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PubMed2000

One-hydrophone method of estimating distance and depth of phonating dolphins in shallow water.

Previous attempts at localizing cetaceans have generally used multiple hydrophone arrays and multichannel recording systems. In this paper, a low-budget localization technique using only one hydrophone is described. The time delays of the signals traveling via the surface and bottom reflection paths to the hydrophone, relative to the direct signal, are used to calculate the distance and the depth of a phonating animal. Only two additional measures, the depth of the bottom and hydrophone, have to be taken. The method requires relatively shallow waters and a flat bottom surface. Echolocating and burst pulsing Hawaiian spinner dolphins (Stenella longirostris) at the Waianae coast of Oahu, Hawaii, were localized over different bottom substrates. A tracking range of up to 100 m was achieved. The accuracy of the method is estimated by the total error differential technique. The relative distance estimation error is below 35% and the absolute depth error below 0.7 m, so that the location method is sufficiently precise for examining source levels in our study area. Because of its simplicity, the method ideally complements sound recordings and visual sightings of marine mammals and could lead to a better understanding of the nature and use of click trains by dolphins.

Animal Communication↗
PubMed2000

Classification of electronically generated phantom targets by an Atlantic bottlenose dolphin (Tursiops truncatus).

Animal behavior experiments require not only stimulus control of the animal's behavior, but also precise control of the stimulus itself. In discrimination experiments with real target presentation, the complex interdependence between the physical dimensions and the backscattering process of an object make it difficult to extract and control relevant echo parameters separately. In other phantom-echo experiments, the echoes were relatively simple and could only simulate certain properties of targets. The echo-simulation method utilized in this paper can be used to transform any animal echolocation sound into phantom echoes of high fidelity and complexity. The developed phantom-echo system is implemented on a digital signal-processing board and gives an experimenter fully programmable control over the echo-generating process and the echo structure itself. In this experiment, the capability of a dolphin to discriminate between acoustically simulated phantom replicas of targets and their real equivalents was tested. Phantom replicas were presented in a probe technique during a materials discrimination experiment. The animal accepted the phantom echoes and classified them in the same manner as it classified real targets.

Acoustics↗
PubMed1965

MOTH SOUNDS AND THE INSECT-CATCHING BEHAVIOR OF BATS.

Captive bats trained to catch mealworms tossed in midair turned away from most of these targets when simultaneously confronted with a recorded train of the ultrasonic pulses generated by an Arctiid moth. When similarly exposed to the recorded echolocation pulses of another bat, presented at the same intensity as the "moth" sounds, they were not so affected.

Animals↗
PubMed1985

Sonar tracking of horizontally moving targets by the big brown bat Eptesicus fuscus.

When following a moving target, echolocating bats (Eptesicus fuscus) keep their heads aimed at the target's position. This tracking behavior seems not to involve predicting the target's trajectory, but is achieved by the bat's pointing its head at the target's last known position. The bat obtains frequent position updates by emitting sonar signals at a high rate. After the lag between head and target positions and the nonunity tracking gain were corrected for, bats' tracking accuracy in the horizontal plane was +/- 1.6 degree.

Animals↗
PubMed1979

Perception of echo phase information in bat sonar.

Echolocating bats (Eptesicus fuscus) can detect changes as small as 500 nanoseconds in the arrival time of sonar echoes when these changes appear as jitter or alternations in arrival time from one echo to the next. The psychophysical function relating the bat's performance to the magnitude of the jitter corresponds to the half-wave rectified cross-correlation function between the emitted sonar signals and the echoes. The bat perceives the phase or period structure of the sounds, which cover the 25- to 100-kilohertz frequency range, as these are represented in the auditory system after peripheral transformation. The acoustic image of a sonar target is apparently derived from time-domain or periodicity information processing by the nervous system.

Animals↗
PubMed1979

Neural axis representing target range in the auditory cortex of the mustache bat.

In echolocating bats, the primary cue for determining distance to a target is the interval between an emitted orientation sound and its echo. Whereas frequency is represented by place in the bat cochlea, no anatomical location represents of primary range. Target range is coded by the time interval between grouped discharges of primary auditory neurons in response to both the emitted sound and its echo. In the frequency-modulated-signal processing area of the auditory cortex of the mustache bat (Pteronotus parnellii rubiginosus), neurons respond poorly or not at all to synthesized orientation sounds or echoes alone but respond vigorously to echoes following the emitted sound with a specific delay from targets at a specific range. These range-tuned neurons are systemically arranged along the rostrocaudal axis of the frequency-modulated-signal processing area according to the delays to which they best respond, and thus represent target range in terms of cortical organization. The frequency-modulated-signal processing area therefore shows odotopic representation.

