Echolocation versus echo suppression in humans

Several studies have shown that blind humans can gather spatial information through echolocation. However, when localizing sound sources, the precedence effect suppresses spatial information of echoes, and thereby conflicts with effective echolocation. This study investigates the interaction of echolocation and echo suppression in terms of discrimination suppression in virtual acoustic space. In the ‘Listening’ experiment, sighted subjects discriminated between positions of a single sound source, the leading or the lagging of two sources, respectively. In the ‘Echolocation’ experiment, the sources were replaced by reflectors. Here, the same subjects evaluated echoes generated in real time from self-produced vocalizations and thereby discriminated between positions of a single reflector, the leading or the lagging of two reflectors, respectively. Two key results were observed. First, sighted subjects can learn to discriminate positions of reflective surfaces echo-acoustically with accuracy comparable to sound source discrimination. Second, in the Listening experiment, the presence of the leading source affected discrimination of lagging sources much more than vice versa. In the Echolocation experiment, however, the presence of both the lead and the lag strongly affected discrimination. These data show that the classically described asymmetry in the perception of leading and lagging sounds is strongly diminished in an echolocation task. Additional control experiments showed that the effect is owing to both the direct sound of the vocalization that precedes the echoes and owing to the fact that the subjects actively vocalize in the echolocation task.     

Neural correlates of natural human echolocation in early and late blind echolocation experts

Background

A small number of blind people are adept at echolocating silent objects simply by producing mouth clicks and listening to the returning echoes. Yet the neural architecture underlying this type of aid-free human echolocation has not been investigated. To tackle this question, we recruited echolocation experts, one early- and one late-blind, and measured functional brain activity in each of them while they listened to their own echolocation sounds.

Results

When we compared brain activity for sounds that contained both clicks and the returning echoes with brain activity for control sounds that did not contain the echoes, but were otherwise acoustically matched, we found activity in calcarine cortex in both individuals. Importantly, for the same comparison, we did not observe a difference in activity in auditory cortex. In the early-blind, but not the late-blind participant, we also found that the calcarine activity was greater for echoes reflected from surfaces located in contralateral space. Finally, in both individuals, we found activation in middle temporal and nearby cortical regions when they listened to echoes reflected from moving targets.

Conclusions

These findings suggest that processing of click-echoes recruits brain regions typically devoted to vision rather than audition in both early and late blind echolocation experts.     

Echolocation of Multiple Targets in 3-D Space from a Single Emission

Using frequency-modulated echolocation sound, bat can capture a moving target in real three-dimensional (3-D) space. It is impossible to locate multiple targets in 3-D space by using only the delay time between an emission and the resultingechoes received at two points (i.e., two ears). To locate multiple targets in 3-D space requires directional information for each target. 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. The frequency of the spectral notch, which is the frequency corresponding to the minimum of the external ear's transfer function (EEDNF), provides a crucial cue for directional localization. In the model we present, a computational model todiscriminate multiple close targets in 3-D space utilizing echoes evoked by a single emission by distinguishing the interference of echoes from each object and the EEDNF corresponding to each target.     

Size does not matter: size-invariant echo-acoustic object classification

Echolocating bats can not only extract spatial information from the auditory analysis of their ultrasonic emissions, they can also discriminate, classify and identify the three-dimensional shape of objects reflecting their emissions. Effective object recognition requires the segregation of size and shape information. Previous studies have shown that, like in visual object recognition, bats can transfer an echo-acoustic object discrimination task to objects of different size and that they spontaneously classify scaled versions of virtual echo-acoustic objects according to trained virtual-object standards. The current study aims to bridge the gap between these previous findings using a different class of real objects and a classification—instead of a discrimination paradigm. Echolocating bats (Phyllostomus discolor) were trained to classify an object as either a sphere or an hour-glass shaped object. The bats spontaneously generalised this classification to objects of the same shape. The generalisation cannot be explained based on similarities of the power spectra or temporal structures of the echo-acoustic object images and thus require dedicated neural mechanisms dealing with size-invariant echo-acoustic object analysis. Control experiments with human listeners classifying the echo-acoustic images of the objects confirm the universal validity of auditory size invariance. The current data thus corroborate and extend previous psychophysical evidence for sonar auditory-object normalisation and suggest that the underlying auditory mechanisms following the initial neural extraction of the echo-acoustic images in echolocating bats may be very similar in bats and humans.     

