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Directional responses by kittens to an auditory stimulus.

Six kittens were tested in a 2-choice maze to determine if they would execute directional approach responses to an auditory stimulus. Tests were done at various postnatal ages (4-10, 11-17, 18-24, and 25-31 days) and comparison was made with responses of 5 kittens tested without auditory stimulation. At the earliest ages no evidence was seen for auditory approach responses but by 18-24 days consistent approch responses were found.

Acoustic Stimulation↗

Morphology of the abdominal wall in the bat, Pteronotus parnellii (Microchiroptera: Mormoopidae): implications for biosonar vocalization.

We investigated the structure of the abdominal wall of Pteronotus parnellii and made comparisons with eight other species of Microchiroptera and one megachiropteran. Similar to other mammals, the abdominal wall of bats consists of the three flank muscles laterally and the m. rectus abdominis ventrally. In Microchiroptera, flank muscles are mostly confined to dorsal portions of the wall. The mm. transversus abdominis and obliquus internus abdominis form the bulk of the wall; the m. obliquus externus is poorly developed. Ventrolaterally, a large portion of the wall is a dense, bilaminar aponeurosis, composed of collagen, elastin, and fibroblasts. The thicker, superficial lamina derives from the mm. obliquus internus and transversus abdominis. The deep lamina is a continuation of the transversalis fascia. Collagen fibers of the two fused laminae are oriented orthogonally, resulting in a resilient, composite fabric. Fascicles of the flank muscles are oriented along the margins of the aponeurosis so that their forces appear to be concentrated onto the aponeurosis. We suggest that this system is adapted for the regulation and generation of intra-abdominal pressure. The abdominal wall of Pteropus, the one megachiropteran examined, lacks the derived aponeurosis and is similar to other mammals. We consider the abdominal wall of Microchiroptera to be analogous to the diaphragma, in that it functions in the regulation of pressure within body cavities and facilitates biosonar vocalization.

Abdominal Muscles↗

Spatial and spectral dependence of the auditory periphery in the northern leopard frog.

We investigated directionalities of eardrum vibration and auditory nerve response in anesthetized northern leopard frogs (Rana pipiens pipiens). Simultaneous measures of eardrum velocities and firing rates from 282 auditory nerve fibers were obtained in response to free-field sounds from eight directions in the horizontal plane. Sound pressure at the external surface of the ipsilateral eardrum was kept constant for each presentation direction (+/- 0.5 dB). Significant effects of sound direction on eardrum velocity were shown in 90% of the cases. Maximum or minimum eardrum velocity was observed more often when sounds were presented from the lateral and posterior fields, or from the anterior and contralateral fields, respectively. Firing rates of 38% of the fibers were significantly affected by sound direction and maximum or minimum firing rate was observed more frequently when sounds were delivered from the lateral fields, or from the anterior and contralateral fields, respectively. Directionality patterns of eardrum velocity and nerve firing also vary with sound frequency. Statistically significant correlation between eardrum velocity and nerve fiber firing rate was demonstrated in only 45% of the fibers, suggesting that sound transmission to the inner ear through extratympanic pathways plays a non-trivial role in the genesis of directionality of auditory nerve responses.

Animals↗

Responses to simulated echoes by neurons in the barn owl's auditory space map.

The natural acoustical environment contains many reflective surfaces that give rise to echoes, complicating the task of sound localization and identification. The barn owl (Tyto alba), as a nocturnal predator, relies heavily on its auditory system for tracking and capturing prey in this highly echoic environment. The external nucleus of the owl's inferior colliculus (ICx) contains a retina-like map of space composed of "space-specific" auditory neurons that have spatially limited receptive fields. We recorded extracellularly from individual space-specific neurons in an attempt to understand the pattern of activity across the ICx in response to a brief direct sound and a simulated echo. Space-specific neurons responded strongly to the direct sound, but their response to a simulated echo was suppressed, typically, if the echo arrived within 5 ms or less of the direct sound. Thus we expect there to be little or no representation within the ICx of echoes arriving within such short delays. Behavioral tests using the owl's natural tendency to turn their head toward a sound source suggested that owls, like their space-specific neurons, similarly localize only the first of two brief sounds. Naive, untrained owls were presented with a pair of sounds in rapid succession from two horizontally-separated speakers. With interstimulus delays of less than 10 ms, the owl consistently turned its head toward the leading speaker. Longer delays elicited head turns to either speaker with approximately equal frequency and in some cases to both speakers sequentially.

Acoustic Stimulation↗

Frequency modulated sound pattern analysis in the lesser bulldog bat: the role of interactions between adjacent frequency elements of complex sounds.

