequencies ranged from 50–400 Hz (median=80 Hz), and best frequencies shifted from 50 to 250 Hz with stimulus level. Best sensitivity of utricular fibers was distributed from –70 to –40 dB re: 1 g (mean=–52 dB), which is about 30 dB less sensitive than saccular fibers. Q50% fell between 0.16 and 11.50 (mean=2.04) at 15 dB above threshold. We observed temporal response patterns of entrained phase-locking, double phase-locking, phase-locked bursting, and non-phase-locked bursting. Most utricular fibers were directionally selective with various directional response profiles, and directional selectivity was stimulus-level dependent. Horizontal best-response axes were distributed in a 152° range while mid-sagittal best-response axes were clustered around the fish longitudinal axis, which is consistent with the horizontal orientation of the utricle and morphological polarizations of utricular hair cells. Therefore, results of this study indicate that the utricle in this vertebrate plays an auditory role in azimuth and that utricular fibers extend the response dynamic range of this species in directional hearing.  相似文献   

5.
Vertebrate hair cell systems receive innervation from efferent neurons in the brain. Here we report the responses of octavolateral efferent neurons that innervate the inner ear and lateral lines in a teleost fish, Dormitator latifrons, to directional linear accelerations, and compare them with the afferent responses from the saccule, the main auditory organ in the inner ear of this species. Efferent neurons responded to acoustic stimuli, but had significantly different response properties than saccular afferents. The efferents produced uniform, omnidirectional responses with no phase-locking. Evoked spike rates increased monotonically with stimulus intensity. Efferents were more broadly tuned and responsive to lower frequencies than saccular afferents, and efferent modulation of the otolithic organs and lateral lines is likely more pronounced at lower frequencies. The efferents had wide dynamic ranges, shallow rate-level function slopes, and low maximum discharge rates. These findings support the role of the efferent innervation of the otolithic organs as part of a general arousal system that modulates overall sensitivity of the peripheral octavolateral organs. In addition, efferent feedback may help unmask biologically relevant directional stimuli, such as those emitted by a predator, prey, or conspecific, by reducing sensitivity of the auditory system to omnidirectional ambient noise.  相似文献   

6.
While particle motion is thought to directly stimulate the inner ear of most fish species, it is difficult to measure and might not be predictable from pressure measurements in a small tank. It is therefore important to replicate experiments conducted relative to pressure measurements using stimuli of known particle motion, to ensure that unmeasured components of the stimulus field do not produce misleading frequency response profiles. The frequency sensitivity of the inner ear of the plainfin midshipman fish, Porichthys notatus, in response to isopressure stimuli has been described. This study now examines the frequency and directional response properties of midshipman saccular afferents in response to whole-body displacements simulating acoustic particle motion. Best frequencies were distributed bimodally, with peaks at 50 Hz and 100 Hz. Most units had cosinusoidally shaped directional response profiles in the horizontal and vertical planes, though some units showed slight deviations from this pattern. A few units (probably saccular efferents) had omnidirectional directional response profiles and did not phase lock to the stimulus waveform. These results are consistent with responses of the midshipman saccular nerve to isopressure stimuli, and strengthen the hypothesis that the frequency sensitivity of the midshipman ear matches the frequency content of behaviorally relevant vocalizations.  相似文献   

7.
We studied the directionality of spike timing in the responses of single auditory nerve fibers of the grass frog, Rana temporaria, to tone burst stimulation. Both the latency of the first spike after stimulus onset and the preferred firing phase during the stimulus were studied. In addition, the directionality of the phase of eardrum vibrations was measured. The response latency showed systematic and statistically significant changes with sound direction at both low and high frequencies. The latency changes were correlated with response strength (spike rate) changes and were probably the result of directional changes in effective stimulus intensity. Systematic changes in the preferred firing phase were seen in all fibers that showed phaselocking (i.e., at frequencies below 500–700 Hz). The mean phase lead for stimulation from the contralateral side was approximately 140° at 200 Hz and decreased to approximately 100° at 700 Hz. These phaseshifts correspond to differences in spike timing of approximately 2 ms and 0.4 ms respectively. The phaseshifts were nearly independent of stimulus intensity. The phase directionality of eardrum vibrations was smaller than that of the nerve fibers. Hence, the strong directional phaseshifts shown by the nerve fibers probably reflect the directional characteristics of extratympanic pathways. Accepted: 23 November 1996  相似文献   

