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Acoustic orientation via sequential comparison in an ultrasonic moth.

Orientation of female lesser wax moths (Achroia grisella) to male calling song was tested on a locomotion-compensator device that withheld all inter-aural acoustic differences from the insect. Under these circumstances, females remained longer in the vicinity of the sound source if they experienced a variable sound level that increased when approaching the source rather than a level that remained constant at all times. Analyses of orientation paths revealed that greater retention near the source was achieved by enhanced turning when the perceived sound level remained unchanged or decreased but retaining the previous heading when the level increased. These findings suggest that acoustic orientation can be supplemented by mechanisms based on sequential, as opposed to instantaneous, comparison of auditory input. Such mechanisms may be valuable when binaural hearing is impaired or asymmetric or in environments where acoustic differences at the two ears are unreliable indications of direction to the sound source.

Animals↗

Living in a "stethoscope": burrow-acoustics promote auditory specializations in subterranean rodents.

Subterranean mammals rely to a great extent on audition for communication and to be alerted to danger. The only hitherto published report on burrow acoustics revealed that in tunnels of blind mole-rats (Spalax ehrenbergi), airborne sounds of 440 Hz propagated best whereas lower and higher frequencies were effectively attenuated. Morpho-functional analyses classify the ear of subterranean mammals as a low-sensitivity and low-frequency device. Concordantly, hearing is characterized by low sensitivity and a restricted frequency range tuned to low frequencies (0.5-4 kHz). Some authors considered the restricted hearing in subterranean mammals vestigial and degenerate due to under-stimulation. In contrast to this view stand a rich (mostly low-frequency) vocal repertoire and progressive structural specializations of the middle and inner ear. Thus, other authors considered these hearing characteristics adaptive. To test the hypothesis that acoustical environment in burrows of different species of subterranean mammals is similar, we measured sound attenuation in burrows of Fukomys mole-rats (formerly known as Cryptomys, cf. Kock et al. 2006) of two differently sized species at different locations in Zambia. We show that in these burrows, low-frequency sounds (200-800 Hz) are not only least attenuated but also their amplitude may be amplified like in a stethoscope (up to two times over 1 m). We suggest that hearing sensitivity has decreased during evolution of subterranean mammals to avoid over-stimulation of the ear in their natural environment.

Acoustics↗

Free-flight phonotaxis in a parasitoid fly: behavioural thresholds, relative attraction and susceptibility to noise.

The phonotactic capacity of tachinid flies to acoustically detect and localize a sound source simulating their cricket host was investigated in a large flight room. Acoustic measurements were performed to estimate the actual stimulus delivered to the flies, revealing highly heterogeneous sound fields. When presented with a simulated cricket song in red or infrared light conditions, the flies readily flew to the sound source and landed on it. Behavioural phonotactic thresholds were established as a function of carrier frequency and were found to coincide well with the frequency of the host's natural song (4.5-5.2 kHz). Experiments revealed that the same range of frequencies is preferentially attractive to the free-flying flies, and that the reliability of signal detection in the presence of noise is best at behaviourally relevant frequencies.

Acoustic Stimulation↗

Representation of sound source direction in the superior colliculus of the guinea pig in a virtual auditory environment.

The deep layers of the superior colliculus (SC) receive visual, auditory, and somatosensory input. A major function of the SC is the control of orientation movements of the eye, head, and pinna. While a topographical map for sound source direction remains elusive in primary auditory structures of mammals, such a map for azimuthal sound source directions has been reported in the deep layers of the SC. Moreover, a gradient of elevation tuning has been also seen in the SC of ferrets and cats. Here we demonstrate that a virtual auditory environment can be used to reveal azimuthal and elevational topography for auditory spatial receptive fields in neurons in the SC of guinea pigs. Individual, head-related transfer functions (HRTF) were measured in ten guinea pigs for 122 directions in the upper hemispheric field and convolved with white noise. Many neurons (39%) in the deep layers showed robust responses to these virtual sounds, and the majority of these neurons had small spatial receptive fields that were restricted to the contralateral hemifield. Best directions varied from 0 degree to 135 degree azimuth along the contralateral side and from --10 degree to 60 degree elevation. Like previous studies using free-field stimulation, a gradient of best azimuth direction was found along the rostral-caudal axis, with rear directions represented caudally and front directions rostrally. The topographical organization for best elevations had not been studied previously in the guinea pig. We found that it roughly followed the mediolateral axis, with preference for high elevations represented medially and low elevations laterally. A similar organization using free-field stimulation has been reported in the ferret.

