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PubMed2002

Linking the sounds of dolphins to their locations and behavior using video and multichannel acoustic recordings.

It is difficult to attribute underwater animal sounds to the individuals producing them. This paper presents a system developed to solve this problem for dolphins by linking acoustic locations of the sounds of captive bottlenose dolphins with an overhead video image. A time-delay beamforming algorithm localized dolphin sounds obtained from an array of hydrophones dispersed around a lagoon. The localized positions of vocalizing dolphins were projected onto video images. The performance of the system was measured for artificial calibration signals as well as for dolphin sounds. The performance of the system for calibration signals was analyzed in terms of acoustic localization error, video projection error, and combined acoustic localization and video error. The 95% confidence bounds for these were 1.5, 2.1, and 2.1 m, respectively. Performance of the system was analyzed for three types of dolphin sounds: echolocation clicks, whistles, and burst-pulsed sounds. The mean errors for these were 0.8, 1.3, and 1.3 m, respectively. The 95% confidence bound for all vocalizations was 2.8 m, roughly the length of an adult bottlenose dolphin. This system represents a significant advance for studying the function of vocalizations of marine animals in relation to their context, as the sounds can be identified to the vocalizing dolphin and linked to its concurrent behavior.

Acoustics↗
PubMed2004

Old world frog and bird vocalizations contain prominent ultrasonic harmonics.

Several groups of mammals such as bats, dolphins and whales are known to produce ultrasonic signals which are used for navigation and hunting by means of echolocation, as well as for communication. In contrast, frogs and birds produce sounds during night- and day-time hours that are audible to humans; their sounds are so pervasive that together with those of insects, they are considered the primary sounds of nature. Here we show that an Old World frog (Amolops tormotus) and an oscine songbird (Abroscopus albogularis) living near noisy streams reliably produce acoustic signals that contain prominent ultrasonic harmonics. Our findings provide the first evidence that anurans and passerines are capable of generating tonal ultrasonic call components and should stimulate the quest for additional ultrasonic species.

Animal Communication↗
PubMed1988

Underwater audiogram of a false killer whale (Pseudorca crassidens).

Underwater audiograms are available for only a few odontocete species. A false killer whale (Pseudorca crassidens) was trained at Sea Life Park in Oahu, Hawaii for an underwater hearing test using a go/no-go response paradigm. Over a 6-month period, auditory thresholds from 2-115 kHz were measured using an up/down staircase psychometric technique. The resulting audiogram showed hearing sensitivities below 64 kHz similar to those of belugas (Delphinapterus leucas) and Atlantic bottlenosed dolphins (Tursiops truncatus). Above 64 kHz, this Pseudorca had a rapid decrease in sensitivity of about 150 dB per octave. A similar decrease in sensitivity occurs at 32 kHz in the killer whale, at 50 kHz in the Amazon River dolphin, at 120 kHz in the beluga, at 140 kHz in the bottlenosed dolphin, and at 140 kHz in the harbor porpoise. The most sensitive range of hearing was from 16-64 kHz (a range of 10 dB from the maximum sensitivity). This range corresponds with the peak frequency of echolocation pulses recorded from captive Pseudorca.

Acoustic Stimulation↗
PubMed1994

Comparison of sonar discrimination: dolphin and an artificial neural network.

The capability of an echolocating dolphin to discriminate differences in the wall thickness of cylinders (3.81 cm o.d. and 12.7 cm length) was determined by Au and Pawloski [J. Comp. Physiol. A 170, 41-47 (1992)]. The dolphin was required to discriminate a standard target from comparison targets of differing wall thicknesses. Performance varied from 96% to 56% correct depending on the wall thickness of the comparison targets. The 75% correct threshold was determined to be wall thickness differences of -0.23 mm for comparison targets with thinner walls and +0.27 mm for comparison targets with thicker walls than the standard. The dolphin performance was unchanged in the presence of artificial broadband masking noise until the echo-energy-to-noise ratio fell below approximately 15 dB. A counterpropagation artificial neural network was used to examine broadband echo features from the same cylinders. Features of the echoes were determined by passing them through a filter bank of constant-Q filters. Echo features of the standard and each comparison target were analyzed in pairs by a neural network having two output nodes. Twenty echoes per target were used in the training set and 30 additional echoes per target were used in the test set. For the noise free condition, the network performed at a comparable level to the dolphin for Q values between 4 and 5. In the presence of noise, Q values between 7 and 8 were needed before the network could perform at a comparable level to the dolphin for echo-energy-to-noise ratios of 10 and 15 dB.(ABSTRACT TRUNCATED AT 250 WORDS)

Acoustic Stimulation↗
PubMed1994

Stroboscopic hearing as a mechanism for prey discrimination in frequency-modulated bats?