8.
This paper reviews and discusses several investigations of the peripheral neural code for the directional axis of acoustical particle motion in the saccule of two fishes: goldfish (Carassius auratus) and toadfish (Opsanus tau). Most saccular afferents are directional in the manner of hair cells, having a cosine-shaped directional response pattern. The saccular sensory epithelia are orientated almost vertically in a parasagittal plane. In the horizontal plane, these epithelia are orientated obliquely with respect to the midline. Hair-cell stereocilia project perpendicularly. Thus, directional response patterns of saccular afferents tend to be orientated in azimuth parallel to the orientation of the epithelia in the head. The oblique angle of the toadfish saccule is greater than that of the goldfish, and the range of best directions in the horizontal plane for each species reflects those differing orientations. The azimuth of acoustical particle motion could be computed by comparing the relative activation of the two saccules, as is the case for the ears of most terrestrial vertebrates. The spatial patterns of saccular hair-cell orientation of most fishes thus appear to have little function in azimuthal source location, but for toadfish are probably most important for determining the elevation of monopole sources.  相似文献   

9.
A striking feature of vestibular hair cells is the polarized arrangement of their stereocilia as the basis for their directional sensitivity. In mammals, each of the vestibular end organs is characterized by a distinct distribution of these polarized cells. We utilized the technique of post-fixation transganglionic neuronal tracing with fluorescent lipid soluble dyes in embryonic and postnatal mice to investigate whether these polarity characteristics correlate with the pattern of connections between the endorgans and their central targets; the vestibular nuclei and cerebellum. We found that the cerebellar and brainstem projections develop independently from each other and have a non-overlapping distribution of neurons and afferents from E11.5 on. In addition, we show that the vestibular fibers projecting to the cerebellum originate preferentially from the lateral half of the utricular macula and the medial half of the saccular macula. In contrast, the brainstem vestibular afferents originate primarily from the medial half of the utricular macula and the lateral half of the saccular macula. This indicates that the line of hair cell polarity reversal within the striola region segregates almost mutually exclusive central projections. A possible interpretation of this feature is that this macular organization provides an inhibitory side-loop through the cerebellum to produce synergistic tuning effects in the vestibular nuclei. The canal cristae project to the brainstem vestibular nuclei and cerebellum, but the projection to the vestibulocerebellum originates preferentially from the superior half of each of the cristae. The reason for this pattern is not clear, but it may compensate for unequal activation of crista hair cells or may be an evolutionary atavism reflecting a different polarity organization in ancestral vertebrate ears.  相似文献   

10.
This paper examines the directionality of tuberous electroreceptor responses and relates them to a polarity bias seen for passive electrolocation by electric fish (Hypopomus). We recorded from Burst Duration Coders (BDCs) while stimulating with 1 kHz single period sine waves with electric fields oriented horizontally in different directions. Electroreceptors have figure-8 directional sensitivity profiles with two, usually unequal lobes of sensitivity separated by 180°. For most units the larger lobe points inward, while for a few, the lobes are symmetrical or the larger lobe points outward. The differences correlate with differences in frequency tuning of the receptors. We can alter, and even reverse, the directional asymmetry of a single unit by changing the frequency of the stimulus. Two general response profiles result, with two corresponding classes of tuning curves. The degree of asymmetry varies with position on the body surface. The asymmetries and the effects of stimulus frequency and of tuning can be modeled with a linear/non-linear/linear cascade filter. The behavioral preference for approaching the head end ( + ) of an electrode is difficult to understand in light of the asymmetry of responses we report for amplitude-coding BDCs but can be understood by reference to the time-coding Pulse Marker (PM) receptors.Abbreviations BDC Burst Duration Coder - EOD electric organ discharge - nALL anterior lateral line nerve - PM Pulse Marker  相似文献   

