Biomedical subjects

S Carlile

Publications and source records attributed to S Carlile.

At least 19 recordsLinked to original sources

PubMed1999

The localisation of spectrally restricted sounds by human listeners.

The two principal binaural cues to sound location are interaural time differences (ITDs), which are thought to be dominant at low frequencies, and interaural level differences (ILDs), which are thought to dominate at mid to high frequencies. The outer ear also filters the sound in a location dependent manner and provides spectral cues to location. In these experiments we have examined the relative contribution of these cues to the auditory localisation performance by humans. Six subjects localised sounds by pointing their face toward the perceived location of stimuli presented in complete darkness in an anechoic chamber. Control stimuli were spectrally flat (400 Hz to 16 kHz), while the relative contribution of location cues in the low frequency channels was determined using noise high passed at 2 kHz and in the high frequency channels using stimuli low passed at 2 kHz. The removal of frequencies below 2 kHz had little effect on either the pattern of systematic errors or the distribution of localisation estimates with the exception of an increase in the size of the standard deviations associated with a few rear locations. This suggests considerable redundancy in the auditory localisation information contained within a broadband sound. In contrast, restricting the target spectrum to frequencies below 2 kHz resulted in a large increase in the cone-of-confusion errors as well as a subject dependent biasing of the front-to-back or back-to-front confusions. These biases and the reduction in localisation accuracy for high pass stimuli at some posterior locations are consistent with a contribution of spectral information at low frequencies.

Acoustic Stimulation↗

PubMed1998

Methods for spherical data analysis and visualization.

A systematic analysis of the localization of objects in extra-personal space requires a three-dimensional method of documenting location. In auditory localization studies the location of a sound source is often reduced to a directional vector with constant magnitude with respect to the observer, data being plotted on a unit sphere with the observer at the origin. This is an attractive form of data representation as the relevant spherical statistical and graphical methods are well described. In this paper we collect together a set of spherical plotting and statistical procedures to visualize and summarize these data. We describe methods for visualizing auditory localization data without assuming that the principal components of the data are aligned with the coordinate system. As a means of comparing experimental techniques and having a common set of data for the verification of spherical statistics, the software (implemented in MATLAB) and database described in this paper have been placed in the public domain. Although originally intended for the visualization and summarization of auditory psychophysical data, these routines are sufficiently general to be applied in other situations involving spherical data.

Animals↗

PubMed1998

Healthcare and the information age: implications for medical education.

The information age, combining rapidly developing information technology and massive growth in biomedical and clinical data, is placing special demands on healthcare workers. Further, radical changes in access to information in our society are affecting the doctor-patient relationship. These changes necessitate a new approach to primary and continuing medical education. A number of imperatives for medical education are identified and some practical changes to a medical curriculum are described.

Computer Communication Networks↗

PubMed1998

Medical problem based learning supported by intranet technology: a natural student centred approach.

In response to the explosion in medical information, there have been considerable recent changes in medical curriculum development. The move to problem based learning (PBL) is, in part, a result of these changes. The Faculty of Medicine at the University of Sydney has exploited a WWW based intranet for the development, delivery, management and evaluation of it's problem based, graduate medical program (GMP). This system has been employed to develop the 72 medical problems that contribute to the first two years of the GMP. The activities of more than 400 members of the faculty have been coordinated using the intranet to develop the wide range of resources to support learning in the program. Daily management of the curriculum is also enabled using Web site posting of bulletins, e-mail and ongoing development of technology training. Coupled with the PBL problems is a formative assessment system that provides questions and feedback that cover the whole range of learning topics. Part of the student and staff evaluation is supported both informally and formally through the use of a 'Feedback' button on each web page and web delivered structured formal evaluations, respectively.

Curriculum↗

PubMed1998

Altered spectral localization cues disrupt the development of the auditory space map in the superior colliculus of the ferret.