The transfer function of a target limits the jitter detection threshold with signals of echolocating FM-bats

The delay jitter discrimination threshold in bats is a disputed subject. Some investigators have obtained results indicating that bats are able to discriminate alternations in delay down to 10 ns, which appears incredible for purely physical reasons. Using actual bat echolocation sequences recorded during an easy detection task to measure simulated delay jitter, it is shown here that jitter detection thresholds in the order of some tens of nanoseconds are actually physically realizable. However, if the transfer function of the target simulating apparatus is not perfect, the lowest thresholds are in the order of hundreds of nanoseconds and variable between individual bats. This phenomenon is shown to arise as a consequence of the variation in signal parameters from call to call. When the transfer function from a real jitter experiment was artificially applied to the echoes, the jitter detection thresholds again were several hundred nanoseconds. This is the first study to point out a limiting role of the transfer function of a system faced with variations in echolocation signal parameters, something that should be considered in evaluating all sonar systems with variable signal structure.     

Discrimination Contours for Moving Sounds Reveal Duration and Distance Cues Dominate Auditory Speed Perception

Evidence that the auditory system contains specialised motion detectors is mixed. Many psychophysical studies confound speed cues with distance and duration cues and present sound sources that do not appear to move in external space. Here we use the ‘discrimination contours’ technique to probe the probabilistic combination of speed, distance and duration for stimuli moving in a horizontal arc around the listener in virtual auditory space. The technique produces a set of motion discrimination thresholds that define a contour in the distance-duration plane for different combination of the three cues, based on a 3-interval oddity task. The orientation of the contour (typically elliptical in shape) reveals which cue or combination of cues dominates. If the auditory system contains specialised motion detectors, stimuli moving over different distances and durations but defining the same speed should be more difficult to discriminate. The resulting discrimination contours should therefore be oriented obliquely along iso-speed lines within the distance-duration plane. However, we found that over a wide range of speeds, distances and durations, the ellipses aligned with distance-duration axes and were stretched vertically, suggesting that listeners were most sensitive to duration. A second experiment showed that listeners were able to make speed judgements when distance and duration cues were degraded by noise, but that performance was worse. Our results therefore suggest that speed is not a primary cue to motion in the auditory system, but that listeners are able to use speed to make discrimination judgements when distance and duration cues are unreliable.     

Characteristics of the acoustic field of interfering reflections and the echolocation hearing of the dolphin

A model of the acoustic field of interfering reflections from steel cylinders was developed. Analysis of the model showed the availability of great potential resources for a decrease of the influence of unwanted echoes and hence for increasing the signal-to-clatter ratio. The conformity of the available models of the echolocation hearing of the dolphin to the acoustic field of the clatter was considered. The participation of mandidle mental foramens in conducting the echo to the cochlea was considered. In this case the hearing aperture is determined by the dimensions of mental foramens, while the hearing base is determined by the distance between the mental foramens of the left and right mandible halves. There are good reasons to believe that the optimal dimensions of the aperture and the base of echolocation hearing of Odontoceti essentially increase the effectiveness of defense of their sonar from reverberation.     

ECHOLOCATION IN DOLPHINS WITH A DOLPHIN-BAT COMPARISON

ABSTRACT

Dolphins possess a highly sophisticated auditory system and a keen capability for echolocation. Signals are emitted in the form of high intensity, short duration, broadband exponentially decaying pulses. The frequency spectra of echolocation signals used by many dolphins are dependent on the output intensity of the signals and not on any fine tuning by the animals. When the output intensity is low, the center frequency of the click tends to be low. As the output intensity increases, the center frequency also tends to increase. The pulses propagate from the dolphin's melon in a relatively narrow beam, and echoes are received via the lower jaw, with a slightly wider beam. Echo- locating dolphins can detect targets at ranges of approximately 100 plus meters, depending on the size of the targets. Target discrimination experiments have shown that dolphins can discriminate the shape, size, material composition and internal structure of targets from the echoes. The broadband short duration properties of the signal allow the echoes to have high temporal resolution, so that within the structure of the echoes a considerable amount of information on the properties of the target can be conveyed. A brief comparison between the bat and dolphin sonar system will also be made. Bats typically emit much longer signals and a wider variety of different types of signals than dolphins. Signals used by some bats are suited to detecting Doppler shift, whereas the dolphin signal is designed to be tolerant of Doppler effects.     