11.
Single units of the goldfish torus semicircularis (TS) were recorded in response to pure tones. Response areas (RA) were obtained by recording the number of spikes evoked by tones in a range of frequencies and levels within the units' dynamic range. RAs gave estimates of best sensitivity (BS), characteristic frequency (CF), most excitatory frequency at each level (BF), and Q10dB. Peri-stimulus-time histograms (PSTH), interspike interval histograms (ISIH), and period histograms were obtained at various frequencies and levels to describe the units' temporal response patterns.The distribution of CF is nonuniform with modes at 155, 455, and 855 Hz. The distribution of the coefficient of synchronization to standard tones is also nonuniform, revealing a dichotomy between units with little or no phase-locking and those that phase-lock strongly. PSTHs for units without significant phase-locking vary widely and include patterns resembling those of the mammalian auditory brainstem. Compared with saccular afferents, torus units tend to have lower spontaneous rates, greater sensitivity, and sharper tuning. Unlike saccular afferents, BF is independent of level for most torus units. Some torus units are similar to saccular afferents while others reveal significant transformations of information between the periphery and the midbrain.Abbreviations BF best frequency - BS best sensitivity - CF characteristic frequency - ISIH inter-spike interval histogram - PSTH peri-stimulus-time histogram - RA response area - TS torus semicircularis  相似文献   

12.
The plainfin midshipman fish, Porichthys notatus, is a vocal species of teleost fish that generates acoustic signals for intraspecific communication during social and reproductive behaviors. All adult morphs (females and males) produce single short duration grunts important for agonistic encounters, but only nesting males produce trains of grunts and growls in agonistic contexts and long duration multiharmonic advertisement calls to attract gravid females for spawning. The midshipman fish uses the saccule as the main acoustic endorgan for hearing to detect and locate vocalizing conspecifics. Here, I examined the response properties of evoked potentials from the midshipman saccule to determine the frequency response and auditory threshold sensitivity of saccular hair cells to behaviorally-relevant single tone stimuli. Saccular potentials were recorded from the rostral, medial and caudal regions of the saccule while sound was presented by an underwater speaker. Saccular potentials of the midshipman, like other teleosts, were evoked greatest at a frequency that was twice the stimulus frequency. Results indicate that midshipman saccular hair cells of non-reproductive adults had a peak frequency sensitivity that ranged from 75 (lowest frequency tested) to 145 Hz and were best suited to detect the low frequency components (≤105 Hz) of midshipman vocalizations.  相似文献   

13.
Single-unit recordings were made from areas in the midbrain (torus semicircularis) of the oyster toadfish. We evaluated frequency tuning and directional responses using whole-body oscillation to simulate auditory stimulation by particle motion along axes in the horizontal and mid-sagittal planes. We also tested for bimodality in responses to auditory and hydrodynamic stimuli. One recording location in each animal was marked by a neurobiotin injection to confirm the recording site. Recordings were made in nucleus centralis, nucleus ventrolateralis, and the deep cell layer. Most units were frequency-selective with best frequencies between 50 and 141 Hz. Suppression of activity was apparent in 10% of the cells. Bimodality was common, including inhibition and suppression of background activity by auditory or hydrodynamic stimulation. The majority of the cells were directionally selective with directional response patterns that were sharpened compared with those of primary saccular afferents. The best directional axes were arrayed widely in spherical space, covering most azimuths and elevations. This representation is adequate for the computation of the motional axis of an auditory stimulus for sound source localization.Abbreviations BF best frequency - DCL deep cell layer - DON descending octaval nucleus - DRP directional response pattern - FFT fast Fourier transform - LL lateral lemniscus - NC nucleus centralis - NVL nucleus ventrolateralis - PVC periventricular cells - R coefficient of synchronization - TS torus semicircularis - Z Rayleigh statistic  相似文献   

14.
Auditory receptors of the locust (Locusta migratoria) were investigated with respect to the directionality cues which are present in their spiking responses, with special emphasis on how directional cues are influenced by the rise time of sound signals. Intensity differences between the ears influence two possible cues in the receptor responses, spike count and response latency. Variation in rise time of sound pulses had little effect on the overall spike count; however, it had a substantial effect on the temporal distribution of the receptor's spiking response, especially on the latencies of first spikes. In particular, with ramplike stimuli the slope of the latency vs. intensity curves was steeper as compared to stimuli with steep onsets (Fig. 3). Stimuli with flat ramplike onsets lead to an increase of the latency differences of discharges between left and right tympanic receptors. This type of ramplike stimulus could thus facilitate directional hearing. This hypothesis was corroborated by a Monte Carlo simulation in which the probability of incorrect directional decisions was determined on the basis of the receptor latencies and spike counts. Slowly rising ramps significantly improved the decisions based on response latency, as compared to stimuli with sudden onsets (Fig. 4). These results are compared to behavioural results obtained with the grasshopper Ch. biguttulus. The stridulation signals of the females of this species consist of ramplike pulses, which could be an adaptation to facilitate directional hearing of phonotactically approaching males.Abbreviations HFR high frequency receptor - ILD interaural level difference - LFR low frequency receptor - SPL sound pressure level - WN white noise  相似文献   