Acoustic Stimulation↗

The contribution of general and specific motor inhibitory sets to the so-called auditory inhibition of return.

The detection of sounds that come from a region of space recently exposed to acoustic stimulation is often slower than the detection of sounds coming from regions of space previously unexposed to acoustic stimulation. The relative increase in reaction time (RT) to targets in recently stimulated locations is generally termed "inhibition of return" (IOR). This term alludes to the possibility that spatial attention is biased against returning to recently visited locations, thus favoring the sampling of new sources of information. However, auditory IOR effects found in paradigms where subjects have to detect a first sound (cue) without making an overt response to it, and then respond as fast as possible to a second sound (target), may be due to a purely motor inhibition carried over from cue to target. Such motor inhibition has been shown to be maximal when cue and target belong to the same category, such as when they occupy the same spatial position. We have assessed the possible contribution of this motor inhibition to auditory IOR effects by having subjects respond to both cues and targets randomly presented in a right location and a left location. Reaction time to targets preceded by cues at the same location was longer than reaction times to targets preceded by cues at the opposite location (IOR effect). Compared to a condition in which subjects responded only to targets, the IOR effect was smaller, but still significant, in the double response condition, suggesting that such an effect depends on both motor inhibition and other factors, possibly related to covert spatial orienting and oculomotor control. A second experiment indicated that the IOR effect component independent of motor inhibition was slightly but significantly greater when space was relevant to the task because subjects had to report the positions of both cues and targets, compared to when space was irrelevant to the task because subjects were not required to report stimulus positions.

Acoustic Stimulation↗

Enhancement of visual perception by crossmodal visuo-auditory interaction.

Neurophysiological studies have shown in animals that a sudden sound enhanced perceptual processing of subsequent visual stimuli. In the present study, we explored the possibility that such enhancement also exists in humans and can be explained through crossmodal integration effects, whereby the interaction occurs at the level of bimodal neurons. Subjects were required to detect visual stimuli in a unimodal visual condition or in crossmodal audio-visual conditions. The spatial and the temporal proximity of multisensory stimuli were systematically varied. An enhancement of the perceptual sensitivity (d') for luminance detection was found when the audiovisual stimuli followed a rather clear spatial and temporal rule, governing multisensory integration at the neuronal level.

Adult↗

Inharmonicity detection. Effects of age and contralateral distractor sounds.

Detection of mistuned partials in otherwise harmonic complex tones was investigated in naïve subjects of three different age groups. Signals were presented at constant sensation level to compensate for differences in hearing sensitivity and to specifically examine age-related changes in inharmonicity perception. Performance was measured under two conditions, monaural signal presentation and dichotic signal-noise presentation, with the latter aiming at the influence of contralateral distractor sounds. Stimuli were complex tones with ten harmonics and 125-Hz fundamental frequency. Mistuning detection was measured for the first, second, fourth, and eighth harmonic. In a three-interval, three-alternative forced-choice procedure, subjects were required to distinguish a complex tone containing one mistuned partial from two reference tones, with all partials at their harmonic frequencies. Thresholds were measured as the amount of frequency shift necessary for the mistuning to be detected. Performance deteriorated moderately with age for the two higher partials tested, but not for the lower ones. Thresholds for dichotic signal/noise presentation did not differ significantly from monaural ones in any of the age groups. Results are discussed in relation to hypotheses of harmonicity perception in auditory scene analysis and with respect to the investigation of patients suffering form respective deficits due to acquired brain lesions.

