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[Ultrasonic echolocation of soft tissues in dogs during decompression].

Ultrasonic location of femoral soft tissues of anesthetized dogs was performed using a modified unit UZKAP-3. The animals were exposed to 0.5 mPa for 2 hours and then to decompression to sea level pressure. The concomitant echographic changes pointed to the formation of extra- and intravascular gas bubbles in the above tissues. The bubbles could be identified visually in the nearest echographic band because the instrumentation used made it possible to eliminate the background image of structural inhomogeneities of the tissue layer.

Animals↗
PubMed1993

What ears do for bats: a comparative study of pinna sound pressure transformation in chiroptera.

Using a moveable loudspeaker and an implanted microphone, we studied the sound pressure transformation of the external ears of 47 species of bats from 13 families. We compared pinna gain, directionality of hearing and interaural intensity differences (IID) in echolocating and non-echolocating bats, in species using different echolocation strategies and in species that depend upon prey-generated sounds to locate their targets. In the Pteropodidae, two echolocating species had slightly higher directionality than a non-echolocating species. The ears of phyllostomid and vespertilionid species showed moderate directionality. In the Mormoopidae, the ear directionality of Pteronotus parnellii clearly matched the dominant spectral component of its echolocation calls, unlike the situation in three other species. Species in the Emballonuridae, Molossidae, Rhinopomatidae and two vespertilionids that use narrow-band search-phase echolocation calls showed increasingly sharp tuning of the pinna to the main frequency of their signals. Similar tuning was most evident in Hipposideridae and Rhinolophidae, species specialized for flutter detection via Doppler-shifted echoes of high-duty-cycle narrow-band signals. The large pinnae of bats that use prey-generated sounds to find their targets supply high sound pressure gain at lower frequencies. Increasing domination of a narrow spectral band in echolocation is reflected in the passive acoustic properties of the external ears (sharper directionality). The importance of IIDs for lateralization and horizontal localization is discussed by comparing the behavioural directional performance of bats with their bioacoustical features.

Animals↗
PubMed2003

Mysterious Mystacina: how the New Zealand short-tailed bat (Mystacina tuberculata) locates insect prey.

The New Zealand short-tailed bat Mystacina tuberculata evolved in the absence of terrestrial mammals and initially with few potential predators. Unusual among bats, it is well adapted for the capture of prey on the ground. Bats from Fiordland, New Zealand had relatively low wing loadings and aspect ratios adapted for flight in cluttered habitats. We predicted that M. tuberculata would locate prey in air (uncluttered space) by echolocation. Echolocation call sequences associated with prey capture (terminal buzzes) were heard in the field, and bats detected and localized prey suspended on fishing line by echolocation in a flight cage. The bats emitted brief, multiharmonic echolocation calls at low duty cycle during search phase, and 64% of calls contained most energy in the fundamental harmonic. Approach- and terminal-phase calls were also broadband and multiharmonic. We predicted that bats would not use echolocation to locate prey hidden on the ground in leaf litter (cluttered space). Bats seemed unable to locate hidden prey precisely from the air and instead hunted for such prey while crawling. Echolocation calls were emitted at a low repetition rate on the ground, suggesting that here echolocation was used for orientation and not for prey detection. We experimentally removed cues available to the bats and showed that bats located mealworms in leaf litter by listening for prey-generated noises and possibly by olfaction.

Animals↗
PubMed1990

Audiovocal interactions during development? Vocalisation in deafened young horseshoe bats vs. audition in vocalisation-impaired bats.

Horseshoe bats (Rhinolophus rouxi) were deafened in their 3rd-5th postnatal week. Subsequently their vocalisations were monitored to evaluate the impact of audition on the development of echolocation pulses. Hearing impairment affected the echolocation pulses as follows: the frequency of the constant frequency (CF) component was altered by between +4 kHz and -14 kHz, and the dominance of the second harmonic of the pulses was neutralised by a relative increase in intensity of the first and third harmonics. A second experiment focused on possible influences of acoustical self-stimulation with echolocation pulses on the establishment of auditory fovea representation in the inferior colliculus (IC). Frequency control of echolocation pulses was disrupted by larynx denervation. Thereafter, the bats produced multiharmonic echolocation signals (4-11 harmonics) varying in frequency. IC tonotopy, however, as monitored by stereotaxic electrophysiology, showed the same developmental dynamics as seen in control specimens (Fig. 10). Both experiments indicate that throughout postnatal development echolocation pulses are under auditory feedback control, whereas maturation of the auditory fovea and shifts in its frequency tuning represent an innate process. The significance of this postnatal development might be the adjustment of the vocal motor system of each bat to the frequency of its 'personal' auditory fovea.