15.
1. Pure tone displacement sensitivity and bandwidth were measured from the saccule of the ear in two anabantid species (Trichogaster trichopterus and Helostoma temincki) using microphonic potentials with a 1 microV RMS threshold for the second harmonic of the stimulus frequency. 2. Saccular microphonics were recorded in both species from 80 to 1600 Hz, with lowest thresholds between 100 and 200 Hz. The overall microphonic response curves (sensitivity and bandwidth) of the two species were statistically similar to one another with an analysis of variance, although there were statistically different thresholds at 100 and 800 Hz. 3. The hair cell orientation patterns of the saccular epithelia differ in the two species. Consequently, the comparative sizes of the saccular sensory epithelium and numbers of sensory hair cells were examined. The saccular sensory epithelium of Helostoma is about 40% larger and contains nearly 50% more hair cells than the saccular epithelium of a comparably sized Trichogaster. 4. An extracranial air bubble, located in the suprabranchial chamber, is found in both species. The bubble has direct access to the saccular chamber in Trichogaster through a foramen which is absent in Helostoma. Despite the difference in morphology and the larger numbers of sensory hair cells in Helostoma, hearing sensitivity and bandwidth is similar in the two species. Although the structural differences in the auditory periphery do not affect pure tone sensitivity and bandwidth, other aspects of fish hearing such as frequency discrimination, discrimination of signals in the presence of noise, and/or sound localization ability may be affected by these structural differences.  相似文献   

16.
Two different kinds of mechanoreceptive hairs (smooth and feathered) on the second antennae of the freshwater crayfish, Orconectes virilis, have been investigated for their stimulus coding propertics. These mechanoreceptors show a great deal of non-linear behaviour both in threshold and in directionality. An effective appraoch for the investigation of such systems is noise analysis in the frequency domain. This method has been used here to calculate zero-, first- and second-order kernels. Sensory cells reveal different first- and second-order kernels, depending on which type of hair is being stimulated. The first-order kernel has a pronounced peak in the frequency response at 110 Hz if a feathered hair is stimulated and at 60 Hz if a smooth hair is stimulated. The second-order kernel shows a number of pronounced peaks in the frequency response between 40 and 110 Hz, but only if a feathered hair is stimulated. Smooth hair stimulation results in less sharp peaks but in higher gain for the same range of stimulus frequencies.  相似文献   

17.
Hair cells in the inner ear display a characteristic polarization of their apical stereocilia across the plane of the sensory epithelium. This planar orientation allows coherent transduction of mechanical stimuli because the axis of morphological polarity of the stereocilia corresponds to the direction of excitability of the hair cells. Neuromasts of the lateral line in fishes and amphibians form two intermingled populations of hair cells oriented at 180° relative to each other, however, creating a stimulus-polarity ambiguity. Therefore, it is unknown how these animals resolve the vectorial component of a mechanical stimulus. Using genetic mosaics and live imaging in transgenic zebrafish to visualize hair cells and neurons at single-cell resolution, we show that lateral-line afferents can recognize the planar polarization of hair cells. Each neuron forms synapses with hair cells of identical orientation to divide the neuromast into functional planar-polarity compartments. We also show that afferent neurons are strict selectors of polarity that can re-establish synapses with identically oriented targets during hair-cell regeneration. Our results provide the anatomical bases for the physiological models of signal-polarity resolution by the lateral line.  相似文献   