Adult↗

Predictiveness of a visual distractor modulates saccadic responses to auditory targets.

We are faster to orient our eyes toward a visual target that also produces a sound. Conversely, the response to an auditory target is prolonged if a visual distractor is presented at a spatially incongruent position. Here, participants exhibited faster saccadic reaction times when an auditory target was more likely to be presented opposite to a visual distractor than when the stimuli only rarely occurred in spatial disparity. In contrast to experiments with visual targets and an auditory distractor, a spatially congruent visual distractor did not facilitate the response to an auditory target. We interpret the results in terms of an ocular inhibition process to suppress an automatic orienting response to the location of the visual distractor. This process is shown to be modulated by the predictability of target location.

Acoustic Stimulation↗

The timing of stimulus localisation and the Simon effect: an ERP study.

The Simon effect, an acceleration of responses at the same side that a stimulus is presented, is assumed to be the consequence of an automatic response activation evoked by the processing of the irrelevant stimulus location. This activation has been reported to decline as responses become slower. Consequently, the Simon effect decays over time. However, it remains unclear when this activation starts and what process initiates it. Up to now, the decaying hypothesis and its temporal properties have been based on indirect evidence. In the present study we tested the timing of the decay of the Simon effect more directly by combining a localisation task and a Simon task in an EEG study. It can be shown that the response activation is evoked by visual spatial processing, and that the size of the Simon effect steadily decreases as a function of the time between this localisation process and the manual response. However, this finding only holds if the encoding of relevant stimulus features follows the localisation process unequivocally.

Acoustic Stimulation↗

Spatial auditory attention is modulated by tactile priming.

Previous studies have shown that cross-modal processing affects perception at a variety of neuronal levels. In this study, event-related brain responses were recorded via whole-head magnetoencephalography (MEG). Spatial auditory attention was directed via tactile pre-cues (primes) to one of four locations in the peripersonal space (left and right hand versus face). Auditory stimuli were white noise bursts, convoluted with head-related transfer functions, which ensured spatial perception of the four locations. Tactile primes (200-300 ms prior to acoustic onset) were applied randomly to one of these locations. Attentional load was controlled by three different visual distraction tasks. The auditory P50m (about 50 ms after stimulus onset) showed a significant "proximity" effect (larger responses to face stimulation as well as a "contralaterality" effect between side of stimulation and hemisphere). The tactile primes essentially reduced both the P50m and N100m components. However, facial tactile pre-stimulation yielded an enhanced ipsilateral N100m. These results show that earlier responses are mainly governed by exogenous stimulus properties whereas cross-sensory interaction is spatially selective at a later (endogenous) processing stage.

Acoustic Stimulation↗

Spatial and non-spatial auditory processing in the lateral intraparietal area.

We tested the responses of neurons in the lateral parietal area (area LIP) for their sensitivity to the spatial and non-spatial attributes of an auditory stimulus. We found that the firing rates of LIP neurons were modulated by both of these attributes. These data indicate that, while area LIP is involved in spatial processing, non-spatial processing is not restricted to independent channels.

Acoustic Stimulation↗

Activity-dependent modulation: a non-linearity in the unilateral vestibulo-ocular reflex pathways.

It is well established that the vestibulo-ocular reflex (VOR) depends not only on sensory stimulation but also on the behavioral context associated with the stimulation. Recent modeling studies suggested that including a non-linearity in the activation function of the VOR neurons achieves the desired context-dependence for the VOR without resorting to currently assumed complex cortical computations. With the non-linearity, neurons operate as non-linear summers of incoming activity with sensitivities modulated by their activation levels. In this study we examined whether such a non-linearity exists in the unilateral VOR pathways in behaving monkeys. Acoustic clicks were employed to evoke unilateral VOR responses during fixation, head motion and smooth pursuit. We found that the click-evoked unilateral VOR responses did not simply sum in a linear manner with the eye movements initiated by head or target motion. Instead, the same acoustic click evoked larger eye movements if the ongoing eye movements were in the same direction. We also showed that the interaction between the ongoing eye movement and the click-evoked response was close to being multiplicative. These results revealed a previous unknown non-linearity in the unilateral VOR pathways, which may have important implications on the neural implementation of the context-dependence for the VOR.

