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PubMed1990

Discrimination of jittered sonar echoes by the echolocating bat, Eptesicus fuscus: the shape of target images in echolocation.

1. Behavioral experiments with jittering echoes examined acoustic images of sonar targets in the echolocating bat, Eptesicus fuscus, along the echo delay or target range axis. Echo phase, amplitude, bandwidth, and signal-to-noise ratio were manipulated to assess the underlying auditory processes for image formation. 2. Fine delay acuity is about 10 ns. Calibration and control procedures indicate that this represents temporal acuity rather than spectral discrimination. Jitter discrimination curves change in phase when the phase of one jittering echo is shifted by 180 degrees relative to the other, showing that echo phase is involved in delay estimation. At an echo detectability index of about 36 dB, fine acuity is 40 ns, which is approximately as predicted for the delay accuracy of an ideal receiver. 3. Compound performance curves for 0 degrees and 180 degrees phase conditions match the crosscorrelation function of the echoes. The locations of both 0 degrees and 180 degrees phase peaks in the performance curves shift along the time axis by an amount that matches neural amplitude-latency trading in Eptesicus, confirming a temporal basis for jitter discrimination.

Acoustic Stimulation↗

PubMed2005

Echolocation calls and communication calls are controlled differentially in the brainstem of the bat Phyllostomus discolor.

BACKGROUND: Echolocating bats emit vocalizations that can be classified either as echolocation calls or communication calls. Neural control of both types of calls must govern the same pool of motoneurons responsible for vocalizations. Electrical microstimulation in the periaqueductal gray matter (PAG) elicits both communication and echolocation calls, whereas stimulation of the paralemniscal area (PLA) induces only echolocation calls. In both the PAG and the PLA, the current thresholds for triggering natural vocalizations do not habituate to stimuli and remain low even for long stimulation periods, indicating that these structures have relative direct access to the final common pathway for vocalization. This study intended to clarify whether echolocation calls and communication calls are controlled differentially below the level of the PAG via separate vocal pathways before converging on the motoneurons used in vocalization. RESULTS: Both structures were probed simultaneously in a single experimental approach. Two stimulation electrodes were chronically implanted within the PAG in order to elicit either echolocation or communication calls. Blockade of the ipsilateral PLA site with iontophoretically application of the glutamate antagonist kynurenic acid did not impede either echolocation or communication calls elicited from the PAG. However, blockade of the contralateral PLA suppresses PAG-elicited echolocation calls but not communication calls. In both cases the blockade was reversible. CONCLUSION: The neural control of echolocation and communication calls seems to be differentially organized below the level of the PAG. The PLA is an essential functional unit for echolocation call control before the descending pathways share again the final common pathway for vocalization.

Animal Communication↗

PubMed2007

Echolocation and passive listening by foraging mouse-eared bats Myotis myotis and M. blythii.

The two sibling mouse-eared bats, Myotis myotis and M. blythii, cope with similar orientation tasks, but separate their trophic niche by hunting in species-specific foraging microhabitats. Previous work has shown that both species rely largely on passive listening to detect and glean prey from substrates, and studies on other bat species have suggested that echolocation is ;switched off' during passive listening. We tested the hypothesis that mouse-eared bats continuously emit echolocation calls while approaching prey. Echolocation may be needed for orientation while simultaneously listening for prey. Because these sibling species forage in different microhabitats and eat different prey, we also compared their echolocation behaviour and related it to their ecology. Both species used echolocation throughout prey approach, corroborating a functional role for echolocation during gleaning. Captive bats of both species emitted similar orientation calls, and pulse rate increased during prey approach. Between the search to approach phases, call amplitude showed a sudden, dramatic drop and bats adopted ;whispering echolocation' by emitting weak calls. Whispering echolocation may reduce the risks of masking prey-generated sounds during passive listening, the mouse-eared bats' main detection tactic; it may also avoid alerting ultrasound-sensitive prey. In several cases M. myotis emitted a loud buzz made of 2-18 components when landing. We hypothesise that the buzz, absent in M. blythii at least when gleaning from the same substrate, is used to assess the distance from ground and refine the landing manoeuvre. Our findings have implications for niche separation between sibling species of echolocating bats, support a role for echolocation during passive listening and suggest a functional role for buzzes in landing control.