Animals↗
PubMed1978

Echo detection and target-ranging neurons in the auditory system of the bat Eptesicus fuscus.

Some of the neurons in the nucleus intercollicularis and auditory cortex of the echolocating bat Eptesicus fuscus respond selectively to sonar echoes occurring with specific echo delays or pulse-echo intervals. They do not respond for a wide range of other types of sounds or for sonar echoes at longer or shorter pulse-echo intervals; they may, therefore, be specialized for detection and ranging of sonar targets.

Animals↗
PubMed1979

Harmonic-sensitive neurons in the auditory cortex of the mustache bat.

Human speech and animal sounds contain phonemes with prominent and meaningful harmonics. The biosonar signals of the mustache bat also contain up to four harmonics, and each consists of a long constant-frequency component followed by a short frequency-modulated component. Neurons have been found in a large cluster within auditory cortex of this bat whose responses are facilitated by combinations of two or more harmonically related tones. Moreover, the best frequencies for excitation of these neurons are closely associated with the constant-frequency components of the biosonar signals. The properties of these neurons make them well suited for identifying the signals produced by other echolocating mustache bats. They also show how meaningful components of sound are assembled by neural circuits in the central nervous system and suggest a method by which sounds with important harmonics (or formants) may be detected and recognized by the brain in other species, including humans.

Action Potentials↗
PubMed1975

Responses to cold in the midbrain sonar center of hibernating and tropical bats.

The central gray matter of the bat midbrain, when electrically stimulated, causes the animal to produce a string of species-specific biosonar cries. Changes in this response with progressive cooling were studied in tropical homeothermic bats and in temperate hibernating bats. The species of hibernators chosen often move between hibernacula in the winter, flying and echolocating at low body temperatures (Tb). It was found that the midbrain "sonar center" exhibits a differential response to cooling that depends on the thermal propensities of the animal and its natural environment. Tropical bats followed a Q10 similar to that reported for other nonhibernating mammals and ceased responding at Tb 14-15 degrees C. Temperate zone-hibernating bat brains showed a relative insensitivity to temperature change and still responded at Tb 4-5 degrees C. Individual sonar cries within a string showed that duration was correlated with temperature but amplitude was unaffected. The study provides data for the functional separation of some parameters of biosonar and gives further evidence for differential nervous function in eurythermal versus stenothermal animals.

Adaptation, Physiological↗
PubMed2002

Selectivity for echo spectral interference and delay in the auditory cortex of the big brown bat Eptesicus fuscus.

The acoustic environment for an echolocating bat can contain multiple objects that reflect echoes so closely separated in time that they are almost completely overlapping. This results in a single echo with a spectrum characterized by deep notches due to interference. The object of this study was to document the possible selectivity, or lack thereof, of auditory neurons to the temporal separation of biosonar signals on a coarse (ms) and fine (micros) temporal scale. We recorded single-unit activity from the auditory cortex of big brown bats while presenting four protocol designs using wideband FM signals. The protocols simulated a pair of partially overlapping echoes where the separation between the first and second echo varied between 0 and 72 micros, a pulse followed by a single echo at varying delay from 0 to 30 ms, a pulse followed at a fixed delay by a pair of partially overlapping echoes that had a varying temporal separation of 0-72 micros, and a pulse followed, with a varying delay between 0 and 30 ms, by a pair of echoes that themselves had a fixed temporal separation on a microsecond time scale. About half of the cortical units showed increased spike counts to pairs of partially overlapping echoes at particular separations (6-72 micros) compared with a baseline stimulus at 0-micros separation. For many neurons tested with a pulse followed by two overlapping echoes, we observed a sensitivity to the coarse delay between the pulse and pair of overlapping echoes and to the separation between the two echoes themselves. The sensitivity to the partial overlap between the two echoes was not tuned to a single temporal separation. For bats, this means that the absolute range to the closest reflector and range between reflectors may be jointly encoded across a small population of single units. There are several possible neuronal mechanisms for encoding the separation between two nearby echoes based on the sensitivity to spectral notches.

Acoustic Stimulation↗
PubMed2004

Ambiguities in sound-duration selectivity by neurons in the inferior colliculus of the bat Molossus molossus from Cuba.