Acoustic Stimulation↗

Spatiotemporal interactions between audition and touch depend on hand posture.

We report two experiments designed to assess the consequences of posture change on audiotactile spatiotemporal interactions. In Experiment 1, participants had to discriminate the direction of an auditory stream (consisting of the sequential presentation of two tones from different spatial positions) while attempting to ignore a task-irrelevant tactile stream (consisting of the sequential presentation of two vibrations, one to each of the participant's hands). The tactile stream presented to the participants' hands was either spatiotemporally congruent or incongruent with respect to the sounds. A significant decrease in performance in incongruent trials compared with congruent trials was demonstrated when the participants adopted an uncrossed-hands posture but not when their hands were crossed over the midline. In Experiment 2, we investigated the ability of participants to discriminate the direction of two sequentially presented tactile stimuli (one presented to each hand) as a function of the presence of congruent vs incongruent auditory distractors. Here, the crossmodal effect was stronger in the crossed-hands posture than in the uncrossed-hands posture. These results demonstrate the reciprocal nature of audiotactile interactions in spatiotemporal processing, and highlight the important role played by body posture in modulating such crossmodal interactions.

Acoustic Stimulation↗

Assessing the effect of visual and tactile distractors on the perception of auditory apparent motion.

In this study we investigated the effect of the directional congruency of tactile, visual, or bimodal visuotactile apparent motion distractors on the perception of auditory apparent motion. Participants had to judge the direction in which an auditory apparent motion stream moved (left-to-right or right-to-left) while trying to ignore one of a range of distractor stimuli, including unimodal tactile or visual, bimodal visuotactile, and crossmodal (i.e., composed of one visual and one tactile stimulus) distractors. Significant crossmodal dynamic capture effects (i.e., better performance when the target and distractor stimuli moved in the same direction rather than in opposite directions) were demonstrated in all conditions. Bimodal distractors elicited more crossmodal dynamic capture than unimodal distractors, thus providing the first empirical demonstration of the effect of information presented simultaneously in two irrelevant sensory modalities on the perception of motion in a third (target) sensory modality. The results of a second experiment demonstrated that the capture effect reported in the crossmodal distractor condition was most probably attributable to the combined effect of the individual static distractors (i.e., to ventriloquism) rather than to any emergent property of crossmodal apparent motion.

Acoustic Stimulation↗

Saltation in pitch perception.

Sensory saltation is a spatiotemporal illusion in which the location of a brief stimulus is displaced towards a subsequent one following closely in time and space. This study investigated in three experiments whether or not saltation is present in spectral pitch, a non-spatial dimension. Employing the "symmetrical-rabbit" paradigm, listeners judged the continuity of sequences of six short tones, differing in pitch (Exp. 1). Furthermore, the "reduced-rabbit" paradigm consisting of only three short tones was used in combination with an objective two-alternative forced-choice task (Exp. 2) and a subjective judgment task (Exp. 3). All findings indicated displacements in pitch towards subsequent tones when the interstimulus interval between the tones was short, and the frequency separation was small. This suggests a saltation-like illusion for non-spatial stimulus parameters. Possible explanations are discussed in view of the supramodal characteristic of the phenomenon.

Acoustic Stimulation↗

Influence of head-to-trunk position on sound lateralization.