Animals↗
PubMed2003

Can two streams of auditory information be processed simultaneously? Evidence from the gleaning bat Antrozous pallidus.

A tenet of auditory scene analysis is that we can fully process only one stream of auditory information at a time. We tested this assumption in a gleaning bat, the pallid bat (Antrozous pallidus) because this bat uses echolocation for general orientation, and relies heavily on prey-generated sounds to detect and locate its prey. It may therefore encounter situations in which the echolocation and passive listening streams temporally overlap. Pallid bats were trained to a dual task in which they had to negotiate a wire array, using echolocation, and land on one of 15 speakers emitting a brief noise burst in order to obtain a food reward. They were forced to process both streams within a narrow 300 to 500 ms time window by having the noise burst triggered by the bats' initial echolocation pulses as it approached the wire array. Relative to single task controls, echolocation and passive sound localization performance was slightly, but significantly, degraded. The bats also increased echolocation interpulse intervals during the dual task, as though attempting to reduce temporal overlap between the signals. These results suggest that the bats, like humans, have difficulty in processing more than one stream of information at a time.

Acoustic Stimulation↗
PubMed2002

Support for the allotonic frequency hypothesis in an insectivorous bat community.

The allotonic frequency hypothesis proposes that certain insectivorous bat species can prey upon moths that can hear bat echolocation calls by using echolocation frequencies outside the sensitivity range of moth ears. The hypothesis predicts that the peak frequencies of bat echolocation calls are correlated with the incidence of moths in the diets of these bats. The aim of this study was to test this prediction on a bat community dominated by bats using low duty cycle echolocation calls, i.e. aerial foraging, insectivorous species using frequency modulated calls. The community consisted of nine species, two molossids, Sauromys petrophillus and Tadarida aegyptiaca, five vespertilionids, Eptesicus capensis, Eptesicus hottentotus, Miniopteris schreibersii, Myotis tricolor, and Myotis lesueuri, one rhinolophid, Rhinolophus clivosus, and one nycterid, Nycteris thebaica. The insect fauna in the habitat used by the bat community was suited to the testing of the allotonic frequency hypothesis because more than 90% of the moths comprising the insect fauna were tympanate. These included Pyralidae (3.8%), Geometridae (44.9%), Notodontidae (3.8%), Arctiidae (4.6%), Lymantriidae (0.8%) and Noctuidae (32.4%). As predicted, peak echolocation frequency was correlated with the incidence of moths in the diets of these nine species (r=0.98, df=7, P<0.01). Furthermore, multivariate analysis revealed that echolocation frequency (t=9.91, n=129, P<0.001) was a better predictor of diet than forearm length (t=5.51, n=129, P<0.001) or wing area (t=-3.41, n=129, P<0.001). This suggests that the selection pressure exerted by moth hearing might have acted directly on call frequency and secondarily on body size and wing morphology, as part of the same adaptive complex. It is unlikely that dietary differences were due to temporal and spatial differences in the availability of prey because the pattern of differences in skull morphology of the nine species supported our dietary analyses. The skull morphology of a bat represents a historical record of the kind of diet it has become adapted to over its evolutionary history. These results suggest that prey defences may mediate other factors structuring bat communities, e.g. competition. Competition may be reduced for those species of bats that can circumvent prey defences.

Acoustics↗
PubMed2001

The synchronisation of signal emission with wingbeat during the approach phase in soprano pipistrelles (Pipistrellus pygmaeus).

Previous studies have shown that, during search flight in bats, wingbeat, respiration and echolocation are synchronised in a 1:1 relationship. An efficiently integrated locomotor-respiratory system enables bats to produce intense echolocation signals at little or no cost above that required for flight. In this study, we investigated the coupling of wingbeat with echolocation in the laboratory during approach flight in soprano pipistrelles (Pipistrellus pygmaeus) using high-speed digital video at 250 and 500 frames s(-)(1). We found that soprano pipistrelles most commonly produce single or double pulses per wingbeat. Single pulses per wingbeat occurred in two alternative positions: immediately before the end of the upstroke or after the start of the downstroke. Double pulses per wingbeat were emitted in the same wingbeat positions on the upstroke and the downstroke, as in single pulses per wingbeat. We suggest that, during approach flight, the coupling of more than one echolocation signal with a single wingbeat and expiratory cycle allows echolocation to remain energetically economic. When soprano pipistrelles approached a Perspex disc target, an increase in mean repetition rate was achieved by producing an extra pulse per wingbeat. Finally, we hypothesise that the bat's approach to potentially interesting targets in the same horizontal plane as it's flight path, i.e. during flapping flight, may be characterised by the production of double pulses per wingbeat, resulting in a unique pattern of echolocation pulse intervals.