Spectral localization cues provided by the outer ear are utilized in the construction of the auditory space map in the superior colliculus (SC). The role of the outer ear in the development of this map was examined by recording from the SC of anesthetized, adult ferrets in which the pinna and concha had been removed in infancy. The acoustical consequences of this procedure were assessed by recording outer ear impulse responses via a probe-tube microphone implanted in the wall of the ear canal. Both monaural and binaural spectral cues normally show a number of asymmetric features within the horizontal plane, which allow azimuthal locations on either side of the interaural axis to be discriminated. These features were eliminated or altered by chronic pinnectomy. The responses of auditory units in the SC to noise bursts presented in the free field were examined at sound levels of approximately 10 and 25 dB above unit threshold. After bilateral pinnectomy, the representation of auditory space was severely degraded at both sound levels. In contrast to normal ferrets, many units had bilobed azimuthal response profiles, indicating that they were unable to resolve sound locations on either side of the interaural axis. There was also much less order in the distribution of best azimuths or elevations of those units that were tuned to a single direction. Some units were tuned to locations that extended much further into the hemifield ipsilateral to the recording side than the normal range of best azimuths. Unilateral removal of the outer ear, which disrupts the monaural spectral cues for one side only, had a much smaller effect on the development of the auditory representation. At supra- and near-threshold sound levels, the representation of sound azimuth in the SC on both sides of the brain was less scattered than that found after bilateral pinna removal. Nevertheless, units with bilobed responses, broader tuning, and inappropriate best azimuths were observed in both the left and right SC of ferrets in which the left pinna and concha had been removed in infancy. These data illustrate that the localization cues provided by the outer ear play a critical role in the development of the auditory space map in the SC. In contrast to other manipulations of either auditory or visual inputs, the map does not appear to adapt to the changes in spectral cues brought about by pinna removal, suggesting that residual binaural cues are, by themselves, insufficient for its normal maturation.

Acoustic Stimulation↗

PubMed1997

The nature and distribution of errors in sound localization by human listeners.

Measurement of localization performance will reflect errors that relate to the sensory processing of the cues to sound location and the errors associated with the method by which the subject indicates the perceived location. This study has measured the ability of human subjects to localize a short noise burst presented in the free field with the subject indicating the perceived location by pointing their nose towards the source. Subjects were first trained using a closed loop training paradigm which involved instantaneous feedback as to the accuracy of head pointing which resulted in the reduction of residual localization errors and a rapid acquisition of the task by the subjects. Once trained, 19 subjects localized between 4 and 6 blocks of 76 target locations. The data were pooled and the distribution of errors associated with each target location was examined using spherical methods. Errors in the localization estimates for about one third of the locations were rotationally symmetrical about their mean but the remaining locations were best described by an elliptical distribution (Kent distributed). For about one half of the latter locations the orientations of the directions of the greatest variance of the distributions were not aligned with the azimuth and elevation coordinates used for describing the spatial location of the targets. The accuracy (systematic errors) and the distribution of the errors (variance) in localization for our population of subjects were also examined for each test location. The size of the data set and the methods of analysis provide very reliable measures of important baseline parameters of human auditory localization.

Acoustic Stimulation↗

PubMed1996

The development of topographically-aligned maps of visual and auditory space in the superior colliculus.

The role of the superior colliculus in attending and orienting to sensory stimuli is facilitated by the presence within this midbrain nucleus of superimposed maps of different sensory modalities. We have studied the steps involved in the development of topographically-aligned maps of visual and auditory space in the ferret superior colliculus. Injections of fluorescent beads into the superficial layers showed that the projection from the contralateral retina displays topographic order on the day of birth (PO). Recordings made from these layers at the time of eye opening, approximately 1 month later, revealed the presence of an adult-like map of visual space. In contrast, the auditory space map in the deeper layers emerged gradually over a much longer period of postnatal life. In adult ferrets in which one eye had been deviated laterally just before eye opening, the auditory spatial tuning of single units recorded in the contralateral superior colliculus was shifted by a corresponding amount, so that the registration of the visual and auditory maps was maintained. Chronic application of the NMDA-receptor antagonist MK801 disrupted the normal development of the auditory space map, but had no effect on the visual map in either juvenile or adult animals, or on the auditory map once it had matured. These findings indicate that visual cues may play an instructive role, possibly via a Hebbian mechanism of synaptic plasticity, in the development of appropriately tuned auditory responses, thereby ensuring that the neural representations of both modalities share the same coordinates. Changes observed in the auditory representation following partial lesions of the superficial layers at PO suggest that these layers may provide the source of the visual signals responsible for experience-induced plasticity in auditory spatial tuning.

