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PubMed2002

Spectral determination of responses to species-specific calls in the dorsal nucleus of the lateral lemniscus.

This study evaluated how neurons in the dorsal nucleus of the lateral lemniscus (DNLL) in Mexican free-tailed bats respond to both tone bursts and species-specific calls. Up to 20 calls were presented to each neuron, of which 18 were social communication and 2 were echolocation calls. We also measured excitatory response regions (ERRs): the range of tone burst frequencies that evoked discharges at a fixed intensity. Neurons were unselective for one or another call in that each neuron responded to any call so long as the call had energy that encroached on its ERR. Additionally, responses were evoked by the same set of calls, and with similar spike counts, when they were presented normally or reversed. By convolving activity in the ERRs with the spectrogram of each call, we showed that responses to tones accurately predicted discharge patterns evoked by species-specific calls. DNLL cells are remarkably homogeneous in that neurons having similar BFs responded to each of the species-specific calls with similar response profiles. The homogeneity was further illustrated by the ability to accurately predict the response profiles of a particular DNLL cell to species-specific calls from the ERR of another similarly tuned DNLL cell. Thus DNLL neurons tuned to the same or similar frequencies responded to species-specific calls with latencies and temporal discharge patterns that were so similar as to be virtually interchangeable. What this suggests is that DNLL responses evoked by complex sounds can be largely explained by a simple summation of the excitation in each neuron's ERR. Finally, superimposing the spectrograms of each call on the responses evoked by that call revealed that the DNLL population response re-creates both the spectral and the temporal features of each signal.

Acoustic Stimulation↗
PubMed2003

Echoes of bat-pollinated bell-shaped flowers: conspicuous for nectar-feeding bats?

Nectar-feeding glossophagine bats searching for flowers are guided by their echolocation system as well as olfactory cues in detecting and recognizing nectar sources. Therefore, chiropterophilous plants, which depend on these bats as pollinators, may be expected to have evolved acoustically conspicuous flowers that facilitate detection. As it is poorly understood how bats manage to find and recognize flowers acoustically, we investigated the echoes of some of the flowers pollinated by bats. Echoes of bell-shaped bat-pollinated flowers have characteristic features with respect to the echoes they reflect to a calling bat and differ from the echoes of leaves or other objects in their surroundings: the echoes are comparatively long and of complex spectral composition. Owing to the specific shape of the flowers, characteristic 'spectral directional patterns' result when the spectra of the echoes are plotted against the angle of sound incidence. We suggest that bats are able to recognize such flowers - and probably other objects as well - not only by a characteristic spectral composition of the echo but also by comparing sequential echoes, at the same time taking into account their exact calling position relative to the object.

Adaptation, Biological↗
PubMed2006

Extreme diving of beaked whales.

Sound-and-orientation recording tags (DTAGs) were used to study 10 beaked whales of two poorly known species, Ziphius cavirostris (Zc) and Mesoplodon densirostris (Md). Acoustic behaviour in the deep foraging dives performed by both species (Zc: 28 dives by seven individuals; Md: 16 dives by three individuals) shows that they hunt by echolocation in deep water between 222 and 1885 m, attempting to capture about 30 prey/dive. This food source is so deep that the average foraging dives were deeper (Zc: 1070 m; Md: 835 m) and longer (Zc: 58 min; Md: 47 min) than reported for any other air-breathing species. A series of shallower dives, containing no indications of foraging, followed most deep foraging dives. The average interval between deep foraging dives was 63 min for Zc and 92 min for Md. This long an interval may be required for beaked whales to recover from an oxygen debt accrued in the deep foraging dives, which last about twice the estimated aerobic dive limit. Recent reports of gas emboli in beaked whales stranded during naval sonar exercises have led to the hypothesis that their deep-diving may make them especially vulnerable to decompression. Using current models of breath-hold diving, we infer that their natural diving behaviour is inconsistent with known problems of acute nitrogen supersaturation and embolism. If the assumptions of these models are correct for beaked whales, then possible decompression problems are more likely to result from an abnormal behavioural response to sonar.

Activity Cycles↗
PubMed2006

Lack of generalization of object discrimination between spatial contexts by a bat.

