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Tonic efferent-induced cochlear damping in roosting and echolocating mustached bats.

The activity of the medial olivocochlear (MOC) efferent system in mustached bats, Pteronotus p. parnellii, was studied by monitoring changes in the mechanical properties of the cochlea. The changing properties were expressed by the decay time (DT) of cochlear microphonic potentials produced by transient-induced ringing (Henson et al., 1995). Tape-recorded roost noise (biosonar and communication sounds) produced sudden, marked decreases in DT when presented to the contralateral ear of animals adapted to the quiet. When the animals were first removed from their roosts the DT was relatively short (1.2-1.5 ms) but this gradually lengthened by about 0.5-1.0 ms as they rested in a quiet chamber. The time required to reach a stable, quiet-adapted state after noise exposure varied with SPL and exposure time; in many experiments recovery was in the range of 90-120 min. When quiet-adapted bats were isolated and allowed to fly and echolocate for 20 min, the DTs measured within a few minutes after the end of the flight were also short and only slowly returned to longer preflight values. The administration of a single dose of gentamicin, which blocks MOC effects, greatly reduced the amount of suppression (damping) observed after periods of noise and echolocation sound exposure. We conclude that tonic MOC activity is induced by the natural vocalizations and roost noise and this activity probably regulates and protects the highly resonant cochlear partition.

Adaptation, Physiological↗

Evidence for echolocation in the oldest known bats.

The earliest-known bats are represented by excellent fossil material, including virtually complete skeletons of Icaronycteris index from the early Eocene (50 Myr BP) of western Wyoming and Palaeochiropteryx tupaiodon from the middle Eocene (45 Myr BP) 'Grube Messel' of western Germany. These taxa have been closely allied with Recent Microchiroptera, a suborder of diverse bats noted for their powers of ultrasonic echolocation. A problem with this relationship is the alleged absence in the Eocene forms of specializations in the auditory region and other aspects of the skeletal system. It has been proposed, therefore, that the oldest bats are members of a group more primitive and possibly ancestral to the Microchiroptera and the visually oriented Megachiroptera. Previously undescribed specimens now show, however, that Icaronycteris and Palaeochiropteryx share special basicranial features with microchiropterans which suggest comparable refinement of ultrasonic echolocation. These results support the theory that a sophisticated sonar system was present in the earliest records of microchiropteran history.

Animals↗

Brain structures of echolocating and nonecholocating bats, derived in vivo from magnetic resonance images.

Magnetic resonance images of the brain of five species of wild bats, including three species of Microchiroptera, one species of echolocating Megachiroptera and one species of nonecholocating Megachiroptera, were obtained in vivo. The relative volumes of the inferior colliculus and the superior colliculus to the brainstem were derived from the magnetic resonance images and compared among different species. In general, the relative size of the inferior colliculus was much larger in Microchiropterans than in Megachiropterans, and in echolocating Megachiropterans than in nonecholocating Megachiropterans. The relative size of the superior colliculus was similar in these two suborders. Agreeing with the previous results and consistent with the current hypothesis that Megachiropterans originated from Microchiropterans, the results suggest that the inferior colliculus of Megachiropterans tends to degenerate during the process of evolution, as these fruit bats use more vision and smell than hearing when they forage. The results also demonstrate that magnetic resonance imaging can be used to study the neuroanatomy of wild bats noninvasively.

Animals↗

Beaked whales echolocate on prey.

Beaked whales (Cetacea: Ziphiidea) of the genera Ziphius and Mesoplodon are so difficult to study that they are mostly known from strandings. How these elusive toothed whales use and react to sound is of concern because they mass strand during naval sonar exercises. A new non-invasive acoustic ording tag was attached to four beaked whales(two Mesoplodon densirostris and two Ziphius cavirostris) and recorded high-frequency clicks during deep dives. The tagged whales only clicked at depths below 200 m, down to a maximum depth of 1267 m. Both species produced a large number of short, directional, ultrasonic clicks with significant energy below 20 kHz. The tags recorded echoes from prey items; to our knowledge, a first for any animal echolocating in the wild. As far as we are aware, these echoes provide the first direct evidence on how free-ranging toothed whales use echolocation in foraging. The strength of these echoes suggests that the source level of Mesoplodon clicks is in the range of 200-220 dB re 1 microPa at 1 m. This paper presents conclusive data on the normal vocalizations of these beaked whale species, which may enable acoustic monitoring to mitigate exposure to sounds intense enough to harm them.