Animals↗

PubMed1996

Masking produced by broadband noise presented in virtual auditory space.

An attempt has been made to relate the masking effects studied under dichotic listening conditions to masking seen in the free field. Rather than use a free-field masking paradigm combined with monaural and binaural listening conditions, broadband maskers presented in virtual auditory space (VAS) have been used. Two virtual locations were tested: One was the right interaural axis (+90 degrees from the anterior midline) and the other was 40 degrees right of the anterior midline. Narrow-band (critical bandwidth) dichotic and diotic maskers were also derived from the VAS masker by bandpass filtering around the test frequency. This procedure preserved the interaural differences within the critical band about the test frequency but removed information outside the critical band. Using a diotic target tone of 0.6 kHz with a narrow-band masker centered on 0.6 kHz there was an increase in signal detection in the dichotic conditions when compared to that attributable to either ear alone. Furthermore, there was no further advantage in signal detection at this target frequency when a broadband VAS masker was used. This suggests that for low-frequency targets, the binaural differences within the critical band about the target frequency are sufficient for effective unmasking. In contrast, for a target frequency of 4 kHz, a dichotic narrow-band masker resulted in a reduction in detection compared to that attributable to either ear. However, detection improved to the level attributable to the far ear when a broadband VAS masker was used. This suggests that information outside the critical band is involved in the unmasking of high-frequency targets.

Adult↗

PubMed1994

Responses of neurons in the ferret superior colliculus to the spatial location of tonal stimuli.

Using multi-unit recordings, we compared the azimuthal spatial selectivity of auditory neurons in the deep layers of the ferret superior colliculus (SC) to broadband and tonal stimuli. Responses to noise were tuned at different sound levels to a single location, which varied topographically along the rostrocaudal axis of the nucleus to form a map of sound azimuth. Frequency response profiles tended to be multi-peaked, so the spatial tuning was examined at two or more frequencies in each case. Some of the azimuthal response profiles obtained with tonal stimuli were bilobed, as expected from the spatially ambiguous cues available at individual frequencies, although the rest were tuned to a single region of space. The preferred sound directions usually varied with the frequency used, and the range of auditory best positions at each recording site was significantly greater with tones than with noise. Comparison with the acoustical properties of the auditory periphery suggested that the near-threshold positional selectivity of many of the tonal responses may be determined by the monaural directionality of the outer ear. When the sound level was raised by 20 dB so that both ears were stimulated at all speaker locations, the range of tonal best positions obtained at each frequency increased and some of the units responded best to pure tones located in the ipsilateral hemifield. The lack of topographic order in the distribution of tonal spatial selectivity along the rostrocaudal axis of the SC indicates the need for a broadband input, incorporating the spectral localization cues provided by the outer ear, in the construction of a neural map of auditory space.

Acoustic Stimulation↗

PubMed1994

Measuring the human head-related transfer functions: a novel method for the construction and calibration of a miniature "in-ear" recording system.