A stereotyped approach phase vocalization response of Noctilio albiventris to artificial echoes simulating a virtual approaching object was used to assess the ability of the bat to analyze and extract distance information from the artificial echoes. The performance of the bats depended on the temporal pattern of frequency change of the continuously sweeping frequency modulated (FM) component of the signals. When the bats were presented with a CF/FM signal containing a time-reversed upward FM sweep, they responded with approach phase behavior at a performance level that was significantly below that seen with a CF/FM signal containing a naturally structured downward FM sweep. When the FM sweep was divided into a series of brief pure tone steps, the extent to which the bats showed a difference in their capability to process upward versus downward FM sweeps depended on the difference in frequency between the pure tone steps. The bats effectively processed downward but not upward FM sweeps when the difference in frequency between pure tone frequency elements of the FM sweeps was from about 100-200 Hz, but they effectually processed both downward and upward FM sweeps when the tonal elements composing the FM sweeps were separated by more than about 200 Hz. This suggests that the ability of the bats to effectively process downward but not upward FM sweeps is based on local interactions between adjacent frequency elements of the complex sounds.

Animals↗

Echo SPL influences the ranging performance of the big brown bat, Eptesicus fuscus.

Four bats of the species Eptesicus fuscus were trained in a two-alternative forced-choice procedure to discriminate between two phantom targets that differed in range. The rewarded stimulus was located at a distance of 52.7 cm, while the other unrewarded stimulus was further away. Only one target was presented at a time. In the first experiment we measured the range discrimination performance at an echo SPL of -28 dB relative to the bat's sonar transmission. A 75% correct performance level was arbitrarily defined as threshold and was obtained at a delay difference of 80 microseconds, corresponding to a range difference of 13.8 mm. In the second experiment the delay difference was fixed at 150 microseconds and the echo SPL varied between -8 and -48 dB relative to sonar emissions. The performance of the bats depended on the relative echo SPL. AT -28 dB the bats showed the best performance. It deteriorated at an increase of the relative echo SPL to -18 dB and -8 dB. The performance also deteriorated when the relative echo SPL was reduced to -38 dB and -48 dB. Only at low relative echo SPLs did the bats partially compensate for the reduction in echo SPL and increased the SPL of their emitted signals by a few dB. Our results support the hypothesis that neurons exhibiting paradoxical latency shift may be involved in encoding target range. This hypothesis predicts a decrease in performance at high echo SPLs as we found it in our experiments. The observed reduction in performance at very low echo SPLs may be due to a decrease in S/N ratio.

Animals↗

Labile cochlear tuning in the mustached bat. I. Concomitant shifts in biosonar emission frequency.

The cochlea of the mustached bat (Pteronotus parnellii) has sharp tuning characteristics and pronounced resonance within a narrow band near the second harmonic, constant frequency (CF2) component of the animal's biosonar signals. That fine frequency discrimination occurs within this narrow band is evident from Doppler-shift compensation, whereby bats in flight lower the frequency of emitted CF2s to maintain returning echoes within this band. This study examined various factors capable of producing shifts in both the cochlear resonance frequency (CRF) and CF2s emitted by stationary bats and bats actively Doppler-shift compensating on a pendulum. Each of three experimental factors shifted the CRF in a reversible manner. Changes in body temperature produced an average CRF shift of 39 +/- 18 Hz/degrees C. The CRF increased with flight by 150 +/- 100 Hz and returned to baseline values within 10 min after flight. Contralateral sound exposure produced smaller (100 +/- 20 Hz), rapid shifts in the CRF, suggesting that a mechanism different from the temperature- and flight-related shifts was involved. Changes in the CRF induced by temperature and flight were accompanied by shifts in the emitted CF2 of stationary and moving bats. Coupled with a companion study of associated shifts in neural tuning, the concomitant changes in CRF and CF2 provide evidence of cochlear tuning lability in the mustached bat.

Acoustic Stimulation↗

Evidence for perception of fine echo delay and phase by the FM bat, Eptesicus fuscus.

The big brown bat, Eptesicus fuscus, can perceive small changes in the delay of FM sonar echoes and shifts in echo phase, which interact with delay. Using spectral cues caused by interference, Eptesicus also can perceive the individual delays of two overlapping FM echoes at small delay separations. These results have been criticized as due to spectral artifacts caused by overlap between stimulus echoes and extraneous sounds (Pollak 1993). However, no amplitude or spectral variations larger than 0.05 dB accompany delay or phase changes produced by the electronic apparatus. No reverberation falls in the narrow span of delays required to produce the bat's performance curve from echo interference cues. Consistent differences in the durations of sonar sounds for 6 bats that perform the same in the experiments demonstrate that overlap between stimulus echoes and extraneous echoes is not necessary, and changes in the amount of echo overlap have no effect on performance. Noise-induced random variations in echo spectra outweigh putative spectral artifacts, and deliberately-introduced spectral "artifacts" do not improve performance overall but instead yield new time-frequency images. Amplitude-latency trading of perceived delay, proposed as a demonstration that the latency of neural discharges encodes delay (Pollak et al. 1977), confirms that the bat's fine delay and phase perception depends on a temporal neural code. The perceived delays depend on stimulus delays, not the delays of extraneous sounds. The rejected criticisms are based on physiological results with random-phase FM stimuli which are irrelevant to neural coding of fine echo delay and phase.