A hypothesis was proposed that bats employing frequency-modulated (FM) echolocation pulses could utilize dynamic information of a flying insect to discriminate prey on the basis of "stroboscopic hearing." To test this hypothesis, single unit recordings were made from the inferior colliculus (IC) of the little brown bat, Myotis lucifugus. Response characteristics of IC units to trains of modulated and unmodulated sound pulses were analyzed at various pulse repetition rates that corresponded to a bat's pulse emission rates during the different stages of its target directed flight. The results show that amplitude modulation (AM) across a train of sound pulses was faithfully encoded in the units' discharge pattern when the pulse repetition rate was different from the AM frequency. When the AM frequency was integer multiples of the pulse repetition rate, the stimulus amplitude was reduced drastically under these conditions. Consequently, the discharge of an IC unit diminished precipitously, or if there was a phase delay between the pulse onset and the modulating sinusoidal waveform the unit fired to each modulation cycle with more or less the same vigor as if the modulation was absent. These data indicate that the across-pulse amplitude modulation becomes undetectable when the AM frequency is integer multiples of the pulse repetition rate. It is interpreted that FM bats can potentially employ a "stroboscopic hearing" strategy for discriminating insects on the basis of the wing-beat frequency of the prey.

Acoustic Stimulation↗
PubMed1994

Acoustic information available to bats using frequency-modulated sounds for the perception of insect prey.

Through the present study, the acoustic information available to an echolocating bat that uses brief frequency-modulated (FM) sonar sounds for the pursuit and capture of insect prey has been characterized. Computer-generated sonar pulses were broadcast at tethered insects, and the returning echoes were recorded on analog tape at high speed for off-line analyses. Echoes from stationary and fluttering insects were displayed using time waveform, spectrogram, power spectrum, and cross-correlation representations. The results show echo signatures for the different insect species studied, which change with the angle of incident sound. Sequences of echoes from fluttering insects show irregular changes in sound amplitude and time-frequency structure, reflecting a random temporal relation between the changing wing position and the arrival of incident sound. A set of recordings that controlled the temporal relation between incident sound and insect wing position suggests that information about the spatial profile of a flying insect could be enhanced if the bat were to produce a sequence of sounds that synchronized briefly with the moving target's wing-beat cycle. From this study, it has been proposed that the FM bat receives stroboscopic-like glimpses of fluttering prey whose spatial representation depends on the operation of the bat's sonar receiver.

Animals↗
PubMed1995

Time and frequency domain characteristics of sperm whale clicks.

Regular clicks from diving sperm whales, both large bull males and smaller females, were recorded in deep oceanic water off the Azores and subsequently sampled to computer disks for digital analysis. A total of 8540 clicks were marked and analyzed. Simple temporal analysis of the interclick intervals during feeding dives revealed mean click rates for male sperm whales of 1.1713 s-1 and 1.9455 s-1 for females. Fourier analysis showed distinctive peaks in the spectra of bull male sperm whales at 400 Hz and 2 kHz which were stable over extended periods of up to 20 mins. The clicks contained higher frequency components with energy ranging up to at least 12 kHz but not concentrated at any sharply defined frequency. The clicks of smaller female sperm whales showed similar spectral peaks, shifted to 1.2 and 3 kHz, respectively, but these peaks were less pronounced than those in the male click spectra and less stable with time. Higher frequencies were also present up to at least 15 kHz. The previously reported multiple pulse structure of sperm whale clicks is confirmed, but digital filtering reveals this structure to be frequency dependent. Analysis using the short-time Fourier transform confirms the complex time-frequency structure of individual clicks. The frequencies at which the multiples emerge in male and female clicks supports the idea of air cavities in the sperm whale head acting as sound reflectors, although the magnitude of the second pulse at high frequencies suggests some form of off axis distortion. It is also possible that air cavity resonance in the head of the sperm whale may act to reinforce the high-frequency components of the click, and that such components may have superior range and resolution performance in terms of echolocation.

Animal Communication↗
PubMed1999

Perception of complex tones and its analogy to echo spectral analysis in the bat, Megaderma lyra.