A method for the construction of a small "in-ear" system for recording the human-free field-to-eardrum and headphone-to-eardrum transfer functions is described. Customized in-ear inserts were obtained by a simple ear printing and electroplating method, resulting in a thin (< 0.25 mm) outer shell that minimized obstruction of the entrance at the ear canal. The insert can be used to position a microphone probe tube deep within the auditory canal. The effects of this recording system on the sound field in the ear canal were calibrated using a model head equipped with a second internal microphone close to the eardrum. Transfer functions were recorded for 343 different stimulus locations in free space and for a headphone sound source. For the free-field stimuli the presence of the recording system resulted in a small attenuation with maximum effects around 3.5 and 12.5 kHz (-1.5 and -2.0 dB, respectively). Passing the data through an auditory filter model reduced the averaged attenuation to less than -1.4 dB. Phase was undistorted up to 2.5 kHz. These results suggests that the perturbations produced by the insert are unlikely to be perceptually relevant.

Acoustic Stimulation↗

PubMed1994

Monaural and binaural spectrum level cues in the ferret: acoustics and the neural representation of auditory space.

1. The role of the structures of the outer ear in producing monaural and binaural spectral cues to sound location was examined acoustically in the ferret. A probe microphone was introduced across the wall of the external auditory canal and its responses to digitally constructed wideband signals were recorded for a large number of free field locations. 2. In the intact animal the patterns of both monaural and binaural cues were asymmetrical for horizontal locations about the interaural axis. For anterior sound locations the monaural transformations demonstrated relative gains at middle and high frequencies and a location-dependent frequency notch. Changing elevation resulted in variations in the corner frequencies of these spectral features. Additionally, there was greater front-back asymmetry in the binaural spectral cues for locations in lateral space when compared with locations near the midline. 3. Surgical removal of the pinna and concha (pinnectomy) eliminated all the major front-back asymmetrical features in the horizon monaural and binaural spectral transformations as well as the elevation-dependent variations in the monaural spectra. Thus the residual transformations were ambiguous for sound locations in lateral space, resulting in "cones of confusion" centered on the interaural axis. 4. These cues were reflected in the topographic representation of auditory space in the deeper layers of the superior colliculus (SC). Previous studies have shown that spatial tuning at near-threshold sound levels is based on monaural pinna cues, whereas binaural inputs are utilized at higher levels that stimulate both ears. In the intact ferret we examined statistically the topography of the representation of sound azimuth for near-threshold and suprathreshold stimuli and the alignment of the auditory and visual representations in the SC. The distributions of auditory best positions within the SC for near- and suprathreshold stimulus levels were statistically indistinguishable, suggesting that both monaural and binaural cues are integrated in this neural representation of space. 5. Pinnectomy resulted in a large increase in the number of auditory units that responded best to two distinct locations in space. One lobe of the response was tuned appropriately in terms of the position of the unit within the SC, demonstrating that the residual acoustical cues are sufficient for the construction of a topographic representation of auditory space. However, the second region of space, thereby producing an ambiguous representation.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

PubMed1993

Changes induced in the representation of auditory space in the superior colliculus by rearing ferrets with binocular eyelid suture.

There have been conflicting reports concerning the importance of visual experience in the development of auditory localization mechanisms. We have examined the representation of auditory space in the superior colliculus of adult ferrets that were visually deprived by binocular eyelid suture from postnatal days 25-28, prior to natural eye opening, until the time of recording. This procedure attenuated the transmission of light by a factor of a least 20-25 and blurred the image so that, as long as the eyelids were still fused, the responses of visual units in the superficial layers of the superior colliculus were labile and very poorly tuned. After the eyelids were opened, the representation of the visual field in these layers appeared to be normal. Acoustically responsive units were, as usual, almost exclusively restricted to the deeper layers of the superior colliculus. However, unlike normal animals, where responses occurring only at stimulus onset predominate, most of these units exhibited sustained or multi-peaked discharge patterns. The degree of spatial tuning of individual units recorded from the normal and deprived groups of animals was not significantly different in either azimuth or elevation. Normally orientated maps of both sound azimuth and elevation were also found in the visually deprived ferrets. However, abnormalities were present in the topography and precision of these representations and consequently in their alignment with the overlying visual map. In particular, an increase was observed in the proportion of auditory units with spatially ambiguous receptive fields, in which the maximum response occurred at two distinct locations. These results indicate that patterned visual experience is not required for establishing at least a crude map of auditory space in the superior colliculus, but suggest that it may play a role in refining this representation during development.