Animals↗
PubMed2005

Phylogenetic relationships amongst swifts and swiftlets: a multi locus approach.

We recently reconstructed the troublesome swiftlet phylogeny using cytochrome-b mitochondrial DNA sequences. The relationship of the giant swiftlet (Hydrochous gigas) with swiftlets of the genus Aerodramus was, however, unresolved. In an attempt to clarify this issue, we now incorporated mitochondrial 12S rRNA and nuclear beta-fibrinogen intron 7 nuclear DNA sequences with the cyt-b sequences of six swiftlet, two swift, and one hummingbird outgroup species. A partition homogeneity (PH) test, used to determine the congruence of phylogenetic signal between two sets of sequences, suggested that cyt-b and Fib7 sequences were incongruent and therefore should not be combined. However, further analyses revealed that the apparent incongruence was probably due to the high amount of variation in cyt-b sequences. Separate and combined analyses of the three sequences unambiguously placed H. gigas as the sister-group of Aerodramus and supported monophyly of the swiftlets. These results were supported by analyses of combined NADH dehydrogenase subunit-2 (ND2) and cyt-b sequences of H. gigas in combination with sequences previously published by other workers. Recently, it was shown that the pygmy swiftlet (C. troglodytes)--in our phylogenetic analyses consistently placed with other, non-echolocating, Collocalia species--is in fact able to echolocate. Echolocation thereby lost its value to distinguish between different swiftlet genera. Furthermore, the phylogenetic distribution of echolocation can be explained either by its single evolution at the base of the swiftlets, with subsequent loss, or by independent evolution in Aerodramus and C. troglodytes. Because yet unpublished data suggest that only the auditory nuclei in swiftlet brains show adaptations to echolocation, the latter explanation seems the more likely one.

Animals↗
PubMed2000

Detection distances of bottom-set gillnets by harbour porpoises (Phocoena phocoena) and bottlenose dolphins (Tursiops truncatus).

Many odontocetes die annually in gillnet fisheries. Why they become entangled is not yet clear. Maybe some species detect the nets too late to avoid collision. Therefore, the target strength of 11 types of bottom-set gillnets was measured under 0 and 45 degrees angles of incidence. From these target strengths and from knowledge on the echolocation abilities of two odontocete species (harbour porpoises, bottlenose dolphins), the detection ranges of the nets by these small cetaceans could be estimated. The 90% detection range by echolocating harbour porpoises, approaching the nets at right (perpendicular) angles under low noise level conditions, varied between 3 and 6 m depending on the net type. For bottlenose dolphins, under high noise conditions, the 90% detection range varied between 25 and 55 m. At other angles of approach, the estimated detection ranges are shorter. The study suggests that echolocating bottlenose dolphins can detect nets in time to avoid collision, whereas echolocating harbour porpoises cannot in most cases. Suggestions for future research to reduce small cetacean bycatch by improving the nets' detectability by echolocation are given.

Animals↗
PubMed2002

Periaqueductal gray and the region of the paralemniscal area have different functions in the control of vocalization in the neotropical bat, Phyllostomus discolor.

The periaqueductal gray matter and the region of the paralemniscal area were neuroanatomically delineated in the brain of the neotropical bat Phyllostomus discolor[Wagner (1843) Arch. Naturgesch., 9, 365-368] and were probed with electrical microstimulation for eliciting vocalizations. In a well-delimited rostral portion of the periaqueductal gray exclusively, communication calls could be triggered at low stimulation currents. Communication calls as well as echolocation calls could be elicited at the dorsal and ventral edges of this area. Pharmacological stimulation with microdialysed kainic acid in this particular periaqueductal gray area demonstrated that neurons and not fibres of passage are activated for triggering vocalization. Solely echolocation calls were emitted upon electrical microstimulation or with microdialysed kainic acid in the region of the paralemniscal area. The periaqueductal gray appears to be involved in vocal pathways that control both communication calls and echolocation calls, while the region of the paralemniscal area seems to be specialized for control of echolocation calls only. Respiration is similarly influenced by stimulation in the periaqueductal gray and the region of the paralemniscal area. Periaqueductal gray and paralemniscal area interact differently with the final common pathway for vocalization, and may represent different functional organization in the vocal controlling pathways for communication calls and echolocation calls.