This study examines duration selectivity in auditory neurons of the inferior colliculus of the bat Molossus molossus (Molossidae, Chiroptera) from Cuba. Three main types of duration selectivity, short-, band-, and long-pass, as previously described in other species, are present in M. molossus. The range of best durations in the inferior colliculus of this species approximates the durations of their echolocation calls, suggesting that, as has been shown in other species of bats and frogs, the filter mechanism that produces duration tuning is selective for species-specific sounds relevant to behavior. Duration coding in M. molossus is not unambiguous because approximately 30% of the short- and band-pass neurons respond best to two different stimulus durations. This bimodal duration selectivity could be explained by time delayed excitatory inputs that coincide with an inhibitory rebound. In addition, the effect of stimulus intensity on duration selectivity was tested. For most of the neurons (78%), duration selectivity was affected by absolute sound pressure level and/or small changes of sound pressure. In this respect, the processing of stimulus duration by collicular neurons seems to be more complex in M. molossus than in other species studied so far.

Acoustic Stimulation↗
PubMed1985

Inhibition and level-tolerant frequency tuning in the auditory cortex of the mustached bat.

For echolocation the mustached bat, Pteronotus parnellii, emits complex orientation sounds (pulses), each consisting of four harmonics with long constant-frequency components (CF1-4) followed by short frequency-modulated components (FM1-4). The CF signals are best suited for target detection and measurement of target velocity. The CF/CF area of the auditory cortex of this species contains neurons sensitive to pulse-echo pairs. These CF/CF combination-sensitive neurons extract velocity information from Doppler-shifted echoes. In this study we electrophysiologically investigated the frequency tuning of CF/CF neurons for excitation, facilitation, and inhibition. CF1/CF2 and CF1/CF3 combination-sensitive neurons responded poorly to individual signal elements in pulse-echo pairs but showed strong facilitation of responses to pulse-echo pairs. The essential components in the pairs were CF1 of the pulse and CF2 or CF3 of the echo. In 68% of CF/CF neurons, the frequency-tuning curves for facilitation were extremely sharp for CF2 or CF3 and were "level-tolerant" so that the bandwidths of the tuning curves were less than 5.0% of best frequencies even at high stimulus levels. Facilitative tuning curves for CF1 were level tolerant only in 6% of the neurons studied. CF/CF neurons were specialized for fine analysis of the frequency relationship between two CF sounds regardless of sound pressure levels. Some CF/CF neurons responded to single-tone stimuli. Frequency-tuning curves for excitation (responses to single-tone stimuli) were extremely sharp and level tolerant for CF2 or CF3 in 59% of CF1/CF2 neurons and 70% of CF1/CF3 neurons. Tuning to CF1 was level tolerant in only 9% of these neurons. Sharp level-tolerant tuning may be the neural basis for small difference limens in frequency at high stimulus levels. Sharp level-tolerant tuning curves were sandwiched between broad inhibitory areas. Best frequencies for inhibition were slightly higher or lower than the best frequencies for facilitation and excitation. We thus conclude that sharp level-tolerant tuning curves are produced by inhibition. The extent to which neural sharpening occurred differed among groups of neurons tuned to different frequencies. The more important the frequency analysis of a particular component in biosonar signals, the more pronounced the neural sharpening. This was in addition to the peripheral specialization for fine frequency analysis of that component. The difference in bandwidth or quality factor between the excitatory tuning curves of peripheral neurons and the facilitative and excitatory tuning curves of CF/CF neurons was larger at higher stimulus levels.(ABSTRACT TRUNCATED AT 400 WORDS)

Acoustic Stimulation↗
PubMed1988

Delay-tuned combination-sensitive neurons in the auditory cortex of the vocalizing mustached bat.