The effect of horizontal head position on the lateralization of dichotic sound stimuli was investigated in four experiments. In experiment 1, subjects adjusted the interaural level difference (ILD) of a stimulus (band-pass noise) to the subjective auditory median plane (SAMP) while simultaneously directing the beam of a laser attached to the head to visual targets in various directions. The adjustments were significantly correlated with head position, shifting in a direction toward the side to which the head was turned. This result was replicated in experiment 2, which employed a two-alternative forced-choice method, in which stimuli of different ILD were presented and left/right judgments were made. In both experiments, the average magnitude of the shift of the SAMP was about 1 dB over the range of head positions from straight ahead to 60 degrees to the side. The shift of the SAMP indicates that any shift in head position induces a change in sound lateralization in the opposite direction, i.e., the intracranial sound image is shifted slightly to the left when the head is directed to the right and to the right when the head is to the left. In experiments 3 and 4, the effect of head position was compared with that of eye position by using the same methods as in experiment 2. Both shifts in SAMP, induced by either head- or eye-position changes, are in the same direction and, on average, of about the same magnitude (experiment 3), and head- and eye-position effects compensate approximately for each other during variations of head position when the gaze remains fixed to a visual target in space (experiment 4).

Acoustic Stimulation↗

Unilateral testing of utricular function.

A modified rotatory chair test is reported in which radial acceleration, generated by eccentric displacement of the subject during constant angular velocity, is exploited as a unilateral stimulation to the otolith organs. During constant angular rate rotation, the test subject is displaced laterally on the rotating turntable by 3.5 cm, so that one labyrinth becomes aligned with the rotatory axis while the second - eccentric - labyrinth is solely exposed to the altered gravito-inertial acceleration (GIA). Previously reported results showed that the direction of the response is independent of the direction of turntable rotation, ruling out any canal influence, and indicated that in a normal population the response, measured in one eye, was symmetrical for displacement of the left and right labyrinths. This mode of stimulus thus appears to elicit a unilateral otolith-ocular response (OOR). Examination of this unilateral OOR was extended in the present study; comparative testing with head-tilt to gravity, i.e. involving bilateral stimulation to the otolith organs, was carried out. Movements of both eyes were recorded (by three-dimensional video-oculography), in order to examine response conjugacy. To verify the specificity of the unilateral stimulus, tests were performed with patients who had previously undergone unilateral section of the vestibular nerve as treatment for acoustic neuroma. The eccentric displacement profile (EDP) and head-tilt stimulus each included ten cycles of left-right oscillation in order to permit signal averaging. In the normal subjects (n=12) the torsional component of the OOR proved to be both labyrinth-symmetrical and conjugate, during both bilateral and unilateral otolith stimulation. OOR gain (ocular torsion/GIA tilt) was higher for bilateral than unilateral stimulation. Bilateral OORs, obtained from three of the five unilaterally deafferented patients, proved less symmetrical and conjugate than in the normals. Unilateral OORs in all five patients were characteristically asymmetrical, with little or no response during stimulation of the diseased labyrinth.

Acceleration↗

Visuomotor adaptation without vision?

In 1995, an aftereffect following treadmill running was described, in which people would inadvertently advance when attempting to run in place on solid ground with their eyes closed. Although originally induced from treadmill running, the running-in-place after-effect is argued here to result from the absence of sensory information specifying advancement during running. In a series of experiments in which visual information was systematically manipulated, aftereffect strength (AE), measured as the proportional increase (post-test/pre-test) in forward drift while attempting to run in place with eyes closed, was found to be inversely related to the amount of geometrically correct optical flow provided during induction. In particular, experiment 1 (n=20) demonstrated that the same aftereffect was not limited to treadmill running, but could also be strongly generated by running behind a golf-cart when the eyes were closed (AE=1.93), but not when the eyes were open (AE=1.16). Conversely, experiment 2 (n=39) showed that simulating an expanding flow field, albeit crudely, during treadmill running was insufficient to eliminate the aftereffect. Reducing ambient auditory information by means of earplugs increased the total distances inadvertently advanced while attempting to run in one place by a factor of two, both before and after adaptation, but did not influence the ratio of change produced by adaptation. It is concluded that the running-in-place aftereffect may result from a recalibration of visuomotor control systems that takes place even in the absence of visual input.

Adaptation, Psychological↗