Neural coding of echo-envelope disparities in echolocating bats

The effective use of echolocation requires not only measuring the delay between the emitted call and returning echo to estimate the distance of an ensonified object. To locate an object in azimuth and elevation, the bat’s auditory system must analyze the returning echoes in terms of their binaural properties, i.e., the echoes’ interaural intensity and time differences (IIDs and ITDs). The effectiveness of IIDs for echolocation is undisputed, but when bats ensonify complex objects, the temporal structure of echoes may facilitate the analysis of the echo envelope in terms of envelope ITDs. Using extracellular recordings from the auditory midbrain of the bat, Phyllostomus discolor, we found a population of neurons that are sensitive to envelope ITDs of echoes of their sonar calls. Moreover, the envelope-ITD sensitivity improved with increasing temporal fluctuations in the echo envelopes, a sonar parameter related to the spatial statistics of complex natural reflectors like vegetation. The data show that in bats envelope ITDs may be used not only to locate external, prey-generated rustling sounds but also in the context of echolocation. Specifically, the temporal fluctuations in the echo envelope, which are created when the sonar emission is reflected from a complex natural target, support ITD-mediated echolocation.     

Integration of Semi-Circular Canal and Otolith Cues for Direction Discrimination during Eccentric Rotations

Humans are capable of moving about the world in complex ways. Every time we move, our self-motion must be detected and interpreted by the central nervous system in order to make appropriate sequential movements and informed decisions. The vestibular labyrinth consists of two unique sensory organs the semi-circular canals and the otoliths that are specialized to detect rotation and translation of the head, respectively. While thresholds for pure rotational and translational self-motion are well understood surprisingly little research has investigated the relative role of each organ on thresholds for more complex motion. Eccentric (off-center) rotations during which the participant faces away from the center of rotation stimulate both organs and are thus well suited for investigating integration of rotational and translational sensory information. Ten participants completed a psychophysical direction discrimination task for pure head-centered rotations, translations and eccentric rotations with 5 different radii. Discrimination thresholds for eccentric rotations reduced with increasing radii, indicating that additional tangential accelerations (which increase with radius length) increased sensitivity. Two competing models were used to predict the eccentric thresholds based on the pure rotation and translation thresholds: one assuming that information from the two organs is integrated in an optimal fashion and another assuming that motion discrimination is solved solely by relying on the sensor which is most strongly stimulated. Our findings clearly show that information from the two organs is integrated. However the measured thresholds for 3 of the 5 eccentric rotations are even more sensitive than predictions from the optimal integration model suggesting additional non-vestibular sources of information may be involved.     

Evidence of effects of human disturbance on alert response in Père David's deer (Elaphurus davidianus)

To understand effects of human disturbance on alert response of Père David's deer, we carried out an experiment in the Dafeng Père David's Deer Reserve (32°59′–33°03′N, 120°47′–120°53′E), China. In the spring and summer, we observed alert responses (including stare, walking away, and flee) of deer and recorded the intensity of tourist disturbance in a small display pen using a laser‐range finder to measure the alert distance of a free‐ranging group in a large enclosure. We also recorded the pattern of head orientation when deer were resting in these two deer groups. After statistical analysis, we found that: 1) in small pen, the frequency of alert response was significantly different among different intensities of human disturbance; strong disturbance resulted in higher frequency of alert response; 2) stare distance in the free‐ranging group in summer was significantly longer than that in spring, but the distance of walking away and the distance of flee showed no significant difference between the two seasons; and 3) in free‐ranging group, there was no significant directional difference in head orientation, whereas in display group, there was a significant directional difference in head orientation. We suggest that: 1) under the captive situation, human disturbance may be one of the factors that affect alert response in Père David's deer; and 2) Père David's deer adopted different alert response to adapt to human disturbance under different circumstance. We recommended that relationships between alert response and human disturbance should be considered in ex situ conservation of this field extinct deer. Zoo Biol 26:461–470, 2007. © 2007 Wiley‐Liss, Inc.     

Why do shrews twitter? Communication or simple echo-based orientation

Shrews are very vocal animals. We tested behaviourally whether the high-pitched laryngeal ‘twittering’ calls of as-yet unclear function serve for communication or echo-based orientation. We used a representative species from each of the two largest phylogenetic groups of shrews. In both species, experimental manipulation of substrate density, but not of the likelihood of conspecific presence, affected the shrews'' call rate when exploring an unknown environment. This adaptation of call rate to the degree of habitat clutter parallels bat echolocation and suggests that shrews may use the echoes and reverberations of their calls for identifying routes through their habitat or for probing habitat type. To assess the acoustic feasibility of shrew echo orientation, we ensonified shrew habitats in the field with an ‘artificial shrew’ (small speaker mounted close to a sensitive microphone). The data showed that shrew-like calls can indeed yield echo scenes useful for habitat assessment at close range, but beyond the range of the shrews'' vibrissae.     