The gleaning bat Megaderma lyra emits broadband echolocation sounds consisting of multiple frequency components. The present study investigates into which perceptual qualities the spectral characteristics of echoes may be translated in the auditory system of M. lyra. Three bats were trained in a 2-AFC behavioral experiment to classify nine complex tones, which spectrally resembled M. lyra's sonar calls, into two perceptual categories. Then the bats' spontaneous responses to unknown complex tones were recorded. The results show that the animals based their classifications of the complex tones on a sound quality which was mediated by their broadband frequency spectra. The bats used the training stimuli as spectral templates and classified the test stimuli according to their broadband spectral similarity with the learned patterns. Assuming that passive hearing and echo processing are governed by similar perceptual qualities and subject to similar limitations, the perceptual mode which was used by the bats to compare the multicomponent spectral patterns in the reported experiments could serve as a powerful tool for the spectral analysis of M. lyra's multicomponent echoes. The analogy between the perception of complex tones and echo spectral analysis in M. lyra is theoretically elaborated in the "formant-mode" model.

Animals↗
PubMed1999

Killer whale (Orcinus orca) hearing: auditory brainstem response and behavioral audiograms.

Killer whale (Orcinus orca) audiograms were measured using behavioral responses and auditory evoked potentials (AEPs) from two trained adult females. The mean auditory brainstem response (ABR) audiogram to tones between 1 and 100 kHz was 12 dB (re 1 mu Pa) less sensitive than behavioral audiograms from the same individuals (+/- 8 dB). The ABR and behavioral audiogram curves had shapes that were generally consistent and had the best threshold agreement (5 dB) in the most sensitive range 18-42 kHz, and the least (22 dB) at higher frequencies 60-100 kHz. The most sensitive frequency in the mean Orcinus audiogram was 20 kHz (36 dB), a frequency lower than many other odontocetes, but one that matches peak spectral energy reported for wild killer whale echolocation clicks. A previously reported audiogram of a male Orcinus had greatest sensitivity in this range (15 kHz, approximately 35 dB). Both whales reliably responded to 100-kHz tones (95 dB), and one whale to a 120-kHz tone, a variation from an earlier reported high-frequency limit of 32 kHz for a male Orcinus. Despite smaller amplitude ABRs than smaller delphinids, the results demonstrated that ABR audiometry can provide a useful suprathreshold estimate of hearing range in toothed whales.

Animals↗
PubMed2000

Ultrasound sensitivity in the cricket, Eunemobius carolinus (Gryllidae, Nemobiinae).

Extracellular recordings from the cervical connectives in both long- and short-winged E. carolinus reveal auditory units that are sensitive to frequencies > 15 kHz with best sensitivity at 35 kHz (79 dB SPL threshold). Stimuli in this frequency range also elicit a startle response in long-winged individuals flying on a tether. For single-pulse stimuli, startle and neck connective thresholds decrease with increasing ultrasound duration, consistent with the operation of an exponential integrator with a approximately 32.5-ms time constant. There is evidence for adaptation to long duration pulses (> 20 ms) in the neck connectives, however, as it is more difficult to elicit responses to the later stimuli of a series. For paired-pulse stimuli consisting of 1-ms pulses of 40 kHz, temporal integration was demonstrated for pulse separations < 5 ms. For longer pulse separations, startle thresholds were elevated by 3 dB and appear to be optimally combined. Startle thresholds to 5 ms frequency modulated (FM) sweeps (60-30 kHz) and pure tone pulses (40 kHz) did not differ. The characteristics and sensitivity of this ultrasound-induced startle response did not differ between males and females. As in some other tympanate insects, ultrasound sensitivity in E. carolinus presumably functions in the context of predation from echolocating bats.

Animals↗
PubMed2001

Hydrodynamic trail-following in harbor seals (Phoca vitulina).

Marine mammals often forage in dark or turbid waters. Whereas dolphins use echolocation under such conditions, pinnipeds apparently lack this sensory ability. For seals hunting in the dark, one source of sensory information may consist of fish-generated water movements, which seals can detect with their highly sensitive whiskers. Water movements in the wake of fishes persist for several minutes. Here we show that blindfolded seals can use their whiskers to detect and accurately follow hydrodynamic trails generated by a miniature submarine. This shows that hydrodynamic information can be used for long-distance prey location.

Acoustics↗
PubMed1966

Visual and nonvisual auditory systems in mammals. Anatomical evidence indicates two kinds of auditory pathways and suggests two kinds of hearing in mammals.