Animals↗

Echolocation calls of bats are influenced by maternal effects and change over a lifetime.

The greater horseshoe bat, Rhinolophus ferrumequinum, is a model species in echolocation studies, and emits calls containing long constant-frequency (CF) components. The bats have auditory systems tuned sharply to frequencies close to the resting CF (RF) values. Call frequency and neural processing are both flexible within individual bats which use this mode of echolocation. The simple structure of the calls makes them ideal for sonographic analysis. Here, in a large-scale and long-term analysis of changes in the vocalizations of bats we show that: (i) the calls of R. ferrumequinum aged 1-28 years vary seasonally and over a lifetime in a predictable manner; and (ii) an infant's RF is at least partly determined by the RF of its mother. We consider the relative importance of genetic and learning factors upon the correlation between RFS of mothers and offspring.

Aging↗

Listening for bats: the hearing range of the bushcricket Phaneroptera falcata for bat echolocation calls measured in the field.

The hearing range of the tettigoniid Phaneropterafalcata for the echolocation calls of freely flying mouseeared bats (Myotis myotis) was determined in the field. The hearing of the insect was monitored using hook electrode recordings from an auditory interneuron, which is as sensitive as the hearing organ for frequencies above 16 kHz. The flight path of the bat relative to the insect's position was tracked by recording the echolocation calls with two microphone arrays, and calculating the bat's position from the arrival time differences of the calls at each microphone. The hearing distances ranged from 13 to 30 m. The large variability appeared both between different insects and between different bat approaches to an individual insect. The escape time of the bushcricket, calculated from the detection distance of the insect and the instantaneous flight speed of the bat, ranged from 1.5 to more than 4s. The hearing ranges of bushcrickets suggest that the insect hears the approaching bat long before the bat can detect an echo from the flying insect.

Animals↗

Sperm whale behaviour indicates the use of echolocation click buzzes "creaks" in prey capture.

During foraging dives, sperm whales (Physeter macrocephalus) produce long series of regular clicks at 0.5-2 s intervals interspersed with rapid-click buzzes called "creaks". Sound, depth and orientation recording Dtags were attached to 23 whales in the Ligurian Sea and Gulf of Mexico to test whether the behaviour of diving sperm whales supports the hypothesis that creaks are produced during prey capture. Sperm whales spent most of their bottom time within one or two depth bands, apparently feeding in vertically stratified prey layers. Creak rates were highest during the bottom phase: 99.8% of creaks were produced in the deepest 50% of dives, 57% in the deepest 15% of dives. Whales swam actively during the bottom phase, producing a mean of 12.5 depth inflections per dive. A mean of 32% of creaks produced during the bottom phase occurred within 10 s of an inflection (13x more than chance). Sperm whales actively altered their body orientation throughout the bottom phase with significantly increased rates of change during creaks, reflecting increased manoeuvring. Sperm whales increased their bottom foraging time when creak rates were higher. These results all strongly support the hypothesis that creaks are an echolocation signal adapted for foraging, analogous to terminal buzzes in taxonomically diverse echolocating species.

Acoustics↗

A blind mobility aid modeled after echolocation of bats.