Animals↗
PubMed1998

One tone, two ears, three dimensions: a robotic investigation of pinnae movements used by rhinolophid and hipposiderid bats.

Bats, which echolocate using broadband calls, are believed to employ the passive acoustic filtering properties of the head and pinnae to provide spectral cues which encode 3-D target angle. Microchiropteran species whose calls consist of a single, constant frequency harmonic (i.e., some species in the families Rhinolophidae and Hipposideridae) may create additional acoustic localization cues via vigorous pinna movements. In this work, two types of echolocation cues generated by moving a pair of receivers aboard a model sensor head are investigated. In the first case, it is supposed that a common 3-D echolocation principle employed by all bats is the creation of alternative viewing perspectives, and that constant frequency (CF) echolocators use pinna movement rather than morphology to alter the acoustic axes of their perceptual systems. Alternatively, it is possible rhinolophids and hipposiderids move their ears to create dynamic cues--in the form of frequency and amplitude modulations--which vary systematically with target elevation. Here the use of binaural and monaural timing cues derived from amplitude modulated echo envelopes are investigated. In this case, pinna mobility provides an echolocator with a mechanism for creating dramatic temporal cues for directional sensing which, unlike interaural timing differences, do not degrade with head size.

Animals↗
PubMed1999

Evolutionary adaptations of cochlear function in Jamaican mormoopid bats.

Mormoopid bat species have their echolocation system adapted to different hunting strategies. To study the corresponding mechanical properties of their inner ear, we measured distortion-product otoacoustic emissions to assess cochlear sensitivity and tuning. Mormoops blainvillii, Pteronotus macleayii and P. quadridens use frequency-modulated echolocation signals, sometimes preceded by a short narrowband signal component. Their distortion-product otoacoustic emission-threshold curves are most sensitive between 30 and 50 kHz and show no adaptation to the narrowband echolocation components. In contrast, the constant-frequency bat P. parnellii always uses long constant-frequency call components. Its inner ear is maximally sensitive at 62 kHz, the echo-frequency of the dominant constant-frequency component, and pronounced insensitivities at 61 and 93 kHz (CF2 and CF3 call frequency) are the major evolutionary change in comparison to its relatives. Furthermore, in P. parnellii, the optimum cochlear frequency separation is minimal at 62 and 93 kHz, associated with enhanced cochlear tuning, while for the other mormoopids there is no indication of enhanced tuning. The phylogeny of mormoopids, assessed by mitochondrial DNA analysis, shows a close relationship between the Pteronotus species. This suggests that major cochlear redesign, associated with the acquisition of echolocation-call specific cochlear processing in P. parnellii, has occurred within a relatively short evolutionary time scale.

Acoustic Stimulation↗
PubMed1993

Psychophysical frequency modulation thresholds in a FM-bat, Tadarida brasiliensis.

Echolocating bats hunting flying insects discriminate complex temporal patterns of acoustic stimuli. For bats using frequency modulated sonar calls (FM bats), there are no behavioral data on the perception of sinusoidally frequency modulated (SFM) stimuli. Discrimination performance for SFM stimuli of varying modulation depth was measured in 4 Tadarida brasiliensis in a two-alternative, forced choice procedure. A center frequency of 40 kHz was modulated with rates between 10 and 2000 Hz. It was found that discrimination performance improved from a mean threshold modulation depth of 3.05 kHz at a modulation rate of 2000 Hz to 1.58 kHz at a modulation rate of 10 Hz. Psychoacoustical modulation depth thresholds of T. brasiliensis are thus distinctly larger than those observed in bat species emitting constant frequency (CF) components followed by an FM-sweep, in active echolocation experiments. The modulation thresholds of T. brasiliensis are discussed in connection with the ability of bats to discriminate insect wingbeats. A comparison between non-echolocating mammals and the FM bat T. brasiliensis shows that the ability to echolocate is not reflected in the modulation thresholds.

Acoustic Stimulation↗
PubMed2006

Keeping up with bats: dynamic auditory tuning in a moth.