Animals↗

Complex sound analysis in the lesser bulldog bat: evidence for a mechanism for processing frequency elements of frequency modulated signals over restricted time intervals.

A stereotypical approach phase vocalization response of the lesser bulldog bat, Noctilio albiventris, to artificial echoes simulating a virtual approaching object was used to assess the ability of the bat to analyze and extract distance information from the artificial echoes. The performance of the bat was not significantly different when presented with naturally structured CF/FM echoes containing FM elements that sweep continuously from about 75-55 kHz in 4 ms or with CF/FM echoes containing FM components constructed from a series of 98 pure tone frequency steps, each with a duration of 0.04 ms. The performance of the bat remained unchanged when the duration of the tone steps was increased up to 0.08 ms but declined sharply to a level that was significantly below that seen with a naturally structured echo when the steps were 0.09 ms or longer. The performance of the bat depended on the duration of the individual tone steps, which could not exceed a specific upper limit of about 0.08 ms. The study suggests that the bats have adaptations for processing individual narrow band segments of FM signals over specific time intervals.

Animals↗

Complex sound analysis in the FM bat Eptesicus fuscus, correlated with structural parameters of frequency modulated signals.

Big brown bats, Eptesicus fuscus, were presented with artificial frequency modulated (FM) echoes that simulated an object becoming progressively closer to the bat. A stereotyped approach phase behavioral response of the bat to the virtual approaching target was used to determine the ability of the bat to analyze FM signals for target distance information. The degree to which the bats responded with approach phase behavior to a virtual approaching target was similar when they were presented with either a naturally structured artificial FM echo or an artificial FM echo constructed from a series of brief pure tone steps. The ability of the bats to respond to an FM signal structured from a sequence of pure tone elements depended on the number of pure tone steps in the series; the bats required the presentation of tone-step FM signals containing about 83 or greater pure tone elements. Moreover, the duration of the individual tone steps of the tone-step FM signals could not exceed a specific upper limit of about 0.05 ms. Finally, it appears that the bats were able to independently resolve individual tone steps within the tone-step FM signals that were separated by about 450 Hz or more.

Animals↗

Detection of frequency modulation in the FM-bat Phyllostomus discolor.

In a two-alternative forced-choice procedure lesser spear-nosed bats, Phyllostomus discolor, had to discriminate between a pure tone stimulus and a sinusoidally frequency-modulated signal generated at the same carrier frequency as the tone. Modulation depths of the SFM stimuli were reduced until the animals' performance dropped below the 75%-correct level which was used to determine difference limens for detection of frequency modulation (FMDL). The dependence of FMDLs on modulation and carrier frequency was systematically investigated. For a carrier frequency of 18.5 kHz, average FMDLs increased from 95 Hz at a modulation frequency of 10 Hz to 820 Hz at a modulation frequency of 2000 Hz which corresponds to Weber ratios (2 delta f/f) of 0.005 and 0.044 respectively. Further, difference limens were found to increase linearly in proportion to carrier frequency throughout a major part (9-74 kHz) of the species' hearing range. In comparison to other mammals, P. discolor has a pronounced capability for frequency discrimination which might be related to the extensive use of individually distinct frequency-modulated communication calls and audio-vocal learning.

Acoustic Stimulation↗

Frequency and space representation in the inferior colliculus of the FM bat, Eptesicus fuscus.

The tonotopic organization and spatial sensitivity of 217 inferior collicular (IC) neurons of Eptesicus fuscus were studied under free field stimulation conditions. Acoustic stimuli were delivered from a loudspeaker placed 21 cm ahead of the bat to determine the best frequency (BF) and minimum threshold (MT) of isolated IC neurons. A BF stimulus was then delivered as the loudspeaker was moved horizontally across the frontal auditory space of the bat to locate the best azimuthal angle (BAZ) at which the neuron had its lowest MT. The stimulus was then raised 3 dB above the lowest MT to determine the horizontal extent of the auditory space within which a sound could elicit responses from the neurons. This was done by moving the loudspeaker laterally at every 5 degrees or 10 degrees until the neuron failed to respond. These measurements also allowed us to redetermine the BAZ at which the neuron fired maximal number of impulses. Electrodes were placed evenly across the whole IC surface and IC neurons were sampled as many as possible within each electrode penetration. Tonotopic organization and spatial sensitivity were examined among all 217 IC neurons as a whole as well as among IC neurons sequentially sampled within individual electrode penetrations. The whole population of 217 IC neurons is organized tonotopically along the dorsoventral axis of the IC. Thus, low frequency neurons are mostly located dorsally and high frequency neurons ventrally with median frequency neurons intervening in between. The BAZ of these 217 IC neurons tend to shift from lateral to medial portions of the contralateral frontal auditory space with increasing BF.(ABSTRACT TRUNCATED AT 250 WORDS)

Acoustic Stimulation↗

Coding of small sinusoidal frequency and amplitude modulations in the inferior colliculus of 'CF-FM' bat, Rhinolophus ferrumequinum.