1. FM-FM neurons in the auditory cortex of the mustached bat are sensitive to a pair of frequency-modulated (FM) sounds that simulates an FM component of the orientation sound and an FM component of the echo. These neurons are tuned to particular delays between the two FM components, suggesting an encoding of target range information. The response properties of these FM-FM neurons, however, have previously been studied only with synthesized orientation sounds and echoes delivered from a loud-speaker as substitutes for the bat's own orientation sounds and corresponding echoes. In this study, the combination sensitivity and delay tuning of FM-FM neurons were examined while the bat was actively vocalizing. 2. When the bat produced orientation sounds in an anechoic environment, or synthesized single FM echoes were delivered to a silent bat, the FM-FM neurons showed weak or no response. In contrast, when synthesized FM echoes were delivered with a particular delay from the FM component of the vocalized orientation sounds, the FM-FM neurons exhibited strong facilitative responses. 3. In both the vocalizing bats and the silent bats with substituted synthesized orientation sounds, all FM-FM neurons tested responded preferentially to the same echo harmonic (FM2, FM3, or FM4). 4. In vocalizing bats, FM-FM neurons showed maximum response to an echo FM component delivered with a particular delay (best delay) from an FM component in the orientation sound. Best delays measured with vocalized orientation sounds were nearly the same as those measured with synthesized orientation sounds. 5. The equivalent effect of a vocalized orientation sound and a synthesized FM1 component on the activity of FM-FM neurons indicates that, during echolocation, the FM1 component in the vocalized orientation sound stimulates the auditory system and conditions the FM-FM neurons to be sensitive to echoes with particular delays from the vocalized orientation sounds. 6. The amount of vocal self-stimulation to the inner ear by the bat's own vocalized sounds was measured by recording cochlear microphonic potentials (CMs). Spectral analysis of CM indicated that the amount of vocal self-stimulation by each harmonic of an orientation sound was equivalent to a sound of 70 dB sound pressure level (SPL) for the first harmonic (H1), 91 dB SPL for H2, 83 dB SPL for H3, and 70 dB SPL for H4, when the amplitude of the vocalized sound was 117 dB SPL at 5 cm in front of the bat's mouth.

Animals↗
PubMed1993

Differences in response properties of neurons between two delay-tuned areas in the auditory cortex of the mustached bat.

1. The orientation sound (pulse) of the mustached bat, Pteronotus parnellii parnellii, consists of four harmonics (H1-4), each containing a long constant-frequency component (CF1-4) followed by a short frequency-modulated component (FM1-4). The auditory cortex of this species contains several "combination-sensitive" areas: FM-FM, dorsal fringe (DF), ventral fringe (VF), CF/CF, and H1-H2. The FM-FM, DF, and VF areas each consist of neurons tuned to particular delays of echo FMn (n = 2, 3, or 4) from pulse FM1, and have an echo-delay (target-range) axis. This delay axis is from 0.4 to approximately 18 ms in the FM-FM area, to approximately 9 ms in the DF area, and to approximately 5 ms in the VF area. Therefore we hypothesized that the VF area was more specialized for the processing of range information in the terminal phase of echolocation than was the FM-FM area. The aim of our present studies was to find differences in response properties between neurons with best delays shorter than 6 ms in the VF and FM-FM areas and thus to test our hypothesis. 2. In the terminal phase of target-directed flight, the rate of pulse emission becomes higher, pulse duration (in particular, CF duration) becomes shorter, echo delay becomes shorter, and echoes (both the CF and FM components) are less Doppler shifted. Therefore, a "temporal-pattern-simulating (TPS)" stimulus was designed to mimic the train of pulse-echo pairs that would be heard by the bat during the terminal phase, and responses of single neurons to the TPS stimulus and other types of stimuli were recorded from the VF and FM-FM areas of the auditory cortex of unanesthetized bats with a tungsten-wire microelectrode. 3. Best delays of the neurons studied range between 0.9 and 5.5 ms (2.64 +/- 0.72 ms, N = 181) for the VF area, and between 0.6 and 6.0 ms (3.64 +/- 1.14, N = 144) for the FM-FM area. More neurons in the VF area than those in the FM-FM area showed no response or a poor response to the TPS stimulus. Therefore VF neurons are less suited than neurons in the FM-FM area for processing target ranges in the terminal phase of target-directed flight. Facilitative delay-tuning curves were commonly sandwiched between inhibitory delay-tuning curves. The lack of response or poor response to the TPS stimulus can be explained by this inhibition.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
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↗
PubMed2000

Reorganization of the frequency map of the auditory cortex evoked by cortical electrical stimulation in the big brown bat.