A new model of a mobility aid for the blind was designed using microprocessor and ultrasonic devices. This mobility aid was evaluated based on psychophysical experiments. In this model, a downswept FM ultrasound signal is emitted from a transmitting array with broad directional characteristics in order to detect obstacles. The ultrasound reflections from the obstacles are picked up by a two-channel receiver. The frequency of the emitted ultrasound is swept from 70 to 40 kHz within 1 ms, so it has almost the same characteristics as the ultrasound a bat produces for echolocation. The frequency of the reflected ultrasound wave is down converted by about 50:1 by using a microcomputer with A/D and D/A converters. These audible waves are then presented binaurally through earphones. In this method obstacles may be perceived as localized sound images corresponding to the direction and the size of the obstacles. From the results of psychophysical experiments, it was found that downswept FM ultrasound was superior for the recognition of small obstacles compared to other ultrasonic schemes. With it a blind person can recognize a 1-mm-diameter wire. It was also proved that the blind could discriminate between several obstacles at the same time without any virtual images. This mobility aid, modeled after the bat's echolocation system, is very effective at detecting small obstacles placed in front of the head.

Animals↗

A model of echolocation of multiple targets in 3D space from a single emission.

Bats, using frequency-modulated echolocation sounds, can capture a moving target in real 3D space. The process by which they are able to accomplish this, however, is not completely understood. This work offers and analyzes a model for description of one mechanism that may play a role in the echolocation process of real bats. This mechanism allows for the localization of targets in 3D space from the echoes produced by a single emission. It is impossible to locate multiple targets in 3D space by using only the delay time between an emission and the resulting echoes received at two points (i.e., two ears). To locate multiple targets in 3D space requires directional information for each target. The frequency of the spectral notch, which is the frequency corresponding to the minimum of the external ear's transfer function, provides a crucial cue for directional localization. The spectrum of the echoes from nearly equidistant targets includes spectral components of both the interference between the echoes and the interference resulting from the physical process of reception at the external ear. Thus, in order to extract the spectral component associated with the external ear, this component must first be distinguished from the spectral components associated with the interference of echoes from nearly equidistant targets. In the model presented, a computation that consists of the deconvolution of the spectrum is used to extract the external-ear-dependent component in the time domain. This model describes one mechanism that can be used to locate multiple targets in 3D space.

Animals↗

A computational sensorimotor model of bat echolocation.

A computational sensorimotor model of target capture behavior by the echolocating bat, Eptesicus fuscus, was developed to understand the detection, localization, tracking, and interception of insect prey in a biological sonar system. This model incorporated acoustics, target localization processes, flight aerodynamics, and target capture planning to produce model trajectories replicating those observed in behavioral insect capture trials. Estimates of target range were based on echo delay, azimuth on the relative intensity of the echo at the two ears, and elevation on the spectral pattern of the sonar return in a match/mismatch process. Flapping flight aerodynamics was used to produce realistic model trajectories. Localization in all three spatial dimensions proved necessary to control target tracking and interception for an adequate model of insect capture behavior by echolocating bats. Target capture using maneuvering flight was generally successful when the model's path was controlled by a planning process that made use of an anticipatory internal simulation, while simple homing was successful only for targets directly ahead of the model bat.

Animals↗

Echolocation signals of wild Atlantic spotted dolphin (Stenella frontalis).

An array of four hydrophones arranged in a symmetrical star configuration was used to measure the echolocation signals of the Atlantic spotted dolphin (Stenella frontalis) in the Bahamas. The spacing between the center hydrophone and the other hydrophones was 45.7 cm. A video camera was attached to the array and a video tape recorder was time synchronized with the computer used to digitize the acoustic signals. The echolocation signals had bi-modal frequency spectra with a low-frequency peak between 40 and 50 kHz and a high-frequency peak between 110 and 130 kHz. The low-frequency peak was dominant when the signal the source level was low and the high-frequency peak dominated when the source level was high. Peak-to-peak source levels as high as 210 dB re 1 microPa were measured. The source level varied in amplitude approximately as a function of the one-way transmission loss for signals traveling from the animals to the array. The characteristics of the signals were similar to those of captive Tursiops truncatus, Delphinapterus leucas and Pseudorca crassidens measured in open waters under controlled conditions.

Animals↗

The influence of flight speed on the ranging performance of bats using frequency modulated echolocation pulses.