Examination of the structural organization of the auditory system of the brain stem shows that the system is composed of a number of separate ascending pathways. This suggests that there may be at least two auditory systems, analogous to the rod and cone pathways in vision. We examined this possibility by investigating the variation in relative size of the medial and lateral superior olivary nuclei in a number of different mammalian species. The lateral superior olive is present in the hedgehog (an insectivore), cat (acarnivore), and squirrel monkey a(primate), but the medial superior olive is absent in the hedgehog. In a group of animals of the same taxonomic order (rodents) the lateral superior olive was present in all species examined, but the medial superior olive was almost wholly absent in the mouse and very prominent in the chinchilla and guinea pig. The absence of the medial superior olive in some animals is surprising because recent anatomical and physiological work has implicated the nucleus in auditory localization. Because of this implication, the medial and lateral olivary nuclei were examined in three species of bat and one dolphin, all echolocating animals. The medial superior olive was absent in these animals, and the lateral superior olive was prominent. These observations support the idea that the medial and lateral superior olives are nuclei on two different ascending auditory systems. It was also noted that the medial superior olive was always well developed in animals with well-developed eyes, and this suggested that the nucleus is in some way related to the visual system. We examined this idea by studying the relation between the numbers of cells in the medial superior olive and in the nucleus of the 6th cranial nerve (one of the motor nuclei concerned with eye movement) in a number of mammalian species. An approximately linear function was found between the sizes of the 6th nucleus and of the medial superior olive in three primates with cone-cell retinas (squirrel monkey, man, and macaque) and four rodents with rod-cell retinas (mouse, rat, guinea pig, and chinchilla). The cell numbers for the ground squirrel (a rodent with cone-cell retina) fitted an extension of the primate curve, and the cell numbers for the cat (in whose retina rods predominate) fitted an extension of the rodent curve. Thus, it is clear that the medial superior olive is related to the visual system, and that it is present in animals with cone-cell fovea and retina (diurnalanimals) and animals with rod-cell retina (that is, nocturnal animals) having good vision. In nonvisual nocturnal animals the nucleus is small or absent. The medial superior olive is probably not concerned with auditory localization in the psychophysical sense but is probably concerned with the movement of head and eyes in the direction of a sound in space. Localization in the psychophysical sense and fine auditory discrimination probably depend upon the ascending pathway which includes the lateral superior olive.

Anatomy, Comparative↗
PubMed1987

Auditory pathways to the frontal cortex of the mustache bat, Pteronotus parnellii.

In primates, certain areas of the frontal cortex play a role in guiding movements toward visual or auditory objects in space. The projections from auditory centers to the frontal cortex of the bat Pteronotus parnellii were examined because echolocating bats utilize auditory cues to guide their movements in space. An area in the frontal cortex receives a direct projection from a division of the auditory thalamus, the suprageniculate nucleus, which in turn receives input from the anterolateral peri-olivary nucleus, an auditory center in the medulla. This pathway to the frontal cortex bypasses the main auditory centers in the midbrain and cortex and could involve as few as four neurons between the cochlea and the frontal cortex. The auditory cortex is also a major source of input to the frontal cortex. This area of the frontal cortex may link the auditory and motor systems by its projections to the superior colliculus.

Animals↗
PubMed1984

Initiation of behavior by single neurons: the role of behavioral context.

Flying crickets avoid sources of ultrasound, possibly echolocating bats, by making rapid steering movements that turn them away from the stimulus. Electrical stimulation of a single, identified sensory interneuron (Int-1) elicits avoidance steering; depressing its response to ultrasound abolishes avoidance steering. Int-1 is necessary and sufficient for this behavior but only while the cricket is in flight. Thus, the sufficiency of Int-1 for eliciting this behavior is contingent on behavioral context.

Acoustic Stimulation↗
PubMed2006

Bats: important reservoir hosts of emerging viruses.

Bats (order Chiroptera, suborders Megachiroptera ["flying foxes"] and Microchiroptera) are abundant, diverse, and geographically widespread. These mammals provide us with resources, but their importance is minimized and many of their populations and species are at risk, even threatened or endangered. Some of their characteristics (food choices, colonial or solitary nature, population structure, ability to fly, seasonal migration and daily movement patterns, torpor and hibernation, life span, roosting behaviors, ability to echolocate, virus susceptibility) make them exquisitely suitable hosts of viruses and other disease agents. Bats of certain species are well recognized as being capable of transmitting rabies virus, but recent observations of outbreaks and epidemics of newly recognized human and livestock diseases caused by viruses transmitted by various megachiropteran and microchiropteran bats have drawn attention anew to these remarkable mammals. This paper summarizes information regarding chiropteran characteristics and information regarding 66 viruses that have been isolated from bats. From these summaries, it is clear that we do not know enough about bat biology; we are doing too little in terms of bat conservation; and there remain a multitude of questions regarding the role of bats in disease emergence.