Animals↗

PubMed2006

Hearing in large (Eidolon helvum) and small (Cynopterus brachyotis) non-echolocating fruit bats.

Comparing the hearing abilities of echolocating and non-echolocating bats can provide insight into the effect of echolocation on more basic hearing abilities. Toward this end, we determined the audiograms of two species of non-echolocating bats, the straw-colored fruit bat (Eidolon helvum), a large (230-350 g) African fruit bat, and the dog-faced fruit bat (Cynopterus brachyotis), a small (30-45 g) bat native to India and Southeast Asia. A conditioned suppression/avoidance procedure with a fruit juice reward was used for testing. At 60 dB SPL, the hearing range of E. helvum extends from 1.38 to 41 kHz with best sensitivity at 8k Hz; the hearing range of C. brachyotis extends from 2.63 to 70 kHz with best sensitivity at 10 kHz. As with all other bats tested so far, neither species was able to hear below 500 Hz, suggesting that they may not use a time code for perceiving pitch. Comparison of the high-frequency hearing abilities of echolocating and non-echolocating bats suggests that the use of laryngeal echolocation has resulted in additional selective pressure to hear high frequencies. However, the typical high-frequency sensitivity of small non-echolocating mammals would have been sufficient to support initial echolocation in the early evolution of bats, a finding that supports the possibility of multiple origins of echolocation.

Animals↗

PubMed1992

Spatial processing within the mustache bat echolocation system: possible mechanisms for optimization.

1. The directionality of an echolocation system is determined by the acoustic properties of both the emitter and receiver, i.e., by the radiation pattern of the emitted pulse and the directionally of the external ears. We measured the directionality of the echolocation system of the greater mustache bat (Pteronotus parnellii) at the 30 kHz, 60 kHz and 90 kHz harmonics of its echolocation pulse by summing, at points throughout the frontal sound field, the echo attenuation due to the spread of pulse energy and the attenuation due to the spread of pulse energy and the attenuation due to the directionality of its external ears. The pulse radiation pattern at the 3 harmonics was measured by comparing the output of a microphone moved throughout the frontal sound field against a second reference microphone at the center of the field. External ear directionality at the 3. harmonics was measured by presenting free-field sounds throughout the frontal sound field, and recording the intensity thresholds of cochlear microphonic potentials, and the intensity thresholds of monaural neurons in the inferior colliculus tuned to one of the 3 harmonics. 2. When compared with ear directionality, the echolocation system was found to be more directional for the center of the sound field in several respects. At all harmonics, attenuation of sounds originating in the peripheral part of the field was increased by 10 to 13 dB. Areas of maximum sound intensity contracted toward the center of the field. Also, the isointensity contours of the echolocation system were more radially symmetrical about the center of the field. 3. At 60 kHz, sound intensity along the azimuth within the echolocation system was nearly constant 26 degrees to either side of the center of the field. This suggests that the radiation pattern of the echolocation pulse and the directionality of the external ears complement one another to produce an acoustic environment at the center of the sound field in which stimulus intensity is stabilized to allow more effective analysis of various aspects of the echolocation target. In particular, we suggest that this intensity stabilization may allow the bat to more effectively resolve the interaural intensity differences it uses to localize prey. 4. Predictions of the azimuthal spatial tuning of binaurally sensitive neurons in the inferior colliculus within the echolocation system were compared with their spatial tuning when only ear directionality is considered.(ABSTRACT TRUNCATED AT 400 WORDS)

Acoustic Stimulation↗

PubMed1991

No cost of echolocation for bats in flight.