Animals↗

PubMed1992

The effect of acute hypoxia on the latency of the human auditory brainstem evoked response.

Recent studies have shown a decrease in the amplitude and an increase in the threshold of the cat's auditory brainstem evoked response (ABER) during severe hypoxia (PaO2 of 20 to 30 Torr). In this study we have examined the effects of euoxia (end tidal PO2 100 Torr) and mild hypoxia (end tidal PO2 of 45 to 50 Torr) on the latency of the ABER in 6 human subjects. Hypoxia resulted in a blood O2 saturation of between 75 to 85% and caused a significant prolongation of the latency of wave V of the ABER by 0.185 +/- 0.045 ms (Mean +/- S.D; p < 0.01). The prolongation of the ABER during severe hypoxia has previously been attributed to a change in peripheral sensitivity. Using the stimulus level/response latency relationship obtained for each subject under normal breathing conditions, the change in latency produced by mild hypoxia can be interpreted as a mean shift in auditory sensitivity of 5.1 +/- 3.4 dB. These results suggest that the auditory system is sensitive to much smaller changes in blood O2 saturation than previously thought.

Adult↗

PubMed1992

The effects of chronic hypoxia on human auditory system sensitivity.

We have examined the effects of prolonged periods of hypoxia produced at high altitudes on the latency of the auditory brain-stem evoked response (ABER) in 9 subjects at around sea level, 3,500 m, and 4,370 m. Following an ascent from 1,300 m to 3,500 m over 24 h, the mean blood O2 saturation fell to 86.5 +/- 1.2% (+/- S.E.M.) and was associated with a mean prolongation of latency of wave V of the ABER of 0.34 +/- 0.10 ms (p = 0.011 two-tailed paired t-test). Using the stimulus-level/response latency relation determined at around sea-level for each subject, this prolongation of wave V corresponded to a mean reduction in sensitivity of 9.1 dB +/- 1.6 dB. Over a period of 72 h, blood O2 saturation improved slightly (mean 88.1% +/- 1.8%) and mean wave V latency returned to control values. A second rapid ascent to 4,370 m reduced blood O2 to below prerecovery levels (82.5% +/- 1.7%), but in this case there were no significant changes in auditory sensitivity (p = 0.79 two-tailed paired t-test). These data show that mild hypoxia results in an initial decrease in auditory sensitivity. However, the recovery of sensitivity with more prolonged exposure suggests that the auditory system can compensate for chronic mild hypoxia.

Adult↗

PubMed1991

The auditory periphery of the ferret: postnatal development of acoustic properties.

The development of the acoustics of the auditory periphery of the ferret was examined by measuring the spectral transfer functions (STFs) and the directional characteristics of the outer ears of animals ranging in age from postnatal day 32 (P32) to P54. Using an impulse response technique the STFs were obtained from up to 250 locations throughout free space. The directional responses were calculated for frequencies between 1 kHz and 30 kHz. The low frequency roll-off of the STF decreased with increasing age from around 15 kHz at P32 to an adult value of around 8 kHz by P51. The directional responses of the outer ear of the immature ferrets differed significantly from adult animals in a fashion that was consistent with the smaller size of the auditory periphery. However, by P51 the responses were generally within the normal adult range. The implications of the relatively rapid development of the acoustics of the auditory periphery are discussed in terms of the development of mechanisms subserving sound localization.

Acoustic Stimulation↗

PubMed1990

Auditory brainstem of the ferret: maturation of the brainstem auditory evoked response.