Discrimination and generalization are important elements of cognition in the daily lives of animals. Nectar-feeding bats detect flowers by olfaction and probably vision, but also use echolocation and echo-perception of flowers in immediate target surroundings. The echo received from an interference-rich flower corolla is a function of a bat's own relative position in space. This raises the question how easily a free-flying bat will generalize an echo stimulus from a learning situation to a new spatial context where differences in relative flight approach trajectories may lead to an unfamiliar spectral composition of the self-generated echoes. We trained free-flying Glossophaga soricina in echoacoustic discrimination in a two-alternative forced-choice (2-AFC) paradigm at location A. We then tested at location B for spontaneous transfer of discrimination ability. Bats did not spontaneously transfer the discrimination ability acquired at A to location B. This lack of spontaneous generalization may have been caused by factors of the underlying learning mechanisms. 2-AFC tasks may not be representative of the natural foraging behaviour of flower-visiting bats. In contrast to insect-eating bats that constantly evaluate the environment to detect unpredictable prey, the spatial stability of flowers may allow flower visitors to rely on spatial memory to guide foraging. The 2-AFC task requires the disregard (learned irrelevance) of salient spatial location cues that are different at each new location. In Glossophaga, a conjunction between spatial context and 2-AFC discrimination learning may have inhibited the transfer of learned irrelevance of spatial location in the 2-AFC task to new spatial locations. Alternatively, the bats may have learnt the second discrimination task completely anew, and were faster only because of an acquired learning set. We suggest a dissociation between 2-AFC task acquisition and novel object discrimination learning to resolve the issue.

Animals↗
PubMed2001

Ultrasound avoidance behaviour in the bushcricket Tettigonia viridissima (Orthoptera: Tettigoniidae).

The responses of female Tettigonia viridissima to simulated bat echolocation calls were examined during tethered flight. The insects responded with three distinct behaviours, which occurred at graded stimulus intensities. At low intensities (threshold 54 dB SPL), T. viridissima responded by steering away from the sound source (negative phonotaxis). At intensities approximately 10 dB higher, beating of the hindwing was interrupted, although the insect remained in the flight posture. A diving response (cessation of the wingbeat, closure of the forewings and alignment of the legs against the body) occurred with a threshold of 76 dB SPL. Considering these thresholds, we estimate that the diving response occurs at approximately the sound amplitude at which many aerial-hawking bats first receive echoes from the insect. The other behaviours probably occur before the bat detects the insect and should therefore be interpreted as early avoidance behaviours. The repertoire of startle responses in T. viridissima, with directional and non-directional components, is similar to those of crickets and moths, but quite different from those described for another bushcricket (Neoconocephalus ensiger), which shows only a non-directional response. This supports the conclusion that bat-evasive behaviours are not conserved within the Tettigoniidae, but instead are shaped by the ecological constraints of the insects.

Animals↗
PubMed1975

Sound localization by the bottlenose porpoise Tursiops truncatus.

1. Sound localization was measured behaviourally for the Atlantic bottlenose porpoise (Tursiops truncatus) using a wide range of pure tone pulses as well as clicks simulating the species echolocation click. 2. Measurements of the minimum audible angle (MAA) on the horizontal plane give localization discrimination thresholds of between 2 and 3 degrees for sounds from 20 to 90 kHz and thresholds from 2-8 to 4 degrees at 6, 10 and 100 kHz. With the azimuth of the animal changed relative to the speakers the MAAs were 1-3-1-5 degrees at an azimuth of 15 degrees and about 5 degrees for an azimuth of 30 degrees. 3. MAAs to clicks were 0-7-0-8 degrees. 4. The animal was able to do almost as well in determining the position of vertical sound sources as it could for horizontal localization. 5. The data indicate that at low frequencies the animal may have been localizing by using the region around the external auditory meatus as a detector, but at frequencies about 20 kHz it is likely that the animal was detecting sounds through the lateral sides of the lower jaw. 6. Above 20 kHz, it is likely that the animal was localizing using binaural intensity cues. 7. Our data support evidence that the lower jaw is an important channel for sound detection in Tursiops.

Animals↗
PubMed1980

Coding of directional information by single neurones in the S-segment of the FM bat, Myotis lucifugus.

1. Response parameters of S-segment neurones of the FM bat Myotis lucifugus were measured as a sound was delivered from different azimuthal angles around the animal's head. 2. The response parameters investigated were the amplitude and threshold of the evoked potential (N3) of the S-segment, together with the threshold, latency and number of impulses (per stimulus pulse) of single units. 3. All the neurones studied had their lowest thresholds either at 20-40 degrees contralateral, or 20-40 degrees ipsilateral or at the front (0 degrees). 4. The amplitude of the sound affected the relationship between stimulus direction and the amplitude of a non-monotonic N3, and the relationship between stimulus direction and the number of impulses of a non-monotonic single unit. It had so such effects with a monotonic N3 and a monotonic single unit. 5. From a study of N3 amplitudes and numbers of impulses of single neurones, it appeared that an azimuthal difference as small as 3 degrees could be easily coded at a 95% correct level with stimuli presented at around 20 degrees ipsilateral, 20 degrees contralateral, and at the front. 6. The inter-aural pressure difference (IPD), which is considered an essential cue for echolocation in Myotis (Shimozawa et al. 1974), changed linearly with angle from 0 to 40 degrees lateral at a rate of 0.4 dB/degree for sounds between 33.5 and 49.0 kHz. 7. Assuming the just-detectable IPD to be 0.5 dB (as in man), the minimum detectable azimuthal difference of Myotis around the median plane would be 1.25 degrees.