Many species of bat use ultrasonic frequency modulated (FM) pulses to measure the distance to objects by timing the emission and reception of each pulse. Echolocation is mainly used in flight. Since the flight speed of bats often exceeds 1% of the speed of sound, Doppler effects will lead to compression of the time between emission and reception as well as an elevation of the echo frequencies, resulting in a distortion of the perceived range. This paper describes the consequences of these Doppler effects on the ranging performance of bats using different pulse designs. The consequences of Doppler effects on ranging performance described in this paper assume bats to have a very accurate ranging resolution, which is feasible with a filterbank receiver. By modeling two receiver types, it was first established that the effects of Doppler compression are virtually independent of the receiver type. Then, used a cross-correlation model was used to investigate the effect of flight speed on Doppler tolerance and range-Doppler coupling separately. This paper further shows how pulse duration, bandwidth, function type, and harmonics influence Doppler tolerance and range-Doppler coupling. The influence of each signal parameter is illustrated using calls of several bat species. It is argued that range-Doppler coupling is a significant source of error in bat echolocation, and various strategies bats could employ to deal with this problem, including the use of range rate information are discussed.

Acceleration↗

Discrimination of complex synthetic echoes by an echolocating bottlenose dolphin.

Bottlenose dolphins (Tursiops truncatus) detect and discriminate underwater objects by interrogating the environment with their native echolocation capabilities. Study of dolphins' ability to detect complex (multihighlight) signals in noise suggest echolocation object detection using an approximate 265-micros energy integration time window sensitive to the echo region of highest energy or containing the highlight with highest energy. Backscatter from many real objects contains multiple highlights, distributed over multiple integration windows and with varying amplitude relationships. This study used synthetic echoes with complex highlight structures to test whether high-amplitude initial highlights would interfere with discrimination of low-amplitude trailing highlights. A dolphin was trained to discriminate two-highlight synthetic echoes using differences in the center frequencies of the second highlights. The energy ratio (delta dB) and the timing relationship (delta T) between the first and second highlights were manipulated. An iso-sensitivity function was derived using a factorial design testing delta dB at -10, -15, -20, and -25 dB and delta T at 10, 20, 40, and 80 micros. The results suggest that the animal processed multiple echo highlights as separable analyzable features in the discrimination task, perhaps perceived through differences in spectral rippling across the duration of the echoes.

Acoustic Stimulation↗

An echolocation model for range discrimination of multiple closely spaced objects: transformation of spectrogram into the reflected intensity distribution.

Using frequency-modulated echolocation, bats can discriminate the range of objects with an accuracy of less than a millimeter. However, bats' echolocation mechanism is not well understood. The delay separation of three or more closely spaced objects can be determined through analysis of the echo spectrum. However, delay times cannot be properly correlated with objects using only the echo spectrum because the sequence of delay separations cannot be determined without information on temporal changes in the interference pattern of the echoes. To illustrate this, Gaussian chirplets with a carrier frequency compatible with bat emission sweep rates were used. The delay time for object 1, T1, can be estimated from the echo spectrum around the onset time. The delay time for object 2 is obtained by adding T1 to the delay separation between objects 1 and 2 (extracted from the first appearance of interference effects). Further objects can be located in sequence by this same procedure. This model can determine delay times for three or more closely spaced objects with an accuracy of about 1 micros, when all the objects are located within 30 micros of delay separation. This model is applicable for the range discrimination of objects having different reflected intensities and in a noisy environment (0-dB signal-to-noise ratio) while the cross-correlation method is hard to apply to these problems.

Animals↗

Invariance of evoked-potential echo-responses to target strength and distance in an echolocating false killer whale.