A longitudinal study of developmental changes in the brainstem auditory evoked response (BAER) was made on 19 ferrets between postnatal days 25 (P25) and 50. Responses to free-field click stimuli were recorded from anaesthetized animals, and compared with data obtained from 8 adult ferrets. A reproducible BAER was first recordable on P27, although the response onset was generally later in smaller animals. BAER onset preceded eye opening, which started on P32. Adult-like thresholds were observed in all animals by P40, but the age at which they were attained was also dependent on size. The BAER in the adult ferret consists of 4 main vertex-positive peaks occurring in the first 5 ms following transient acoustic stimulation. In the youngest animals the presence of an additional peak (between II and III) and the slurring of peaks III and IV were consistent features. The individual peaks undergo an asymmetrical pattern of development, with mean peak I latency attaining an adult value at P40, while mean peak IV latency is still 115% of the mean adult value at that age. BAERs could routinely be recorded using high stimulus presentation rates (greater than 40/s), though an increase in absolute and interpeak latencies occurred, the extent of which decreased with age. The pattern of BAER development in the ferret is compared with that in other species, and the concept of the 'silent period' (period between conception and onset of hearing) as a standard unit of auditory development is introduced.

Acoustic Stimulation↗

PubMed1990

The auditory periphery of the ferret. I: Directional response properties and the pattern of interaural level differences.

The transformations of sound by the auditory periphery of the ferret have been investigated using an impulse response technique for a large number of sound locations surrounding the animal. Individual frequencies were extracted from the detailed spectral transformation functions (STFs) obtained for each stimulus location and, using sophisticated spatial interpolation routines, were used to calculate the directional response of the periphery at that frequency. The strength of the directional response was directly related to the analysis frequency. Furthermore, as the analysis frequency was increased to 20 kHz, the orientation of the directional response increased in elevation from the horizon (E0 degrees) to about E30 degrees, while the azimuthal location remained fairly constant at 30 degrees to 40 degrees from the midline. For analysis frequencies above 20 kHz, the response became increasingly directional toward the ipsilateral interaural axis. The interaural level differences (ILDs) were also calculated for all animals studied. ILDs increased from around 5 to 25 dB over the range of frequencies from 3-24 kHz. The two-dimensional patterns of iso-ILD contours were roughly concentric and centered on the interaural axis for frequencies below 16 kHz. For higher frequencies, there was a tendency for the ILD contours to be centered on more anterior and inferior locations. The increased directionality of the auditory periphery with increasing analysis frequency, together with the presence of sharp nulls in the response at high analysis frequencies, is consistent with a diffractive effect produced by the aperture of the pinna. However, this simple model does not predict the directional responses over the low to middle frequency range.

Animals↗

PubMed1990

The auditory periphery of the ferret. II: The spectral transformations of the external ear and their implications for sound localization.

In the previous paper the directional response characteristics of the ferret auditory periphery were examined. In this study further measurements of the spectral transfer functions (STFs) of the auditory periphery were obtained at locations close to the tympanic membrane. There was considerable variation in the STFs recorded from different animals and between recordings made at each end of the auditory canal in the same animal. However, calculation of the so called "location dependency function" demonstrated that changes in the location of the stimulus produced the same pattern of changes in the STFs in all recordings. Changes in the spectral transformation for azimuth locations in the ipsilateral auditory field were examined by calculating the horizon STF. The gain transformations of frequencies below 20 kHz were found to be asymmetrical about the interaural axis so that maximum gain was obtained for anterior stimulus locations. In contrast, the maximum gain for frequencies above 20 kHz was obtained for stimulus locations about the interaural axis, and movement of the stimulus location into either the anterior or posterior fields produced symmetrical reductions in gain. These changes were related to the directional properties of the periphery examined in the previous paper [S. Carlile, J. Acoust. Soc. Am. 88, 2180-2195 (1990)]. The spatial resolution of the monaural information provided by the peripheral STFs is dependent on the rate of change of the transformations as a function of azimuthal displacement of the stimulus location. This was examined by calculating the unsigned first spatial derivative for each frequency in the horizon STF.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