Animals↗
PubMed2007

Vocal premotor activity in the superior colliculus.

Chronic neural recordings were taken from the midbrain superior colliculus (SC) of echolocating bats while they were engaged in one of two distinct behavioral tasks: virtual target amplitude discrimination (VTAD) and real oscillating target tracking (ROTT). In the VTAD task, bats used a limited range of sonar call features to discriminate the amplitude category of echoes, whereas in the ROTT task, the bat produced dynamically modulated sonar calls to track a moving target. Newly developed methods for chronic recordings in unrestrained, behaving bats reveal two consistent bouts of SC neural activity preceding the onset of sonar vocalizations in both tasks. A short lead bout occurs tightly coupled to vocal onset (VTAD, -5.1 to -2.2 ms range, -3.6 +/- 0.7 ms mean lead time; ROTT, -3.0 to + 0.4 ms range, -1.2 +/- 1.3 ms mean lead time), and this activity may play a role in marking the time of each sonar emission. A long lead bout in SC activity occurs earlier and spreads over a longer interval (VTAD, -40.6 to -8.4 ms range, -22.2 +/- 3.9 ms mean lead time; ROTT, -29.8 to -7.1 ms range, -17.5 +/- 9.1 ms mean lead time) when compared with short lead events. In the goal-directed ROTT task, the timing of long lead event times vary with the bat's sonar call duration. This finding, along with behavioral studies demonstrating that bats adjust sonar call duration as they track targets at changing distance, suggests the bat SC contributes to range-dependent adjustments of sonar call duration.

Action Potentials↗
PubMed2006

A neural correlate of stochastic echo imaging.

Bats quickly navigate through a highly structured environment relying on echolocation. Large natural objects in the environment, like bushes or trees, produce complex stochastic echoes, which can be characterized by the echo roughness. Previous work has shown that bats can use echo roughness to classify the stochastic properties of natural objects. This study provides both psychophysical and electrophysiological data to identify a neural correlate of statistical echo analysis in the bat Phyllostomus discolor. Psychophysical results show that the bats require a fixed minimum roughness of 2.5 (in units of base 10 logarithm of the stimulus fourth moment) for roughness discrimination. Electrophysiological results reveal a subpopulation of 15 of 94 recorded cortical units, located in an anterior region of auditory cortex, whose rate responses changed significantly with echo roughness. It is shown that the behavioral ability to discriminate differences in the statistics of complex echoes can be quantitatively predicted by the neural responses of this subpopulation of auditory-cortical neurons.

Acoustic Stimulation↗
PubMed2004

Combination sensitivity and processing of communication calls in the inferior colliculus of the Moustached Bat Pteronotus parnellii.

Many animals use complex communication calls in social behaviors. In some species we know the features in the calls that elicit particular behaviors, but we do not understand how the auditory system encodes the calls. Nor do we understand the mechanisms underlying neural selectivity to calls. Our studies of the auditory midbrain of the Moustached Bat Pteronotus parnellii have revealed a neural mechanism important for generating selective responses to calls. Neurons that integrate information across different frequencies show selectivity to communication calls. "Combination sensitivity" may be a common mechanism for encoding complex sounds because it is also important for encoding echolocation signals.

Animal Communication↗
PubMed2003

Synchronization of a nonlinear oscillator: processing the cf component of the echo-response signal in the cochlea of the mustached bat.