Frequency discrimination of brief tonal steps as a function of frequency in the lesser bulldog bat

In a two-alternative, forced-choice task lesser bulldog bats were trained to distinguish between a pure tone pulse and a pulse composed of a series of brief tonal steps oscillating between two different frequencies. The tone-step pulse gradually approximates the pure tone pulse as the frequency difference between the steps becomes progressively smaller. Frequency difference limens for the brief tonal frequency steps were determined for a broad range of ultrasonic frequencies. The variation in tone-step difference limens with frequency appears to be correlated to the frequency structure of the bat's short-constant-frequency/frequency-modulated echolocation sound. There was a marked decline in the value of the relative frequency difference limens (Weber ratio) over a fairly narrow range of frequencies above the constant frequency and a sharp increase in threshold above this range. The relative thresholds for frequency discrimination were small and uniform over the frequency range of the frequency-modulated sweep and increased for frequencies below the frequency- modulated sweep. Thus, the most accurate frequency-discrimination abilities occur over a narrow frequency range around the frequency of the constant-frequency component of returning echoes. Frequency discrimination over the range of frequencies of the frequency-modulated component is relatively good. Accepted: 20 March 1999     

A modeling approach to explain pulse design in bats

In this modeling study we wanted to find out why bats of the family Vespertilionidae (and probably also members of other families of bats) use pulses with a certain bandwidth and duration. Previous studies have only speculated on the function of bandwidth and pulse duration in bat echolocation or addressed this problem by assuming that bats optimize echolocation parameters to achieve very fine acuities in receiving single echoes. Here, we take a different approach by assuming that bats in nature rarely receive single echoes from each pulse emission, but rather many highly overlapping echoes. Some echolocation tasks require individual echoes to be separated to reconstruct reflection points in space. We used an established hearing model to investigate how the parameters bandwidth and pulse duration influence the separation of overlapping echoes. Our findings corroborate the following previously unknown or unsubstantiated facts: 1. Broadening the bandwidth improves the bat's lower resolution limit. 2. Increasing the sweep rate (defined by bandwidth and pulse duration) improves acuity of each extracted echo. 3. Decreasing the sweep rate improves the probability of frequency channels being activated. Since facts 2 and 3 affect sweep rate in an opposing fashion, an optimum sweep rate will exist, depending on the quality of the returning echoes and the requirements of the bat to improve acuity. The existence of an optimal sweep rate explains why bats are likely to use certain combinations of bandwidth and pulse duration to obtain such sweep rates.     

The Effect of Self-Efficacy on Visual Discrimination Sensitivity

Can subjective belief about one''s own perceptual competence change one''s perception? To address this question, we investigated the influence of self-efficacy on sensory discrimination in two low-level visual tasks: contrast and orientation discrimination. We utilised a pre-post manipulation approach whereby two experimental groups (high and low self-efficacy) and a control group made objective perceptual judgments on the contrast or the orientation of the visual stimuli. High and low self-efficacy were induced by the provision of fake social-comparative performance feedback and fictional research findings. Subsequently, the post-manipulation phase was performed to assess changes in visual discrimination thresholds as a function of the self-efficacy manipulations. The results showed that the high self-efficacy group demonstrated greater improvement in visual discrimination sensitivity compared to both the low self-efficacy and control groups. These findings suggest that subjective beliefs about one''s own perceptual competence can affect low-level visual processing.     

Discrimination of insect wingbeat-frequencies by the batRhinolophus ferrumequinum