Brain auditory evoked potentials (AEPs) were recorded in a false killer whale Pseudorca crassidens trained to accept suction-cup EEG electrodes and to detect targets by echolocation. AEP collection was triggered by echolocation pulses transmitted by the animal. The target strength varied from -22 to -40 dB; the distance varied from 1.5 to 6 m. All the records contained two AEP sets: the first one of a constant latency (transmission-related AEP) and a second one with a delay proportional to the distance (echo-related AEP). The amplitude of echo-related AEPs was almost independent of both target strength and distance, though combined variation of these two parameters resulted in echo intensity variation within a range of 42 dB. The amplitude of transmission-related AEPs was independent of distance but dependent on target strength: the less the target strength, the higher the amplitude. Recording of transmitted pulses has not shown their intensity dependence on target strength. It is supposed that the constancy of echo-related AEP results from variation of hearing sensitivity depending on the target strength and release of echo-related responses from masking by transmitted pulses depending on the distance.

Animals↗

Acoustic features of objects matched by an echolocating bottlenose dolphin.

The focus of this study was to investigate how dolphins use acoustic features in returning echolocation signals to discriminate among objects. An echolocating dolphin performed a match-to-sample task with objects that varied in size, shape, material, and texture. After the task was completed, the features of the object echoes were measured (e.g., target strength, peak frequency). The dolphin's error patterns were examined in conjunction with the between-object variation in acoustic features to identify the acoustic features that the dolphin used to discriminate among the objects. The present study explored two hypotheses regarding the way dolphins use acoustic information in echoes: (1) use of a single feature, or (2) use of a linear combination of multiple features. The results suggested that dolphins do not use a single feature across all object sets or a linear combination of six echo features. Five features appeared to be important to the dolphin on four or more sets: the echo spectrum shape, the pattern of changes in target strength and number of highlights as a function of object orientation, and peak and center frequency. These data suggest that dolphins use multiple features and integrate information across echoes from a range of object orientations.

Acoustics↗

Measurements of atmospheric attenuation at ultrasonic frequencies and the significance for echolocation by bats.

The absorption of sound propagating through the atmosphere under laboratory conditions of 25 degrees C and 50% relative humidity was measured at frequencies from 30 to 200 kHz. The attenuating effect on the passage of ultrasonic sounds through air ranged from 0.7 dB/m at 30 kHz. These measurements confirm theoretical expectations and earlier observations that atmospheric attenuation is progressively more severe at higher frequencies and that the atmosphere acts as a low-pass filter for conducting sounds in the frequency range used for echolocation by bats. Different species of bats use different portions of this range of frequencies, and bats emitting sonar signals predominantly above 100 kHz encounter especially severe attenuation of over 3 dB/m. With the greatly restricted operating distances for echolocation at such high frequencies, bats using these higher frequencies must be under compelling ecological pressures of a higher priority than long-range detection of targets.

Animals↗

Target detection in reverberation by an echolocating Atlantic bottlenose dolphin (Tursiops truncatus).

An experiment was conducted to determine the ability of an echolocating Atlantic bottlenose dolphin (Tursiops truncatus) to detect targets in the presence of reverberation. Reverberation was induced by a clutter screen consisting of forty-eight 5.1-cm-diam cork spheres spaced 15.2 cm apart and arranged in a rectangular array, located behind the targets. Hollow aluminum cylinders having the same outer diameter and wall thickness and three different lengths were used as targets. The dolphin was trained to station in a hoop, 6 m from the targets, and to echolocate the target upon the reception of an audio cue. Only one of the targets would be presented in a target-present trial. Data were collected on the animal's detection performance as a function of the separation distance between the clutter screen and the targets. The animal's performance for the smallest cylinder varied from 91% to 55% correct as the separation distance decreased from 10.2 to 0 cm. The target strength of the clutter screen and the cylinders were measured both in terms of the energies and the maximum peak-to-peak amplitudes of the incident and reflected echoes. These measurements indicated that when the target-clutter screen separation distance was 0 cm, the dolphin's 50% correct detection occurred at an energy echo-to-reverberation ratio of approximately 0.25 dB and at peak-to-peak echo-to-reverberation ratio of 2.6 dB. The results also indicated that the dolphin's performance varied almost linearly with the echo-to-reverberation ratio. The animal's pulse emissions were monitored by a microprocessor system and the results are presented in terms of the average number of clicks and the average response latency, as a function of the separation distance.

Animals↗