Cochlear microphonic potential (CM) was recorded from the CF2 region and the sparsely innervated zone (the mustached bat's cochlea fovea) that is specialized for analyzing the Doppler-shifted echoes of the first-harmonic (approximately 61 kHz) of the constant-frequency component of the echolocation call. Temporal analysis of the CM, which is tuned sharply to the 61 kHz cochlear resonance, revealed that at the resonance frequency, and within 1 msec of tone onset, CM is broadly tuned with linear magnitude level functions. CM measured during the ongoing tone and in the ringing after tone offset is 50 dB more sensitive, is sharply tuned, has compressive level functions, and the phase leads onset CM by 90 degrees: an indication that cochlear responses are amplified during maximum basilar membrane velocity. For high-level tones above the resonance frequency, CM appears at tone onset and after tone offset. Measurements indicate that the two oscillators responsible for the cochlear resonance, presumably the basilar and tectorial membranes, move together in phase during the ongoing tone, thereby minimizing net shear between them and hair cell excitation. For tones within 2 kHz of the cochlear resonance the frequency of CM measured within 2 msec of tone onset is not that of the stimulus but is proportional to it. For tones just below the cochlear resonance region CM frequency is a constant amount below that of the stimulus depending on CM measurement delay from tone onset. The frequency responses of the CM recorded from the cochlear fovea can be accounted for through synchronization between the nonlinear oscillators responsible for the cochlear resonance and the stimulus tone.

Acoustic Stimulation↗
PubMed2003

The development of a single frequency place in the mammalian cochlea: the cochlear resonance in the mustached bat Pteronotus parnellii.

Cochlear microphonic potentials (CMs) were recorded from the sharply tuned, strongly resonant auditory foveae of 1- to 5-week-old mustached bats that were anesthetized with Rompun and Ketavet. The fovea processes Doppler-shifted echo responses of the constant-frequency component of echolocation calls. During development, the frequency and tuning sharpness of the cochlear resonance increases, and CM ringing persists for longer after the tone. CM is relatively insensitive at tone onset and grows linearly with increased stimulus level. During the tone, the CM is more sensitive and grows compressively with increased stimulus level and phase leads onset CM by 90 degrees for frequencies below the resonance. CM during the ringing is also sensitive and compressive and phase leads onset CM by 180 degrees below the resonance and lags it by 180 degrees above the resonance. Throughout postnatal development, CMs measured during the tone and in the ringing increase both in sensitivity and compression. The cochlear resonance appears to be attributable to interaction between two oscillators. The more broadly tuned oscillator dominates the onset response, and the narrowly tuned oscillator dominates the ringing. Early in development, mechanical coupling between the oscillators results in a relatively broadly tuned system with several frequency modes in the CM at tone onset and in the CM ringing. Beating occurs between the resonance and the stimulus response during the tone and between two components of the narrowly tuned oscillator at tone offset. At maturity, the CM has three modes for frequencies within 10 kHz of the resonance at tone onset and a single, sharply tuned mode in the ringing.

Acoustic Stimulation↗
PubMed2004

The emergence of temporal hyperacuity from widely tuned cell populations.

Typically, individual neural cells operate on a millisecond time scale yet behaviorally animals reveal sub-microsecond acuity. Our model resolves this huge discrepancy by using populations of many widely tuned cells to attain sub-microsecond resolution in a temporal discrimination task. An echolocating bat uses its auditory system to locate objects and it demonstrates remarkable temporal precision in psychophysical tasks. Auditory cells were simulated using realistic parameters and connected in three ascending layers with descending projections from auditory cortex. Coincidence detection of firing collicular cells at thalamus and subsequent integration of multiple inputs at cortex, produce an estimate of time represented as the mean of the active cortical population. Multiple estimates allow the model bat to use memory to recognize predictable change in stimuli values. The best performance is produced using cortical feedback and a computation of target time based on combining the current and previous estimates. Temporal hyperacuity is attained through population coding of physiologically realistic cells but depends on the inherent properties of the psychophysical task.

Acoustic Stimulation↗
PubMed2005

Effects of backward masking on the responses of the inferior collicular neurons in the big brown bat, Eptesicus fuscus.

Temporal features of sound convey information vital for behaviors as diverse as speech recognition by human and echolocation by bats. However, auditory stimuli presented in temporal proximity might interfere with each other. Although much progress has been made in the description of this phenomenon from psychophysical studies, the neural mechanism responsible for its formation at central auditory structures especially at the inferior colliculus (IC), a midbrain auditory nucleus which practically receives massive bilateral projections from all the major auditory structures in the brainstem, remains unclear. This study was designed to investigate it in vivo by using electrophysiological recording from the inferior collicular neurons of the big brown bat, Eptesicus fuscus. In our results, the responses of 12 (38%, n= 31) neurons to the test sound (leading sound) were obviously inhibited by the masker (lagging sound). The inhibitory effects in these neurons were correlated with the inter-stimulus level difference (SLD) and the inter-stimulus onset asynchrony (SOA) interval. The strength of backward masking increased with the masker intensity increasing, the test sound intensity decreasing and the SOA interval shortening. There were no obvious effects of backward masking on the responses of many other neurons (52%, 16/31), and yet in a part of these neurons, the neural inhibition of responses to the test sound was observed at the special SLD and the special SOA intervals. Moreover, few of the 31 sampled IC neurons (10%, 3/31) displayed facilitating responses to the test sound at the special SLD and the special SOA intervals. These data demonstrate that a lot of IC neurons are involved in the generation of the backward masking of acoustical perception. It is conjectured that the temporal dynamic integration between the leading inhibitory inputs evoked by the masker sound and the excitatory inputs evoked by the test sound might play a key role in shaping the acoustical response characteristics of the IC neurons.