Echolocation has evolved in relatively few animal species. One constraint may be the high cost of producing pulses, the echoes of which can be detected over useful distances. The energy cost of echolocation in a small (6 g) insectivorous bat, when hanging at rest, was recently measured at 0.067 Joules per pulse, implying a mean cost for echolocation in flight of 9.5 x basal metabolic rate (range 7 to 12x). Because flight is very costly, whether the costs of echolocation and flying are additive is an important question. We measured the energy costs of flight in two species of small echolocating Microchiroptera using a novel combination of respirometry and doubly-labelled water. Flight energy expenditure (adjusted for body mass) was not significantly different between echolocating bats and non-echolocating bats and birds. The low cost of echolocation for flying vertebrates may have been a significant factor favouring its evolution in these groups.

Animals↗

PubMed2006

Echolocation calls of Poey's flower bat (Phyllonycteris poeyi) unlike those of other phyllostomids.

Unlike any other foraging phyllostomid bat studied to date, Poey's flower bats (Phyllonycteris poeyi-Phyllostomidae) emit relatively long (up to 7.2 ms), intense, single-harmonic echolocation calls. These calls are readily detectable at distances of at least 15 m. Furthermore, the echolocation calls contain only the first harmonic, which is usually filtered out in the vocal tract of phyllostomids. The foraging echolocation calls of P. poeyi are more like search-phase echolocation calls of sympatric aerial-feeding bats (Molossidae, Vespertilionidae, Mormoopidae). Intense, long, narrowband, single-harmonic echolocation calls focus acoustic energy maximizing range and favoring detection, which may be particularly important for cruising bats, like P. poeyi, when flying in the open. Flying in enclosed spaces, P. poeyi emit short, low-intensity, frequency-modulated, multiharmonic echolocation calls typical of other phyllostomids. This is the first report of a phyllostomid species emitting long, intense, single-harmonic echolocation calls with most energy in the first harmonic.

Animals↗

PubMed2007

Do Swiftlets have an ear for echolocation? The functional morphology of Swiftlets' middle ears.

The Oilbird and many Swiftlet species are unique among birds for their ability to echolocate. Echolocaters may benefit from improved hearing sensitivity. Therefore, morphological adaptations to echolocation might be present in echolocating birds' middle ears. We studied the functional morphology of the tympano-ossicular chain of seven specimens of four echolocating Swiftlet species and one specimen each of five non-echolocating species. Three dimensional (3D) reconstructions were made from micro-Computer-Tomographic (muCT) scans. The reconstructions were used in functional morphological analyses and model calculations. A two dimensional (2D) rigid rod model with fixed rotational axes was developed to study footplate output-amplitudes and to describe how changes in the arrangement of the tympano-ossicular chain affect its function. A 3D finite element model was used to predict ossicular-chain movement and to investigate the justification of the 2D approach. No morphological adaptations towards echolocation were found in the middle-ear lever system or in the mass impedance of the middle ear. A wide range of middle-ear configurations result in maximum output-amplitudes and all investigated species are congruent with these predicted best configurations. Echolocation is unlikely to depend on adaptations in the middle ear tympano-ossicular chain.

Animals↗

PubMed2006

The evolution of echolocation in bats.

Recent molecular phylogenies have changed our perspective on the evolution of echolocation in bats. These phylogenies suggest that certain bats with sophisticated echolocation (e.g. horseshoe bats) share a common ancestry with non-echolocating bats (e.g. Old World fruit bats). One interpretation of these trees presumes that laryngeal echolocation (calls produced in the larynx) probably evolved in the ancestor of all extant bats. Echolocation might have subsequently been lost in Old World fruit bats, only to evolve secondarily (by tongue clicking) in this family. Remarkable acoustic features such as Doppler shift compensation, whispering echolocation and nasal emission of sound each show multiple convergent origins in bats. The extensive adaptive radiation in echolocation call design is shaped largely by ecology, showing how perceptual challenges imposed by the environment can often override phylogenetic constraints.

Animals↗

PubMed2005

A nuclear DNA phylogenetic perspective on the evolution of echolocation and historical biogeography of extant bats (chiroptera).