In a search phase of echolocation, big brown bats, Eptesicus fuscus, emit biosonar pulses at a rate of 10/s and listen to echoes. When a short acoustic stimulus was repetitively delivered at this rate, the reorganization of the frequency map of the primary auditory cortex took place at and around the neurons tuned to the frequency of the acoustic stimulus. Such reorganization became larger when the acoustic stimulus was paired with electrical stimulation of the cortical neurons tuned to the frequency of the acoustic stimulus. This reorganization was mainly due to the decrease in the best frequencies of the neurons that had best frequencies slightly higher than those of the electrically stimulated cortical neurons or the frequency of the acoustic stimulus. Neurons with best frequencies slightly lower than those of the acoustically and/or electrically stimulated neurons slightly increased their best frequencies. These changes resulted in the over-representation of repetitively delivered acoustic stimulus. Because the over-representation resulted in under-representation of other frequencies, the changes increased the contrast of the neural representation of the acoustic stimulus. Best frequency shifts for over-representation were associated with sharpening of frequency-tuning curves of 25% of the neurons studied. Because of the increases in both the contrast of neural representation and the sharpness of tuning, the over-representation of the acoustic stimulus is accompanied with an improvement of analysis of the acoustic stimulus.

Animals↗
PubMed2000

Interdependence of spatial and temporal coding in the auditory midbrain.

To date, most physiological studies that investigated binaural auditory processing have addressed the topic rather exclusively in the context of sound localization. However, there is strong psychophysical evidence that binaural processing serves more than only sound localization. This raises the question of how binaural processing of spatial cues interacts with cues important for feature detection. The temporal structure of a sound is one such feature important for sound recognition. As a first approach, we investigated the influence of binaural cues on temporal processing in the mammalian auditory system. Here, we present evidence that binaural cues, namely interaural intensity differences (IIDs), have profound effects on filter properties for stimulus periodicity of auditory midbrain neurons in the echolocating big brown bat, Eptesicus fuscus. Our data indicate that these effects are partially due to changes in strength and timing of binaural inhibitory inputs. We measured filter characteristics for the periodicity (modulation frequency) of sinusoidally frequency modulated sounds (SFM) under different binaural conditions. As criteria, we used 50% filter cutoff frequencies of modulation transfer functions based on discharge rate as well as synchronicity of discharge to the sound envelope. The binaural conditions were contralateral stimulation only, equal stimulation at both ears (IID = 0 dB), and more intense at the ipsilateral ear (IID = -20, -30 dB). In 32% of neurons, the range of modulation frequencies the neurons responded to changed considerably comparing monaural and binaural (IID =0) stimulation. Moreover, in approximately 50% of neurons the range of modulation frequencies was narrower when the ipsilateral ear was favored (IID = -20) compared with equal stimulation at both ears (IID = 0). In approximately 10% of the neurons synchronization differed when comparing different binaural cues. Blockade of the GABAergic or glycinergic inputs to the cells recorded from revealed that inhibitory inputs were at least partially responsible for the observed changes in SFM filtering. In 25% of the neurons, drug application abolished those changes. Experiments using electronically introduced interaural time differences showed that the strength of ipsilaterally evoked inhibition increased with increasing modulation frequencies in one third of the cells tested. Thus glycinergic and GABAergic inhibition is at least one source responsible for the observed interdependence of temporal structure of a sound and spatial cues.

Acoustic Stimulation↗
PubMed2001

Serotonin effects on frequency tuning of inferior colliculus neurons.

We investigated the modulatory effects of serotonin on the tuning of 114 neurons in the central nucleus of the inferior colliculus (ICc) of Mexican free-tailed bats and how serotonin-induced changes in tuning influenced responses to complex signals. We obtained a "response area" for each neuron, defined as the frequency range that evoked discharges and the spike counts evoked by those frequencies at a constant intensity. We then iontophoretically applied serotonin and compared response areas obtained before and during the application of serotonin. In 58 cells, we also assessed how serotonin-induced changes in response areas correlated with changes in the responses to brief frequency-modulated (FM) sweeps whose structure simulated natural echolocation calls. Serotonin profoundly changed tone-evoked spike counts in 60% of the neurons (68/114). In most neurons, serotonin exerted a gain control, facilitating or depressing the responses to all frequencies in their response areas. In many cells, serotonergic effects on tones were reflected in the responses to FM signals. The most interesting effects were in those cells in which serotonin selectively changed the responsiveness to only some frequencies in the neuron's response area and had little or no effect on other frequencies. This caused predictable changes in responses to the more complex FM sweeps whose spectral components passed through the neurons' response areas. Our results suggest that serotonin, whose release varies with behavioral state, functionally reconfigures the circuitry of the IC and may modulate the perception of acoustic signals under different behavioral states.

Acoustic Stimulation↗