Acoustic Stimulation↗
PubMed1984

Preference of a revolving target to a stationary one by the big brown bat, Eptesicus fuscus.

Utilizing a three-ramp platform, we studied the detection of a revolving and a stationary target in the presence of background clutter by trained Eptesicus fuscus. During the test, the mean amplitude of echo from either target was always larger than that of the background echoes at the bat-to-target distance of 30, 70 and 100 cm. The amplitude of the echo reflected back from a revolving target was modulated between a maximum and a minimum value. An electric motor was used to revolve a target. The frequency contents of the motor noise were mostly below 1 kHz. While the total percent response of approaching either target is always more than 90% at every bat-to-target distance tested, the bats approach a revolving target more frequently than a stationary one. Echolocation pulses emitted by the bats during the test were recorded and analyzed. The bats shortened their pulse durations and interpulse intervals and lowered the frequency contents as they entered into the crawling phase from the searching phase. Potential interference of background echoes and ambient noise with the performance of the bats is discussed. The preference of a revolving target to a stationary one by the bats is perhaps due to the fact that a revolving target has a higher releasing value than a stationary one does.

Animals↗
PubMed1982

Encoding of target range and its representation in the auditory cortex of the mustached bat.

The time course of acoustic events is a critical element for the recognition of biologically meaningful sounds. Echolocating bats analyze the time intervals between their emitted biosonar pulses and the echoes returning from objects to assess target distance (range). In this study, we have explored the auditory cortex of the mustached bat, Pteronotus parnellii rubiginosus, using pairs of acoustic stimuli mimicking the multiharmonic biosonar signals (pulses) used by this species and their echoes. A discrete field of auditory cortex dorsorostral to the tonotopically organized primary field contains neurons which are insensitive to pure tone, frequency-modulated (FM), or noise stimuli presented singly. Rather, they respond strongly to pairs of stimuli, specifically, the fundamental FM component of the pulse paired with an FM component of one of the higher harmonics of the echo. We call these neurons FM1-FMn facilitation neurons. There are three separate longitudinal clusters in this cortical area containing FM1-FM2, FM1-FM3, and FM1-FM4 neurons, respectively. Moreover, FM1-FMn neurons are specifically sensitive to the time delay between the two FM components, i.e., the time delay of the echo from the pulse. Thus, they can decode target range. Two types of delay-sensitive neurons were found. Tracking neurons, whose response to echo delay varied according to repetition rate and stimulus duration, were found rarely. Delay-tuned neurons, which were tuned to specific time delays (best delays) of the echo from the pulse, were much more evident. Both types of neurons are organized into columns with similar best delays, and the best delay of delay-tuned neurons was found to increase systematically along the cortical surface in the rostrocaudal direction. This area, therefore, contains a neural representation of target range along this best delay axis. Such an axis exists in each of the clusters of FM1-FM2, FM1-FM3, and FM1-FM4 neurons. This is a new type of cortical organization which is not tonotopic but which represents an important acoustic cue related to the time course of acoustic events.

Animals↗
PubMedDate not supplied

[Features of the reactions of neurons of the cochlear nuclei of the pointed-eared bat Myotis oxygnathus to ultrasonic stimuli].

Studies of the reactions of neurones in the cochlear nuclei in the bat Myotis oxygnathus to ultrasonic stimulation show that 61% of the investigated neurones in a frequency range 20-40 kHz change tonic response for repetitive bursts, this change being dependent on the filling frequency and the intensity of stimulation. It is suggested that peculiarities of the neuronal activity result from the effect of a special inhibitory system presented in bats which uses in echolocation a wide range frequency-modulated impulses.

Animals↗
PubMed1995

A computational model of auditory perception.

Building functional models of the auditory system that incorporate neurophysiological, behavioral findings is essential to uncovering mechanisms underlying auditory perception. This paper presents results from modeling the echolocation ability of the FM bat, Myotis lucifugus, using MATLAB on a Sun Sparc-10 as a computational and modeling platform. It is shown that the adaptability and versatility of such computational software is ideal for modeling this complex biological system.

Animals↗