Bats (Order Chiroptera), the only mammals capable of powered flight and sophisticated laryngeal echolocation, represent one of the most species-rich and ubiquitous orders of mammals. However, phylogenetic relationships within this group are poorly resolved. A robust evolutionary tree of Chiroptera is essential for evaluating the phylogeny of echolocation within Chiroptera, as well as for understanding their biogeographical history. We generated 4 kb of sequence data from portions of four novel nuclear intron markers for multiple representatives of 17 of the 18 recognized extant bat families, as well as the putative bat family Miniopteridae. Three echolocation-call characters were examined by mapping them onto the combined topology: (1) high-duty cycle versus low-duty cycle, (2) high-intensity versus low-intensity call emission, and (3) oral versus nasal emission. Echolocation seems to be highly convergent, and the mapping of echolocation-call design onto our phylogeny does not appear to resolve the question of whether echolocation had a single or two origins. Fossil taxa may also provide insight into the evolution of bats; we therefore evaluate 195 morphological characters in light of our nuclear DNA phylogeny. All but 24 of the morphological characters were found to be homoplasious when mapped onto the supermatrix topology, while the remaining characters provided insufficient information to reconstruct the placement of the fossil bat taxa with respect to extant families. However, a morphological synapomorphy characterizing the Rhinolophoidea was identified and is suggestive of a separate origin of echolocation in this clade. Dispersal-Vicariance analysis together with a relaxed Bayesian clock were used to evaluate possible biogeographic scenarios that could account for the current distribution pattern of extant bat families. Africa was reconstructed as the center of origin of modern-day bat families.

Amino Acid Sequence↗

PubMed1997

Responses of bat inferior collicular neurons to recorded echolocation pulse trains.

Under free field stimulation conditions, we studied the responses of inferior collicular neurons of the big brown bat, Eptesicus fuscus, to echolocation pulse trains which were recorded during the entire process of hunting. The entire series of recorded echolocation pulse trains was edited in different sequences according to echolocation pulses of different hunting phases. When stimulated with all edited sequences of echolocation pulse trains, the temporal characteristics and the relative position of the echolocation pulses of a specific hunting phase affect the number of impulses of each inferior collicular neuron studied. When stimulated with two different intensities, more than 59% of inferior collicular neurons studied discharged selectively to the echolocation pulses of a specific hunting phase such that the number of impulses discharged to the echolocation pulses of the most and least preferred hunting phases differed by at least 50%. Passible mechanisms for this selective response are discussed.

Animals↗

PubMed1993

An audio-vocal interface in echolocating horseshoe bats.

The control of vocalization depends significantly on auditory feedback in any species of mammals. Echolocating horseshoe bats, however, provide an excellent model system to study audio-vocal (AV) interactions. These bats can precisely control the frequency of their echolocation calls by monitoring the characteristics of the returning echo; they compensate for flight-induced Doppler shifts in the echo frequency by lowering the frequency of the subsequent vocalization cells (Schnitzler, 1968; Schuller et al., 1974, 1975). It was the aim of this study to investigate the neuronal mechanisms underlying this Doppler-shift compensation (DSC) behavior. For that purpose, the neuronal activity of single units was studied during spontaneous vocalizations of the bats and compared with responses to auditory stimuli such as playback vocalizations and artificially generated acoustic stimuli. The natural echolocation situation was simulated by triggering an acoustic stimulus to the bat's own vocalization and by varying the time delay of this artificial "echo" relative to the vocalization onset. Single-unit activity was observed before, during, and/or after the bat's vocalization as well as in response to auditory stimuli. However, the activity patterns associated with vocalization differed from those triggered by auditory stimuli even when the auditory stimuli were acoustically identical to the bat's vocalization. These neurons were called AV neurons. Their distribution was restricted to an area in the paralemniscal tegmentum of the midbrain. When the natural echolocation situation was stimulated, the responses of AV neurons depended on the time delay between the onset of vocalization and the beginning of the simulated echo. This delay sensitivity disappeared completely when the act of vocalization was replaced by an auditory stimulus that mimicked acoustic self-stimulation during the emission of an echolocation call. The activity of paralemniscal neurons was correlated with all parameters of echolocation calls and echoes that are relevant in context with DSC. These results suggest a model for the regulation of vocalization frequencies by inhibitory auditory feedback.