Summary Five Greater Horseshoe bats,Rhinolophus ferrumequinum, were trained in a two-alternative forced-choice procedure to discriminate between artificial echoes of insects fluttering at different wingbeat rates. The stimuli were electronically produced phantom targets simulating fluttering insects with various wingbeat frequencies (Figs. 3, 4). Difference thresholds for wingbeat rates of 50 Hz and 100 Hz were determined. For an S+ of 50 Hz the difference threshold values lay between 2.8 and 4.6 Hz for individual bats; with an S+ of 100 Hz they increased to between 9.8 and 12.0 Hz (Figs. 5, 6, Table 1).Three bats, previously trained to discriminate between a S+ of 50 Hz and a S– with a lower wingbeat rate, were tested with higher frequency stimuli. When they had to decide between their old S+ of 50 Hz and either a 60 or 70 Hz echo two bats continued to select the 50 Hz stimulus while the third bat now preferred the faster fluttering insects (Table 2).During the discrimination task the echolocation behavior of the bats was monitored. When the phantom targets were presented all bats increased their duty-cycle of sound emission from about 40% to sometimes near 70%. They did so by either emitting longer echolocation calls or by increasing the sound repetition rate (Figs. 7, 8).The results show that Greater Horseshoe bats can determine the wingbeat rate of flying insects with an accuracy between 6 and 12%. Possible cues for flutter rate determination by cf-fm bats from natural and artificial insect echoes are discussed.Abbreviations DC duty-cycle - PD pulse duration - PI pulse interval - cf constantfrequency - fm frequency modulation     

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)     

Different Auditory Feedback Control for Echolocation and Communication in Horseshoe Bats

Auditory feedback from the animal''s own voice is essential during bat echolocation: to optimize signal detection, bats continuously adjust various call parameters in response to changing echo signals. Auditory feedback seems also necessary for controlling many bat communication calls, although it remains unclear how auditory feedback control differs in echolocation and communication. We tackled this question by analyzing echolocation and communication in greater horseshoe bats, whose echolocation pulses are dominated by a constant frequency component that matches the frequency range they hear best. To maintain echoes within this “auditory fovea”, horseshoe bats constantly adjust their echolocation call frequency depending on the frequency of the returning echo signal. This Doppler-shift compensation (DSC) behavior represents one of the most precise forms of sensory-motor feedback known. We examined the variability of echolocation pulses emitted at rest (resting frequencies, RFs) and one type of communication signal which resembles an echolocation pulse but is much shorter (short constant frequency communication calls, SCFs) and produced only during social interactions. We found that while RFs varied from day to day, corroborating earlier studies in other constant frequency bats, SCF-frequencies remained unchanged. In addition, RFs overlapped for some bats whereas SCF-frequencies were always distinctly different. This indicates that auditory feedback during echolocation changed with varying RFs but remained constant or may have been absent during emission of SCF calls for communication. This fundamentally different feedback mechanism for echolocation and communication may have enabled these bats to use SCF calls for individual recognition whereas they adjusted RF calls to accommodate the daily shifts of their auditory fovea.     

Improved Discrimination of Visual Stimuli Following Repetitive Transcranial Magnetic Stimulation

Background

Repetitive transcranial magnetic stimulation (rTMS) at certain frequencies increases thresholds for motor-evoked potentials and phosphenes following stimulation of cortex. Consequently rTMS is often assumed to introduce a “virtual lesion” in stimulated brain regions, with correspondingly diminished behavioral performance.

Methodology/Principal Findings

Here we investigated the effects of rTMS to visual cortex on subjects'' ability to perform visual psychophysical tasks. Contrary to expectations of a visual deficit, we find that rTMS often improves the discrimination of visual features. For coarse orientation tasks, discrimination of a static stimulus improved consistently following theta-burst stimulation of the occipital lobe. Using a reaction-time task, we found that these improvements occurred throughout the visual field and lasted beyond one hour post-rTMS. Low-frequency (1 Hz) stimulation yielded similar improvements. In contrast, we did not find consistent effects of rTMS on performance in a fine orientation discrimination task.

Conclusions/Significance

Overall our results suggest that rTMS generally improves or has no effect on visual acuity, with the nature of the effect depending on the type of stimulation and the task. We interpret our results in the context of an ideal-observer model of visual perception.     

Highly Directional Sonar Beam of Narwhals (Monodon monoceros) Measured with a Vertical 16 Hydrophone Array

Recordings of narwhal (Monodon monoceros) echolocation signals were made using a linear 16 hydrophone array in the pack ice of Baffin Bay, West Greenland in 2013 at eleven sites. An average -3 dB beam width of 5.0° makes the narwhal click the most directional biosonar signal reported for any species to date. The beam shows a dorsal-ventral asymmetry with a narrower beam above the beam axis. This may be an evolutionary advantage for toothed whales to reduce echoes from the water surface or sea ice surface. Source level measurements show narwhal click intensities of up to 222 dB pp re 1 μPa, with a mean apparent source level of 215 dB pp re 1 μPa. During ascents and descents the narwhals perform scanning in the vertical plane with their sonar beam. This study provides valuable information for reference sonar parameters of narwhals and for the use of acoustic monitoring in the Arctic.