Many night-flying insects evolved ultrasound sensitive ears in response to acoustic predation by echolocating bats . Noctuid moths are most sensitive to frequencies at 20-40 kHz , the lower range of bat ultrasound . This may disadvantage the moth because noctuid-hunting bats in particular echolocate at higher frequencies shortly before prey capture and thus improve their echolocation and reduce their acoustic conspicuousness . Yet, moth hearing is not simple; the ear's nonlinear dynamic response shifts its mechanical sensitivity up to high frequencies. Dependent on incident sound intensity, the moth's ear mechanically tunes up and anticipates the high frequencies used by hunting bats. Surprisingly, this tuning is hysteretic, keeping the ear tuned up for the bat's possible return. A mathematical model is constructed for predicting a linear relationship between the ear's mechanical stiffness and sound intensity. This nonlinear mechanical response is a parametric amplitude dependence that may constitute a feature common to other sensory systems. Adding another twist to the coevolutionary arms race between moths and bats, these results reveal unexpected sophistication in one of the simplest ears known and a novel perspective for interpreting bat echolocation calls.

Acoustic Stimulation↗
PubMed1996

Does behavior reflect phylogeny in swiftlets (Aves: Apodidae)? A test using cytochrome b mitochondrial DNA sequences.

Swiftlets are small insectivorous birds, many of which nest in caves and are known to echolocate. Due to a lack of distinguishing morphological characters, the taxonomy of swiftlets is primarily based on the presence or absence of echolocating ability, together with nest characters. To test the reliability of these behavioral characters, we constructed an independent phylogeny using cytochrome b mitochondrial DNA sequences from swiftlets and their relatives. This phylogeny is broadly consistent with the higher classification of swifts but does not support the monophyly of swiftlets. Echolocating swiftlets (Aerodramus) and the nonecholocating "giant swiftlet" (Hydrochous gigas) group together, but the remaining nonecholocating swiftlets belonging to Collocalia are not sister taxa to these swiftlets. While echolocation may be a synapomorphy of Aerodramus (perhaps secondarily lost in Hydrochous), no character of Aerodramus nests showed a statistically significant fit to the molecular phylogeny, indicating that nest characters are not phylogenetically reliable in this group.

Animals↗
PubMed2005

Male sperm whale acoustic behavior observed from multipaths at a single hydrophone.

Sperm whales generate transient sounds (clicks) when foraging. These clicks have been described as echolocation sounds, a result of having measured the source level and the directionality of these signals and having extrapolated results from biosonar tests made on some small odontocetes. The authors propose a passive acoustic technique requiring only one hydrophone to investigate the acoustic behavior of free-ranging sperm whales. They estimate whale pitch angles from the multipath distribution of click energy. They emphasize the close bond between the sperm whale's physical and acoustic activity, leading to the hypothesis that sperm whales might, like some small odontocetes, control click level and rhythm. An echolocation model estimating the range of the sperm whale's targets from the interclick interval is computed and tested during different stages of the whale's dive. Such a hypothesis on the echolocation process would indicate that sperm whales echolocate their prey layer when initiating their dives and follow a methodic technique when foraging.

Acoustics↗
PubMed2002

Functional organization of the pallid bat auditory cortex: emphasis on binaural organization.

This report maps the organization of the primary auditory cortex of the pallid bat in terms of frequency tuning, selectivity for behaviorally relevant sounds, and interaural intensity difference (IID) sensitivity. The pallid bat is unusual in that it localizes terrestrial prey by passively listening to prey-generated noise transients (1-20 kHz), while reserving high-frequency (<30 kHz) echolocation for obstacle avoidance. The functional organization of its auditory cortex reflects the need for specializations in echolocation and passive sound localization. Best frequencies were arranged tonotopically with a general increase in the caudolateral to rostromedial direction. Frequencies between 24 and 32 kHz were under-represented, resulting in hypertrophy of frequencies relevant for prey localization and echolocation. Most neurons (83%) tuned <30 kHz responded preferentially to broadband or band-pass noise over single tones. Most neurons (62%) tuned >30 kHz responded selectively or exclusively to the 60- to 30-kHz downward frequency-modulated (FM) sweep used for echolocation. Within the low-frequency region, neurons were placed in two groups that occurred in two separate clusters: those selective for low- or high-frequency band-pass noise and suppressed by broadband noise, and neurons that showed no preference for band-pass noise over broadband noise. Neurons were organized in homogeneous clusters with respect to their binaural response properties. The distribution of binaural properties differed in the noise- and FM sweep-preferring regions, suggesting task-dependent differences in binaural processing. The low-frequency region was dominated by a large cluster of binaurally inhibited neurons with a smaller cluster of neurons with mixed binaural interactions. The FM sweep-selective region was dominated by neurons with mixed binaural interactions or monaural neurons. Finally, this report describes a cortical substrate for systematic representation of a spatial cue, IIDs, in the low-frequency region. This substrate may underlie a population code for sound localization based on a systematic shift in the distribution of activity across the cortex with sound source location.

Animals↗