Adaptation, Physiological↗

PubMed2002

Orienting responses and vocalizations produced by microstimulation in the superior colliculus of the echolocating bat, Eptesicus fuscus.

An echolocating bat actively controls the spatial acoustic information that drives its behavior by directing its head and ears and by modulating the spectro-temporal structure of its outgoing sonar emissions. The superior colliculus may function in the coordination of these orienting components of the bat's echolocation system. To test this hypothesis, chemical and electrical microstimulation experiments were carried out in the superior colliculus of the echolocating bat, Eptesicus fuscus, a species that uses frequency modulated sonar signals. Microstimulation elicited pinna and head movements, similar to those reported in other vertebrate species, and the direction of the evoked behaviors corresponded to the site of stimulation, yielding a map of orienting movements in the superior colliculus. Microstimulation of the bat superior colliculus also elicited sonar vocalizations, a motor behavior specific to the bat's acoustic orientation by echolocation. Electrical stimulation of the adjacent periaqueductal gray, shown to be involved in vocal production in other mammalian species, elicited vocal signals resembling acoustic communication calls of E. fuscus. The control of vocal signals in the bat is an integral part of its acoustic orienting system, and our findings suggest that the superior colliculus supports diverse and species-relevant sensorimotor behaviors, including those used for echolocation.

Acoustics↗

PubMed2000

Moth hearing in response to bat echolocation calls manipulated independently in time and frequency.

We measured the auditory responses of the noctuid moth Noctua pronuba to bat echolocation calls which were manipulated independently in time and frequency. Such manipulations are important in understanding how insect hearing influences the evolution of echolocation call characteristics. We manipulated the calls of three bat species (Rhinolophus hipposideros, Myotis nattereri and Pipistrellus pipistrellus) that use different echolocation call features by doubling their duration or reducing their frequency, and measured the auditory thresholds from the A1 cells of the moths. Knowing the auditory responses of the moth we tested three predictions. (i) The ranking of the audibility of unmanipulated calls to the moths should be predictable from their temporal and/or frequency structure. This was supported. (ii) Doubling the duration of the calls should increase their audibility by ca. 3 dB for all species. Their audibility did indeed increase by 2.1-3.5 dB. (iii) Reducing the frequency of the calls would increase their audibility for all species. Reducing the frequency had small effects for the two bat species which used short duration (2.7-3.6 ms) calls. However, the relatively long-duration (50 ms), largely constant-frequency calls of R. hipposideros increased in audibility by 21.6 dB when their frequency was halved. Time and frequency changes influence the audibility of calls to tympanate moths in different ways according to call design. Large changes in frequency and time had relatively small changes on the audibility of calls for short, largely broadband calls. Channelling energy into the second harmonic of the call substantially decreased the audibility of calls for bats which use long-duration, constant-frequency components in echolocation calls. We discuss our findings in the contexts of the evolution of both bat echolocation call design and the potential responses of insects which hear ultrasound.

Animals↗

PubMed2004

Echolocation signals of free-ranging killer whales (Orcinus orca) and modeling of foraging for chinook salmon (Oncorhynchus tshawytscha).