18.
Summary The response dynamics of cercal afferents in the cockroach, Periplaneta americana, were determined by means of a cross-correlation technique using a Gaussian white noise modulation of wind as a stimulus. The white noise stimulus could evoke sustained firing activity in most of the afferents examined (Fig. 1). The spike discharges were unitized and then cross-correlated with the stimulus to compute 1st- and 2nd-order Weiner kernels. The Ist-order kernels from a total of 28 afferents were biphasic and closely matched the time differential of a pulse (Figs. 1, 3 and 4). The amplitude and waveform of the kernels depended on the stimulus angle in such a way that the kernels were the mirror image of those on the polar opposite side (Figs. 2 and 3). The 2nd-order kernels were also differential. They had 2 diagonal peaks and 2 off-diagonal valleys in a 2-dimensional plot with 2 time axes (Figs. 1, 5 and 6). This 4-eye configuration was basically invariant irrespective of the stimulus angle, although the kernels varied in amplitude when the stimulus angle was changed. The time between the peak and a following trough of the 1st-order kernel was constant and had a mean of 4.6±0.1 ms, whereas the time between 2 diagonal peaks of the 2nd-order kernels was 4.7±0.1 ms (Figs. 4 and 6), suggesting that wind receptors (filiform sensilla) on cerci act as a band-pass filter with a peak frequency of about 106 Hz. The peak time, however, varies from 2.3 to 6.9 ms in both kernels, which may reflect the spatial distribution of the corresponding hairs on the cercus. The summation of the 1st- (linear) and 2nd-order (nonlinear) models precisely predicted the timing of the spike firing (Fig. 8). Thus, these 2 lower-order kernels can totally characterize the response dynamics of the wind receptors. The nonlinear response explains the directional sensitivity of the sensory neurons, while the differentiating 1st-order kernel explains the velocity sensitivity of the neurons. The nonlinearity is a signal compression in which one of the diagonal peaks of the 2nd-order kernel always offsets the downward phase of the 1st-order kernel (Fig. 7) and obviously represents a half-wave rectification property of the wind receptors that are excited by hair movement in only one direction and inhibited by hair movement in the polar opposite direction.  相似文献   

19.
The inner ears of a few fishes in the teleost superorder Ostariophysi are structurally unlike those of most other teleosts. Scanning electron microscopy was used to determine if other ostariophysans share these unusual features. Examined were the families Cyprinidae, Characidae, and Gymnotidae (all of the series Otophysi), and Chanidae (of the sister series Anotophysi), representing the four major ostariophysan lineages, the auditory organs of which have not yet been well described. Among the Otophysi, the saccular and lagenar otolith organs are similar to those reported for other ostariophysans. The lagena is generally the larger of the two organs. The saccular sensory epithelium (macula) contains long ciliary bundles on the sensory hair cells in the caudal region, and short bundles in the rostral region. The saccule and the lagena each have hair cells organized into two groups having opposing directional orientations. In contrast, Chanos, the anotophysan, has a saccular otolith larger than the lagenar otolith, and ciliary bundles that are more uniform in size over most of its saccular macula. Most strikingly, its saccular macula has hair cells organized into groups oriented in four directions instead of two, in a pattern very similar to that in many nonostariophysan teleosts. We suggest that the bi-directional pattern seen consistently in the Otophysi is a derived development related to particular auditory capabilities of these species.  相似文献   

20.
Summary European grassfrogs (Rana temporaria) were stimulated with pulsed sinusoidal, vertical vibrations (10–300 Hz) and the responses of 46 single midbrain neurons were recorded in awake, immobilized animals.Most units (40) had simple V-shaped excitatory vibrational tuning curves. The distribution of best frequencies (BF's) was bimodal with peaks at 10 and 100 Hz and the thresholds ranged from 0.02 to 1.28cm/s2 at the BF.Twenty-three neurons showed phasic-tonic and 11 neurons phasic responses. The dynamic range of seismic intensity for most neurons was 20–30 dB.In contrast to the sharp phase-locking in peripheral vibration-sensitive fibers, no phase-locking to the sinusoidal wave-form was seen in the midbrain neurons. The midbrain cells did not respond at low stimulus intensities (below 0.01–0.02 cm/s2) where a clear synchronization response occurs in saccular fibers.Six midbrain neurons had more complex response characteristics expressed by inhibition of their spontaneous activity by vibration or by bi-and trimodal sensory sensitivities.In conclusion, the vibration sensitive cells in the midbrain of the grassfrog can encode the frequency, intensity, onset and cessation of vibration stimuli. Seismic stimuli probably play a role in communication and detection of predators and the vibration-sensitive midbrain neurons may be involved in the central processing of such behaviorally significant stimuli.Abbreviation BF best frequency  相似文献   

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