Single neurons in the inferior colliculus of the Greater Horseshoe bat, Rhinolophus ferrumequinum, showed two broad categories of response patterns to sinusoidally frequency (SFM) or amplitude (SAM) modulated stimuli. Tonic responding cells (best excitatory frequency (BEF) between 10 and 90 kHz) showed a rough sinusoidal modulation of the discharge pattern to SFM. Transient responding neurons generally showing on- or off-responses to pure tones, (BEF between 65 and 88 kHz), displayed highly synchronized discharge patterns to SFM-cycles (Fig. 1). Modulation rates between 20 and 100 Hz were most effective and some neurons encoded modulation rates up to 350 Hz (Figs. 2 and 3). The SFM responses were best synchronized to the modulation envelope for center frequencies in the upper portion of the tuning curve (Figs. 4 and 5). Sharply tuned neurons with BEF around 80 kHz had the lowest threshold for modulation depth (+/- 10 Hz or 0.025%) (Fig. 6). In general, SAMs evoked the same type of response patterns and were encoded down to modulation index of 3% (Fig. 7). The fine frequency and amplitude discriminations for periodical modulations by collicular neurons is discussed as related to the detection and discrimination performance of bats, when preying on flying insects in clustered surroundings.

Acoustic Stimulation↗

Coding of sinusoidally amplitude modulated acoustic stimuli in the inferior colliculus of the rufous horseshoe bat, Rhinolophus rouxi.

Single neuron responses to sinusoidally amplitude modulated (SAM) signals were studied in the inferior colliculus of the horseshoe bat, Rhinolophus rouxi. 57% of the neurons responded to SAM stimuli with periodical discharges synchronized to the modulation cycle. The proportion of cells driven by amplitude modulated signals was independent of the best frequency of the neurons. Best modulation frequencies were at or below 100 Hz in about 70% of the neurons. Synchronized activity could be elicited by modulation frequencies up to 400 Hz. Best SAM responses were observed at stimulus intensities 10 dB above threshold. Generally the BMF of a neuron did not change with intensity. The BMF decreased with decreasing modulation depth of the amplitude modulation. A trend for a topographical organization of neurons according to best modulation frequencies was detected. The results did not reveal any significant specialization of the bat's auditory system for coding of amplitude modulations as compared to other mammals.

Animals↗

A comparative study of the physiological properties of the inner ear in Doppler shift compensating bats (Rhinolophus rouxi and Pteronotus parnellii).

Cochlear microphonic (CM) and evoked neural (N-1) potentials were studied in two species of Doppler shift compensating bats with the aid of electrodes chronically implanted in the scala tympani. Potentials were recorded from animals fully recovered from the effects of anesthesia and surgery. In Pteronotus p. parnellii and Rhinolophus rouxi the CM amplitude showed a narrow band, high amplitude peak at a frequency about 200 Hz above the resting frequency of each species. In Pteronotus the peak was 25-35 dB higher in amplitude than the general CM level below or above the frequency of the amplitude peak. In Rhinolophus the amplitude peak was only a few dB above the general CM level but it was prominent because of a sharp null in a narrow band of frequencies just below the peak. The amplitude peak and the null were markedly affected by body temperature and anesthesia. In Pteronotus high amplitude CM potentials were produced by resonance, and stimulated cochlear emissions were prominent in Pteronotus but they were not observed in Rhinolophus. In Pteronotus the resonance was indicated by a CM afterpotential that occurred after brief tone pulses. The resonance was not affected by the addition of a terminal FM to the stimulus and when the ear was stimulated with broadband noise it resulted in a continual state of resonance. Rapid, 180 degree phase shifts in the CM were observed when the stimulus frequency swept through the frequency of the CM amplitude peak in Pteronotus and the frequency of the CM null in Rhinolophus. These data indicate marked differences in the physiological properties of the cochlea and in the mechanisms responsible for sharp tuning in these two species of bats.

Anesthesia, General↗

Acoustic localization in an ostariophysian fish.

Under approximate free field conditions acoustic localization could be demonstrated in an ostariophysian fish, the ide (Leuciscus idus). The efficient vibration links of both succuli with the unpaired swimbladder (via the Weberian ossicles) do apparently not prclude directional hearing in this important group of freshwater fish.

Animals↗