Fish-eating "resident"-type killer whales (Orcinus orca) that frequent the coastal waters off northeastern Vancouver Island, Canada have a strong preference for chinook salmon (Oncorhynchus tshawytscha). The whales in this region often forage along steep cliffs that extend into the water, echolocating their prey. Echolocation signals of resident killer whales were measured with a four-hydrophone symmetrical star array and the signals were simultaneously digitized at a sample rate of 500 kHz using a lunch-box PC. A portable VCR recorded the images from an underwater camera located adjacent to the array center. Only signals emanating from close to the beam axis (1185 total) were chosen for a detailed analysis. Killer whales project very broadband echolocation signals (Q equal 0.9 to 1.4) that tend to have bimodal frequency structure. Ninety-seven percent of the signals had center frequencies between 45 and 80 kHz with bandwidths between 35 and 50 kHz. The peak-to-peak source level of the echolocation signals decreased as a function of the one-way transmission loss to the array. Source levels varied between 195 and 224 dB re: 1 microPa. Using a model of target strength for chinook salmon, the echo levels from the echolocation signals are estimated for different horizontal ranges between a whale and a salmon. At a horizontal range of 100 m, the echo level should exceed an Orcinus hearing threshold at 50 kHz by over 29 dB and should be greater than sea state 4 noise by at least 9 dB. In moderately heavy rain conditions, the detection range will be reduced substantially and the echo level at a horizontal range of 40 m would be close to the level of the rain noise.

Animals↗

PubMed2004

Echolocation signals of dusky dolphins (Lagenorhynchus obscurus) in Kaikoura, New Zealand.

An array of four hydrophones arranged in a symmetrical star configuration was used to measure the echolocation signals of the dusky dolphin (Lagenorhynchus obscurus) near the Kaikoura Peninsula, New Zealand. Most of the echolocation signals had bi-modal frequency spectra with a low-frequency peak between 40 and 50 kHz and a high-frequency peak between 80 and 110 kHz. The low-frequency peak was dominant when the source level was low and the high frequency peak dominated when the source level was high. The center frequencies in the dusky broadband echolocation signals are among the highest of dolphins measured in the field. Peak-to-peak source levels as high as 210 dB re 1 microPa were measured, although the average was much lower in value. The levels of the echolocation signals are about 9-12 dB lower than for the larger white-beaked dolphin (Lagenorhynchus albirostris) which belongs to the same genus but is over twice as heavy as the dusky dolphins. 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 wave form and spectrum of the echolocation signals were similar to those of other dolphins measured in the field.

Animals↗

PubMed2005

Instrumenting free-swimming dolphins echolocating in open water.

Dolphins within the Navy Marine Mammal Program use echolocation to effectively locate underwater mines. They currently outperform manmade systems at similar tasks, particularly in cluttered environments and on buried targets. In hopes of improving manmade mine-hunting sonar systems, two instrumentation packages were developed to monitor free-swimming dolphin motion and echolocation during open-water target detection tasks. The biosonar measurement tool (BMT) is carried by a dolphin and monitors underwater position and attitude while simultaneously recording echolocation clicks and returning echoes through high-gain binaural receivers. The instrumented mine simulator (IMS) is a modified bottom target that monitors echolocation signals arriving at the target during ensonification. Dolphin subjects were trained to carry the BMT in open-bay bottom-object target searches in which the IMS could serve as a bottom object. The instrumentation provides detailed data that reveal hereto-unavailable information on the search strategies of free-swimming dolphins conducting open-water, bottom-object search tasks with echolocation.

Animals↗

PubMed1995

Sensory integration in the bottlenosed dolphin: immediate recognition of complex shapes across the senses of echolocation and vision.

In matching-to-sample tests, a bottlenosed dolphin (Tursiops truncatus) was found capable of immediately recognizing a variety of complexly shaped objects both within the senses of vision or echolocation and, also, across these two senses. The immediacy of recognition indicated that shape information registers directly in the dolphin's perception of objects through either vision or echolocation, and that these percepts are readily shared or integrated across the senses. Accuracy of intersensory recognition was nearly errorless regardless of whether the sample objects were presented to the echolocation sense and the alternatives to the visual sense (E-V matching) or the reverse, with samples presented to the visual sense and alternatives to the echolocation sense (V-E matching). Furthermore, during V-E matching, the dolphin was equally facile at recognition whether the sample objects exposed to vision were "live," presented in air in the real world, or were images displayed on a television screen placed behind an underwater window. Overall, the results suggested that what a dolphin "sees" through echolocation is functionally similar to what it sees through vision.

Animals↗