Animals↗
PubMed1999

Timing in the auditory system of the bat.

Echolocating bats use audition to guide much of their behavior. As in all vertebrates, their lower brainstem contains a number of parallel auditory pathways that provide excitatory or inhibitory outputs differing in their temporal discharge patterns and latencies. These pathways converge in the auditory midbrain, where many neurons are tuned to biologically important parameters of sound, including signal duration, frequency-modulated sweep direction, and the rate of periodic frequency or amplitude modulations. This tuning to biologically relevant temporal patterns of sound is created through the interplay of the time-delayed excitatory and inhibitory inputs to midbrain neurons. Because the tuning process requires integration over a relatively long time period, the rate at which midbrain auditory neurons respond corresponds to the cadence of sounds rather than their fine structure and may provide an output that is closely matched to the rate at which motor systems operate.

Animals↗
PubMed2006

Neural mechanisms underlying selectivity for the rate and direction of frequency-modulated sweeps in the inferior colliculus of the pallid bat.

This study describes mechanisms that underlie neuronal selectivity for the direction and rate of frequency-modulated sweeps in the central nucleus of the inferior colliculus (ICC) of the pallid bat (Antrozous pallidus). This ICC contains a high percentage of neurons (66%) that respond selectively to the downward sweep direction of the bat's echolocation pulse. Some (19%) are specialists that respond only to downward sweeps. Most neurons (83%) are also tuned to sweep rates. A two-tone inhibition paradigm was used to describe inhibitory mechanisms that shape selectivity for sweep direction and rate. Two different mechanisms can create similar rate tuning. The first is an early on-best frequency inhibition that shapes duration tuning, which in turn determines rate tuning. In most neurons that are not duration tuned, a delayed high-frequency inhibition creates rate tuning. These neurons respond to fast sweep rates, but are inhibited as rate slows, and delayed inhibition overlaps excitation. In these neurons, starting a downward sweep within the excitatory tuning curve eliminates rate tuning. However, if rate tuning is shaped by duration tuning, this manipulation has no effect. Selectivity for the downward sweep direction is created by an early low-frequency inhibition that prevents responses to upward sweeps. In addition to this asymmetry in arrival times of low- and high-frequency inhibitions, the bandwidth of the low-frequency sideband was broader. Bandwidth influences the arrival time of inhibition during an FM sweep because a broader sideband will be encountered sooner. These findings show that similar spectrotemporal filters can be created by different mechanisms.

Acoustic Stimulation↗
PubMed2004

Differential temporal coding of rhythmically diverse acoustic signals by a single interneuron.

The omega neuron 1 (ON1) of the cricket Teleogryllus oceanicus responds to conspecific signals (4.5 kHz) and to the ultrasonic echolocation sounds used by hunting, insectivorous bats. These signals differ in temporal structure as well as in carrier frequency. We show that ON1's temporal coding properties vary with carrier frequency, allowing it to encode both of these behaviorally important signals. Information-transfer functions show that coding of 4.5 kHz is limited to the range of amplitude-modulation components that occur in cricket songs (<32 Hz), whereas coding of 30-kHz stimuli extends to the higher pulse rates that occur in bat sounds ( approximately 100 Hz). Nonlinear coding contributes to the information content of ON1's spike train, particularly for 30-kHz stimuli with high intensities and large modulation depths. Phase locking to sinusoidal amplitude envelopes also extends to higher AM frequencies for ultrasound stimuli. ON1s frequency-specific behavior cannot be ascribed to differences in the shapes of information-transfer functions of low- and high-frequency-tuned receptor neurons, both of which are tuned more broadly to AM frequencies than ON1. Coding properties are nearly unaffected by contralateral deafferentation. ON1's role in auditory processing is to increase binaural contrast through contralateral inhibition. We hypothesize that its frequency-specific temporal coding properties optimize binaural contrast for sounds with both the spectral and temporal features of behaviorally relevant signals.

Acoustic Stimulation↗